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August 20, 1975

ATOLL RESEARCH
BULLETIN

191. BIBLIOGRAPHY OF THE NATURAL HISTORY
OF THE BAHAMA ISLANDS
by William T. Gillis, Roger Byrne,

and Wyman Harrison

Issued by
THE SMITHSONIAN INSTITUTION
Washington, D.C., U.S.A.

ATOLL RESEARCH BULLETIN
NO. 191

BIBLIOGRAPHY OF THE NATURAL HISTORY
OF THE BAHAMA ISLANDS
by William T. Gillis, Roger Byrne,
and Wyman Harrison

Issued by
THE SMITHSONIAN INSTITUTION
Washington, D.C., U.S.A.

August 20, 1975

ACKNOWLEDGMENT

The Atoll Research Bulletin is issued by the Smithsonian
Institution as a part of its Tropical Biology Program. It
is sponsored by the National Museum of Natural History, with
the production and distribution handled by the Smithsonian
Press. The editing is done by the Tropical Biology staff,
Botany Department, Museum of Natural History.

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

The sole responsibility for all statements made by authors
of papers in the Atoll Research Bulletin rests with them, and
statements made in the Bulletin do not necessarily represent
the views of the Smithsonian nor those of the editors of the
Bulletin.

Editors

F. R. Fosberg
M.-H. Sachet

Smithsonian Institution
Washington, D.C. 20560

D. R. Stoddart

Department of Geography
University of Cambridge
Downing Place
Cambridge, England

GO NA ESN st cs

Introduction
Bahama literature

Guide to abbreviations used in literature
citations

A. Anthropology

B. Botany (including Bacteria, Fungi,
and General Agriculture)

C. Climate and Meteorology

ES Entomology and Arachnology

FE. Folklore

G. Geology, Geophysics, and Geodesy
H. Herpetology

i. Ichthyology

K. Geography

L. Mammalogy (including Sea Mammals)

M. Medicine, Human and Veterinary, and
Public Health

N. General Natural History, including
Reports and Semi-Popular Accounts

0. Ornithology
Pe Oceanography
Q. Tourist items related to Natural History

Re Paleontology

S\. Soil Science
te Turks and Caicos Islands
W. Novels and Other Stories with Mention

of Natural History

Xe: Industries Based on Natural Resources

Page

78

80
85
90
99
102
104

105

107

109

D4

a NSA.

NGS

De

Malacology

Zoology, General and Miscellaneous
Invertebrate

110

NS)

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7

BIBLIOGRAPHY OF THE NATURAL HISTORY
OF THE BAHAMA ISLANDS

by William T. Gillis! Roger Byrne” and Wyman Harrison”

INTRODUCTION

This bibliography has been compiled as a project adjunct to the
preparation of a new vascular flora of the Bahamas and Turks and Caicos
Islands by the first author. In an attempt to understand the distribu-
tional patterns of organisms as related to the geological history of
the Bahama Platform, notes were recorded from various publications
dealing with the natural history of the Bahama Islands.4 Portions of
this bibliography are thus the result of such study of the biogeography
of this region. The second and third authors had independently compiled
a Bahama bibliography published in part by Fang and Harrison (1972).
This work had resulted from dissertation research by Byrne on recent
vegetational change in the eastern Bahamas. Harrison had been working
on geological and geomorphological problems in the northern islands.

At a late stage of compilation, we proposed to co-operate and pool our
resources. This paper is the result. Compilation efforts ceased on
15 March 1974.

eenold Arboretum of Harvard University, 22 Divinity Ave., Cambridge,

MA 02138 and Dept. of Biology, University of Miami, Coral Gables,
FL 33124, U.S.A. Present address: Biology Dept., Hope College,
Holland, MI 49423, U.S.A.

anept. of Geography, University of California, Berkeley, CA 94720, U.S.A.

“Erindale College, University of Toronto, 3359 Mississauga Road, Toronto,
Ontario, Canada. Present address: Great Lakes Program, Energy and
Environmental Systems Division, Argonne National Laboratory, 9700 South
Cass Ave., Argonne, IL 60439, U.S.A.

4

We construe the term ''Bahamas'' to signify the entire archipelago includ-
ing both the Commonwealth of the Bahamas and the Crown Colony of the
Turks and Caicos Islands.

(Manuscript received July 1974--Eds.)

ae

Three other bibliographies of the natural history of the Bahamas
were of value in attempting to draw up as exhaustive a bibliography as
possible. The work was greatly assisted by the Land Resource Bibliog-
raphy No. 1, Bahamas, which was compiled initially by N. W. Posnett
(1968) and later revised by Posnett and P. M. Reilly (1971) for the
Land Resources Division, Foreign and Commonwealth Office, Overseas
Development Administration in Tolworth, Surrey. Their bibliographies
were intended as references for those working with land resource sur-
veys. They were essentially "in-house documents" designed for use on
surveys of natural resources. A number of entries are confidential and
not available to the public; others are of a nature to have been mimeo-
graphed and distributed "in house," again not generally available to
the scientific community. Such entries have not been repeated herein.
With permission of Messrs. Posnett and Reilly and their colleagues, we
have extracted items from their bibliographies in compiling this one. ©
We owe extraordinary thanks to them for their co-operation and permis-
sion to use their material herein.

Boersma (1968) and Fang and Harrison (1972) were most excellent
resources additional to those already at hand. The former lists a
number of references difficult to find, for instance, those of the
United States Naval Underwater Ordnance Station, the United States
Naval Oceanographic Office, and the United States Naval Research Lab.
It is unlikely that these will be easy to obtain through any but the
most specialized libraries or from the publishers.

Unpublished reports were not repeated herein because of the futil-
ity in trying to obtain them for reference. The bibliographies cited
above should be consulted for such entries. Theses and dissertations,
on the other hand, have been included because of their being generally
available either on inter-library loan from the University libraries
where they are deposited, or on microfilm from University Microfilms,
North Zeeb Road, Ann Arbor, Michigan 48103, U.S.A.

Items have been included only if the region concerned is specifi-
cally mentioned. For papers dealing with nearby Florida or for general
works on the topics mentioned, consult Boersma (1968). No attempt has
been made to include all monographs written about organisms that happen
to have one or more members inhabiting the Bahamas. One exception is
that a special attempt was made to search for references dealing with
insects and arachnids because of the small number of references in this
area. On the other hand, such policy was not followed for such groups
as vascular plants. The recent paper by Gillis (1974) has surveyed
botanical literature for monographic studies of Bahama vascular plants.
The interested person is referred to this paper in Rhodora; its rather
extensive bibliography is not repeated herein.

Searches were made through indexing periodicals such as Zoological
Record, the Bibliography of North American Geology, Applied Science and
Technology Index (1970-72), Abstracts of North American Geology,
Abstracts of Entomology, Entomology Abstracts, Aquatic Biology Abstracts,
Aquatic Sciences and Fisheries Abstracts, Biological Abstracts

(1969 - 15 March 1973), Blackwelder's Guide to the Taxonomic Literature
of Vertebrates, Engineering Societies Library, Engineering Index via

OLE (on-line service prepared by the National Science Library, 1972-73),
Science Citation Index, and the index issues (where available) of jour-
nals likely to contain research papers on the Bahamas. A MEDLARS search
was made by Countway Library, Harvard Medical School, for papers dealing
with this region.

The first author systematically checked journals whose subject
matter deals with the Bahamas. Then he examined the "Literature cited"
or "Bibliography" section for each paper of concern leading to addi-
tional references. Undoubtedly there are omissions, but we admit to
having had the benefit of the four previous bibliographies. Individuals
who have given us references are too numerous to mention by name. Of
special assistance were colleagues in various departments of Harvard
University and other naturalists whose work involves the Bahamas.

We should like to acknowledge with grateful appreciation the follow-
ing persons who assisted us far beyond the usual expectations for librar-
ians: Mrs. Patricia Hall and her staff at the Libraries of the Harvard
University Herbaria; Mrs. Ruth Hill and her staff at the Library of the
Museum of Comparative Zoology at Harvard; Mrs. Lorraine Spencer Garry of
the Science and Medicine Library of the University of Toronto Library;
Mrs. Glenn White of the National Agricultural Library, Bethesda, Mary-
land; Miss Phyllis Edwards and her staff at the Botany Department of the
British Museum (Natural History), London; Messrs. R. Desmond and D. V.
Wilson and their staff at the Library of the Royal Botanic Gardens, Kew;
Miss Isabel Hamilton of the Nassau Public Library; and Mrs. D. Gail
Saunders of the Department of Public Records, Ministry of Education and
Culture, Nassau. Other libraries of special value to us were those of
the Marine Biological Laboratory, Woods Hole, Massachusetts, the New York
Public Library, and the Widener Library of Harvard University. We are
deeply grateful to Mrs. Alan Kindig for typing final copy of the manu-
script.

Finally and most importantly, we should like to acknowledge a gen-
erous anonymous grant to the Arnold Arboretum of Harvard University under
whose auspices this work was compiled. A further gift to the Arboretum
from Mrs. Charles S. Bird of East Walpole, Massachusetts has made it
possible to distribute this bibliography more widely.

The continuing assistance and co-operation of our friends in the
governments of the Commonwealth of the Bahamas and of the Turks and
Caicos Islands have made it possible to have access to much information.
We especially acknowledge Mr. Oris S. Russell, O.B.E. Permanent Secretary
of External Affairs (formerly Permanent Secretary of Agriculture and
Fisheries) and to Mr. Claude E. M. Smith, current Director of Agriculture
and Fisheries, government of the Bahamas. ~

BAHAMA LITERATURE

Some journals involved in the Bahama literature are unique. The
Journal of the Bahama Society for the Diffusion of Knowledge appeared
in 20 issues between 1835 and 1837, published apparently in Nassau.
The parent society was entitled the Bahama Society for the Diffusion of
Useful Knowledge, but the word "Useful" was dropped from the title of
its journal. Most of the articles were published without a by-line;
furthermore, no editor was ever indicated in published lists of officers.
As a consequence, we cite such entries under "Anonymous" at the beginning
of each alphabetical listing. Camplete copies of this journal are rare;
the only Western Hemisphere copies we know of are at the Library of
Congress, the Harvard University Herbaria Libraries, the Department of
Public Records (Archives) in Nassau, and in our personal collections
(an incomplete copy exists at the Nassau Public Library). All but one
of these was deposited in its respective library as a result of the
compilation of this bibliography.

The Bahamian government itself occasionally issued reports that had
a limited circulation. There was the Bulletin of the Agricultural Depart-
ment, Bahamas, edited by W. Munro Cunningham. Vol. I, No. 1 appeared in
January 1906. After that it was printed more or less on a monthly basis
until at least March 1910 (Vol. V, No. 1). A set is on file at the

Library of the Royal Botanic Gardens, Kew.

There was once also the Communications of the Nassau Agricultural
Society, a loyalist publication which was issued on an irregular basis
around 1801-2. In effect, it was the predecessor of the Journal of the
Society for the Diffusion of Knowledge and the Bulletin of the Agricul-
tural Department, Bahamas. The only copy we know of extant and complete

is in the British Museum (Natural History).

Charles Johnson Maynard of Newton, Massachusetts issued two journals
privately, the contents of which dealt chiefly with his researches in the
Bahamas. These are the nearest we have ever come to finding one-man pub-
lishing ventures. According to local stories, the only contribution to
the journal not made by Maynard himself was the paper on which he printed.
He wrote most of the contributed articles, made the ink, and printed
copies himself. One of these journals is Contributions to Science
(1889-96), especially noted for his monographic study of the land snail
genera Cerion and Strophia. Contributions to Science lasted for 3 vol-
(1908-20), which lasted for 12 volumes. ~ Copies of both journals, espe-
cially the latter, are rare (see also Batchelder, 1951).

During the latter years (1969-70) in which Bahama Airways was the
flag carrier of the Bahamas, an in-flight magazine ''Flamingo" was avail-
able in the seat pocket for passengers to read. A number of quasi nat-
ural history articles appeared. Because of difficulty in the reader's
attempt to retrieve copies of this magazine through the usual library
channels, its contents are not indexed herein. Similarly, Clipper
Magazine, the in-flight magazine of Pan American World Airways

(Vol. 9, No. 6 for December 1969-January 1970) contains articles on the
Bahamas. The same is true for most of the contents of Bahamian Review
Magazine, a commercial tourist-oriented propaganda magazine currently
available.© In the late 1940's and early 1950's, another magazine of
the same type, Nassau, Magazine of the Bahamas had a brief existence.
The ephemeral Canada-West Indies Magazine had a Bahamas-emphasis issue
in 1945. Because of the only peripheral relation of any of the articles
to natural history, they have not been indexed herein.

Of greater value, but of great scarcity in libraries, are the
Colonial Reports made periodically to the British Colonial Office by
officials stationed in the colonies. Attention is called to them as a
whole for the information they contain on the natural history and indus-
try of the region, but none is indexed herein. The Nassau Guardian,
daily newspaper issued in Nassau, has published a number of documents
related to the Islands, their history, politics, and resources. A list
of such documents was compiled by the Guardian in 1910, but such docu-
ments -- those issued both before and since that date -- are available
in few libraries outside the Bahamas. The Guardian has played a more
lasting role in natural history. For more than 15 years, Mrs. Leslie
(Helen Burns) Higgs has contributed a column to the Guardian every
Thursday relating to plants, gardening, and agriculture. In addition,
for the past year, the first author (W. T. Gillis) has contributed a
monthly column to Conch News, weekly newspaper of the Turks and Caicos
Islands. This deals with native plants. In the 1960's, Mr. Nixon
Smiley of Miami, Florida contributed a large number of articles on the
Bahamas in his featured column in the Miami Herald. None of these news-
paper reports is indexed herein.

For ease of reference, entries are categorized by subject matter.
Where the content of a reference relates to more than one area of inter-
est, it is repeated in these several areas. For example,

Rosén, Nils. 1911. Contribution to the fauna of the Bahamas
I. A general account of the fauna, with remarks on the physiography of
the islands. II. The Reptiles. III. The fishes. Lunds Univ. Rrssk.
Nebeenta, 2, Bd. 7 (5): 3-72 1s indexed under G, H, I, N, and Z
(Categories identified below). ng EAs 7

Various references dealing with oceanography and marine geology
completely overlap these two related fields, so they are included under
both categories. Based on the compilers' judgment, we may have omitted
salient papers from one or another of these categories. Hence, both G -
Geology, Geodesy, and Geophysics and P - Oceanography should be consulted
for references in these areas. Generally speaking, papers dealing with
properties of the water and sea bottom are included under P; those
dealing with compaction and accretion into rock are under G. There is
some overlap similarly with G - Geology, Geodesy, and Geophysics and

K - Geography.

2D 0. Box N-494, Nassau, N.P. Bahamas.

Somewhat over 90% of the published references have been examined
personally and the pagination verified, especially against citations in
other listings. Unless an author is named, a paper is listed as "Anony-
mous.'' Agencies are considered to be publishers, not authors. Because
of the small number of published works on the Turks and Caicos Islands
generally, we have indexed such items both with their related disciplines
and also separately under T.

Topics follow alphabetically according to their content: (This is
the sequence of listing in the bibliography)

N 54S 439M WO 4 = BAe) Gath aw

Anthropology, including archaeology and history

Botany, including fungi, bacteria, and agriculture
Climate and meteorology

Entomology and Arachnology

Folklore dealing with natural history

Geology, including Geodesy and Geophysics

Herpetology

Ichthyology

Geography

Mammalogy

Medicine, both Human and Veterinary, including Public Health
General natural history, including semi-popular accounts
Ornithology

Oceanography

Tourist information related to natural history
Paleontology -

Soil "science

Turks and Caicos Islands

Novels and other stories with mention of natural history
Industries based on natural resources

Malacology

Zoology, general and invertebrate

GUIDE TO ABBREVIATIONS USED IN LITERATURE CITATIONS

Abhand. Abhandlungen

Abst. Abstract(s)

Acad. Academy, Academic, Académie
Adv. Advancement

Afd. Afhandlingar

Afr. African, Africain(e)

Alum Alumni

Ag. Agriculture, Agricultural
Anat Anatomical

Ann. Annual, Annals, Annales
Anthrop Anthropologist, Anthropology
Antiq Antiquity

Appl Applied, Applicata

Arbor Arboretum

Arch Archives

Ark. Arkiv

Rrssk. Rrsskrift

ASB Association of Southeastern Biologists
Assn. Association

Astron. Astronomy

Bd. Board, Band

Berl. Berlin

Bibl. Bibliography

Biblioth. Bibliothéque, Bibliothek
Blatt. Blatter

Boll. Bollettino

Bot. Botany, Botanical, Botanische, Botanique
Brasil. Brasileira

Britt: British

Buel Bulletin

Bus. Business

Calack: California

Can. Canadian

Carib. Caribbean

Chem. Chemical

Chron. Chronic

Cree, Circular

Cai Gaal

Gol: Colonial

Colle Collection(s)

Comm. Commission

Comp. Comparative

Conf. Conference

Cong. Congress

Cons. Conservation, Conservatoire
Contr. Contributions

Cosmochim. Cosmochimica

Ces Comptes Rendus

Dept. Department, Departemento
Die£. Diffusion

Dis. Diseases

Diss. Dissertation

Div. Division

Ecol. Ecological

Econ. Economic

Ed’. Education

Eng. England

Engin. Engineering

Emmet. Entomology, Entomological
Env. Environmental

Epidem. Epidemiology

Even. Evening

Exch. Exchange

Exped.

Fl:
Found.
Fr.

Gard.
Gart.
Geochin.
Geof.
Geogr.
Geol.
Geophy .
Gesam.
Gesell.

Helv.
Herb.
Hist.
HMSO

Hort.

Hydrobiol.

Ind.
Inst.
Int.
Is.

Lyc.

Mag.
Malac.
Malak.
Mamm.
Mar.
Mass.
Med.
Mem.
Meteorol.
Micro.
Midl.
Mineral.

Expedition

Flora
Foundation
Frangais (e)

Garden, Gardeners

Garten

Geochimica

Geofisica

Geography, Geographical, Geographic
Geology, Geological, Geologists, Geologicae
Geophysical, Geophysics

Gesamten

Gesellschaft(en)

Helveticae

Herbarum, Herbarium, Herbier

History, Historia

Her (His) Majesty's Stationery Office
Horticulture, Horticultural
Hydrobiologie

Industrial, Industries

Institute, Instituto, Institution
International

Island(s)

Journal
Jahrbucher

Knowledge
Koniglich

Laboratory

Lepidoptera, Lepidopterists
Limnological

Linnaean

Lyceum

Magazine

Malacological
Malakozoologische

Mammalogy

Marine

Massachusetts

Medical, Medicine

Memoir(s), Memorias
Meteorologische, Meteorological
Microscopical

Midland

Mineralogists, Mineralogische

Massachusetts Institute of Technology

Miscellaneous
Mus. Museum, Muséum
Monatsch. Monatschrift
Monog. Monograph(s)
Mycol. Mycologia
Mycopath. Mycopathologia
Nat. Natural, Naturalist, Naturae
Natl. National
Naturgesch. Naturgeschichte
Naturw. Naturwissenschaftliche
Neerl. Neerlandische, Néerlandais
Nor. Norvegica
Not. Notulae
Notiz. Notizblatt
Nov. Novitates
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N.Y. New York
NZ. New Zealand
Obs. Observatory
Occ. Occasional
Oceanog. Oceanographic
Off. Office. Official
Ord. Ordnance
Ont. Orto
Paleont. Paleontology
Parasit. Parasitology, Parasitological
Pecrol: Petrology, Petroleum
Pharm. ‘Pharmacological
Phas. Philosophical
Phila. Philadelphia
Polat. Pola tical
Preserv. Preservation
Proc. Proceedings
Prof. Professional
Protistenk. Protistenkunde
Protozool. Protozoological
Psych. Psychology
Pub. Public, Publication(s), Publicazioni
Quart. Quarterly
Rec. Record(s), Recueil
Ref. Reference
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Rept. Report
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SE Southeastern

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Sp. Species, Specierum

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UZSi; United States (of America)
U.S.G.P.0. United States Government Printing Office
Univ. University, Universidad, Universitat
U.S.G.S United States Geological Survey
U.S.D.A. United States Department of Agriculture
Veg. Vegetabile

Verte Veterinary

Vol. Volume

Wash. Washington

W.I. West Indies

Wiss. Wissenschaften

Yearb. Yearbook

Zeitsch. Zeitschrift

Zool. Zoology, Zoologische

LITERATURE CITED IN INTRODUCTION

Batchelder, C. F. 1951. A bibliography of the published writings of
Charles Johnson Maynard (1845-1929). J. Soc. Bibl. Nat. Hist.
2. (7): 227-260.

Boersma, Anne. 1968. Bibliography on the Bahama Islands. M.1.T.
Exp. Astron. Jah. Rept. No. RN-37. 60 pp.

11

Fang, Carol and W. Harrison. 1972. Bahamas Bibliography. A list of
citations for scientific, engineering and historical articles .

pertaining to the Bahama Islands. Spec. Sci. Rept. No. 56.
Warpinia-Inst. Mar.*sci., Gloucester Point, Virginia 23062).

Gillis, William T. 1974. Name changes for the seed plants in the
Bahama flora. Rhodora 76: 67-138.

Posnett, N. W. 1968. Land resource bibliography No. 1, Bahamas.
Land Resources Division, Directorate of Overseas Surveys,
Tolworth, Surrey.

---------- and P. M. Reilly. 1971. Land resource bibliography.
I. Bahamas. Land Resources Division, Foreign and Commonwealth
Office, Overseas Development Administration. London.

13

A. ANTHROPOLOGY, ARCHAEOLOGY, AND HISTORY

Anonymous. 1970. Commonwealth of the Bahamas. Statistical Abstract.
Dept. of Statistics, Cabinet Office. Nassau, Bahamas. 211 pp.

Anson, Peter F. 1957. The hermit of Cat Island: the life of Fra
Jerome Hawkes. P. J. Kennedy. New York, N. Y. 286 pp.

Bain, G. L. 1959. The early history of the Bahama Islands to 1730.
M.A. thesis. University of London.

Bethel, Arnold Talbot. 1914. The early settlers of the Bahama
Islands. A. B. Vance. Jacksonville, Florida. 116 pp.

Beven, Kathy Sue, ed. 1972. Cultural anthropology survey of San
Salvador Island, the Bahamas. College Center of the Finger
Lakes, Corning, N. Y. 164 pp.

Booy, Theodore de. 1912. Lucayan remains on the Caicos Islands.
Amer. Anthrop. 14: 81-105.

wooo ---- 1913. Lucayan artifacts from the Bahamas. Amer. Anthrop.
15: 1-7. (Contr. Heye Mus. No. 1).

---------- 1919, On the possibility of determining the first land-
fall of Columbus by archeological research. Hispanic Amer. Hist.
Rev. 2: 55-61.

Bradley, Wendell P. 1969. They live by the wind. Part 4 - The
fishermen of the Bahamas. Alfred A. Knopf, Inc. New York,
N- Y.> pp. 211-267.

Brooks, W. K. 1889. On the Lucayan Indians. Natl. Acad. Sci. Mem.
4: 213-222 + 12 pl. (Art. X).

Bullen, Ripley P. 1959. Similarities in pottery decoration from
Florida, Cuba, and the Bahamas. 33rd Int. Cong. Americanists
2: 107-110.

Byrne, Anthony Roger. 1972. Man and the variable vulnerability of
island life: a study of recent vegetation change in the Bahamas.
Unpublished dissertation, University of Wisconsin, Dept. of
Geography. 311 pp.

---------- 1973. Man and the variable vulnerability of island life:
a study of recent vegetation change in the Bahamas. Dissert.
ADSer yoo Gl 1) 22..pp'.

14

Byrne, Anthony Roger and John C. Munday, Jr. 1972. Photodensity and
the impact of shifting agriculture on subtropical vegetation: a
case study in the Bahamas. Proc. 8th Int. Symp. remote sensing
of environment, 2-6 Oct. 1972. No. 195600-1-X. Env. Res. Inst.
Michigan, Ann Arbor, Michigan. pp. 1311-1326.

Churton, Edward Townson (Bishop). 1887. The island missionary of the
Bahamas. Practical addresses and notes; intended chiefly for
ordinands. J. Masters and Co., London. 108 pp. 1888 = 2ndved:
The island missionary of the Bahamas. A manual of instruction and
routine... J. Masters and Co.” Londons I1282pp-

Clavel, M. 1904. Items of folk-lore from Bahama Negroes. J. Amer.
Folk-lore 17: 36-38.

Craton, Michael. 1962. A history of the Bahamas. Collins. London.
520) pps 1968= "Second ed. 'Collins) “London. 520" ppe

Crowley, Daniel J. 1958-59. L'héritage africaine dans les Bahamas.
Présence Afr. N.S. 23 (Dec.-Jan.): 41-58.

Dupuch , Sir Etienne. 1967: The Tribune stony; (Ernest, Benne ltde
London. 162 pp.

Fang, Carol and Wyman Harrison. 1972. Bahamas Bibliography. A list
of citations for scientific, engineering and historical articles
pertaining to the Bahama Islands. Spec. Sci. Rept. 56, Virginia
Institute of Marine Sciences. Gloucester Point, VA. 23062.

Gallagher, Patrick. 1961. Bahamas (in News and Notes). Amer. Antiq.
PAE AGT

Goggin, J. M. 1939. An anthropological reconnaissance of Andros
Island, Bahamas. Amer. Antiq. 5: 21-26.

---------- 1946. The Seminole Negroes of Andros Island, Bahamas.
Florida Hist. Quart. 24: 201-206.

Granberry, Julian. 1955a. A survey of Bahamian archeology. M.S.
thesis. University of Florida. Gainesville, Florida. 375 pp.

mo-------- 1955b. An anthropological reconnaissance of Bimini,
Bahamas. Amer. Antiq. 22: 378-381.

ma -------- 1956. The cultural position of the Bahamas in Caribbean
archeology. Amer. Antiq. 22: 128-134.

Hart, Miss. 1823-24. Letters from the Bahama Islands, written in
1823-24. H.C. Carey and I.vlea. London.- 20% pp.

IS

Hoffman, Charles A. Jr. 1967. Bahama prehistory: cultural adaptation
to an island environment. Ph.D. Dissertation, Dept. of Anthropology,
University of Arizona, Tucson, Arizona. 133 pp.

---------- 1970. The palmetto Grove site on San Salvador, Bahamas.
Florida State Mus. Contr. (Soc. Sci. 16): 1-26.

Holmes, W. H. 1894. Caribbean influence in the prehistoric art of
Sseuthern states. Amer. Antiq. © 7: 71-79.

Joyce, T. A. 1919. Notes on a wooden stool from the island of
Eleuthera, Bahamas. Man 19: 1-2.

Krieger, Herbert W. 1937. The Bahama Islands and their prehistoric
population. Explorations and Field-Work, Smithsonian Inst.
(for 1936): 93-98.

Kuczynski, R. R. 1953. Demographic survey of the British Colonial
Empire. Auspices of Royal Institute of International Affairs,
Vol. 3. West Indian and American Territories. Oxford Univ. Press.

McCutcheon, E. S. 1927. The island song book. Nassau.

MacLaury, J. C. 1970. Archeological investigations on Cat Island,
Bahamas. Florida State Mus. Contr: ~(Soc. Sci. No. 16): 27-50.

Manwaring, G. E. 1957. Woodes Rogers, privateer and governor.
A. Deans Peggs, ed. Margate Press. Nassau. 48 pp. (Reprint of
the introduction to 1928 edition of Woodes Rogers's "Cruising
voyage round the world.").

Maynard, Charles Johnson. 1890. Some inscriptions found in Hartford
Cave, Rum Key, Bahamas. Contr. Sci. 1: 167-171.

--=------- 1893. Traces of the Lucayan Indians in the Bahamas.
SOatr. oCL.. 2525-54.

---------- 1915. Some traces of the Lucayan Indians in the Bahamas.
Rec. Walks and Talks with Nature 7: 196; 197-200.

Mazess, R. B. 1967. Skin color in Bahamian Negroes. Human Biol.
39: 145-154.

Miller, W. Hubert. 1945. The colonization of the Bahamas, 1647-1670.
William and Mary Quart. 2: 33-46.

Mills, T. Wesly. 1887. The study of a small and isolated community
in the Bahama Islands. Amer. Nat. 21: 875-885.

Newton, Bertram A. 1968. A history of Red Bays, Andros. (mimeographed) .
Published privately by the author.

16

Norton, Graham. 1967. People of the Out Islands. Geog. Mag. 40:
704-728.

Otterbein, Keith F. 1959. Setting of fields: a form of Bahamian
obeah.~ Phila. Anthrop) Soe. Bulle) (as 2015 73

---------- 1963a. The family organization of Andros Islanders: a case
study of mating system and household composition of a community in
the Bahama Islands. Ph.D. dissertation. University of Pittsburgh.
Pittsburgh, Pa.

---------- 1963b. The household composition of Andros Islanders. Soc.
and Econ’. Stud). 32) 78-365.

Pascoe, C. F. 1901. Two hundred years of the S.P.G. (Society for the
Propagation of the Gospel in Foreign Parts, 1701-1900) .), SoReGe
Westminster. 1429 pp. Chapter xxviii - The Bahamas. pp 216-227b.

Peggs, A. Deans. 1955. A short history of the Bahamas, 2nd ed. Dean
Peggs’ Research! Fund..._Margate Press. )Nassau.., 2nd ed. - 19575957) pp’.

Peters, Thelma. 1946. Blockade-running in the Bahamas during the Civil
War. ) lequesiza)S.: 16-29).

Porter, Kenneth W. 1945. Notes on Seminole Negroes in the Bahamas.
Florida Hist. Quart. 24: 56-60.

Rainey, Froelich. 1941. Excavations in the Ft. Liberté Region, Haiti.
Yale Univ... Publ. Anthrop:. .235: 1-48.

Rawson, William. 1868. Report on the Bahamas hurricane of October 1866
with a description of the city of Nassau, N.P. Nassau Guardian.
Nassau. 29 pp.

Reid, Ira DeA. 1942. The John Canoe festival. A New World Africanism.
Phylon (Atlanta Univ. Rev. Race and Culture, 4th quarter) 3: 349-370.

Rodgers, William B. 1966. Development and specialization: a case from
the Bahamas. Ethnology 5: 409-414.

a Nadal and C. H. Wallace. 1969. Development and changes in popula-
tion distribution in the Out Island Bahamas, Anthropologica
(Ottawa) 11: 189-201.

Rolle, Kermit and Gwen Ellingsen. 1966 (reprinted 1969). Out island
lore. Litho Graphic Press, Hicksville, N. Y. 536 pp.

Rouse, Irving. 1960. The entry of man into the West Indies. Yale
Univ. Publ: Anthrop. No. 61. "26: pp.

Sadler, H. E. 1970. Turks Island landfall. Privately offered for
sale. Grand Turk. 200 pp.

d7

Sharer, Cyrus Jewett. 1955. The population growth of the Bahama
Islands. Ph.D. dissertation. University of Michigan. Ann Arbor,
Michigan. 126 pp + viii.

Siebert, Wilbur H. 1913. The legacy of the American Revolution to the
British West Indies and Bahamas. In History of the American loy-
abists: (Ohio State Unive Publ: 17 (27): 3=50°

Smiley, Nixon. 1972. The lost tribe of Andros. Tropic (Miami Herald
Mag.) for February 20, 1972. pp. 14-18.

Symonette, Michael A. 1973. Discovery of a nation. An illustrated
history of the Bahamas. Privately published. Nassau. 40 pp.

Valentine, J. Manson. 1973. Culture pattern seen. Muse News (Publi-
cation of the Miami Museum of Science) 4: 314-315; 331-334.

Wright, James Martin. 1905. History of the Bahama Islands, with a
special study of the abolition of slavery in the colony. Ph.D.
dissertation, Johns Hopkins Univ., Baltimore, Md. published in
Shattuck, George - Bahama Islands (1905): 419-583.

woo------- 1915. Wrecking system of the Bahama Islands. Polit. Sci.
Quart. 30: 618-644.

See also entries under Folklore.

B. BOTANY
(Including Bacteria, Fungi, and General Agriculture)

Anonymous, 1802. Communications on different subjects addressed to the
Bahama Agricultural Society. Nassau. 63 pp.

---------- {Sss5a-)) Cocos mnuciteras J2;Bahama Soc soDx£E «= “Knowl.

---------- 1835b. On the medicinal and other uses of plants growing in
the Bahama Islands. J. Bahama Soc. Diff. Knowl. No. 3: 23-30;
Nowe4eeSl=373 Now 5S: 44-46.

corte e---- 1836a. Yam potato. J. Bahama Soc. Diff. Knowl. No. 13:
146-147.

---------- 1836b. (untitled - deals with medicinal properties of 12
plants, extracted from Beach's "American practice of medicine.")
J. Bahama Soc. Diff. Knowl. No. 9: 95-100.

---------- 1836c. The process of curing the palmetto. J. Bahama Soc.
Dift- Knowl: No dom eli 2%

18

Anonymous, 1838. Series of short articles on various plants and their
uses: sweet fennel, garlic, ginger, parsley, Cayenne pepper, orange,
lemon, wax myrtle, hoarhound, English plantain, mustard, elder.

J. Bahama Soc. Diff. Knowl. No. 9: 95-100.
---------- 1887. Sabicu wood. Kew Bull. 1887 (12): 4-5.

---------- 1888. Colonial fruits: Bahama Islands. Kew Bull. 1888:

---------- 1907. Agave seen by Columbus found. Discovery 1: 30-32.

a--------- 1959. Botanizing in the Bahamas. Fairchild Trop. Gard.

---------- 1967. Ming-like tree from nearby Bahamas is introduced to
Florida) by Garden. “/Fairchild Trop. Gard. Bull. 17)/54%

---------- 1970. Cat Island. Where have all the cascarillas gone?
Bahamian Rev. 12: 22.

---------- 1971. Local names of fruits and vegetables in the English-
speaking Caribbean. Cajanus 4 (2): supplement. 32 pp.

Adderley, Lincoln. 1964. Two Bahamian equitant oncidiums. I.
Oncidium lucayanum. Gard. J. 14: 141-142; 186-188.

Agassiz, Alexander. 1888. Three cruises of the United States Coast
and Geodetic Survey Streamer "Blake." Vol. 1, chapt. 5. Relations

of the American and West Indian fauna and flora. Bull. Mus. Comp.
ZOOS 14 ell Sst

Ames, Oakes. 1910. A new Ponthieva from the Bahamas. Torreya 10: 90-91.

Anderson, R. F. and W. J. Stambough. 1966. The dying pines on Great
Abaco Island, Bahamas. Internal Report, Owens-Illinois Co.

Anthony, Emilia C. 1902. Fern hunting in Nassau. Fern Bull. 10: 65-68.

Asprey, G. F. 1960. Vegetation in the Caribbean area. Carib. Quart.
5: 245-263.

Bailey, L. H. 1939. Coccothrinax, of Florida. Gentes Herb.’ 4: 220-225.

-~--------- 1944. Revision of the American palmettoes. Gentes Herb.
6: 366-459.

---------- and H. E. Moore. 1949. Palms uncertain and new. Gentes
Herb. 8: 93-205.

Balgooy, M. M. J. van. 1969. A study on the diversity of island
floras. Blumea 17: 139-178.

19

Barben, A. de la. 1835. On the cultivation of tobacco. J. Bahama
sec.: Di ft .-Knowl . No. 6: 52-53.

Barrett, Wilfredo H. G. and Lamberto Golfari. 1962. Descripcidén de
dos nuevos variedades del "Pino del Caribe."' Carib. For. 23: 59-71.

Beard, J. S. 1949. The natural vegetation of the Windward and Leeward
Islands. Oxford For. Mem. No. 21.

Bentham, George. 1854. On the tree supplying the Sabict wood of Cuba.
J. Bot. Hooker 6: 235-237.

Bounds, John Howard. 1968. Forestry in the Bahamas. Forest Farmer
29> 10-145; 21-22...

Brace, L. J. K. 1929. Note on the occurrence of Oxypolis filiformis
in the Bahamas. Torreya 29: 16-17.

Britton, N. L. 1890. Review of the provisional list of the plants of
the Bahama Islands. Bull. Torrey Bot. Club 17: 187-188.

---------- 1903. A new Waltheria from the Bahamas. Torreya 3: 105-106.

e--------- 1904a. Explorations in Florida and the Bahamas. J.N.Y. Bot.
Gard. 5: 129-136. b

corcccr--- 1904b. Report on exploration of the Bahamas. J.N.Y. Bot.
Gard. 5: 201-209.

eorcn----- 1904c. Savia bahamensis. Torreya 4: 104-105.
o--------- 1904d. Notes on the flora of the Bahamas. Torreya 4: 190.

---------- 1904e. On Pisonia obtusata and its allies. Bull. Torrey
Bot. Club 31: 614-615.

---------- 1905a. Contributions to the flora of the Bahama Islands -
ie Bile IN Yew BOtee iGard.s 5, 440—45 5

eo---n---- 1905b. Explorations in the Bahamas. J.N.Y. Bot. Gard.
6: 78-85.

---------- 1906a. Contributions to the flora of-the Bahama Islands.
iby a Bulds: NeY; Bot. Gard. 4:,. 115-128.

---------- 1906b. Contributions to the flora of the Bahama Islands.
Ei. Budd. N.Y -) Bot. Gard’, 44, 23573143.

---------- 1907a. Contributions to the flora of the Bahama Islands.
[V.7 BulleeN -Yi. Bot.» Gards.5 +) 511-318.

20

Britton, N. L. 1907b. Report on the continuation of the botanical
exploration of the Bahama Islands. J.N.Y. Bot. Gard. 8: 71-81.

teat 1913. Four undescribed West Indian sedges. Torreya 13:
Zio ely.

---------- and C. F. Millspaugh. 1920. The Bahama flora. Privately
published. ~ New York; N."Y.~ Reprinted "1962." Hatner Publes¢or
New York) Nea v1" 695, ppr

Bunt; J... S.,.K. E:* Cooksey, M-oAz Heeb, ©.9 €. Lee, and B, Pevaaydlon-
1970. Assay of algal nitrogen fixation in the marine subtropics
by acetylene reduction. Nature 227: 1163-1164.

Byrne, Anthony Roger. 1972. Man and the variable vulnerability of
island life: a study of recent vegetation change in the Bahamas.
Ph.D. dissertation. University of Wisconsin, Madison, Wisconsin.
Si ppe

---------- 1973. Man and the variable vulnerability of island life: a
study of recent vegetation change in the Bahamas. Diss. Abst.
$5 CM) 2 pps

---------- and John C. Munday, Jr. 1972. Photodensity and the impact
of shifting agriculture on subtropical vegetation: a case study in
the Bahamas. Proc. 8th int. symposium remote sensing of environ-
ment, 2-6 Oct. 1972. No. 195600-1-X. Environ. Res. Inst. Michigan.
Ann Arbor, Michigan. pp. 1311-1326.

Catesby, Mark. 1730-47. Natural History of Carolina, Florida and the
Bahama Islands. London. 2 vols. 220 plates. (For publication,
see J..S0C. Bibb. Nat... Hist. 3: 3528.)

Clough, Garret €C. and George Fulk. 1971. The vertebrate fauna and the
vegetation of East Plana Cay, Bahama Islands. Atoll Res. Bull.
S587 3

Coker, William S. 1905. Vegetation of the Bahama Islands. in: Shattuck,
George B. The Bahama Islands. Geographical Society of Baltimore.
Johns Hopkins Press. pp. 185-270 + pl. xxxiii-xlvii.

Committee report. 1889. Report of the committee consisting of Messrs.
W. Carruthers, W. F. R. Weldon, J. G. Baker, G. M. Murray, and W. T.
Thistelton-Dyer (Secretary), appointed for the purpose of exploring
the flora of the Eahamas. ‘Rept. Brit. Assn. Adv. Sci. 58: 36-503.

Correll, Donovan S. 1974. Flora of the Bahama Islands - new additions.
Fairchild Trops- Gard. Bully 292 a2

Daniell, William F. 1863. On the cascarilla and other species of
Croton of the Bahamas and other West Indian Islands. Pharm. J.
Trans, Pls 42) l44=15037.2226-251F:

Ja

Dawes, Clinton J. and Harold J. Humm. 1969. A new variety of Halimeda
lacrimosa Howe. Bull. Mar. Sci. 19: 428-431.

Dolley, Charles S. 1889. The botany of the Bahamas. Proc. Acad. Nat.
pew. Phila. 1889: P31-134.

Drew, G. E. 1912. Report of investigations on marine bacteria carried
on at Andros Island, Bahamas, British West Indies in May 1912.
Carnegie Inst. Washington Yearbook 11: 136-144.

a--------- 1914. On the precipitation of calcium carbonate in the sea
by marine bacteria, and on the action of denitrifying bacteria in
tropical and temperate seas. Papers Tortugas Lab, Carnegie Inst.
Washington Publ. 182: 7-45.

Duncombe, Alfred. 1835. On the caper plant. J. Bahama Soc. Diff.
Knowl. No. 5: 43-44.

Durrell, Zoé C. 1972. .The innocent island - Abaco in the Bahamas.
Durrell Publications. Distributed by Stephen Greene Press.
Brattleboto, Vermont. 157 pp.

Dyer, W. Thiselton. 1888. Flora of the Bahamas. Nature 37: 565-566.

Baton, 0.,C. 1875. A list of the marine algae, collected by Dr. Edward
Palmer on the coast of Florida and at Nassau, Bahama Islands, March-
August 1874. New Haven, Connecticut. 6 pp.

---------- and W. A. Setchell. 1886. A list of plants from Abaco
Island, Bahamas. Johns Hopkins Univ. Cir. 6: 46-47.

Eegers, (H.F.A.?). 1888. in letter to W. A. Thiselton Dyer. Flora of
the Bahamas. Nature 37: 565-566.

Evans, Alexander W. 1911. The Hepaticae of the Bahama Islands. Bull.
Torrey Bot..Club 38: 205-221.

Galstoff, P. S. 1940. Wasting disease causing mortality of sponges in
the West Indies and Gulf of Mexico. Proc. 8th Amer. Sci. Cong.
Seal -421.,

Gardiner, John, L. J. K. Brace, and Charles S. Dolley. 1889. Provisional
list of the plants of the Bahama Islands. Proc. Acad. Nat. Sci.
Phila. 41: 349-426.

Gathorn, William. 1836. Method of curing tobacco in the district of
Holquin, in the Island of Cuba. Method of curing tobacco at Vuelta
Bajo (to the Leeward of Havana). J. Bahama Soc. Diff. Knowl.

No. 10: 102-107.

Gillis, William T. 1970. Isle’ of Columbus. Fairchild Trop. Gard.
Budi. 253 5=7 .

22

Gillis, William T. 1974a. Name changes for the seed plants in the
Bahama flora. Rhodora 76: 67-138.

se 1974b. Phantoms in the flora of the Bahamas. Phytologia
29: pp. 154-166.

---------- ,» Richard A. Howard and George R. Proctor. 1973. Additions
to the Bahama flora since Britton and Millspaugh - I. Rhodora 75:
411-425.

aSocsa---= and William T. Stearn. 1974. Typification and correct names
of Leucaena and Lysiloma species in the Bahama flora. Taxon 23:
185-189.

---------- and George R. Proctor. 1974. Caesalpinia subgenus
Guilandina in the Bahamas, J. Arnold Arbor. 55: 425-430.

Griffiths, Thomas A. 1972. Bahama Bay-rush - a missing link? Bates
College Alumni Mag.

Grisebach, A. H. R. 1859-64. Flora of the British West Indian Islands.
Lovell Reeve and Co. London. 789 pp. Reprinted 1963. J. Cramer,
Weinheim, Weldon and Wesley and Hafner Publishing Co., New York and
Codicote. (For dates of publication etc. see Stearn, W. T., 1965,
in J. Arnold Arbor. 46: 263.).

~--------- 1865. Die geographische Verbreitung der Pflanzen Westindiens.
Abhand. Konigl. Gesell. Wiss. Gottingen 12: 3-80.-

Guppy, H. B. 1917. Plants, seeds, and currents in the West Indies and
Azores. Williams and Norgate. London.

Hall We H: >.1837/" The £an palm; the-olive tree. J. Bahama Soc. Disti.
Knowl. No. 20: 211-212.

Hamilton, William. 1835. (No title - subject deals with quality of
tannins from Divi-divi in the Bahamas.). J. Bahama Soc. Diff.
Knowl. No. 6: 53-59.

---------- 1836a. The pita plant. J. Bahama Soc. Diff. Knowl. No. 8:
~--------- 1836b. On the manufacture of sugar from beet root. J.

Bahama Soc. Diff. Knowl. No. 18: 187-192.
---------- 1836c. <Cochineal. J. Bahama Soc. Diff. Knowl. No: Us-720l-202°

Hannau, Hans W. and Jeanne Garrard (undated, issued in 1970). Flowers
of the Bahamas. Argos, Inc. Miami, Florida. 64 pp.

Hardie, Lawrence A. 1969. Algal crusts from the Bahamas (abs.).
Amer. Assn. Petrol. Geol. Bull. 53: 721.

Be)

Harshberger, John W. 1903. Notes on the strand flora of Great Inagua,
Haiti, and Jamaica. Torreya 3: 67-70.

Herrick, F. H. 1886. Notes on the flora of Abaco and adjoining
islands. Johns Hopkins Univ. Circ. 6: 46-47.

Hickman, Denis and Peggy Hickman. Undated. Guide to the tropical
gardens of the Royal Victoria Hotel. Royal Victoria Hotel.
Nassau. :

Higgs, Helen Burns. 1956. Flowers of Nassau. Miami Post Publ. Co.,
Miami, Florida. Privately published. 19 pp.

---------- (as Mrs. Leslie Higgs) 1969. Bush medicine in the Bahamas.
Privately published by the author with original paintings. 20 pp.

---------- 1973. Vegetable growing in the Bahamas. When, how, and
what to plant. Privately published by Mrs. Higgs and the Nassau
Guardian. Nassau, Bahamas.

Hill, Steven R. 1974. Additions to the Bahama flora. Rhodora.
(in press).

Hitchcock, A. S. 1893. Plants of the Bahamas, Jamaica,,and Grand
Cayman. Ann. Rept. Missouri Bot. Gard. 4: 47-179 + pl. 11-14.

---------- 1898. List of cryptogams collected in the Bahamas, Jamaica,
-and Grand Cayman. Rept. Missouri Bot. Gard. 9: 111-120.

Hooker, Joseph D. 1881. Bahama flora. Rept. Progress and Conditions,
Royal Gardens, Kew for 1880. 30 pp.

---------- hea.) (Crotoneelucerta. (Curtis Bot. Mag. 53: pl. 7515.

Howard, Richard A. 1950. Vegetation of the Bimini Island group,
Bahamas, B.W.I. Ecol. Monog. 20: 317-349.

aa-s------ i775 lherveretation of the Antilles y Chapt. lin WVegeta-
tion and vegetational history of northern Latin America. Alan
Graham, ed. Elsevier Publ. Co. Amsterdam. pp. 1-38.’

a--------- and Henry F. Dunbar. 1964. Additions to the flora of
Inagua, the Bahamas. Rhodora 66: 6-15.

Howe, Marshall A. 1904a. Notes on Bahaman algae. Bull. Torrey Bot.
Shubert; Sse 9'5=100%

---------- 1904b. Collections of marine algae from Florida and the
Bahamas. J.N.Y. Bot. Gard. 5: 164-166.

24

Howe, Marshall A. 1905a. Phycological studies - I. New Chlorophyceae
from Florida and the Bahamas. Bull. Torrey Bot. Club 32: 241-252.

---------- 1905b. Phycological studies - II. New Chlorophyceae, new
Rhodophyceae, and miscellaneous notes. Bull. Torrey Bot. Club 32:
563-571 (separately available as Contr. N.Y. Bot. Gard. No. 72).

coeccc---- 1906. Some photographs of the silk-cotton tree (Ceiba
pentandra) with remarks on the early records of its occurrence in
America. Torreya 6: 217-231.

---------- 1909. Phycological studies - IV. The genus Neomeris and
notes on other Siphonales. Bull. Torrey Bot. Club 36: 74-104.
(Separately available as Contr. N.Y. Bot. Gard. No. 120).

alata and Percy Wilson. 1968. Report on the botanical explora-
tion of the Bahama and Caicos Islands. J.N.Y. Bot. Gard. 9: 41-50.

Humm, Harold J. 1961. The marine algae of Bimini, Bahamas. Cyanophyta
(Abs.). ASB Bull. 8: 32. }

Jackson, W. P. U. 1946. Plant dermatitis in the Bahamas. Brit. Med.
J. 1946 (2): 298.

Kellogg, Royal S. 1951. Yellow pine in the Bahamas. J. Forestry
49: 795-796.

Kohlmeyer, J. and E. Kohlmeyer. 1971.’ Marine fungi from tropical
America and Africa. Mycologia 63: 831-861.

Lewis, Walter H. 1971. Additions to the flora of the Bahama Islands,
Rhodora 73: 46-50.

Little, E. L., Jr. 1947. The name of the wild dilly of Florida.
Rhodora 49: 289-293.

Long, Robert W. and Olga Lakela. 1971. Flora of tropical Florida.
Univ. Miami Press. Miami, Florida. 962 pp.

Luer, Carlyle A. 1972. The native orchids of Florida. New York
Botanical Garden. New York, N. Y. 293 pp.

McCallan, E. A. 1939. Report on the development of agriculture in the
Bahamas. Nassau Guardian. Nassau. 107 pp.

McCallum, May F. 1970. Aerobic bacterial flora of the Bahama Bank.
J. Appl. Bacteriology 33: 533-542.

Mann, A. 1935. Diatoms in bottom deposits from the Bahamas and the
Florida Keys. Carnegie Inst. Washington Publ. 452: 121-128 (Paper
Tortugas Lab No. 19).

25

March, E. W. 1949. Pine forests of the Bahamas. Empire Forestry
Rev. 28: 33-37.

Margolis, Stanley and Robert W. Rex. 1971. Endolithic algae and
micrite envelope formation in Bahamian odlites as revealed by
scanning electron microscopy. Geol. Soc. Amer. Bull. 82: 843-851.

Meyers, Samuel P. 1957. Taxonomy of marine Pyrenomycetes. Mycologia
49: 475-528.

Millar, Robert. 1835. On the cultivation of cotton in the Bahamas.
J. Bahama Soc. Diff. Knowl. No. 6: 48-51.

------- --- 1836. On the cultivation of cotton. J. Bahama Soc. Diff.
Knowl. No. 17: 184-185.

Millspaugh, C. F. 1904. A new Bahaman Euphorbia. Torreya 4: 172.

---------- 1906. Praenunciae Bahamenses. I. Contributions to a flora
of the Bahamian Archipelago. Field Mus. Bot. Ser. Publ. 106,
Vot= II.) No. 32 137-184:

---------- 1909. Praenunciae Bahamenses. II. Contributions to a flora
of the Bahamian Archipelago. Field Mus. Publ. 136, Bot. Ser.
Vols £i,* No. 7: 289-321.

Monty, Claude Leopold Victor. 1966. Geological and environmental
Significance of Cyanophyta (abs.). Diss. Abst. Sec. B. 27: 211B.

---------- 1967. Distribution and structure of Recent stromatolitic
algal mats, eastern Andros Island, Bahamas. Ann. Soc. Géol.
Belgique Bull. 90: B-55-100.

Morris, D. 1896. Memo on the cascarilla bark. In: Bahama flora,

1862-1900, miscellaneous reports, Vol. 108, pp. 40-51. Library,
Royal Botanic Gardens, Kew.

Morrow, Loring and Norton H. Nickerson. 1973. Salt concentrations in
ground waters beneath Rhizophora mangle and Avicennia germinans.
Rhodora 75: 102-106.

Morton, Julia F. 1965. Can the red mangrove provide food, feed, and
fererlizer? Econ. Bot. 19: 113-123.

Morton, Kendal and Julia Morton. 1946. Fifty tropical fruits of
Nassau. Text House, Inc. Coral Gables, Florida.

Murray, G. 1888-89. Catalogue of the marine algae of the West Indian
region. J. Bot. 26: 193-196; 237-243; 303-307; 331-338; 358-363;
27: 237-242; 257-262; 298-305.

26

Nash, George V. 1905a. Botanical exploration of the Inagua Islands,
Bahamas. J.N.Y. Bot. Gard. 6: 1-19.

---------- 1905b. Further explorations in the Republic of Haiti.
J.N.Y. Bot. Gard. 6: 170-191.

---------- 1905c. A trip to the Inaguas. Plant World 8: 91-98 (also
issued as a separate).

Nesbitt, C. R. 1854. Vegetable fibres of the Bahamas. J. Bot. 6:
237-241.

Neumann, A. C. and L. S. Lind. 1969. Algal production and lime ~_.
deposition in the Bight of Abaco: a budget. Geol. Soc. Amer.
Spec. ‘Papers 121: 219 (abst. )).

Nickerson, Norton H. and Joseph W. Tripp. 1973. Floral dimorphism in
Rachicallis americana (Jacq.) Hitch. (Saltwater-bush). Rhodora
7S AVIA1 3%.

Northcroft, George J. H. 1902. Sketches.of Summerland - giving some
account of Nassau and the Bahama Islands. Nassau Guardian. Nassau.
Chapt. 14 - Flora (pp. 152-173); Chapt. 15 - Soil and agriculture
(pp. 174-185).

Northrop, Alice R. 1902. Flora of New Providence and Andros (Bahama
Islands). ‘Mem. Torrey Bot. Club 12: 1-98 + 19 pl. (Reprinted? ini
Northrop, John I. 1910. A naturalist in the Bahamas. Memorial
volume. Columbia Univ. Press. New York. pp. 119-211.)

Orpurt, Philip A. 1964. The microfungal flora of bat cave soil from
Eleuthera Island, the Bahamas. Can. J. Bot. 42: 1629-1633.

Pawsey, R. G. 1967. Pine mortality on Great Abaco Island, and examina-
tion of pine on Grand Bahama and Andros Islands. Internal Rept.,
Ministry of Overseas Development. Government of the Bahamas.
Nassau.

Plate, L. 1906. Pyrodinium bahamense, Ng... nN. isp. Die Leuchtperadanee
des 'Feueresees' von Nassau, Bahamas. Arch. Protistenk. 7: 411-427.

Popenoe, John. 1966a. Bahaman plants at the Fairchild Tropical Garden.
Fairchild) Trop. Gard. 7Bulil..” 21) 5-9);

haters 1966b. Sweetwood bark. Fairchild Trop. Gard. Bull. 21: 6-7.
Sel 1973. Moujean tea. Fairchild Trop. Gard. Bull. 28: 12-14.
Proctor, G. R. 1954-55. Notes on the vegetation of the Turks and

Caicos Islands. Nat. ‘Hist. Soc. Jamaica ‘6: 249-152; (170-1745
199-203.

PATE

Rabley, Margaret B. 1971. An introduction to some wild flowers of the
Bahamas and Caribbean. Collins and Co. London and Glasgow.
(Coloring book).

Read, Robert W. 1966. Coccothrinax inaguensis - a new species from
the Bahamas. Principes 10: 29-35.

---------- 1967. Taxonomy of the cycads of the West Indies and Florida.
Butch Cont. on Cycdad toxicity. “Section Ill: "6 pp-

Rock, Barrett Nelson. 1972. The woods and flora of the Florida Keys:
"Pinnatae". Smithsonian Contr. Bot. 5: 1-35.

Rogers, C. M. 1968. The Linum bahamense complex. Rhodora 70: 439-441.

Rudd, Velva. 1969. Mimosa bahamensis, a Bahama - Yucatan disjunct.
Phytologia 18: 143-147.

Russell, Oris S. 1958. Preliminary report on the flora of the Exuma
Cays. in Prospectus for new national trust. Published by Bahamas
National Trust. pp. 16-21.

Sampson, H. C. Report on the development of agriculture in the Bahamas.
Empire marketing Board Publ. 40. H.M.S.0O. London. 35 pp.

Sawyer, William H. 1955. Medicinal uses of plants by native Inaguans.
Sci. Monthly 80: 371-376.

pseliger; H. H., W. H. Briggley, and E. Swift. 1969. Absolute values

of photon emission from the marine dinoflagellates Pyrodinium
bahamense, Gonyaulax polyedra and Pryocystis lunula. Phytochemistry
and Photobiology 10: 227-232.

Semple, John C. 1970. The distribution of pubescent leaved individuals
of Conocarpus erectus (Combretaceae). Rhodora 72: 544-547.

Slee, M. U. 1970. Crossability values within the slash-Caribbean Pinus
species complex. Euphytica 19: 184-189.

Small, J. K. 1910a. Report on botanical exploration in Andros,
Bahamas. J.N.Y. Bot. Gard. 11: 88-101.

---------- 19T0b.) Exploration an Andros: ~Torreya 10:°151-153. *
ma-------- 1916. Pithecolobium guadalupense. Addisonia 1: 50-52.

Smiley, Nixon. 1968. Bahama plants as ornamentals. Fairchild Trop.
Gard: Budd 255, 71 2..

28

Smith, Robert R., et al. 1971-72. A study of the near shore and
terrestrial flora of San Salvador Island. 3 vol. College Center
of the Finger Lakes. Corning, New York. Includes chapters as
follows:

Bush medicine of San Salvador Island.

study of the cultivated plants on San Salvador Island.
comparative algae study of Grahams Harbour and French Bay.
vegetative study of the peninsular area and Cutt Cay.

vegetative study of the Watling Castle ruins.

vegetative study of the Nairn homestead.

vegetative study of the Fortune Hill plantation site.

An algae study of Bonefish Bay.

The cacti of San Salvador Island.

A vegetative study of dumps and waste areas of San Salvador Island.

rerrrre,

---------- 1973. Tropical Botany (A text for use in San Salvador
Island, Bahamas). 2 vol. College Center of the Finger Lakes.
Corning, New York.

Stearn, W. T. 1958. A key to West Indian mangroves. Kew Bull.
19582 35237;

---------- 1958. Publication of Catesby's 'Natural History of Carolina,
Florida, .and the, Bahama = Isiiands' . ~J.-=Soc. Biol; Nat. Hisit swe.

~--------- 1964. Catharanthus roseus, the correct name for the
Madagascar periwinkle. Lloydia 27: 196-200. °

---------- 1965. Grisebach's 'Flora of the British West Indian Islands',
a biographical and bibliographical introduction. J. Arnold Arbor.
46: 243-285. .

$s-------- 1968. Jamaican and other species of Bumelia (Sapotaceae) .
J. Arnold Arbor. 49: 280-289.

Stimson, William R. 1967. Additions to the flora of the Bimini Island
group, Bahama Islands. Rhodora 69: 60.

Stout, A. B. 1920. Conference notes for April (algae in the Bahamas -
525 spp.); 6 new since Britton and Millspaugh. J.N.Y. Bot. Gard.

21: 97-98.

Taylor, Norman. 1921. Endemism in the Bahama flora. Ann. Bot. 35:
525-5526

Taylor, William Randolph. 1960. Marine algae of the eastern tropical
and subtropical coasts of the Americas. Univ. Michigan Press.
Ann Arbor, Michigan. 870 pp.

29

Traverse, Alfred and Robert N. Ginsburg. 1966. Palynology of the
surface sediments of the Great Bahama Bank, as related to water
movement and sedimentation. Mar. Geol. 4: 417-459.

---------- 1967. Pollen and associated microfossils in the marine
surface sediments of the Great Bahama Bank. Rev. Palaeobotany and
Palynology 3: 243-254.

Tynes, Benjamin. 1835. Additional remarks on the qualities of plants
growing in the Bahamas. J. Bahama Soc. Diff. Knowl. No. 7: 47-48.

Urban, Ignatius. 1898-1913. Symbolae Antillanae seu fundamenta
florae Indiae Occidentalis. Berlin. 7 vol.

Volz, P..A. 1971. A preliminary study of keratinophilic fungi from
Abaco Island, the Bahamas. Mycopath. Mycol. Appl. 43: 337-339.

---------- and Enso. meneke.” £972. Ajprefiminary study of fresh water
fungi from Abaco Island, the Bahamas. Mycopath. Mycol. Appl. 46:
1-3.

---------- and Douglas E. Jerger. 1972. A preliminary study of marine
fungi from Abaco Island, the Bahamas. Mycopath. Mycol. Appl. 48:
271-274.

Wall, David and Barrie Dale. 1969. The “hystrichosphaerid" resting
spore of the dinoflagellate Pyrodinium bahamense Plate 1906.
J. Phycology 5S: 140-149.

Weber, Neal A. 1967. The fungus-growing ant, Trachymyrmex jamaicensis,
on Bimini Island, Bahamas (Hymenoptera: Formicidae). Ent. News
78: 107-109.

Wilcox Marcaret S., Anne Forbes, Sally Shure, L: V. Wilcox, Jr. 1971.
APEreld key to Bahamian mangroves. Carib. J. Sci. 11: 155-157.

Wilson, Percy. 1909. Report on the botanical exploration of the islands
of the Salt Key Bank, Bahamas. J.N.Y. Bot. Gard. 10: 173-176.

Wood, J. Ferguson. 1968. Studies of phytoplankton ecology in tropical
and subtropical environments of the Atlantic Ocean. Part 3.
Phytoplankton communities in the Providence Channels and the Tongue
Of pLne) Oceana Bull MaravSGiva 1S 9481-543).

Yaffa, Harold. 1971. The West Indies cruise on the Utowana (Dec. 1931-
Apr. 1932) Allison V. Armour - David G. Fairchild Plant expedition.
Raarehild=inop. Garda Bull. 26: 910-15.

Zobel, Bruce. 1964. Pines of southeastern U. S., Bahamas and Mexico
and their use in Brazil. in Silvicultura em Sao Paulo, Rev. Técnica
do: Florestal do Estado de Sao Paulo; Vol. 3 (3): 303-310.

30

C. CLIMATE AND METEOROLOGY

Anonymous. Undated. Climatological Data, West Indies, 1946-1951.
Weather Bureau, U. S. Dept. of Commerce. Washington, D. C.

---------- 1836a. Meteorological Diary for September 1836. J. Bahama
Soc... Diff. Knowl.” Now ls so

---------- -1836b. Meteorological diary for October 1836. J. Bahama
Soc, Dr£f Knowl Nome 195)

---------- 1836c. Meteorological diary for November 1836. J. Bahama
Soc. Diff. Knowl. No. 19: 204.

---------- 1837. Meteorological diary for December 1836. J. Bahama
S0¢.=-Ditt .- Knowl’ Nov20 3721 3

---------- Undated (but before 1871). General description of the City
of Nassau and Island of New Providence, Bahamas, West Indies with
meteorological tables and other statistics of interest to invalids
and travelers. New York, N. Y. Privately published. 28 pp.
Reprinted and expanded, 1871. Published by William Baldwin and
Gow, sNews YOu Ni. SOs Dis

---------- 1897. Zum Klima der Bahamas-Inseln. Meteorol. Zeitsch. 14:
309. k

~--------- 1928. The Bahama Islands. Information as to trade, soil,
climate, etc. for intending settlers, tourists and business men.
Development Board. Nassau. 24 pp. + map.

erro -- 1959. North Atlantic tropical cyclones. Technical Paper 36.
Weather Bureau, U..S. Dept. of Commerce. U.S.G.P.0. Washington,
Dec

Brooks, C. E. P. 1921. Meteorology of Nassau, Bahamas, 1852-1919.
Quart. J. Royal Meteorol. Soc. 47: 59-62.

Clackson, J. R. 1971. (Mimeographed). Past records of Bahamas rain-
fall. Meteorological Office. Nassau.

Fassig, Oliver L. 1905a. Climate of the Bahama Islands. In: Shattuck,
George B. The Bahama Islands. Geographical Society of Baltimore.
Johns ‘Hopkins Press. ‘pp. 111-125.

ileal 1905b. Exloration of the upper atmosphere at Nassau, New
Providence by means of kites. In: Shattuck, George B. The Bahama
Islands. Geographical Society of Baltimore. Johns Hopkins Press.
pp. 129-143.

31

Fassig, Oliver L. 1907. Uber das Klima der Bahama-Inseln. Meteorol.
Zertseh . 242/558-5590.

Hayes, Nelson. 1962. The roof of the wind. Alvin Redman, Ltd.
London. Originally published 1961 by Doubleday. 216 pp.

Keogh, Terence. 1933. Hurricane in the Bahamas: one man's adventures
is a big wind. Harpers Mag. (May) 166: 697-708.

Kraus, E. B. 1971. The Bahama Bank planetary boundary layer experi-
mene, 17 April-10 May 1971: “Amer. Meteorol. Soc. 52: 969-972.

Lettau, Bernhard. 1971. The effect of the Bahamas-Lesser Antilles
chain on the annual precipitation frequency. Amer. Meteorol. Soc.
Bull. 52: 781-782.

Ludlum, David M. 1963. Early American hurricanes, 1492-1870. Amer.
Meteorol. Soe.>- boston. . «xii + 198) pp:

Moore, Willis L. and Oliver L. Fassig. 1913. Hurricanes of the West
Indies. U. S. Dept. of Agriculture, Weather Bureau Bull. X.
Hearasb oO: Washington,’ D. C.°" 26" pp.

Northcroft, George J. H. 1902. Sketches of Summerland - giving some
account of Nassau and the Bahama Islands. Chapter 10 - Climate.
pp. 94-108. Nassau Guardian. Nassau.

Perkins, R. D. and Paul Enos. 1968. Hurricane Betsy in the Florida-
Bahama area - geologic effects and comparison with Hurricane Donna.
a Geol .° 76: 710-717.

o--------- 1969. Comparison of geologic effects of Hurricanes Donna and
Betsy in the Florida-Bahama area (abst.). Geol. Soc. Amer. Spec.
Paper 121: 460-461.

Rawson, William. 1868. Report on the Bahamas hurricane’ of October
1866 with a description of the city of Nassau, N.P. Nassau
Guardian, Nassau. 29 pp.

Simpson, Joanne. 1967. An experimental approach to cumulus clouds and
hurricanes. Weather 22: 95-114.

Stone, L. H. J. 1953. The weather of the Bahamas. Weather 8: 301-302.

Sugg, Arnold L. 1966. The hurricane season of 1965. Monthly Weather
Rev. 94: 183-191.

Tannehill, Ivan Ray. 1952. Hurricanes, their nature and history,
particularly those of the West Indies and the southern coasts of
the United States. Princeton Univ. Press. Princeton, N. J.
308 pp.

32

Townsend, P. S. 1826. Memoir on the topography, weather, and diseases
of the Bahama Islands. J. Seymour Co. New York, N. Y. 80 pp.

E. ENTOMOLOGY AND ARACHNOLOGY

Arnett, Ross H., Jr. 1953. The Oedemerid beetles of the Bimini Island
group, Bahama Islands, British West Indies. Amer. Mus. Nov. 1646:
1-13.

Baker, E. W. 1968. The genus Pronematus Canestrini. Ann. Ent. Soc.
Amer. 61: 1091-1097.

Banks, Nathan. 1906. Arachnids from the Bahamas. Bull. Amer. Mus.
Nat. Hisit..225 sls5=189%

Barber, Harry G. 1953. A new subfamily, genus, and species belonging
to the family Eniocephalidae (Hemiptera, Heteroptera). Amer. Mus.
Nov. 1614: 1-4. ;

---------- 1954. A report on the Hemiptera, Heteroptera, from the
Bimini Islands, Bahamas, British West Indies. Amer. Mus. Nov.
1682: 1-18.

---------- and P. D. Ashlock. 1960. The Lygaeidae of the Van Voast-
American Museum of Natural History Expedition to the Bahama Islands,

19535. (Hemiptera, Heteroptera). Proc. Ent. Soc. Washingtonwo2:
117-124.

Benjamin, Foster H. 1934. Description of some native Trypetid flies
with notes on their habits. USDA Tech. Bull. 401. USGPO. 96 pp.

Blake, D. H. 1962. Eight new species of Metachroma from the West
Indies (Coleoptera: Chrysomelidae). Proc. Ent. Soc. Washington
64: 175-180.

eorcr-cH- 1970. A review of the beetles of the genus Metachroma
Chevrolat (Coleoptera: Chrysomelidae). Smithsonian Contr. Zool.
57: 1-111. (pp. 49-50 on Bahamas).

Blocker, H. Derrick. 1971. Distribution of Balcutha in the Bahama
Islands, Newfoundland, and Panama (Homoptera: Cicadellidae).
JONG. Ent. .Soce 79en 15s —1607

Bradley, J. C. 1964. Further notes on the American taxa of Campsomeris
(Hymenoptera: Scoliidae). Ent. News. 75: 101-108.

Branch, Nina and E. L. Seabrook. 1959. Culex (Culex) scimitar, a new
species of mosquito from the Bahama Islands (Diptera, Culicidae).
Ent. Soc. Washington Proc..61 3, 216-2138.

5S

Burks, B. D. 1956. The species of Chryseida (Hymenoptera, Euryomidae) .
Bull. brooklyn Ent. soc. Si, l09-116.

Cambridge, F. O. Pickard. 1901. On a collection of spiders from the
Bahama Islands made by J. L. Bonhote, Esq. with characters of a
new genus and species of Mygalomorphae. Ann. Mag. Nat. Hist. ser.
Wee OO Oe.

Cazier, Mont A. 1951. The Buprestidae of the Bahama Islands, British
West Indies (Coleoptera, Buprestidae). Amer. Mus. Nov. 1517: 1-9.

---------- 1952. Additions to the Buprestid fauna of the Bahama Islands,
British West Indies (Coleoptera, Buprestidae). Amer. Mus. Nov.
ro2. 1-10)

---------- and Lionel Lacey. 1952. The Cerambycidae of the Bahama
Islands, British West Indies (Coleoptera). Amer. Mus. Nov. 1588:
1-55.

Chemsak, John A. 1967. Notes on the Cerambycidae of Grand Bahama
isiands.. ~ Pan, Pac. Ent. 43;. W8l=ss.

Chickering, A. M. 1969. The genus Stenoonops (Araneae, Oonopidae) in
Panama and the West Indies. Breviora 339: 1-35.

Clench, Harry K. 1941la. Notes on two Bahaman Lycaenidae, with the
description of a new subspecies. Torreia 7: 3-7.

---------- 1941b. A new race of Hemiargus for the Bahamas. (Lepidoptera:
Lycaeidae). Mem. Soc. Cubana Hist. Nat. 15: 407-408.

---------- 1943. The Lycaenidae of the Bahama Islands (Lepidoptera:
Rhopalocera). Psyche 49: 52-60.

=-=-=----- 1963. A synopsis of West Indian Lycaenidae with remarks on
their zoogeography. J. Res. Lepid. 2: 247-270.

Coffin, T. Homer. 1905. Mosquitoes of the Bahama Islands (Discussion).
in: Shattuck, George B. The Bahama Islands. Geographical Society
of Baltimore. Johns Hopkins Press. pp. 275-289.

Comstock, W. P. 1946. A Saturniid from the Bahamas (Lepidoptera).
eve. Ent. coc. 54: 17 l=172.

a and E. Irving Huntington. 1943. ‘Lycaenidae of the Antilles
(Lepidoptera, Rhopalocera). Ann. N.Y. Acad. Sci. 45: 49-130.

Curran, C. H. 1953. The Asilidae and Mydaidae of the Bimini Islands,
Bahamas, British West Indies (Diptera). Amer. Mus. Nov. 1644: 1-6.

Darlington, P. J. 1953. West Indian Carabidae (Coleoptera): the Bahama
species. Amer. Mus. Nov. 1650: 1-16.

34

Dodge, H. Rodney. 1965. The Sarcophagidae (Diptera) of the West
Indies. I. the Bahama Islands. Ent. Soc. Amer. Ann. 58: 474-497.

Emerson, K. C. 1957. A new species of Mallophaga from a dove.
J. Kansas Ent.) Soc .)'50 250-56).

Foote, Richard H. 1960. The Tephritidae and Otitidae of the Bahama
Islands (Diptera) J:N.Y. Ent.soc. 68: 85-99:

Ford Everett: Jc, Jt. Looe A revision of the genus Petalium LeConte
in the United States, Greater Antilles, and the Bahamas (Coleoptera:
Anobiidae). U.S.D.A. Tech. Bull. 1467: 1-40.

Hampson, Sir George F. 1901. The Lepidoptera-Phalaenae of the Bahamas.
Ann. Mag. Nat. Hist. ser. 7.7: (246-261.

Howard, L. O. 1905. Mosquitoes of the Bahama Islands (Introduction)
in: Shattuck, George B. The Bahama Islands. Geographical Society
of Baltimore. Johns Hopkins Press. pp. 273-274.

---------- » Harrison G. Dyar, and Frederick Knab. 1912-17. The
mosquitoes of North and Central America and the West Indies.
Vol. 1-1912; Vol: 2-1912; Vol. 3-1915; Voll. 4-1917.. ; Publi Carneeale
Inst. Washington No. 159.

James, Maurice T. 1953. The Stratiomyidae (Diptera) of Bimini, British
West Indies. Amer. Mus. Nov. 1613: 1-6.

---------- 1971. Two new species of Phaenicia from the West Indies
(Diptera; Calliphoridae).. Proc. “Ent. Soc. Wash. 73: 38l2385.

Johnson, Charles W. 1908. The Diptera of the Bahamas, with notes and
description ‘of one new species. «Psyche 153, 69-80.

Knab, Frederick and W. W. Yothers.. 1914.- Papaya fruit fly.:° J.) Age.
Res. 2: 447-453 + 2 pl.

Krombein, Karl V. 1953. The wasps and bees of the Bimini Island group,
Bahamas, British West Indies (Hymenoptera: Aculeata). Amer. Mus.
Nov.) 16335: " 1-29'.

Levi, H. W. 1955. The spider genera Oronota and Stemmops in North
America, Central America, and the West Indies (Araneae: Theridiidae).
Ann.’ Ent.’ Socs Amer.) 483) 1553- 342.

a 1957. The spider genera Crustulina and Steatoda in North
America, Central America, and the West Indies (Araneae, Theridiidae).
Bull; Mus..Comp. Zool)...117,5, 367-424.

~--------- 1959. The spider genera Achaeranea, Theridion, and
Sphyrotinus from Mexico, Central America, and the West Indies.
Bull. Mus. Comp. Zool. 121: 57-163.

55

Linley, John R. and John B. Davies. 1971. Sandflies and ‘tourism in
Florida and the Bahamas and Caribbean area. J. Econ. Ent. 64:
264-278.

Llewellyn, Craig H., Andrew Spielman, and Thomas E. Frothingham, et al.
1970. Survival of arboviruses in Aedes albonotatus, a peridomestic
Bahaman mosquito. Proc. Soc. Exp. Biol. Med. 33: 551-554.

Marston, Norman. 1970. Revision of New World species of Anthrax
(Diptera: Bombyliidae), other than the Anthrax albofasciatus
group. Smithsonian Contr. Zool. 43: 1-148.

Martin, Charles H. 1957. The Asilidae of the Bahama Islands with the
description of two new species (Diptera). Amer. Mus. Nov. 1847:
1-7.

Maynard, Charles Johnson. 1888. Notes on the white ant, found on the
Bahamas. Psyche 5: 149-150.

Menke, A. S. 1970. The genus Ammophila in the West Indies (Hymenoptera;
Sphecidae). Proc. Ent. Soc. Washington 72: 236-239.

Metcalf, Zeno Payne. 1954. Homoptera from the Bahama Islands. Amer.
Mus. Nov. 1698: 1-46.

Park, Orlando. 1954. The Pselaphidae of South Bimini Island, Bahamas,
Britrsn West Indies (Coleoptera). Amer. Mus. Nov. 1674: 1-25.

Paulson, Dennis R. 1966. New records of Bahamian Odonata. Quart. J.
Florida Acad. Sei. 29): "97-110.

Peckham, G. W. and E. G. Peckham. 1894. Spiders of the Marptusa group.
Occ Papers Nat. Hast. Soe. Wisconsin 2: 85-156.

Philip, C. B. 1957. New records of Tabanidae (Diptera) in the Antilles.

Amer. Mus. Nov. 1858: 1-16.

---------- 1958. New records of Tabanidae (Diptera) in the Antilles.
Supplemental report. (South Caicos). Amer: Mus. Nov. 1921: 1-7.

Porter, John &. 1967. A’ check list of the mosquitoes of the Greater
Antilles and the Bahama and Virgin Islands. Mosquito News 27:
35-41.

Powell, Jerry A. 1973. A systematic monograph of New World Ethmiid
moths (Lepidoptera: Gelechioidea). Smithsonian Contr. Zool. 120:
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Rehn, James A. G. 1906. The Orthoptera of the Bahamas. Amer. Mus.
Nae ss eBUEL f922 se O7 AS

36

Rindge, Frederick H. 1952. The butterflies of the Bahama Islands,
British West Indies (Lepidoptera). Amer. Mus. Nov. 1563: 1-18.

---------- 1955. The butterflies of the Van Voast-American Museum of
Natural History Expedition to the Bahama Islands, British West
Indies. Amer. Mus. Nov. 1715: 1-20.

Ruckes, Herbert. 1952a. Some Scutelleroid Hemiptera of the Bahama
Islands, British West Indies. Amer. Mus. Nov. 1591: 1-9.

~--------- 1952b. Two new species of Thyanta Stal (Pentatomidae,
Heteroptera). Bull. Ent. Soc. Brooklynt 47: 65-68:

Sabrosky, C. W. 1959. A revision of the genus Pholeomyia in North
America (Diptera: Milichiidae). Amer. Ent. Soc. Ann. 52: 316-331.

Sakimura, K. 1973. Synonymies and collection record of Rhaebothrips
lativentris Karny (Thysanoptera: Phlaeothripidae). Pacific
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Scott, James A. 1970. A list of Antillean butterflies. J. Res.
Lepidoptera 9: 249-256.

~--------- 1972. Biogeography of Antillean butterflies. Biotropica 4:

Selander, Richard B. and John K. Bouseman. 1960. Meloid bettles
(Coleoptera) of the West Indies. Proc. U.S. Natl. Mus. 3428,
Vol. 111: 197-226.

Sharpe, Emily Mary. 1900. On a collection of butterflies from the
Bahamas. Zool. Soc. London Proc. 1900: 197-203.

Smith, Marion R. 1954. Ants of the Bimini Island group, Bahamas,
British West Indies (Hymenoptera, Formicidae). Amer. Mus. Nov.
L67is T=16.

Snyder, Fred M. 1958. Muscidae from the Bahama Islands (Diptera).
Amer. Mus. Nov. 1893: 1-4.

Spielman, Andrew and Albert E. Weyer. 1965. Description of Aedes
(Howardina) albonotatus (Coquillett), a common domestic mosquito
from the Bahamas. Mosquito News 25: 339-343.

Stannard, L. J., Jr. 1955. The species and subspecies of North
American Allothrips (Thysanoptera; Phaeothripidae). Ann. Ent. Soc.
Amer. 48: 151-157.

Strohecker, H. F. 1953. The Gryllacrididae and Gryllidae of the
Bahama Islands, British West Indies. Amer. Mus. Nov. 1618: 1-11.

Bi

Valentine, Barry D. 1955. The Antribidae of the Bahama Islands,
British West Indies (Coleoptera). Amer. Mus. Nov. 1741: 1-11.

Vaurie, Patricia. 1952a. Insect collecting in the Bimini Island group,
Bahama Islands. Amer. Mus. Nov. 1565: 1-24.

---------- 1952b. The checkered beetles of the Bahama Islands, British
West Indies (Coleoptera, Cleridae). Amer. Mus. Nov. 1547: 1-5.

Ward, R. A. 1955. Biting lice of the genus Saemundssonia (Mallophaga;
Philopteridae) occurring on terns. Proc. U.S. Natl. Mus. 105
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Weber, Neal A. 1967. The fungus-growing ant, Trachymyrmex jamaicensis,
on Bimini Island, Bahamas (Hymenoptera: Formicidae). Ent. News
78: 107-109.

Werneck, F. L. 1951. Notas sObre mal6fagos (Gyropidae). Rev. Brasil.
Biole Wl: 303-313.

Westfall, Minter J., Jr. 1960. The Odonata of the Bahama Islands, the
West Indies. Amer. Mus. Nov. 2020: 1-12.

Wheeler, William Morton. 1905. The ants of the Bahamas, with a list of
the known West Indian species. Bull. Amer. Mus. Nat. Hist. 21:
7I—1:35-.

coor ----- 1934. Some ants from the Bahama Islands. Psyche 41: 230-232.

Wilson, Edward 0. 1964. The ants of the Florida Keys. Breviora 210:
1-14.

Wirth, Willis W. 1956. The Ephydridae (Diptera) of the Bahama Islands.
Amer. Mus. Nov. 1817: 1-20.

woccccc--- and F. S. Blanton. 1956. A new species of salt-marsh sand
fly from Florida, the Bahamas, Panama, and Ecuador: its distribu-
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(Diptera, Heleidae). Florida Ent. 39: 157-162.

and R. W. Williams. 1957. The biting midges of the Bermuda
Islands, with descriptions of five new species (Diptera, Heleidae).
Proc. Ent. Soc. Washington 59: 5-14.

Young, Frank N. 1953. The water beetles of the Bahama Islands, British
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=a am = 1963. A new species of Laccophilus from the Bahamas
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38

F. FOLKLORE

Armbrister, Hilda. 1917. Proverbs from Abaco, Bahamas. J. Amer.
Folk-lore 30: 274.

Berlitz, C€,) 1969. The mystery of Atlantis. Grosset) and) Dunlap:
New York Neayi

Clavel, M. 1904. Items of .folk-lore from Bahama Negroes. J. Amer.
Folk-lore 17: 36-38.

Cleare, W. T. 1917.. Four folk-tales from Fortune Island, Bahamas.
J. Amer. Folk-lore 30: 228-229.

Crowley, Daniel J. 1954. Form and style in a Bahamian folktale.
Cartbe Quart 152 Dis 25S.

---------- 1966. I could talk old story good: creativity in Bahamian
folk lore. Folk Studies No. 17, Univ. California Press. 157 pp.

Curry, Robert A. 1928. Bahamian lore. Privately printed. Paris.
125, pp... (original edition, 1010 ‘copies; 2nded. (1930); sl0S0 scopuiese)

Edwards, C. L. 1889. Folk-lore of the Bahama Negroes. Amer. J. Psych.
Za 519-542.

---------- 1891. Some tales from Bahaman folk-lore. J. Amer. Folk-
lore 3: 47-54;, 247-252.

Ferro, Robert and Michael Grumley. 1970. Atlantis - the autobiography
of a search. Doubleday,., Garden ‘City, N.. Y..° 168 pp.

Finlay, H. H. 1925. Folklore from Eleuthera, Bahamas. J. Amer. Folk-
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Fitz-James, James. 1909. Bahamian folk lore. Montreal. 64 pp.

Harrison, Wyman. 1971. Atlantis undiscovered; Bimini, Bahamas.
Nature 230 (No. 5292): 287-289.

Maynard, Charles J. 1893. Folk-lore among the West Indians. Contr.
SCL 2s Va 23.

Olschki, Leonard. 1941. Ponce de Leon's fountain of youth: history of
a geographical myth. Hispanic Amer. His. Rev. 21: 361-385.

Parsons, Elsie Clews. 1917. Riddles from Andros Island, Bahamas.
J. Amer. Folk-lore 30: 275-277.

---------- 1918. Folk tales of Andros Island, Bahamas. Amer. Folk-
lore Soc. Mem. 13: 1-167.

39

Peek, Basil (arranged and illustrated by). 1949. Bahamian Proverbs.
The Providence Press. Nassau. (Reprinted 1971).
G. GEOLOGY, GEOPHYSICS, AND GEODESY
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---------- 1934. Sedimentation on the Great Bahama Bank. Geogr. J.

---------- 1957: Tongue of the Ocean. Sea Frontiers 3: 147.

---------- 1967. Environmental atlas of the Tongue of the Ocean,
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---------- 1970. Deep sea drilling project: Leg 11. Geotimes 15: 14-16.

Andrews, J. E. 1967. The Bahama Canyon system. Ph.D. dissertation.
University of Miami. Coral Gables, Florida. 104 pp.

---------- 1968a. Morphology of the outer end of the Bahama submarine
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---------- 1968b. Development of the Eleuthera Ridge by large turbidity
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---------- 1968c. The Bahama Canyon system (abst.). Diss. Abst.,
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---------- 1970. Structure and sedimentary development of the outer
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---------- » Francis P. Shepard, and Robert J. Hurley. 1970. Great
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Ball, M. 1967a. Tectonic control of the configuration of the Bahama
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---------- 1967b. Carbonate sand bodies of Florida and the Bahamas.
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---------- wake Mu Gaudet.) and’ G. Lest. 21968. oSparker meflection
seismic measurements in Exuma Sound, Bahamas (abst.). Amer. Geophy.
Union Trans. 49: 196-197.

---------- CHG Ar sHanrimisonm, ied Hurley . jand:C2nE . iLerstivet. al’.
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Great Bahama Bank and Exuma Sound. Bull. Mar. Sci. 19: 243-252.

40

Ball,:.M..;.B; B.: DashsiG. GA, Harruson. et als lO712 Retractaen
seismic measurements in the northeastern Bahamas (abst.). Eos
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Banks, J. E. 1967. Geologic history of the Florida-Bahama Platform.
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Bathurst, R. G. C. 1967a. Oolitic films in low energy carbonate sand
grains, Bimini Lagoon, Bahamas. Mar. Geol. 5: 89-110.

---------- 1967b. Subtidal gelatinous mat, sand stabilizer and food,
Great Bahama Bank. J. Geol. 75: 736-738.

---------- 1968. Precipitation of ooids and other aragonite fabrics in
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---------- 1969. , Bimini Lagoon. in: H. G. Multer, ed. Field Gurdegto
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Sada ea 1971. Carbonate sediments and their diagenesis. (Develop-
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Bavendamm, W. 1932. Die Mikrobiologische Kalkfallung in der tropischen
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Beales, F. W. 1957. Bahamites and, their significance .in oi) explona-
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Bemrose, John. 1950. Bahamas airborne magnetometer survey.
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41

Bemrose, John, et al. 1948. The Bahamas airborne-magnetometer survey
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Benjamin, George J. 1970. Diving into the blue holes of the Bahamas.
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---------- 1930b. Exploring the Great Bahama Bank. Discovery 11: 75-78.

---------- 1933a. The algal sediments of Andros Island, Bahamas.
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=--------- 1933b. The precipitation of calcium carbonate on the Great
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a--------- 1933c. The geology and sedimentation of Andros Island and
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ma-------- 1962. Mechanical characteristics and mineralogy of the
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Boon, J. D. 1968. Trend surface analysis of sand tracer distribution
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Bornhold, Brian D. 1970. Carbonate turbidites in Columbus Basin,
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---------- and Orris H. Pilkey. 1971. Bioclastic turbidite sedimenta-
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POWiniC. On, 0R. lL. Chase, and J. B. Hersey. 1967, Geclogical
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42

Bracey, D. R. 1963. Geologic interpretation of marine magnetic data
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Broecker, Wallace S. and Taro Takahashi. 1965. Calcium carbonate
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anne and D. L. Thurber. 1965. Uranium-series of dating of corals
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wanna and J. van Donk. 1970. Insolation changes, ice volumes, and
the 018 record in deep-sea cores. Rev. Geophy. Space Physics 8:
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Brooks, H. K. 1965. Submarine scarps of the Bahama Banks (abst.).
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Bryan, G. M. and R. G. Markel. 1966. Microtopography of the Blake-
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Bubb, John N. and Donald K. Atwood. 1968. Recent dolomitization of
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Buchanan, Hugh and S. Stephen Streeter. 1970. Determination of -
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Buckley, James D. and Eric H. Willis. 1970. Isotopes’ radiocarbon
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Burns, Warren W. 1947. Bahamas oil exploration. Petrdleo
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Busby, R. F. 1962. Submarine geology of the Tongue of the Ocean,
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Cameron, Barry. 1968. Ecological determination of the source area of
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43

Chaplin, C. C. G. 1962. The Hogsty Reef. Frontiers 27: 41-45.

Chilingar, George Varos. 1960. Ca/Mg ratios of calcareous sediments as
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Christman, R. A. 1956. Review - Landforms of the southeastern Bahamas
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Clay, C. S. and P. A. Rona. 1963. On the existence of bottom corruga-
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Cloud, P. E., Jr. 1955. Bahama Banks west of Andros Island. (abst.).
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---------- » ed. 1962. Environment of calcium carbonate deposition
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Coogan, A. H. 1969a. Bahamian and Floridian biofacies. in: H. G.
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eooc------ 1970. Measurements of compaction in oolitic grainstone.
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Craig, G. Y. 1967. Size-frequency distributions of living and dead
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Cunningham, Richard T. 1966. Evaluation of Bahamian oolitic aragonite
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Dana, J. D. 1872. Corals and coral islands. Dodd, Mead, and Co.
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Dietz, Robert S., John C. Holden, and Walter P. Sproll, 1970.
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44

Donahue, J. 1968. Recent carbonate sediment in the Bahamas: a review.
Proc. 5th Carib. Geol. Conf., St. Thomas, Virgin Is. July 1968.

cor ccc---- 1969. Genesis of oolite and pisolite grains - an energy
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Eardley, A. J. 1962. Structural geology of North America. 2nd ed.
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Ellis, Charles W. 1968. Recent carbonate sedimentation and diagenesis
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Field, Richard M. 1928. The Great Bahama Bank. Studies in marine
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---------- et al. 1931. Geology of the Bahamas. Geol. Soc. Amer.
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45

Field, Richard M. and H. H. Hess. 1933a. Borehole in the Bahamas
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---------- and ----------. 1933b. A bore hole in the Bahamas. Amer.
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Fruth, Lester Sylvester, Jr. 1967. Compaction effects and depth-
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» G. R. Orme, and F. A. Donath. 1966. Experimental compac-
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Garman, R. K. 1960. A geological and geochemical investigation of a
deep well on Andros Island, Bahamas. M.S. thesis. Florida State

Univ. Tallahassee, Florida.

---------- and H. G. Goodell. 1962. Geochemistry and petrography of
the Superior 0il Company test well on Andros Island, B.W.I. as
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Garrett, Peter. 1970. Phanerozoic stromatolites; noncompetitive
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Gibson, T. G. and J. Schlee. 1967. Sediments and fossiliferous rocks
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Gifford, J. A. 1971. Geological investigation of submerged feature
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Wenn na, Owen P. Bricker, and Harold R. Wanléss. 1970. Exposure
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Racer namnm-y ---------- » and ----------, and Peter Garrett. 1971.

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46

Ginsburg, Robert N. and H. A. Lowenstam. 1958. The influence of
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---------- and E. L. Martin. 1965. Radiocarbon ages of oolitic sands
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Recent carbonate sediments from Bimini, Bahamas. J. Sed. Petrol.
42: 211-226.

---------- 1973. Carbonate needles; additional organic sources. Geol.
Soc. Amer. Bull. 84: 927-930.

w--------- and Noel H. Watkins. 1969. Environmental and phylogenetic
control of the chemical composition and fine structure of gorgonian
spicules from Bimini, Bahamas. Geol. Soc. Amer. abst. with program,
part 4: 77-78.

58

Stoddart, D. R. and J. R. Cann. 1965. Nature and origin of beach rock.
J. Sed. Petrol. 35: 243-247.

Supko, Peter R. 1969. Diagenetic patterns in subsurface Bahaman rocks,
San Salvador Island (abst.). in: Geology of the American Mediterra-
nean: Gulf Coast Assn. Geol. Soc. Trans. 19: 504. (Also Amer. Assn.
Petrol. Geol. Bull. 53: 2045.)

---------- 1970. Some aspects of the geology of Bimini, Bahamas. in:
Sedimentary environments and carbonate rocks of Bimini, Bahamas.
pp. 5-14C.

---------- 197la. Depositional and diagenetic features in subsurface
Bahamian rocks (abst.). Diss. Abst. 31: 6733B.

---------- 1971b. Aspects of Bahamian dolomites and insights into their
origin (abst.). Amer. Assn. Petrol. Geol. Bull. 55: 366.

---------- 1971c. "Whisker" crystal cement in a Bahamian rock. in: Car-
bonate cements. Johns Hopkins Univ. Studies in Geol. No. 19: 143-
146.

---------- » D. M. Marszalek, and W. D. Bock. 1970. Sedimentary environ-
ments and carbonate rocks of Bimini, Bahamas: Guidebook, 4th Ann.
Field trip, Miami Geol. Soc. 27 pp.

Taft, W. H. 1968. Yellow Bank, Bahamas: a study of modern marine car-
bonate lithification. Amer. Assn. Petrol. Geol. Bull. 52: 551.

---------- » F. Arrington, A. Haimovitz, C. MacDonald, and C. Woolheater.
1968. Lithification of modern carbonate sediments at Yellow Bank,
Bahamas. Bull. Mar. Sci. Gulf Carib. 18: 762-828.

---------- » and J. W. Harbaugh. 1964. Modern carbonate sediments of
southern Florida, Bahamas, and Espiritu Santo Island, Baja Califor-
nia: a comparison of their mineralogy and chemistry. Stanford Univ.
Publ. Univ. Ser. Geol. Sci. 8: 1-133.

Talwani, M., J. L. Worzel, and M. Ewing. 1959. Gravity anomalies and
structure of the Bahamas. Lamont Geol. Obs. Columbia Univ. New
York,aN. Y. -9i.pp:

lalate » 7rocccccco-, and ---------- 1960. Gravity anomalies and struc-
ture of the Bahamas. Trans. 2nd Carib. Geol. Conf., Mayaguez, Puerto
Rico, 4-9 January, 1959. pp. 156-161.

Thorp, Eldon M. 1934. Preliminary remarks on the calcareous shallow-
water marine deposits of Florida and the Bahamas. J. Sed. Petrol.
4: 111-112.

---------- 1936. Calcareous shallow-water marine deposits of Florida and
the Bahamas. Carnegie Inst. Wash. (Papers from the Dept. of Mar.
Biol.) 29: 37-119.

59

Thorp, Eldon M. 1939. Florida and the Bahama marine calcareous deposits.
in: P. D. Trask, et al. Recent marine sediments. Thomas Murby and
Co., London. pp. 283-297.

Thurber, David L., Edward George Purdy, and Wallace S. Broecker. 1958.
Radiocarbon studies of the Bahama Banks (abst.). Geol. Soc. Amer.
Bali. 69> F652.

Till, Roger. 1970. The relationship between environment and sediment
composition (geochemistry and petrology) in the Bimini Lagoon,
Bahamas. J. Sed. Petrol. 40: 376-385.

Trask, Parker D. 1939. Additional note to E. M. Thorp, Florida and
Bahama marine calcareous deposits. in: P. D. Trask. Recent marine
deposits, p. 292-293. Reprinted 1955. Amer. Assn. Petrol. Geol.
Tulsa, Oklahoma.

Traverse, Alfred and Robert N. Ginsburg. 1966. Palynology of the sur-
face sediments of the Great Bahama Bank, as related to water move-
ment and sedimentation. Mar. Geol. 4: 417-459.

---------- and ---------- 1967. Pollen and associated microfossils in
the marine surface sediments of the Great Bahama Bank. Rev. Palaeo-
botany and Palynology 3: 243-254.

Uchupi, Elazar, J. D. Milliman, and Bruce P. Luyendyk, et al. 1971.
Structure and origin of the southeastern Bahamas. Amer. Assn.
Petrol. Geol. Bull. 55: 687-704.

Vaughan, Thomas Wayland. 1913a. Studies of the geology of the Madre-
poraria of the Bahamas and of southern Florida. Carnegie Inst.
Wash. Yearb. 11: 153-162.

aoc o----- 1913b. Remarks on the geology of the Bahama Islands, and on
the formation of the Floridian and Bahaman oolites (abst.). Wash.
Kead. Sci: J. 3: 302-304.

---------- 1914a. Preliminary remarks on the geology of the Bahamas, with
special reference to the origin of the Bahaman and Floridian oolite.
Carnegie Inst. Wash. Publ. 182 (Papers from the Tortugas Lab. Vol.
5): 47-54.

m--------- 1914b. Investigations of the geology and geological processes
of the reef tracts and adjacent areas in the Bahamas and Florida.
Carnegie Inst. Wash. Yearb. 12: 183-184.

---------- 1914c. Geological investigations in the Bahamas and southern
Florida. Carnegie Inst. Wash. Yearb. 13: 227-253.

---------- 1914d. Sketch of the geologic history of the Florida coral
reef tract and comparisons with other coral reef areas. J. Wash.
Acad. Sci. 4: 26-34.

60

Vaughan, Thomas Wayland. 1915. The geologic significance of the growth-
rate of the Floridian and Bahamian shoalwater corals. Wash. Acad.
Scie iJ. 52 59T-G00K

=== --<-55- et al. 1918. Some shoal-water bottom samples from Murray
Island, Australia, and comparisons of them with samples from Florida
and the Bahamas. Carnegie Inst. Wash. Publ. 213 (Papers from Dept.
Mar.“ Brol.” Vol 3:9), 255=297

-~--------- » M. A. Howe, et al. 1919. Contributions to the geology and
paleontology of the West Indies. Carnegie Inst. Wash. Publ. 291.
184 pp.

Walper, Jack L. 1971. Geotectonic evolution and subsidence of Bahama
Platform; discussion. Geol. Soc. Amer. Bull. 82: 1129-1130.

Wanless, Harold R. 1961. Sedimentary structure zonation on tidal levees,
Andros Isiand, Bahamas (abst.). Amer. Assn. Petrol. Geol. Bull. 53:
748. :

Wassall, Harry William, 3rd. 1956. Geological Bibliographies, 1785-1955.
Bahama Islands. 4 pp. U. S. Geol. Surv., Washington, D. C.

a ateieietatetetatan and Howard Dalton. 1959. Oil prospects in the Bahamas.
World Oil 148: 85-89.

Weeks, L. G. 1948. Highlights on 1947 development in foreign petroleum
fields; Bahamas. Amer. Assn. Petrol. Geol. Bull. 32: 1108.

Weyl, P. K. 1963. Book review: Environment of calcium carbonate deposi-
tion west of Andros Island, Bahamas, by P. E. Cloud (1962). Limnol.
Oceanog. 8: 494.

Winland, H. Dale and R. K. Matthews. 1969. Origin of recent grapestone
grains, Bahama Islands (abst.). Geol. Soc. Amer. Abst. with pro-
grams, 1965. part 7: 239.

Woodring, W. P. 1954. Caribbean land and sea through the ages. Geol.
Soc. Amer. Bull. 65: 719-782. |

Worzel, John Lamar, William Maurice Ewing, and Charles Lum Drake. 1953.
The Bahama Islands regions. Part 1 of Gravity observations at sea
(abst.). Geol. Soc. Amer. Bull. 64: 1494-1495. Also Amer. Mineral.
39: 349-350 (1954).

Young, R. B. 1934. A comparison of certain stromatolitic rocks in the
dolomite series with modern algal sediments in the Bahamas. Geol.
Soc. Oo. Atr.: Trans. 37: 155-1622

61

H. HERPETOLOGY

Barbour, Thomas. 1904. Batrachia and Reptilia from the Bahamas.
Bull. Mus. Comp. Zool. 46: 55-61.

---------- 1906. Additional notes on Bahama snakes. Amer. Nat. 40:
229-232.

---------- 1910. A note regarding the green Anolis from the northern
Bahamas. Proc. Biol. Soc. Wash. 23: 99-100.

---------- 1914. A contribution to the zoogeography of the West Indies,
with especial reference to amphibians and reptiles. Mem. Mus. Comp.
Zool. 44: 205-359.

a--------- 1921. Sphaerodactylus. Mem. Mus. Comp. Zool. 47: 217-278
20 pl.

wo-------- 1923. Another new Bahaman iguana. New Eng. Zool. Club
Proc. 6: 107-109.

---------- 1930a. A list of Antillean reptiles and amphibians.,
Zoologica 11: 61-116.

---------- 1930b. The Anoles. 1. The forms known to occur on the
neotropical islands. Bull. Mus. Comp. Zool. 70: 105-144.

---------- 1935. A second list of Antillean reptiles and amphibians.
Zoologica 19: 77-141.

eo-------- 1937. Third list of Antillean reptiles and amphibians.
Bull. Mus. Comp. Zool. 82: 77-166.

~--------- 1941. A new boa from the Bahamas. New Eng. Zool. Club.
BEOcer lo: Ol-65,.

---------- and Benjamin Shreve. 1935. Concerning some Bahamian rep-
tiles, with notes on the fauna. Proc. Boston Soc. Nat. Hist. 40:
347-365.

---------- and ---------- 1936. New races of Tropidophis and Ameiva
from the Bahamas. Proc. New Eng. Zool. Club 16: 1-3.

Buden, Donald W. and Albert Schwartz. 1968. Reptiles and birds from the
Cay Sal Bank, Bahama Islands. Quart. J. Florida Acad. Sci. 31:
290-320.

Carr, A. F. 1957. Notes on the zoogeography of the Atlantic sea turtles
of the genus Lepidochebys. Rev. Biol. Trop. 5: 45-61.

62

Carr, Archie, Harold Hirth, and Larry Ogren. 1966. The ecology and
migrations of sea turtles, 6. The hawksbill turtle in the Carib-
bean Sea. Amer. Mus. Nov. 2248: 1-29.

Cochran, Doris M. 1934. Herpetological collections from the West
Indies made by Dr. Paul Bartsch under the Walter Rathbone Bacon
Scholarship, 1928-1930. Smithsonian Misc. Coll. 92: 1-48.

Cope, E. 1885. A contribution to the herpetology of Mexico, appendix
on a collection from New Providence, Bahama Islands. Proc. Amer.
Phil. Soc. Phila. 22: 379-404.

la taal! 1887. List of the Batrachia and Reptilia of the Bahama
Islands. Proc. U. S. Natl. Mus. 10: 436-439.

Crombie, Ronald I. 1972. The presence of Hyla squirella in the Bahamas.
Quart. J. Florida Acad. Sci. 35: 49-52.

Etheridge, Richard. 1965. Pleistocene lizards from New Providence.
Quart. J. Florida Acad. Sci. 28: 349-358.

a-5------- 1966. The systematic relationships of West Indian and South
American lizards referred to the Iguanid genus Leiocephalus.
Copeia 1966: 79-91.

Garman, Samuel. 1887a. On West Indian Teiidae in the Museum of Compara-
tive Zoology at Cambridge. Bull. Essex Inst. 19: 1-12.

---------- 1887b. On West Indian reptiles. Iguanidae. Bull. Essex
Inst. 192:25-50 .

---------- 1887c. West Indian Batrachia in the Museum of Comparative
Zoology. Bull. Essex Inst. 19: 13-16.

---------- 1888. Reptiles and batrachians from the Caymans and Bahamas.
Bull. Essex Inst. 20: 1-13.

Goin, Coleman J. 1955. Description of a new subspecies of the frog
Eleutherodactylus ricordi from the Bahamas. Amer. Mus. Nov. 1708:
1-7.

Hanlon, Robert W. 1964. Reproductive activity of the Bahaman boa
(Epicrates striatus). Herpetologica 20: 143-144.

Hecht, Max. L., Vladimir Walters, and Gordon Ramm. 1955. Observations
on the natural history of the Bahaman pigmy boa, Tropidophis
pardalis, with notes on autohemorrhage. Copeia 1955: 249-251.

eeccco---- 1955. The comparison of recent and fossil amphibian, rep-
tilian, and mammalian faunas in the Bahamas. Year Book Amer. Phil.
Soc. for 1954: 133-135.

63

Hodsdon, L. A. and J. F. W. Pearson. 1943. Notes on the discovery and
biology of two Bahaman fresh-water turtles of the genus Pseudemys.
Quart. J. Florida Acad. Sci. 6: 17-23.

Laska, Anthony L. 1970. The structural niche of Anolis scriptus on
Inagua. Breviora 349: 1-6.

Legler, J. M. 1960. Remarks concerning Bahaman blind snakes (genus
Typhlops). Herpetologica 16: 71-72.

Maglio, Vincent J. 1970. West Indian Xenodontine Colubrid snakes:
their probable origin, phylogeny, and zoogeography. Bull. Mus.
Comp. Zool. 141: 1-54.

Maynard, Charles Johnson. 1889. Review: Reptiles and batrachians from
the Caymans and the Bahamas by Samuel Garman. Contr. Sci. 1: 48.

Neill, Wilfred T. 1958. The occurrence of amphibians and reptiles in
saltwater areas and a bibliography. Bull. Mar. Sci. Gulf Carib.
Ss 1-977.

---------- 1964. Frogs introduced on islands. Quart. J. Fla. Acad.
Siew 2 73) 27-130.

Nesbitt, C. R. 1836. On the Bahama fisheries. J. Bahama Soc. Diff.
Knowl. No. 11: 126-136.

Netting, M. Graham. 1954. To Bimini for boas and exhibit ideas.
Carnegie Mag. 28: 185-188.

---------- and Coleman J. Goin. 1944. Another new boa of the genus
Epicrates from the Bahamas. Ann. Carnegie Mus. 30: 71-76.

Sapte.) G., K. and G. C. Klingel. 1952. The reptiles of Great Inagua
Island, British West Indies. Amer. Mus. Nov. 549: 1-25.

Oliver, J. A. 1948. The anoline lizards of Bimini, Bahamas. Amer.
Mus. Nov. 1383: 1-36.

Rand, A. Stanley. 1969. Competitive exclusion among anoles (Sauria:
Iguanidae) on small islands in the West Indies. Breviora 319: 1-16.

Rebach, J. A. 1971. Comparison of gas exchange and water balance of the

nutria, Myocastor coypus, and the hutia, Geocapromys ingrahami.
Ph.D. dissertation, University of Rhode Island. Kingston, R. I.

Richmond, Neil D. 1955. The blind snakes (Typhlops) of Bimini, Bahama
Islands, British West Indies, with a description of a new species.
Amer. Mus. Nov. 1734: 1-7.

64

Rosen, Nils. 1911. Contribution to the fauna of the Bahamas. I. A
general account of the fauna, with remarks on the physiography of
the rslands. - Il. The reptilies., Til. The fishes. jlunds Unawe

rssk. N.F. Afd. 2, Bd. 7 (Acta Universitatis Lundensis, nova
series): 3-72.

Schoener, Thomas W. 1968. The Anolis lizards of Bimini: resource
partitioning in a complex fauna. Ecology 49: 704-726.

Schwartz, Albert. 1965. Geographic variation in Sphaerodactylus
notatus Baird. Rev. Brol. Trop. 13% 161-185;

---------- 1967. The Leiocephalus (Lacertilia, Iguanidae) of the
southern Bahama Islands. Ann. Carnegie Mus. Art. II, 39: 153-185.

---------- 1968a. The geckos (Sphaerodactylus) of the southern Bahama
Islands. Ann. Carnegie Mus. 39: 227-271.

---------- 1968b. Geographic variation in Anolis distichus Cope
(Lacertilia, Iguanidae) in the Bahama Islands and Hispaniola.
Bull. Mus. Comp. Zool. 137: 255-309.

woecccon-- 1968c. Geographic variation in the New World geckkonid liz-.
ard Tarentola americana Gray. Proc. Biol. Soc. Wash. 81: 123-142.

---------- 1972. A new subspecies of Sphaerodactylus decoratus from
the Bahama Islands. Herpetologica 28: 247-251.

---------- and Robert J. Marsh. 1960. A review of the Pardalis-
Maculatus complex of the boid genus Tropidophis of the West Indies.
Bull. Mus. Comp. Zool. 123: 49-84 + 10 fig.

as-------- and Clarence J. McCoy. 1970. A systematic review of Ameiva
auberi in Cuba and the Bahamas. Ann. Carnegie Mus. 41: 45-168.

---------- and Richard Thomas. 1968 (appeared 1969). A review of
Anolis angusticeps in the West Indies. Quart. J. Florida Acad.
Servs: SE 69%

Smith, F. G. Walton. 1954. Taxonomy and distribution of sea turtles.
Fish Bull) U2tS..789* SaS=S1Ss

Stejneger, Leonhard. 1901. Diagnosis of a new species of iguanoid 1liz-
ard from Green Cay, Bahama Islands. Proc. U. S. Natl. Mus. 23: 471.

---------- 1903. A new species of large iguana from the Bahama Islands.
Proc. Biol sSoc.Wash..016:. 129-152

---------- 1905. Batrachians and land reptiles of the Bahama Islands.
in: Shattuck, G. B. ed. The Bahama Islands. Geographical Society
of Baltimore, Johns Hopkins Press. pp. 329-343.

65

Sutcliffe, Robert. 1952. Results of the Catherwood-Chaplin West Indies
Expedition, 1948. Part vi. Amphibia and reptiles. Not. Nat. 243:
1-8.

Thomas, Richard. 1968a. The Typhlops biminiensis group of Antillean
blind snakes. Copeia 1968: 713-722.

---------- 1968b. Notes on Antillean geckos (Sphaerodactylus).
Herpetologica 24: 46-60.

-o-------- and Albert Schwartz. 1966. The Sphaerodactylus decoratus
complex in the West Indies. Brigham Young Univ. Sci. Bull. Biol.
Sera. -1-26.

Wilcox, Karen, Jeanne Y. Carter, and L. V. Wilcox, Jr. 1973. Range
extension of Cyclura figginsi Barbour in the Bahamas. Carib. J.
Semen 52 211-213;

Williams, Ernest E. 1969. The ecology and colonization as seen in the
zoogeography of anoline lizards on small islands. Quart. Rev. Biol.
44: 345-389.

I. ICTHYOLOGY

Anonymous. 1974. World record fish. Saltwater records. Field and
Stream 78 (10): 120.

Alexander, Elizabeth C. 1961. A contribution to the life history,
biology, and geographical distribution of the bonefish, Albula
vulpes (Linnaeus). Dana Rept. No. 53: 1-51.

Anderson, William W. 1957. Larval forms of the fresh-water mullet
(Agonostomus monticola) from the open ocean off the Bahamas and
South Atlantic coast of the United States. Fish Bull. U. S. 57:
415-425.

Aronson, L. R. 1971. Further studies on orientation and jumping behav-
ior in the gobiid fish, Bathygobius soporator. Ann. N. Y. Acad.
Sci. 188: 378-392.

Austin, H. M. 1969. An unusual association between a jack and a ray.
Underw. Nat. 6: 39.

Bean, Barton A. 1905. Fishes of the Bahama Islands. in: George B.

Shattuck, ed. The Bahama Islands. Geographical Society of Balti-
more. Johns Hopkins Press. Baltimore, Maryland. pp. 293-325.

Bohlke, James E. 1955a. The brotulid fish genus Petrotyx from the
Great Bahama Bank. Not. Nat. 273: 1-6.

w--------- 1955b. On the Bahaman fishes of the family Opisthognathidae.
Not. Nat. 281: 1-6.

=--------- 1955¢. A new genus and species of Ophichthid eels from the
Bahamas. Not. Nat. 282: 1-7.

66

Bohlke, James E. 1957a. A new shallow-water brotulid fish from the
Great Bahama Bank. Not. Nat. 295: 1-8.

---------- 1957b. A review of the blenny genus Chaenopsis, and a
description of a related new genus from the Bahamas. Proc. Phila.
Acad. Nat. Sei. 1093. 81-1105.

---------- 1957c. The Bahaman species of emblemariid blennies. Proc.
Phaila.. Acad’. Nat. Sci, 2h09s (25-57.

---------- 1959a. A new cardinal fish (Apogonidae) from the Bahamas.
Noe. Nat S19 oor

---------- 1959b. A new fish of the genus Hypsoblennius (Blenniidae)
from the Bahamas. Not. Nat. 321: 1-5.

ae ee 1959¢c. Studies ‘on: fishes of the family 'Ophidirdae. (ils aihnee
new species from the! Bahamas... Proc. Acad. Nat. Scils Phitas ig:
57-52:

---------- 1960a. Two new Bahaman species of the clinid fish genus
Paraclinus)., “Not. Natt. 35577, 1-8:

---------- 1960b. Comments on serranoid fishes with disjunct lateral
lines, with the description of a new one from the Bahamas. Not.
Nat. 3303. 1-11).

---------- 1961. Two new Bahaman soles of the genus Symphurus (family
Cynoglossidae). Not. Nat. .344: 1-4.

---------- 1963. The species of the West Atlantic gobioid genus
Psilotras. Not. Nat. 562:"1-10:

---------- 1967. A new sexually dimorphic jawfish (Opistognathus:
Opistognathidae) from the Bahamas. Not. Nat. 407: 1-12.

---------- 1969. Pariah scotius, a new sponge-dwelling gobid fish from
the Bahamas. Not. Nat. 421: 1-7.

Sessee=es5 and C. C. G. Chaplin. 1957. Oral incubation in Bahaman jaw-
fishes Opistognathus whitehursti and O. maxillosus. Science 125
(3245))5935'37,

---------- and -----+----- 1968. Fishes of the Bahamas and adjacent
tropical waters. Phila. Acad. Nat. Sci. Livingston Publ. Co.
Wynnewood, Pennsylvania. 771 pp.

---------- and C. R. Robins. 1959. Studies on fishes of the family
Ophidiidae. II. Three new species from the Bahamas. Proc. Phila. -
Acad... Nat. Sci5 Tides 37-52 =

67

Bohlke, James E. and C. R. Robins. 1968. Western Atlantic seven-spined
gobies, with descriptions of ten new species and a new genus, and
connents jon) Pacitive relatives. Proc. Phila. Acad: Nat. Sci. 120:
54-174.

---------- and ---------- 1969. Western Atlantic sponge-dwelling gobies
of the genus Evermannichthys: their taxonomy, habits and relation-
siips.4) Proce. Phila’ Acad iNat. Sci. 121: 1-24.

---------- and D. G. Smith. 1968. A new xenocongrid eel. from the
Bahamas, with notes on other species in the family. Proc. Phila.
Acad: Nat.:Sci. 120: 25-43.

Breder, €. M., Jr. 1932. An annotated list of fishes from Lake Forsyth,
Andros Island, Bahamas. Amer. Mus. Nov. 551: 1-8.

---------- 1933a. Young tarpon on Andros Island. Bull. N. Y. Zool.
SOC 7956 3 165-67 «

---------- 1933b. The significance of Ca to marine fishes on invading
fresh water (abst.). Anat. Rec. 57 (4): 57.

---------- 1934a. A new Gambusia from Andros Island, Bahamas. Amer.
Mus’. 2Nov.'7195 1-3.

---------- 1934b. Ecology of an oceanic fresh-water lake, Andros Island,
Bahamas, with special reference to its fishes. Zoologica 18: 57-88.

Briggs, John C. 1963. A new clingfish of the genus Gobiesox from the
Bahamas. Copeia 1963: 604-606.

---------- 1969. A new genus and species of clingfish (family
Gobiesocidae) from the Bahama Islands. Copeia 1969: 332-334.

buliss,; Harvey R., Jr. and Paul Struhsaker. 1961. Life history notes
on the rough-tail stingray, Dasyatis centroura (Mitchell). Copeia
1961: 232-234.

Cohen, Daniel M. and C. Richard Robins. 1970. A new ophidioid fish
(genus Lucifuga) from a limestone sink, New Providence Island,
Bahamas. Proc. Biol. Soc. Wash. 83: 133-144.

Colin, Patrick L. 1972. Daily activity patterns and effects of environ-
mental conditions on the behavior of the yellowhead jawfish,
Opistognathus aurifrons, with notes on its ecology. Zoologica 57:
137-169.

Cummings, William C., Bradley D. Brahy, and Juanita Y. Spires. 1966.
Sound production, schooling, and feeding habits of the margate,
Haemulon album Cuvier, off North Bimini, Bahamas. Bull. Mar. Sci.
16: 626-640.

68

DeSylva, D. P. 1963. Systematics and life history of the great barra-
cuda Sphyraena barracuda (Walbaum). Stud. Trop. Oceanog. Miami,
No. 1, 179 pp.

Ellington, A. C. 1959. Poisonous fishes in the Caribbean area.
W. Ind. Fish. Bull. (6): 1-5.

Fish, M. P. and W. H. Mowbray. 1959. The production of underwater
sound by Opsanus sp., a new toadfish from Bimini, Bahamas.
Zoologica 44: 71-76.

Fowler, H. W. 1919. Notes on tropical American fishes. Proc. Phila.
Acad. Nat. Sci. 71: 128-155.

Fowler, Henry R. 1947. Notes on Bahama fishes obtained by Mr. Charles
G. Chaplin in 1947, with descriptions of two new species. Not.
Nat. 199: 1-14.

Garman, S. 1896. Report on the fishes collected by the Bahama Expedi-
tion of the State University of Iowa, under Professor C. C. Nutting
in 1893. Bull. Lab. Nat. Hist. State Univ. Iowa 4: 76-93.

Gilbert, Perry A. and Henry Kritzler. 1960. Experimental shark pens
at the Lerner Marine Laboratory. Science 132: 424.

Greenberg, Jerry and Idaz Greenberg. 1972. The living reef: corals,
and fishes of Florida, the Bahamas, Bermuda and the Caribbean.
Seahawk Books. Miami, Florida. 110 pp.

Hazlett, B. and H. E. Winn. 1962. Sound producing mechanism of the
Nassau grouper, Epinephalus striatus. Copeia 2: 447-449.

Herald, Earl S. 1950. Ichthyocampus pawneei, a new pipefish from the
Bahamas. J. Wash. Acad. Sci. 40: 269.

Hess, Paul W. 1962. Notes on some sharks in the western North Atlantic
and Bahama areas. Copeia 1962: 653-656.

Hubbs, Carl L. 1927. Studies of the fishes of the order Cyprinodontes.
VII. G. manni, a new species from the Bahamas. Copeia 164: 61-65.

---------- and R. R. Miller. 1942. Studies of the fishes of the order
Cyprinodontes. XVIII. Cyprinodon laciniatus, new species, from the
Bahamas. Occas. Papers Mus. Zool. Univ. Michigan 458: 1-11.

Jordon, D. S. 1884. An identification of the figures of fishes in
Catesby's Natural History of Carolina, Florida, and the Bahama
Islands. Proc. U. S. Natl. Mus. 7: 190-199.

---------- and C. H. Bollman. 1889. List of fishes collected at Green
Turtle Cay, in the Bahamas. U. S. Natl. Mus. Proc. 11: 549-553.

69

Jordon, D. S. and B. W. Evermann. 1896-1900. The fishes of North and
Middle America: a descriptive catalogue of the species of fish-
like vertebrates found in the waters of North America, north of
the Isthmus of Panama. 4 vol. U.S. Natl. Mus. 47: i-xl + 1-1240;
1-xxx + 1241-2183; i-xxiv + 2183a-3136; l-ci + 3137-3313; + 392 pl.

Kanazawa, Robert H. 1958. A revision of the eels of the genus Conger
with descriptions of four new species. Proc. U. S. Natl. Mus.
108: 219-267.

Klawe, W. L. 1961. Young scombroids from the waters between Cape
Hatteras and Bahama Islands. Bull. Mar. Sci. Gulf. Carib. 11:
150-157.

Krumholz, Louis A. 1957. Measurements of a large sharpnose mackerel
shark, Isurus oxyrinchus, from Bimini, Bahamas. Copeia 1957: 302.

---------- 1959. Stomach contents and organ weights of some bluefin
tuna, Thunnus thynnus (Linnaeus), near Bimini, Bahamas. Zoologica
(New Y.) 4455 127-151".

---------- 1963. Relationships between fertility, sex ratio and exposure
to predation in populations of the mosquitosish Gambusia manni Hubbs
of Bimini, Bahamas. Int. Rev. Gesam. Hydrobiol. 48: 201-256.

LaMonte, Francesca. 1952. Marine games fishes of the world. Doubleday.
GardensGity, N.Y; 190 pp.

---------- 1956. North American game fishes. Doubleday. Garden City,
Neowk wxiv + 202° pp.

Lee, T. 1889. List of fish taken by steamer "Albatross" among Bahama
Islands and at Nassau fish-market during March and- April, 1886.
Rept. U. S. Fish Comm. part 14: 669-672.

Levi, Eldon J. 1973. Juvenile yellowfin menhaden from the Bahama
Islands. Trans. Amer. Fish Soc. 102: 848-849.

Mead, G. W. and J. E. Bohlke. 1958. Gobionellus stigmalophius, a new
goby from the Gulf of Campeche and the Great Bahama Bank. Copeia
1958: 285-289.

Miller, Robert Rush. 1962. Taxonomic status of Cyprinodon baconi, a
killifish from Andros Island, Bahamas. Copeia 1962: 836-837.

Moss, Frank T. 1973. Let's fish the Bahamas! Pastimes (Eastern Air-
lines passenger magazine) 1 (12): 18-23.

Moulton, James M. 1958. The acoustical behavior of some fishes in the
Bimitisnanedene ibtolee Bulle, li 4) eS 5i— 574:

70

Myrberg, A. A., Jr. 1972. Social dominance and territoriality in the

bicolor damselfish, Eupomacentrus portitus (Poey) (Pisces:
Pomacentridae) . Behauileun 41: 207-231

---------- » Samuel J. Ha, Stanley Walewski, and John C. Banbury. 1972.
Effectiveness of acoustic signals in attracting epipelagic sharks
to an underwater sound source. Bull Mar. Sci. 22: 926-949.

Nelson, Donald R., Richard H. Johnson, and Larry G. Waldrop. 1969.
Responses in Bahamian sharks and groupers to low-frequency, pulsed
sounds: 'S). Calaf. vAcade Sci.) Bull) 683 131-1372

Nesbitt, C. R. 1836. On the Bahama fisheries. J. Bahama Soc. Diff.
Knowl. No. 11: 126-136.

Nichols, J. T. 192la. A new ponacentrid and blenny from the Bahamas.
Amer. Mus. Nov. 26: 1-2.

---------- 1921b. A list of Turks Islands fishes, with a description
of a new flatfish. Amer. Mus. Nat. Hist. Bull. 44: 21-24.

-s-------- and Van Campen Heilner. 1928. A rare sole from the Bahamas.
Amer. Mus. Nov. 326: 1.

Norman, J. R. 1931. A new sole from the Bahamas. Amer. Mus. Nov.
AT Seles

Northcroft, George J. H. 1902. Sketches of Summerland - giving some
account of Nassau and the Bahama Islands. Nassau Guardian. Nassau.
Chapter 12: Corals, shells, and fishes. pp. 123-138.

Owre, Harding B. 1972. Marine biological investigations in the Bahamas.
18. The genus Spadella and other Chaetognatha. Sarsia 49: 49-57.

Parr, Albert Eide. 1930. Teleostean shore and shallow-water fishes
from the Bahamas and Turks Island. Bull. Bingham Oceanog. Coll.
33 La lds te 38) baer,

Peterman, R. M. 1971. A possible function of coloration in coral reef
fishes .. “Copeia 1971: 350-331..

Phillips, Craig. 1959. Bahama pigmy angelfish. Aquarium J. 30: 236-237.

Randall, J. E. 1967. Food habits of reef fishes of the West Indies.
Stud. Trop. Oceanog. Miami No. 5: 665-847.

---------- 1968. Caribbean reef fishes. Reigate. T.F.H. Publications,
inc.. Jersey. .City IN. J) Sl8eppe

Rausch, James P. 1973. Marine zoology (a manual prepared for use on
San Salvador Island, Bahamas) 2 vols. College Center of the Finger
Lakes.” Corning; (N..Y.,7 Vol. I= 147 pp.s.Vol.. 2 - 167 pp.

7p

Rivas, Luis Rene. 1960. The fishes of the genus Pomacentrus in Florida
and the western Bahamas. Quart. J. Florida Acad. Sci. 23: 130-162.

---------- 1963. Subgenera and species groups in the poeciliid fish
genus Gambusia Poey. Copeia 2: 331-347.

Robins, C. R. and J. E. Bohlke. 1964. Two new Bahaman gobiid fishes
of the genera Lythrypnus and Garmannia. Not. Nat. 325: 1-9.

Roman, Erl. 1958. Tournaments can aid science (Bahamas International
Tuna Match). Sea Frontiers 4: 164-170.

---------- 1957. The International Game Fish Conference. Sea Frontiers
3: 48-57.

Rosen, Donn Eric and Reeve M. Bailey. 1963. The Poeciliid fishes
(Cyprinodontiformes), their structure, zoogeography, and systemat-
wes. Aner. Mus. Natt Hist. Bull. 126: 1-176.

Rosén, Nils. 1911. Contribution to the fauna of the Bahamas. I. A
general account of the fauna with remarks on the physiography of
tEpeetstands.y Li. The sReptiles., III. The Fishes. Lunds. Univ.
Arssk. N.F. Afd. 2, Bd. 7 (Acta Univ. Lundensis, nova series): 3-72.

Schultz, L. P. 1945. Emmelichthyops atlanticus, a new genus and species
of fish (family Emmelichthyidae) from the Bahamas, with a key to
related genera. J. Wash. Acad. Sci. 35: 132-136.

Smith, C. Lavett. 1964. Hermaphroditism in Bahama groupers. Nat.
Hist. 73: 42-47.

---------- 1972. A spawning aggregation of Nassau grouper, Epinephelus
striatus (Bloch). Trans. Amer. Fish Soc. 101: 256-261.

---------- and James C. Tyler. 1973. Population ecology of a Bahamian
Suprabenthic shore fish assemblage. Amer. Mus. Nov. 2528: 1-38.

Smith, David G. 1968. The occurrence of larvae of the American eel
(Anguilla rostrata) in the Straits of Florida and nearby areas.
Bull. Mar. Sci. 18: 280-293.

emtth, Ff. iG. Walton. 1957. Tracking the elusive tuna. Sea Frontiers
Seml96-209)

Springer, Stewart and Harvey R. Bullis, Jr. 1960. A new species of
sawshark, Pristiophorus schroederi, from the Bahamas . Bull. Mar.
SCis Gulf Carib. 10: 241-254.

---------- and Richard A. Waller. 1969. Hexanchus vitulus, a new six-
gill shark from the Bahamas. Bull. Mar. Sci. 19: 159-174.

72

Sutcliffe, R. 1970. Muraena acutirostris Abbott 1861, a synonym of
the West Atlantic moray Gymnothorax moringa (Cuvier) (Anguillida,
Muraenidae). Carib. J. Sci. 10: 87-91.

Tibbo, S. N. and L. M. Lauzier. 1961. Larval swordfish (Xiphias
gladius) from three localities in the western Atlantic. Fish Res.
Board Canada J. 26: 3248-3251.

Tortonese, Enrico. 1962. Osservazinni: comparative intorno alla
ittiofauna del Mediterraneo e dell'Atlantico occidentale (Florida
e Isole Bahamas). Natura (Milan) 53: 1-20.

Vesey-Fitzgerald, Brian S. and Francesca LaMonte. 1949. Game fish of
the’ world= ~ Harpex:” "New York, “N., Yo" xvai°+ 446 pp.

Voss, Gilbert L. 1953. A contribution to the life history and biology
of the sailfish Istiophorus americanus Cuv. and Val., in Florida
waters. Bull: Mare Seis Gult Caribs-3: 206-240"

Walters, Vladimir. 1957. Alphestes scholanderi, a new sea bass from
the West Indies. Copeia 1957: 283-286.

---------- and C. R. Robins. 1961. A new toadfish (Batrachoididae)

considered to be a glacial relict in the West Indies. Amer. Mus.
Nov. 2047: 1-24.

Wicklund, Robert. 1968. Some night observations on marine animals in
the Bahamas. Underw. Nat. 5: 24-27.

Woods, L. P. 1965. A new squirrel fish (Adioryx pocoy of the family
Holocentridae from the Bahama Islands. Not. Nat. 377: 1-5.

Yocum, Thomas G. -1971. Pinfish and rockcut goby, fishes new to the
Bahamas. Quart. J. Florida Acad. Sci. 34: 131.
K. GEOGRAPHY

Anonymous. 1926. Maps of the Bahama Islands. Published by Authority
(HMSO?). 20 maps + 8 pp. of index.

---------- 1930. The Great Bahama Bank. Geogr. J. 75: 564-565.
----+----- 1947. Directions for the treatment of geographical names in
the Bahama Islands’: Spee. Publ .-6/7.°, Dept. ,of the Interiors aUres.

Board on Geographical Names.

---------- 1955. British West Indies and Bermuda. U. S. Board on
Geographical Names. Dept. of the Interior.

---------- 1967a. Environmental atlas of the Tongue of the Ocean,
Bahamas. U.S. Naval Oceanog. Off. Spec. Publ. SP-94. ..74 ppl.

73

Anonymous. 1967b. Earth photographs from Gemini III, IV, and V.
U.S.N.A.S.A. Spec. Publ. SP-129. 266 pp.

Babcock, William H. 1920. Antillia and the Antilles. Geogr. Rev. 9:
109-124.

Benjamin, George J. 1970. Diving into the blue holes of the Bahamas.
Natl. Geogr. Mag. 138: 347-363.

Blake, H. A. 1887. The Bahamas. Proc. Royal Geogr. Soc. 9: 579-580.

Bogy, Theodoor de. 1919. The less-known regions of the West Indies.
mormocorgr. 18: 57-62:

Bounds, John Howard. 1966. Land use in the Bahamas. Ph.D. Diss. Dept.
of Geography. Univ. Tennessee. Knoxville, Tennessee. 419 pp.

---------- 1969. The Bahamas. Focus 19 (9): 3-7.

Bracey, D. R. 1968. Structural implications of magnetic anomalies
north of the Bahamas-Antilles Islands. Geophy. 33: 950-961.

Bryan, G. M. and R. G. Markl. 1966. Microtopography of the Blake-
Bahama region. Lamont Geol. Obs. Tech. Rept. TR-8. 59 pp.

Chamberlain, Robert S. 1948. Discovery of the Bahama Channel.
~ Tequesta 8: 109-116.

Conrod, A. C., Anne Boersma, and M. G. Kelley. 1968. Investigation of
visible region instrumentation for oceanographic satellites. M.I.T.
Exp. Astron. Lab. Rept. RE-31: Vol. II. 54 pp.

e--------- » M. G. Kelley, and Anne Boersma. 1968. Aerial photography
for shallow water studies on the western edge of the Bahama Banks.
M.I.T. Exp. Astron. Lab. Rept. RE-42.

Coonley, Davenport, and Mancel. 1902. 119 photographs of the Bahamas
presented to the Society's library. Geogr. J. 19: 539.

“Darlington, Philip Jackson, Jr. 1938. The origin of the fauna of the
Greater Antilles, with discussion of dispersal of animals over
water and through the air. Quart. Rev. Biol. 13: 274-300.

Dierickx, Charles Wallace. 1952. An historical geography of the Bahama
Islands. M.S. thesis. University of Florida. Gainesville, Florida.

---------- 1953. Some geographic problems of the Bahama Islands. Quart.
Js) PloridasAcad. "Sci." 163) 212-222.

Dudley, Ray L. 1947. Flying magnetometer completing 80,000 square mile
survey (of the Bahama Islands). World Oil 127: 247, 251.

74

Fisher, Allan C., Jr. 1959. Cape Canaveral's 6,000-mile shooting
gallery. Natl. Geogr. Mag. 116; 421-471.

King, Wayne. 1962. The occurrence of rafts for disperal of land ani-
mals into the West Indies. Quart. Ji. FloridajAcad., Sciae25ea45—52.

Long, E. John. 1964. Bahamas (revised edition). 1959-first edition.
(Around the World Program-Amer. Geogr. Soc.) Doubleday. Garden
CityNews. 2 O4 appr

Lucas, Charles Prestwood. 1890. Historical geography of the British
Colonies. Vol. 2. West Indies... Ch. 2... The Bahamas i(ppemooo) ce
Clarendon Press. Oxford.

Lynam, Edward. 1948. Early days in Bermuda and the Bahamas. Geogr.
Mage, (213) 20122075.

MacArthur, Robert H. and Edward O. Wilson. 1967. The theory of island
biogeography. (Monographs in Population Biology-1) Princeton Univ.
Press... .Princeton, Neo J:

MacKinnen, Daniel. 1804. A tour through the British West Indies in the
years 1802 and 1803, giving a particular account of the Bahama
Islands... “J <.Whate;. London3,272 pp.

Mitchell, Carleton. 1958. The Bahamas, isles of the blue-green sea.
Natl. Geogr. Mag. 113: 147-203.

---------- and James L. Stanfield. 1967.. The Bahamas; more of sea than
of land. (Nati: Geopr. Mage 152) 218=267,.

Norton, Graham. 1967. People of the Out Islands. Geogr. Mag. 40: 704-
128.

Olschki, Leonard. 1941. Ponce de Leon's fountain of youth: history of
a geographical myth. Hispanic Amer. Hist. Rev. 21: 361-385.

Psuty, Norbert P,..and James:S. Bailey... 1969. Lagoons; islands, off-
shore islands, shorelines and banks. in: Earth resource surveys
from spacecraft, v. 2 (Leetsma’,, R..A., principal investigators oR A.
White, ed.) NASA, Earth Resource Group. Houston, Texas. p. G-7-G-14%

Randall, John E. and Carleton Ray. 1958. Bahamian Land-and-Sea Park.
Sea Frontiers 4: 72-80.

Rowe, George. 1864. The colonial empire of Great Britain, considered
chiefly with reference to its physical geography and industrial
productions. The Atlantic group. Christian Knowledge Society.
London.

75

Schott, Charles A. 1882. An inquiry into the variation of the compass
off the Bahama Islands at the time of the landfall of Columbus in
1492-5 Ul io. Coast. Geodetic Surv: Rept. for 1880... Appendix No. :19.

Selby, M. J. 1970. Very high altitude photography in geography. 2:
Landforms. N.Z. J. Geogr. 48: 4 p. (facing pp. 16-17).

Shattuck, George B. 1903. The expedition to the Bahama Islands of the
Geographical Society of Baltimore. Science 18: 427-432.

---------- 1905. The Bahama Islands. Geographical Society of Baltimore.
Johns Hopkins Press. Macmillan Co. 630 pp.

Simpich, Frederick. 1936. Bahama holiday. Natl. Geogr. Mag. 69: 219-
245.

Smith, C. Lavett. 1965. Survey of the Bahamas. Nat. Hist. 74 (10):
62-65.

Talwani, Manik. 1960a. Gravity anomalies in the Bahamas and their
imterpretation. Ph.D. Diss. Columbia Univ. New York, N. Y. 89 pp.

---------- 1960b. Gravity anomalies in the Bahamas (abst.). Diss. Abst.
21: 852-853.

---------- Jeb. Worzel, and) M. Ewing. 1959. Gravity anomalies and
structure of the Bahamas. Lamont Geol. Obs. Columbia Univ. New
Mork. N.Y... 9 pp.

soo rco---- y -cccc-----, and ---------- 1960. Gravity anomalies and
structure of the Bahamas. Trans. 2nd. Carib. Geol. Conf. Mayaguez,
Puerto Rico, 4-9 January, 1959. pp. 156-161.

Thompson, T. A. 1944. A short geography of the Bahamas. Privately
published. Nassau. Revised ed. printed 1949 by Nassau Daily
Tribune. Nassau.

Townsend, P. S. 1826. Memoir on the topography, weather and diseases
of the Bahama Islands. J. Seymour Co. New York, N. Y. 80 pp.

Worzel, John Lamar, William Maurice Ewing, and Charles Lum Drake. 1953.
The Bahamas Islands regions. Part 1 of Gravity observations at sea
(abst.). Geol. Soc. Amer. Bull. 64: 1494-1495. Also Amer. Mineral.
592, 549-350... 1954.

L. MAMMALOGY (INCLUDING SEA MAMMALS)

Allen, Glover M. 1905. Notes on Bahama bats. Proc. Biol. Soc. Wash.
13s 65-71%

76

Allen, Glover M. 1911. Mammals of the West Indies. Bull. Mus. Comp.
Zool. 54: 175-263.

---------- 1937. Geocapromys remains from Exuma Island (Bahamas).
J. Mamm. 18: 369-370.

---------- and C. C. Sanborn. 1937. Notes on bats from the Bahamas.
J. Mamm. 18: 226-228.

Allen, J. A. 1891. Description of a new species of Capromys from the
Plana Cays, Bahamas. Amer. Mus. Nat. Hist. Bull. 3: 329-336.

Backus, Richard H. 1961. Stranded killer whale in the Bahamas. J.
Mamm. 42: 418-419.

Bangs, Outram. 1898. A new raccoon from Nassau Island, Bahamas. Proc.
Biol. Soc. Wash. 12: 91-92.

Caldwell, David K. and Melba C. Caldwell. 1971. Beaked whales, Ziphius
cavirostris, in the Bahamas. Quart. J. Florida Acad. Sci. 34: 157-
160.

Clough, Garrett C. 1969. The Bahaman Hutia: a rodent refound. Oryx
10: 106-108.

---------- 1972. Biology of the Bahaman Hutia, Geocapromys ingrahami.
J. Mamm. 53: 807-823.

---------- 1973. A most peaceable rodent. Nat. Hist. 82: 66-74.

---------- and George Fulk. 1971. The vertebrate fauna and the vegeta-
tion of East Plana Cay, Bahama Islands. Atoll Res. Bull. 138: 1-17.

Cummings, W. C. 1963. Recent studies at the acoustic-video system:
occurrence of animal sounds at Bimini (abst.). Assn. Is. Mar. Labs.
1963: 5S.

Goldman, E. A. 1950. Raccoons of North and Middle America. N. Amer.
Fauna 60: 1-153.

Harper, Francis. 1930. Notes on certain forms of the house mouse (Mus
musculus), particularly those of eastern North America. J. Mamm.
1149-52;

Hecht, M. K. 1955. The comparison of recent and fossil amphibian,
reptilian, and mammalian faunas in the Bahamas. Yearb. Amer. Phil.
Soc. for )1954>):), 133-135:

Howe, R. J. 1971. Social behavior of the Bahaman hutia (Geocapromys
ingrahami) with an investigation of marking behavior. M.S. Thesis.
Univ. Rhode Island. Kingston, R. I. 71 pp.

77

Howe, R. J. and G. Clough. 1971. The Bahaman hutia in captivity. Int.
Zool. Yearb. JT: 89-95:

Koopman, Karl F. 1951. Fossil bats from the Bahamas. J. Mamm. 32: 229.

---------- » Max K. Hecht, and Emanuel Ledecky-Janecek. 1957. Notes on
the mammals of the Bahamas with special reference to the bats. J.
Mamm. 38: 164-174.

Lawrence, Barbara. 1934. New Geocapromys from the Bahamas. Occ. Paper
Boston Soc. Nat. Hist. 8: 189-196.

McKinley, Daniel. 1959. Historical note on the Bahama raccoon. J. Mamm.
40: 248-249.

MacLeay, W. S. 1829. Notes on the genus Capromys of Demarest. Zool.
J. 4: 269-278.

Miller, G. H. 1931. The red bats of the Greater Antilles. J. Mamn.
12: 409-410.

Miller, Gerrit S. 1903. A new Nataline bat from the Bahamas. Proc.
Biol. Soc. Wash. 16: 119-120.

tae 1905. Mammals of the Bahama Islands. in: Shattuck, George
B., ed. The Bahama Islands. Geographical Society of Baltimore.
Johns Hopkins Press. Baltimore, Maryland. pp. 371-384.

---------- 1929. The characters of the genus Geocapromys Chapman.
Smithsonian Misc. Coll. 82 (3029): 1-3.

Moore, Joseph Curtis. 1958. A beaked whale from the Bahama Islands and

comments on the distribution of Mesoplodon densirostris. Amer. Mus.
Nov. 1897: 1-12.

Nesbitt, C. R. 1836. On the Bahama fisheries. J. Bahama Soc. Diff.
Knowl. No. 11: 126-136.

Rebach, J. A. 1971. Comparison of gas exchange and water balance of
the nutria, Myocastor coypus, and the hutia, Geocapromys ingrahami.
Ph.D. Diss. Univ. Rhode Island. Kingston, R. I.

SenWartez, Albert. 1955-9 The status of the species of the brasiliensis
group of the genus Tadarida. J. Mamm. 36: 106-107.

Shamel, H. H. 1931. Bats from the Bahamas. J. Wash. Acad. Sci. 21:
251-2535

Sherman, H. B. 1954. Raccoons of the Bahama Islands. J. Mamm. 35: 126.

78

Struhsaker, Paul. 1967. An occurrence of the Minke whale, Balaenoptera
acutorostrata, near the northern Bahama Islands. J. Mamm. 48: 483.

M. MEDICINE, HUMAN AND VETERINARY, AND PUBLIC HEALTH

Anonymous. 1835. On the medicinal and other uses of plants growing in
the Bahama Islands. J. Bahama Soc. Diff. Knowl. No. 3: 23-30;
No. 4: 31-37; No. 5: 44-46,

---------- 1836. (untitled-deals with medicinal properties of 12 plants,
extracted from Beach's "American practice of medicine."). J. Bahama
Soc. Diff. Knowl. No; 9: 95-100).

---------- 1971. The pattern cf use of marihuana. Tidsskr. Nor.
Laegeforening 91: 2425. (in Norwegian).

Beveridge, Sir Wilfred W. O. 1927. Report on the public health in
New Providence. Nassau. 67 pp.

Black, F. L. and W. J. Houghton. 1967. The significance of mumps
haemagglutinin inhibition titers in normal populations. Amer. J.
Epidem. 85: 101-107.

Clarkson, M. J. and L. N. Owen. 1959. The parasites of domestic animals
in the Bahama Islands. Ann. Trop. Med. Parasit. 53: 341-346.

Cory, Richard A. S. and Grahame M. Barry. 1962. A tuberculin and x-ray
survey of school children in Nassau. Tubercle 43: 145-150.

Cottman, Evans W. (with Wyatt Blassingame). 1963. Out-Island Doctor.
E. P. Dutton and Co., Inc. New York, N. Y. Also published by
Hodder and Stoughton. London. 248 pp.

Edge, P. Granville. 1944. Malaria and nephritis in the British West
Indies. Carib. Med. J. 6: ..32-43.

Feng, 'P.. C.. LJ. Haynes), K.°E.. Magnus:, and J. R. Plimmer. 1964.
Further pharmacological screening of some West Indian medicinal
plants. J. Pharmaceutical Pharm. 16: 115-117.

Florey, Charles duV., Michael M. Gerassimos, and Paul R. Cuadrado. 1966.
Report of a serological survey of the southern half of Eleuthera
Island, Bahamas. W. I. Med. J. 15: 71-82.

Hanek, G. 1972. Monogenetic trematodes from New Providence Island,
Bahamas. J. Parasit. 58: 1117-1118.

Higgs, Mrs. Leslie. 1969. Bush medicine in the Bahamas. Privately
published by the author with original paintings. 20 pp.

79

Hooper, P. A. and B. E. Leonard. 1965. Pharmacological properties of
some West Indian medicinal plants. J. Pharmaceutical Pharm. 17:
98-107.

Hornburg, C. D. and A. W. Saariner. 1971. Bahamas on a salt free diet.
Water and Wastes Engin. 8: 32-34.

Humphrses, S: V..1957.. A study of hypertension in the Bahamas. S. Afr.
Med. J. 31: 694-699.

Jackson, J. R. 1876. Princewood bark, a febrifuge from the Bahamas.
Pham. J Trans. (london) TIL. 6: 681.

Jackson, W. P. U. 1946. Plant dermatitis in the Bahamas. Brit. Med.
e946 (2): 298.

Johnson, Benjamin C. 1960. A sampling survey study of arterial blood
pressure levels in Nassau, New Providence, Bahamas, 1958 for descrip-
tion of levels of blood pressure in a population in relationship to
age, sex, race, and other factors. Dissert. Abst. 20: 3698-3699.

---------- and Richard D. Remington. 1960. A sampling study of blood
pressure levels in White and Negro residents of Nassau, Bahamas
€abst.) + Circulation 222 768.

---------- and ---------- 1961. A sampling study of blood pressure
levels in White and Negro residents of Nassau, Bahamas. J. Chron.
Dis Si:).59-51.

Leam, G. and I. E. Walker. 1963. The occurrence of Platynosomum fastosum
in domestic cats in the Bahamas. Vet. Rec. 72: 46-47.

Moser, Marvin, Richard Morgan, Malcolm Hale, Sibley W. Hoobler, Richard
Remington, H. J. Dodge, Alice I. Macaulay. 1959. Epidemiology of
hypertension with particular reference to the Bahamas. I. Prelimi-
nary report of blood pressure and review of possible etiologic
factors. Amer. J. Cardiology 4: 727-734.

. Penrose, Clement A. 1905. Sanitary conditions of the Bahama Islands.
in: Shattuck, George B., ed. The Bahama Islands. Geographical
Society of Baltimore. Johns Hopkins Press. Baltimore, Maryland.
pp. 387-416.

Rosen, S. and H. V. Rosen. 1971. High frequency studies in school
children in nine countries. Laryngoscope 81: 1007-1013.

Sawyer, William H. 1955. Medicinal uses of plants by native Inaguans.
Sci. Monthly 80: 371-376.

Schoepf, J. D. 1787. Materia medica americana potissimum regni vegeta-
bilis. Erlangen. xviii +170 pp. Reprinted as Bull. Lloyd Li-
braxy, Nowe (Reprod. Ser. 95). 1905.

80

Smith, Robert R., et al. 1971-72. A study of the near shore and ter-
restrial flora of San Salvador Island. 3 vol. College Center of
the Finger Lakes. Corning, N. Y. 1 chapter on Bush medicine of
San Salvador Island.

Spencer, D. J... 1972. “Suicide in’ the Bahamas. , Int. J4 Soc.) Psychiatry,
LSE OS.

Stafford, J. L., R. K. Hill, and E. L. Demontaigne. 1955. Micro-
filariasis in the Turks’ Islands. W. 1. Med. J.'4: 185-187.

Townsend, P. S. 1826. Memoir on the topography, weather and diseases
of the Bahama Islands. J. Seymour Co. New York, N. Y. 80 pp.

N. GENERAL NATURAL HISTORY INCLUDING
REPORTS AND SEMI-POPULAR ACCOUNTS

Anonymous. 1903. Expedition to the Bahama Islands. Bull. Geogr. Soc.
55) 956059 6

---------- (undated, but probably 1960). The Lerner Marine Laboratory
at Bimini, Bahamas. Amer. Mus. Nat: Hist. (New: York, N. Yoeeo2 spp.

---------- 1971. Commonwealth of the Bahama Islands. Background Notes.
Dept: of StatevRubl. 78329 1-4:

Agassiz, Alexander. 1893. Observations in the West Indies. Amer. J.
Sci. 25ex,. / i345 2 LS5 8-562.

---------- 1894. Reconnaissance of the Bahamas. Bull. Mus. Comp. Zool.
26 = 205).

Allen, Elsa G. 1937. New light on Mark Catesby. Auk 54: 349-363.

Allen, Glover M. and Thomas Barbour. 1904. Narrative of a trip to the
Bahamas. Privately printed. Cambridge, Massachusetts. 10 pp.,
Saples 517 WaMapy wl iets

Andrews,, E. A, R. PB. Bigelow, and T. H. Morgan. 1945. Threevat
Bimini. Sci. Monthly 61: 333-344.

Ballou, H. A. 1910. Legislation in the West Indies for the control of
pests and diseases in imported plants. W. I. Bull. 10: 197-234.
(Bahamas, p. 233).

Batchelder, C. F. 1951. A bibliography of the published writings of
Charles Johnson Maynard (1845-1929). J. Soc. Bibl. Nat. Hist. 2
Gis 227-260"

81

Benjamin, George J. 1970. Diving into the blue holes of the Bahamas.
Natl. Geogr. Mag. 138: 347-363.

Boersma, Anne. 1968. Bibliography on the Bahama Islands. M.1.T.
Exp. Astron. Lab. Rept. No. RN-37. 60 pp.

Bounds, John Howard. 1966. Land use in the Bahamas. Ph.D. Diss.
University of Tennessee. Knoxville, Tennessee. 419 pp.

---------- 1969. The Bahamas. Focus 19 (9): 3-7.

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sono sn---- 1905. Explorations in the Bahamas. J.N. Y. Bot. Gard. 6:

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83

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---------- 1963b. An aerial reconnaissance of the Bahama Islands.
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---------- 1930. Above and below Bahaman waters. Nat. Hist. 30: 593-
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84

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---------- and Alexander Sprunt IV. 1971. Parks and conservation in
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---------- 1905b. Some general considerations relating to the Bahama
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---------- and Benjamin LeRoy Miller. 1905. Physiography and geology
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---------- 1962b. Gem-like Bahamas created from sea's violent ups and
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85

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=<-------- and ---------- 1963. Observations of present-day carbonate
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86

Allen, J. A. 1900. New birds from the Bahamas. Auk. 17: 187.

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---------- 1916. The smaller mockingbird of the northern Bahamas .
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Bonhote, J. Lewis. 1899. A list of birds collected on the Island of
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---------- 1902. Field notes on some Bahama birds. Aviculture Mag.
ser., I, Vol. 8: '278-285.;

---=------- 1903a. On a collection of birds from the northern Islands
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---------- 1903c. Bird migration at some of the Bahama lighthouses.
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Boynton, Alice M. 1908. Bahama news. Bird-lore 10: 192.

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87

Bryant, H. 1863. Description of two birds from the Bahama Islands,
hitherto undescribed. Proc. Boston Nat. Hist. Soc. 9: 279-280.

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Buden, Donald W. and Albert Schwartz. 1968. Reptiles and birds of Cay
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---------- 1908. Camps and cruises of an ornithologist. Hodder and
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a--------- Le9ibes hvst-of the’ birds collected by C. “L.~ Winch in the
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~--t------ 1894. Defense glands in a Bahama species of phasa. Contr.
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Riley, J. H. 1903a. A new subspecies of highthawk from the Bahama
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===------- 1903b. The snowy plover in the Bahamas. Auk 20: 433.

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Vaurie, Charles. 1953. Observations and new records of birds from the
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Wetmore, Alexander. 1937. Bird remains from cave deposits on Great
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Zahl, Paul A. 1947. The flamingos of Andros. Sci. Monthly 64: 277-288.

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Ewing, Maurice at al. 1968a. Leg I of the cruises of the drilling
vessel "Glomar Challenger," Orange, Texas to Hoboken, N. J.,
August to September 1968. Site 4 - Shipboard Site reports.
Univ. California Scripps Inst. Oceanog. pp. 179-213. U.S.G.P.0O.
Washington, D. C.

oocrno---- 1968b. Leg I of the cruises of the drilling vessel ''Glomar
Challenger,"' Orange, Texas to Hoboken, N. J., August to September
1968. Site 5. Shipboard Site reports. Univ. California Scripps
Inst. Oceanog. pp. 214-242. U.S.G.P.0. Washington, D. C.

eoocc----- 1969. Regional aspects of deep-water drilling in the Gulf
of Mexico, east of the Bahamas Platform, and on the Bermuda Rise.
Initial reports. Deep-Sea Drilling Project, Glomar Callenger 1:
624-640. U.S.G.P.0. Washington, D. C.

Friedman, Daniel. 1964. Periodic variations of the temperature and
water currents in the Tongue of the Ocean, Bahamas. U. S. Naval
Res. Lab, Interim Rept. 12 pp.

acer cc--H- 1965. Microtopography of sea floor east of Goulding Cay,
Bahamas. Mem. Naval Res. Lab. No. 1612. Washington, D. C.

Fuglister, F. C. 1947. Average monthly sea surface temperatures of
the western North Atlantic Ocean. Papers Physical Oceanog.
Meteorol. M.I.T. Woods Hole Oceanog. Inst. 10: 1-25.

Galtstof££, P: S. 1943. Copper content of sea water. Ecology 24: 263-
265.

94

Gapurro, L. R. A. 1967. Oceanography from space. Boll. Geof. Teorica
Appl; 9siS=16.

Goedicke, T. R. 1960. Some geological results of underwater sound
measurements in the Bahamas. Tech. Rept. Marine Lab, Univ. Miami.
No. 60.

Gossnett, F. R. 1948. Flare triangulation between Florida and the
Bahamas. J. Coast Geodetic Surv. 1: 39-44.

Guilcher, André. 1969. Pleistocene and Holocene sea level changes.
Earth-Sci. Rev. 5: 68-97.

Guppy, H.-Bi -19175. Plants), seeds; and currents in? the Westeindres vane
Azores. Williams and Norgate. London. 531 pp.

Hannau, Hans W. and Bernd H. Mock. (undated, but appeared in 1973).
Beneath the seas of the West Indies. Caribbean, Bahamas, Florida,
Bermuda. Hastings House. New York, N. Y. 104 pp.

Harrison, W. 1964. Sea level oscillations recorded in subsurface at
Freeport, B.W.al. (abst:). Geol. ‘Soc. Amer: Spec. Paper qo-m7s no.

---------- 1971. Atlantis undiscovered; Bimini, Bahamas. Nature 230
(5292): 287-289.

Hay, W. W. 1967. Bimini Lagoon: model carbonate epeiric sea (abst.).
Amer. Assn. Petrol. Geol. Bull. 51: 468-469.

---------- » Felix Wiedenmayer, and Donald S. Marszalek. 1971. Modern
organism communities of Bimini Lagoon and their relation to the
sediments. in: H. Gray Multer, ed. Field Guide to some carbonate
rock environments: Florida Keys and western Bahamas. pp. 66B-66T.

Hays, Earl. 1960. Yamacraw II, Narrative. Woods Hole Oceanog. Inst.
Ref. No. 60-22, Documentation Library.

Huddell, Howard D. 1967. AUTEC sediment deposition/erosion study,
interim report. U. S. Naval Oceanog. Off. Rept. NOO-IR-67-100.

47 pp.

Hurley, Robert J. 1964. Bathymetry of the Straits of Florida and the
Bahama Islands. Part III. Southern Straits of Florida. Bull. Mar.
Sci. “Gul Carib ysl4 34375-3580.

---------- , V. B. Siegler, and L. K. Fink, Jr.. 1962. Bathymetry of the
Straits of Florida and the Bahama Islands. I. Northern Straits of
Florida. -2BullisMareSci«, Guilt Caribe) 22 2515-321.

Jackson, R. A. 1965. Stability of rubble-mound breakwaters, Nassau
Harbor, Nassau, New Providence, Bahamas. U. S. Army Engin. Water
Exp..Station Tech. Rept. No. 2-097;

95

Jelgersma, S. 1966. Sea level changes in the last 10,000 years. Proc.
Royal Meterol. Soc. Int. Symposium-World Climate, 8,000 to O B.C.,
Imperial College of London. pp. 54-71.

Jensen, Homer. 1948. Some technical aspects of Bahamas airborne-
magnetometer survey (abst.). Oil and Gas J. 46: 114; Geophysics
is: 495;.

Katz, B. 1965. Circulation near the southern Berry Islands, Bahamas.
Tech. Rept. U. S. Atomic Energy Comm. CU-23-65.

Kier, Jerry S. and Orrin H. Pilkey. 1971. The influence of sea level
changes on sediment carbonate mineralogy, Tongue of the Ocean,
Bahamas. Mar. Geol. 11: 189-200.

Kornicker, Louis. S. 1962. Hydrography and sedimentology in the vicinity
of Bimini, Great Bahama Bank (abst.). Texas J. Sci. 14: 415-416.

---------- and E. G. Purdy. 1957. A Bahamian faecal-pellet sediment.
jesoed. Petrol. 272 126-128.

Kumpf, Herman E. 1963. Acoustic-video system for biological research.
IV. Use of underwater television in bio-acoustic research. Proc.
Symposium Marine Bio-acoustics. Bimini, Bahamas. pp. 45-57.

Kupferman, Stuart L. 1971. Fallout as an indicator of sedimentary
processes in a shallow water environment (abst.). Eos (Amer.
Geophy. Union Trans.) 52: 258.

Lucas, Gabriel. 1970. L'effet des mouvements de l'eau et de la
morphologie sous-marine sur la répartition des dépdts. Soc. Géol.
Pea GR. Ser. D 270: 1417-1420.

McCallum, M. F. and K. Guhathakurta. 1970. The precipitation of calcium
carbonate from seawater by bacteria. J. Appl. Bacteriology 33:
649-655.

Magnitzky, A. W. and H. V. French. 1960. Tongue of the Ocean research
experiment. U.S. Naval Oceanog. Off. Rept. TR-94. 132 pp.

Mathewson, R. F. 1963. An aerial reconnaissance of the Bahama Islands.
Assi.) Us. Mar. Labs. Nov. 1963: 25.

Miller, Milton A. 1968. Isopoda and Tanaidacea from buoys in coastal
waters of continental United States, Hawaii, and Bahamas (Crustacea).
ess. Wael. Mus’ Proc. 125. (3652): 53: pp’.

Miner, R. W. 1930. Above and below Bahaman waters. Nat. Hist. 30:
593-605.

96

Nesteroff, W. D. and Gene A. Rusnak. 1962. Sedimentary characteristics
of modern turbidites in the Tongue of the Ocean (Bahamas) compared
to abyssal plain turbidites. in: National Coastal and Shallow
water Research Conf. Proc. 1961. pp. 393.

Neumann, A. Conrad. 1973. Quaternary sea level history of Bermuda and
the Bahamas. AMQUA meeting prospectus 4-5 December 1973. Miami,
Florida.

---------- » Conrad D. Gebelein, and Terence P. Scoffin. 1970. The
composition, structure, and erodability of subtidal mats, Abaco,
Bahamas. J. Sed. Petrol. 40: 274-297.

Newell, Norman D. 1951. Organic reefs and submarine dunes of oolite
sand around Tongue of the Ocean, Bahamas. (abst.). Geol. Soc.
Amer. Bull. 62: 1466.

---------- 1953. Under tropic seas (Bahamas) (abst.). Amer. Assn.
Petrol, “Geol 2sBuli 737 2 iZ5

---------- » John Imbrie, Louis Kornicker, and Edward Purdy. 1956.
Bahamian limestone seas (abst.). Geol. Soc. Amer. Bull. 67: 1820.

Oppenheimer, C. H. 1961. Note on the formation of spherical aragonite
bodies in the presence of bacteria from the Bahamas Bank. Geochim.
Cosmochim. Acta 23: 295-296.

Pilkey, 0. H. 1966. Mineralogy of Tongue of the Ocean sediments.
J. Mar. Res. (Sears Found. Mar. Res.). 24: 276-285.

---------- » James B. Rucker, and Normitsu Watabe. 1968. Possible sea-
level-carbonate mineralogy relationship in Tongue of the Ocean
sediments (abst.).°-Geol.' Soc. Amer. Spec...Paper 10127 571

Randall, John E. and Carleton Ray. 1958. Bahamian Land-and-Sea Park.
Sea Frontiers .4: 72-80.

Rausch, James P. 1972. Survey of marine life. College Center of the
Finger Lakes. Corning, New York. 53 pp.

Rietzel, John. 1959. An echo-sounding survey near Green Cay, Bahamas.
Woods Hole Oceanog. Ref. C-1396. 4 pp.

Richards, Horace G. 1971. Sea level during the past 11,000 years as
indicated by data from North & South America. Quaternaria 14: 7-15.

-~--------- 1972. Some aspects of the marine Quaternary of the Caribbean
area. VI Conf. Geol. Caribe Margarita, Venezuela Memorias.
pp. 426-429.

Richardson, W. S. and J. R. Finlen.. 1967. The transport of the North-
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97

Ross, D. S. 1969. Color enhancement for ocean cartography. in: Oceans
from space. P.-C. Badgley, L. Milroy, and L. Childs, eds. Proc.
Gulf Publ. Co. Houston, Texas. pp. 50-63.

Rucker, James B., Newell T. Stiles, and Roswell F. Busby. 1967. Sea-
floor strength observations from the DRV ALVIN in the Tongue of
the Ocean, Bahamas. SE Geol. 8: 1-8.

Ruggles, G. S. 1963. Temporal and spatial variations of the tempera-
ture and sound speed in the Tongue of the Ocean, Bahamas. Manual
U. S. Naval Oceanog. Off. No. 0-33-63.

aaoo------ 1966. Sound velocimeter profiles from the Tongue of the
Ocean and Exuma Sound, Bahamas, February and March 1962. U. S.
Naval Oceanog. Off. Rept. NOO-IR-O-8-66. 38 pp.

wonc------ and L. Cisney. 1966. Determination of sound speed from
temperature measurements in the Tongue of the Ocean, Bahamas.
U. S. Naval Oceanog. Off. Rept. NOO-IR-O-7-66. 16 pp.

Schram, Thomas A. 1970. Marine biological investigations in the Bahamas.
14. Crypris y, a later developmental stage of Nauplius y Hansen.
Sarsia 44: 9-24.

Scoffin, T. P. 1970. The trapping and binding of subtidal carbonate
sediments by marine vegetation in Bimini Lagoon, Bahamas. J. Sed.
Petrol. 40: 249-273.

Shepard, F. P. and H. E. Suess. 1956. Rate of postglacial rise of sea
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Shidy, L. P. 1905. Tides and bench marks at Nassau, New Providence.
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Society of Baltimore. Johns Hopkins Press. Macmillan Co. Balti-
more, Maryland. pp. 51-96.

Shonting, D. H. 1963. Review of the oceanographic environment of the
Tongue of the Ocean, Bahamas. Part 1. Preliminary discussions.
Tech. Mem. Naval Underw. Ord. Station No. 289.

---------- 1970. On the distribution of temperature, salinity, and
oxygen in the Tongue of the Ocean, Bahamas. Bull. Mar. Sci. 20:
35-56.

Siegler, Violet B. 1961. Bathymetric reconnaissance of Exuma Sound.
Univ. Miami Inst. Mar. Sci. Rept. 61-4: 1-9.

Sisler, Frederick, D. 1962. Microbiology and biochemistry of the sedi-
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98

Smith, C. L. 1940. The Great Bahama Bank. 1. General hydrographic and
chemical factors. 2. Calcium carbonate precipitation. J. Mar. Res.
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----r----- 1966... Descending the Andros Reef. Nat. Hist. -75: 38-43.

Smith, N. R. 1926. Report on a bacteriological examination of "chalky
mud'' and sea-water from the Bahama Banks. Carnegie Inst. Wash.
Publis 3440 7-72%

Spencer, Maria. 1967. Bahama deep’ test. Amer. Assn. Petrol. Geol.
Bull. 512) 265-268.

Steinberg; J. C., WG: Cummings) B. Do Brahy, and’ J. Y. MacBain
(Spires). 1965. Further bio-acoustic studies off the west coast
of North Bimini, Bahamas. Bull. Mar. Sci... 15: 942-963.

stommel:, Henry: L958. The Gulf Stream. Univ. Calatornia Press ag
Berkeley, California and Cambridge Univ. Press. Cambridge. 202 pp.

Storr, J. F. 1964. Ecology and oceanography of the coral-reef tract,
Abaco Island, Bahamas. Geol. Soc. Amer. Spec. Paper 79: 1-98.

Szabo, Barney J. 1967. Radium content in plankton and sea water in
the Bahamas. Geochim. Cosmochim. Acta 31: 1321-1331.

---------- 1968. Trace element content of plankton population from the
Bahamas:.” Carib.) Ja.Sc1. 8: 185-186.

---------- 1971. Concentration of radium-226 in Northeast Providence
Channel and, the Tongue of the Ocean, Bahamas. (Bull. Mar. Seu. e21:
748-753.

Till, Roger. 1970.- The’ relationship ‘between: environment: and sediment
composition (geochemistry and petrology) in the Bimini Lagoon,
Bahamas.’ J. Sed..-Petrol . 40: 367-385.

Tillinghast, William H.’ 1881. Notes on the historical hydrography Jot
the Handkerchief Shoal in the Bahamas. Harvard Univ. Bibliog.
Contre 14.6 pp.

Traganza, Eugene Dewees. 1966. Dynamics of the carbon dioxide system
on the Great Bahama Bank (abst.:). Diss. Abst., Sci. Engin. 27:
61B-62B.

-~--------- and B. J. Szabo. 1967. Calculation of calcium anomalies on |
the Great Bahama Bank from alkalinity and chlorinity data. Limnol.
Oceanog. 12: 281-286.

Traverse, Alfred and Robert N. Ginsburg. 1966. Palynology of the sur-
face sediments of the Great Bahama Bank, as related to water move-
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99

Turekin, K. K. 1957. Salinity variations in sea water in the vicinity
of Bimini, Bahamas, British West Indies. Amer. Mus. Nov. 1822:
Po12.

Uchupi, Elazar. 1966. Shallow structure of the Straits of Florida.
Science 153: 529-531.

Vary, Willard E. 1969. Remote sensing by aerial color photography for
water depth penetration and ocean bottom detail. Proc. 6th Int.
Symposium on Remote Sensing of the Environment 2: 1045-1059.

Wanless, Harold R. 1969. Sedimentary structure zonation on tidal
levees, Andros Island, Bahamas (abst.). Amer. Assn. Petrol. Geol.
Bulle 55: 748.

Wennekens, M. Pat. 1961. Oceanography of the Tongue of the Ocean,
Bahama Islands (abst.). J. Geophy. Res. 66: 1560.

Wiedemann, Hartmut U. 1969. Solutions in intertidal and supratidal
environments of modern carbonate sedimentation; their implication
on diagenesis. Chem. Geol. 4: 393-409.

Welkemson, J. R., R. Peloguin, and I. Pelroth. 19635. Airborne radia-
tion thermometer survey, Tongue of the Ocean, 5 through 9 February
1963. U.S. Naval Oceanog. Off. Tech. Rept. IMR-0-20-63.

Wood, J. Ferguson. 1968. Studies of phytoplankton ecology in tropical
and subtropical environments of the Atlantic Ocean. Part 3. Phyto-
plankton communities in the Providence Channels and the Tongue of
emer Ocean., Bulli: Mar. Sci. V8: 481-543.

Woodring, W. P. 1954. Caribbean land and sea through the ages. Geol.
Soc.wamer. Bull. 65: 719-782.
Q. TOURIST ITEMS RELATED TO NATURAL HISTORY

Anonymous. 1963. Bahamas (published for British Overseas Air Corp.
and Bahamas Airways by Inter-Continental Air Guides) 75 pp.

---------- 1970. Commonwealth of the Bahamas Statistical Abstract.
NEpt Ototatistucs.,, Gabinet Office. Nassau. 21) pp.

=a 1970. Man-made island promises double benefits to Bahamas.
Oceang iid ou o0l

---------- 1973. The West Indies and Caribbean Yearbook. Thomas Skinner
Directories. Croyden. London. 985 pp. Bahamas pp. 33-81. Turks
and Caicos Islands. pp. 191-196.

BarnacteaPod- id 1975..— Grand sBahama., otackpole Press. ~ Harrisburg,
Pennsylvania. 206 pp.

100

Butler, G. Paul and Erica Butler. 1960. The Caribbean, Central
America, the Bahamas, Bermuda. Van Nostrand Co. Princeton,
New Jersey. 438 pp.

Carrington, Vivien. 1972. Civics for the Bahamas. Longman Caribbean.
Longman Group Ltd. Harlow, Essex.

Collins, Doreen. 1961. The Turks and Caicos Islands; some impressions
of an English visitor. Carib. Quart. 7: 163-167.

Colonial Office. 1953. An economic survey of the colonial territories,
1951. Vol. 4. The West Indian and American territories. Colonial
No. 281-4. 274 pp. London.

Craton, Michael. 1964. Sun 'n sixpence: a guide to Nassau and the
Bahama Out Islands. S. P. Dupuch, ed. Nassau.

Dupuch, Etienne, Jr., ed. 1970. Bahamas Handbook and Businessman's
Annual, 1970-71, ed. 9. Etienne Dupuch, Jr. Publications. Nassau.
Published each year.

Edson, Wesley. 1964. Retiring to the Caribbean. Doubleday. Garden
City, New York.

Fodor, Eugene, ed. 1969. Fodor's guide to the Caribbean, Bahamas and
Bermuda, 1969. David McKay Co. New York, N. Y. 607 pp.

Gelhorn, Eleanor Cowles. 1955. McKay's guide to Bermuda, the Bahamas,
and the Caribbean. David McKay Co. New York, N. Y. 404 pp.

Gosset, Renée at al., 1968. Die Inseln der unbegreuzten Moglichkeiten
Bahamas. Weltwoche (Zurich) Jan.: 36.

Grafton, P. 1969. Planning an island in the sun. Chartered Surveyor
102: 157-158.

Grandfield, Alison. (no date, but appeared in 1973). The Bahamas -
island by island. Bahamas Ministry of Tourism. Nassau.

Greenway, J. C. 1938. Flight over the Bahamas. Harvard Alumni Bull.
40: 483-488.

Hannau, Hans W. 1970 (undated). Islands of the Bahamas (in full color).
Hastings House. New York, N. Y. 163 pp. including 48 in color.

---------- (undated, but before 1971). Nassau in the Bahamas. Wilhelm
Andermann Veriag. Munich. 61 pp. + 30 color pl.

---------- (undated but appeared in 1971). The Bahama Islands in full
color. Doubleday and Co., Inc. Garden City, New York. 124 pp.

101

Haweis, Stephen. 1924. The happy side (New Providence). Country Life
45% 35-38 + 2epil.

Hepburn, Andrew H. 1958. Complete guide to the Caribbean and the
Bahamas. (American Travel series No. 7). Houghton Mifflin Co.
Boston. 164 pp.

Jefferson, Peter. 1966. Maison de vacances dans les Iles Bahamas.
Architecture frangaise 27: 90-91.

Kline, Harry. 1973. Bahama Islands-boatman's guide to the land and
the water. (revision of J. Linton Riggs's book "Bahama Islands''-
“th ved.)| Charles Scribners Sons. New York, N..Y. 294 pp.

Lindley, John R. and John B. Davies. 1971. Sandflies and tourism in
Florida and the Bahamas and Caribbean area. J. Econ. Ent. 64:
264-278.

Long, E. John. 1964. Bahamas (revised ed.) (Around the World Program-
American Geographical Society). Doubleday Co. Garden City, New
York. 64 pp.

Lyons, Anne Nason. 1969. Tourism (in the Bahamas). Focus 19 (9): 8-11.

Maemubiany Allister, ed. 1938. The West Indies, past and present, with
British Guiana and Bermuda. W. H. and L. Collingridge. London.
440 pp.

Mogridge, Ian. 1969. Property investment in the Bahamas: a guide to
safe investment and maximum profit. Garnston Press. London.

Morgan, D. J. 1967. Prosperous Bahamas. Geogr. Mag. 40: 715-728.

Moseley, Mary. 1926. Bahamas Handbook. Nassau Guardian. Nassau,
Bahamas.

Moss, Frank T. 1973. Let's fish the Bahamas, Pastimes (Eastern Air-
lines Passenger Mag.) 1 (12): 18-23.

Novotny, J. 1966. Bahamy zyi turistiky (Die Bahama Inseln u. der
Fremdenverkehr). Lidé Ieme (Praha) 15: 185-186.

O'Shanshun, Frank. 1969. Bahama Islands, Cays of the Kingdom. in:
Fodor, Eugene, ed. Fodor's guide to the Caribbean Bahamas and
Bermuda. David McKay Co. New York, N. Y.

Savage, Ernest A. 1934. The libraries of Bermuda, the Bahamas, the
British West Indies, British Guiana, British Honduras, Puerto Rico
and the American Virgin Islands: a report to the Carnegie Corpora-
tion of New York. The Library Association. London. 102 pp.

102

Sever, Bruce B. 1964. Basic data - on the economy of the Bahama
Islands. U. S. Bureau of Int. Commerce. Overseas Business Rept.
OBR 64-38. 9 pp.

Stark, Jc Hen giS9l. Stark's history and guide to the Bahama Islands....
including their history, inhabitants, climate, agriculture, geology,
government. Boston. J. H. Stark Co.

---------- 1943. The Bahamas look to the future. Foreign Commerce
Weekly. 11: 8-10. (Reprinted: U. S. Bureau of Foreign and
Domestic Commerce. Inquiry reference service. War developments
in the West Indies. Washington, D. C. 39 pp.).

Thielen, Benedict. 1964. The Bahamas - golden archipelago. Holiday
36: 60-72 +.

Wilson, Frank I. 1864. Sketches of Nassau. Standard Press. Raleigh,
North Carolina.

Wilson, Henry A. 1891. A winter visit to the Bahamas Islands. Mass.
Hort. Soc. Trans. 1891: 210-229.

R. PALEONTOLOGY

Brodkorb, Pierce. 1959. Pleistocene birds from New Providence Island,
Bahamas. Bull. Florida State Mus. 4: 349-371.

Cloud, Preston E., Jr. and Virgil E. Barnes. 1948. Paldeoecology of
the early Ordovician sea in central Texas. Natl. Res. Council,
Rept. Committee on Treatise Mar. Ecol. and Paleoecology 1947-48.
pp. 29-83.

Corgan, James X. 1969. Pleistocene marine molluscan faunules from the
Little Bahama Bank (abst.). Tennessee Acad. Sci. J. 44: 49.

Craig, G. Y. 1967. Size-frequency distributions of living and dead
populations of pelecypods from Bimini, Bahamas. J. Geol. 75: 34-45.

Dall, William H. 1905a. Notes on the fossils of the Bahamas (abst.).
Science 21: 390-391.

---------- 1905b. Fossils of the Bahama Islands, with a list of the
non-marine mollusks. in: Shattuck, George Burbank, ed. The
Bahama Islands. Geographical Society of Baltimore. Johns Hopkins
Press. Baltimore, Maryland. pp. 23-47.

Etheridge, Richard. 1965. Pleistocene lizards from New Providence.
Quart. J. Florida Acad. Sci. 28: 349-358.

103

Fabricius, Frank. 1964. Aktive Lage- und Ortsveranderung bei de
Koloniekoralle Manicena areolata und ihre palaodkologische
Bedeutung (with English summary). Senckenbergiana Lethaea 34:
299-317.

Gibson, T. G. and J. Schlee. 1967. Sediments and fossiliferous rocks
from the eastern side of the Tongue of the Ocean, Bahamas. Deep-
Sea Res. 14: 691-702.

Hebard, Edgar B. 1967. Pleistocene mollusks from New Providence
Island, Bahamas. Nautilus 81: 41-44.

Hecht, M. K. 1955. The comparison of recent and fossil amphibian,
reptilian, and mammalian faunas in the Bahamas. Yearb. Amer.
Bonk Soc. for) 1954* o133-135..

Jones, J. I. and W. D. Bock. 1963. Trace element distribution in some
living and fossil Foraminifera from south Florida, Bahamian, and
Caribbean waters. Geol. Soc. Amer. Ann. Meeting, 1963, Program 88A.-

Koopman, Karl F. 1951. Fossil bats from the Bahamas. J. Mamm. 32: 229.

Kornicker, Louis S. 1963a. The Bahama Banks: a "living" fossil
environment. .J....Geol. Ed.sll: 17-25,

ror------- 1963b. Ecology and classification of Bahamian Cytherellidae
(Ostracoda). Micropaleontology 9: 61-71.

Lynts, George W. 1971. Analysis of the planktonic Foraminiferal fauna
of core 6275, Tongue of the Ocean, Bahamas. Micropaleontology 17:
152-166.

Pessagno, E. A., Jr. 1969 (?). Mesozoic planktonic Foraminifera and
Radiolaria. in: Initial reports of the Deep Sea Drilling Project.
I. 1, Leg 1 of cruises of the Glomar Challenger, Orange, Texas to
Hoboken, N. J. August to September 1968. U.S.G.P.0: Washington,
Dee. pp. 007-621).

---------- 1971. Jurassic and Cretaceous Hagiastridae from the Blake-
Bahama Basin (Site 5A, JOIDES Leg 1) and the Great Valley sequence,
California Coast Ranges. Bull. Amer. Paleont. 60: 1-83.

= — 1972. Cretaceous Radiolaria: II. Pseudoaulophacidae Riedel
from the Cretaceous of California and the Blake-Bahama Basin
(JOIDES leg I). Bull. Amer. Paleont. 61: 283-325.

Richards, Horace Gardiner. 1954. Pleistocene mollusks from Andros
Island, Bahamas. Nautilus 67: 120-121.

---------- 1971. Sea level during the past 11,000 years as indicated by
data from North and South America. Quaternaria 14: 7-15.

104

Vaughan, Thomas W., M. A. Howe, et al. 1919. Contributions to the
geology and paleontology of the West Indies. Carnegie Inst.
Wash... Publ 2291 184" pp

Wetmore, Alexander. 1937. Bird remains from cave deposits on Great
Exuma Island in the Bahamas. Bull. Mus. Comp. Zool. 80: 427-441.

---------- 1938. Bird remains from the West Indies. I. Records from
cave deposits on Crooked Island, Bahamas. Auk 55: 51-55.

ooo ------ 1955. JA check-list of the fossil and prehistoric: birdsiois
North America and the West Indies. Smithsonian Misc. Coll. 131
(S)ise1- 10S;

Wilson, R:o-L.; R:--E. Bergenback, and?C. Ps Finlayson. ~ 196237) Fossael
coral reefs, Fresh Creek, Andros Island, Bahamas (abst.). Geol.
SOc... Amer. sopec.. Paper 683 182.

Wing, Elizabeth, S. 1969. Vertebrate remains excavated from San
Salvador Island, Bahamas. Carib. J. Sci. 9: 25-29.

S. SOIL SCIENCE

Anonymous. 1928. The Bahama Islands. Information as to trade, soil,
climate, etc. for intending settlers, tourists and business men.
Development Board. Nassau. 24 pp. + 1 map.

Ahmad, N. and Robert L. Jones. 1969. Occurrence of aluminous lateritic
soils (bauxites) in the Bahamas and Cayman Islands. Econ. Geol.
64: 804-808.

Illing, L. V. 1954. Bahaman calcareous sands. Amer. Assn. Petrol.
Geol 7 Bull .-35:— 1-95:

Millard, R. S. 1959. Road problems in the Bahamas. Dept. Sci .=Indust-
Res. Road Res. Lab. Note No. RN/3458/RSI. Nassau. 6 pp.

Mooney, Charles N. 1905. Soils of the Bahama Islands. in: Shattuck,
George Burbank, ed. The Bahama Islands. Geographical Society of
Baltimore. Johns Hopkins Press. Baltimore, Maryland. pp. 147-181.

Northcroft, George J. H. 1902. Sketches of Summerland - giving some
account of Nassau and the Bahamas Islands. Nassau Guardian. Nassau.
Chapter 15-Soils and agriculture, pp. 174-185.

Orpurt, Philip A. 1964. The microfungal flora of bat cave soil from
Eleuthera Island, the Bahamas. Canad. J. Bot. 42: 1629-1633.

Robertson, E. C. 1958. Experimental consolidation of aragonite mud
(absitt.).> Wash Acad: Sci. Jine48 14221435.

105

Sharples, S. P. 1883. Turks Island and the guano caves of the Caicos
Islands. Proc. Boston Soc: Nat. Hist. 22: 242-252.

Storr, John. 1835. On certain manures applicable to the soils of the
Bahamas. J. Bahama Soc. Diff. Knowl. No. 5: 40-43.

T. TURKS AND CAICOS ISLANDS

Anonymous. 1836. (On the salt ponds at East.Caicos). J. Bahama Soc.
Dr£f£. Knowl. No. 17: 182-183.

e--------- 1859. Turk's Island. Salt exported. J. Amer. Geogr. Stat.
BOG (NY esl s, 123,

---------- 1973. The West Indies and Caribbean Yearb. Thomas Skinner
Directories. Croyden. 985 pp. Turks and Caicos Is. pp. 191-196.

Booy, Theodoor de. 1912. Lucayan remains on the Caicos Islands. Amer.
Anthrop. 14: 81-105.

---------- 1918. The Turks and Caicos Islands, British West Indies.
Geogr. Rev. 6: 35/-o1 .

Collins, Doreen. 196la. The Turks and Caicos Islands; some impressions
of an English visitor. Carib. Quart. 7: 163-167.

---------- 1961b. Turks and Caicos: unknown islands in the sun. New
Commonwealth 39: 377-378.

Cory, Charles B. 1891. List of birds collected by C. L. Winch in the
Caicos Islands and Inagua, Bahamas, during January and February,
and in Abaco, in March 1891. Auk 8: 296-298.

Darren. A. 1931. A Turks Island call for attention. Canad. W. 1.
Mag. 20: 108-109.

Doran, E. B. 1958. The Caicos conch trade. Geogr. Rev. 48: 388-401.

Doran, Edwin. 1955. Land forms of the southeastern Bahamas. Dept. of
Geogr Univ. OF Lexas. Publ. 5909. Ausitin, Texas 38 pp. +) 12
maps.

Greenway, James C., Jr. 1963. A name for the hummingbird of the Caicos
istandsi s Proc. N- Eng. Zool. Club 15; 105-106.

Guppy, H. B. 1917. Plants, seeds, and currents in the West Indies and
Azores. Williams and Norgate. London. 531 pp.

Howe, Marshall A. and Percy Wilson. 1908. Report on the botanical
exploration of the Bahama and Caicos Islands. J. N. Y. Bot. Gard.
o 41-50:

106

Isocrates. 1944. Bermuda's claim to Turk's Islands. Bermuda Hist.
Quarta (2504) e252 5 ole SSE

Lewis,, €.. Bo» 1954. (The Wurksiand Garaicos Islands.) Nat. Hast-.] Nowe
in Nat. Hist. Soc. Jamaica No. 66.

Nichols, J.T. 1921.. Aslist of Turks Islands fishes’, with a descrip
tion of a new flatfish., Amex. Mus~ Nat. “dist... Bull. 44321248

Parr, Albert Eide. 1930. Teleostean shore and shallow-water fishes
from the Bahamas and Turks Island. Bull Bingham Oceanog. Coll.
S271 -14824558) fio

Peake, R. E. 18995 On the sunvey by the’S..-S. \UBritanngsial otechemeapile
route between Bermuda, Turk's Islands, and Jamaica, with description
by Sir John Murray of the marine deposits brought home. Proc. Royal
Soc. Edinburgh 22: 409-429.

Proctor, G.,R. 1954-55. Notes on; the vegetation of the durksyand
Caicos, Islands. “Nat... Hist: Soc. Jamaica 6: 149-152. 170-l7Ay:
199-203. ;

Ray, Carleton and Alexander Sprunt IV. 1971. Parks and conservation
in the Turks and Caicos Islands. Administrator Office, Turks and
Caicos Is. Grand Turk.

Sadler; He E.s2197/0.. Turks cistand landfall. Privately offered for
Sale: —Grand Turks «2003p p:

Schneider, E. D. and B. C. Heezen. 1966. Sediments of the Caicos Outer
Ridge, the Bahamas. Geol. Soc. Amer. Bull. 77: 1381-1398.

Sharples, S. P. .1883... Turks, Island.and the guano, caves of the; Caicos
Eslands*) Proc.) Bostom soc. Nat. (Haisit.< 2257 242-2525

Smith, Hosay. (undated but appeared in 1968). A history of the Turks
and Caicos Islands. Published privately. Hamilton, Bermuda.

Stafford, J..L., Rw Ks. Hill,) and, E..L., Demontatene:. 1955.) Microsisierar—
~asis in the Turks, islands: W.-1. Med. J..4% l8S-187,

Verhoog, Pieter. 1954. Columbus landed on Caicos. Proc. U. S. Naval
insite oO OPS sale:

Watkins, F. H.. 1908. Report on the salt industry. Colonial) Rept ag.
Mise. 'No...56.. Turks and Caicos Islands. ~~ London:

107

W. NOVELS AND OTHER STORIES WITH MENTION
OF NATURAL HISTORY

Bell, Major H. MacLachlan. 1934. Isles of June. Williams and
Norgate. London. 226 pp.

Bothwell, Jean. 1965. By sail and wind: the story of the Bahamas.
Abelard-Schuman. London. 152 pp.

Bruce, Peter Henry. 1949. Bahamian Interlude. Richard Kent, ed.
J. Culmer. London.

Burns, A. C. 1949. Colonial civil servant. Allen and Unwin.
London. 339 pp.

Cottman, Evans W. 1963 (with Wyatt Blassingame). Out-Island Doctor.
Bee Duccon and) Co. joinc., New, York, N.Y. 248 .pp.

Cowles, E. 1932. Sand-hills and mountains; memoirs of a civil servant.
Aw. otockwell. London. 158 pp.

Defries, Amelia Dorothy. 1917. Ina forgotten colony. Nassau
Guardian. Nassau. 278 pp.

---------- 1929. The Fortunate Island. Cecil Palmer. London.
bel

Forbes, Rosita. 1939. A unicorn in the Bahamas. Herbert Jenkins.
London. 244 pp.

Ford, Leslie. 1952. The Bahamas murder case. Collins. London.

Hains, T. Jenkins. 1908. Bahama Bill. L. C. Page and Co. Boston.
368 pp.

Hayese Netson. ) 1940. Dildo Cay. Houghton Mifflin Go. Boston, .)329 pp.

---------- 1961. The roof of the wind. Doubleday. Garden City, N. Y.
216 pp. Reprinted 1962. Redman Co. London.

Helweg-Larsen, Kjeld. 1963. Columbus never came. Jarrolds. London.
240 pp.

---------- USo7 eseces of Paradise. Jarrolds. London. 41'84 pp.

---------- 1970. Arawaks and astronauts: twenty years on Eleuthera.
Jarrolds. London. 192 pp.

Hemingway, Ernest. 1952. Old man and the sea. Scribners Sons.
New York, N. Y. 140 pp.

108

Hemingway, Ernest. 1970. Islands in the stream. Scribners Sons.
New York, N. Y. 466 pp.

Langton-Jones, Ronald. 1944. Silent sentinals. (On lighthouses).
Frederick Muller. London. 332 pp.

LeGallienne, Richard. 1918. Pieces of eight: being the authentic
narrative of a treasure discovered in the Bahama Islands in the
year 1903 - now first given to the public. Doubleday, Page and
Co; Garden’ City) Ni Yoo 1SS5pp-

Lester, George. 1897. In sunny isles: chapters treating chiefly of
the Bahama Islands and Cuba. Charles H. Kelly. London. 144 pp.

Lorraine, Adela de. 1827. Letters from the Bahama Islands. H. C. Carey
and I. Lea. Philadelphia. 207 pp.

McKenna, Stephen. 1923. By intervention of Providence. Little, Brown
and Co. Boston. 298 pp.

North, Lionel. 1910. Off Elbow Light; a story of adventure in the
Bahamas. Charles H. Kelly. London. 248 pp.

Northcroft, George J. H. 1902. Sketches of Summerland - some account
of Nassau and the Bahama Islands. Nassau Guardian. Nassau. 309 pp.

Paludan-Muller, Frederick. 1867. The fountain of youth. (Translated
from the Danish by Humphrey William Freeland). Macmillan and Co.
London. 147 pp.

Powles, L. D. 1888. Land of the pink Pearl. (or recollections of life
in the Bahamas). Sampson Low, Marston, Searle, and Rivington.
London. 321 pp.

Rost, Leo. 1973. The conch eaters. Wollstonecraft, Inc. Los Angeles,
California. 266 pp.

Slack, Jack. 1968. Finders losers. Hutchinson. London. 174 pp.

Weigh, Audrey. 1970. Bahama adventure. Abelard-Schuman. London.
72 eppre

Wilder, Robert Ingersoll. 1964. Wind from the Carolinas. W. H. Allen.
London. 639 pp.

Young, Everild, 1966. Eleuthera: the island called ''Freedom.'' Regency
Press. London. 181 pp.

109

X. INDUSTRIES BASED ON NATURAL RESOURCES

Anonymous. 1836a. (On the salt ponds at East Caicos). J. Bahama Soc.
Diff. Knowl. No. 17: 182-183.

---------- 1836b. Bleaching of sponge. J. Bahama Soc. Diff. Knowl.
Nok 16 1742

---------- US59). Turk's) sland) Salt exported: + J. Amer. Geogr: Stat.
Soe.y (NY. Le > 123%

---------- 1906. Canning pine-apples in the Bahamas. W.I. Bull. 7:

---------- 1917. Sponge fishing in the Bahamas. Geogr. Rev. 3: 324.

---------- 1961. Wire-quarried building stones in the Bahamas. Quarry
Managers J. (London) 45: 14.

---------- 1968. Salt, east coast solar source. Chem. Engin. News 56:

2--------- 1970. Aragonite mining firm building Bahamas Island. Ocean

Bounds, John Howard. 1972. Industrialization of the Bahamas. Rev.
Geografica (Méx.) No. 77: 95-113.

Francis, Carlton E. 1967. Bahamas show commercial muscle. New Common-
wealth 45 (4): 142-143.

Hamilton, William. 1836. On the manufacture of sugar from beet root.
J. Bahama Soc. Diff. Knowl. No. 18: 187-192.

Hornburg, C. D. and A. W. Saariner. 1971. Bahamas on a salt free diet.
Water and Wastes Engin. 8: 32-34.

Miller, Daniel Newton, Jr. 1961. Early diagenetic dolomite associated
with salt extraction process, Inagua, Bahamas. J. Sed. Petrol. 31:
473-476.

Newbitt, C. R. 1854. Vegetable fibres of the Bahamas. J. Bot. 6:
237-241.

Northrop, John I. 1891. Cultivation of sisal in the Bahamas. Popular
Sci. Monthly 38: 606-619. (Reprinted in: John I. Northrop. 1910.
A naturalist in the Bahamas. Memorial volume. Columbia Univ.
Press sa uNew Yorks, IN. 9p 2 b2-224)).

Preston, George. 1890. Reports on the fibre industries of Yucatan and
the Bahamas. Eyre and Spottiswoode. London. 28 pp. Also Great
Britain Colonial Office. Ann. Ser. No. 85. Parliamentary Papers
48. 1890.

110

Rae, James McClure. 1891. Report on the fibre industry in the Bahamas.
Nassau Guardian. Nassau. 17 pp.

Roberts, E., ed. 1967. Land in the Bahamas. Which (Feb.): 36-41.

Roman, Erl. 1958. Tournaments can aid science (Bahamas International
Tuna Match) Sea Frontiers 4 (3): 164-170.

---------- 1957. The International Game Fish Conference. Sea Frontiers
5.4) 48-57 |

Rowe, George. 1864. The colonial empire of Great Britain, considered
.chiefly with reference to its physical geography and industrial
productions. The Atlantic group. Christian Knowledge Society.
London .

Stephens, William M. 1969. Bimini's concrete wreck. Oceans Mag. 2:
20-27.

Stubbs, G. C. and A. C. Langlois. 1954. Water supply of Nassau,
Bahamas. J. Amer. Water Works Assn. 46: 220-230.

Triviz, P. F. 1968. Bahamas site of new refinery. Sugar y Azucar
(New York) 63: 44.

Watkins, F. H. 1908. Report on the salt industry. Colonial rept. -
misc. No. 56. Turks and Caicos Islands. London.

Y. MALACOLOGY

Anonymous. 1944. A complete set of Bahamian shells, collected and
classified by the Bahamas Conchological Society from its origin,
March 15th, 1949 to January lst, 1944. Privately published,
Bahamas Conchological Society. Nassau. 11 pp.

Baker, Frank Collins. 1903. A partial list of the marine molluscs of
San Salvador (Watling Island), Bahamas. Nautilus 17: 57.

Bartsch, P. 1913. New Mollusks from the Bahama Islands. Proc. U. S.
Natl. Mus. 462; 107-109)'+"3 pl;

---------- 1946. The operculate land mollusks of the family Annulariidae
of the Island of Hispaniola and the Bahama Archipelago. Bull. U. S.
Natl. Mus. 192: 1-264.

Bendall, Wilfred. 1895. A list of the land Mollusca of the Island of
New Providence, Bahamas, with an enumeration of the species recorded
from the other islands. Proc. Malac. Soc. London 1: 292-295.

Bland, Thomas. 1873. On the physical geography of, and the distribution
of terrestrial Mollusca in, the Bahama Islands. Ann. Lyc. Nat. Hist.
New York. 10: 311-324.

111

Bland, Thomas. 1861. On the geographical distribution of the genera
and species of land shells of the West Indian Islands with a cata-
logue of the species of each island. Ann. Lyc. Nat. Hist. New
VYorkG7istvi-valo-+ (9-35).

Brown, N. J. 1886. Notes on the Mollusca of the Bahamas. Concholo-
gists' Exchange (Naturalist) 1: 12-13.

Clapp, G. H. 1913. Land shells collected on the Bimini Islands, Gun
and Cat Cays, Bahamas. Nautilus 27: 63-64.

Clench, William J. 1933. Notes and descriptions of land mollusks from
the Bahamas, based mainly upon collections obtained during the
Utowana expeditions of 1932 and 1933. West Indian Mollusks No. 7.
Proc iN: Eng.) Zool... Club. i13%.77-1001,

---------- 1934. Notes and description of new Cerions from Hispaniola
and the Bahamas, based mainly upon collections obtained during the
Utowana expedition of 1934. Proc. Boston Soc. Nat. Hist. 40:
205-218.

woo ------- 1935. A new species of Cerion from Long Island, Bahamas,
and a note on Cerion milleri (Pfeiffer). Nautilus 49: 49-50.

---------- 1936. A new Cerion from the Bahamas. Nautilus 49: pl eeilkss

ooo ------- 1937a. Shells of Mariguana Island with a review of the
Bahama Helicinidae and descriptions of new Bahama species. Proc.
N. Eng. Zool. Club 16: 57-70.

oeeoae--- 1937b. Descriptions of new land and marine shells from the
Bahama Islands. Proc. N. Eng. Zool. Club 16: 17- 26.

---------- 1937c: Marine bivalves from Little and Great Abaco, Grand
Bahama and Eleuthera, Bahama Islands. Mem. Soc. Cubana Hist. Nat.
11: 31-42.

atelier 1937d. Three new species of Cerions from Long Island,
Bahamas. Nautilus 51: 19-23.

wo-------- 1938a. Origin of the land and freshwater mollusk fauna of
the Bahamas, with a list of species occurring on Cat and Little
‘San Salvador Islands. Bull. Mus. Comp. Zool. 80: 481-541.

---------- 1938b. Land and freshwater mollusks of Grand Bahama and the
Abaco Islands, Bahama Islands. Mem. Soc. Cubana Hist. Nat. 12:
303-333.

---------- 1940. Land and freshwater mollusks of Long Island, Bahama
Islands. Mem. Soc. Cubana Hist. Nat. 14: 3-17 + 1 pl.

LZ

Clench, William J. 1942. Land shells of the Bimini Islands, Bahama
Islands. Proc. N. Eng. Zool. Club 19: 53-67.

---------- 1952. Land and freshwater mollusks of Eleuthera Island,
Bahama Islands. Rev. Soc. Malacologica "Carlos de la Torre" 8:
S7- UNG nd) ply.

---------- 1956. A new Cerion from Bimini, Bahamas (Mollusca). Amer.
Mus. Nov. 1794: 1-3.

---------- 1959a. Land and freshwater mollusks of Great and Little
Inagua, Bahama Islands. Bull. Mus. Comp. Zool. 121: 29-53 + 1 pl.

---------- 1959b. Origin of the land and freshwater molluscs of the
Bahamas. Rept. Amer. Malac. Union 1959: 4-5.

---------- 1961. Land and freshwater mollusks of Caicos, Turks, Ragged
Islands and Islands on the Cay Sal Bank, Bahamas. Occ. Papers on
Mollusks 2: 229-259.

---------- 1963. Land and freshwater mollusks of the Crooked Island
group, Bahamas. Bull. Mus. Comp. Zool. 128: 395-413 + 3 pl.

---------- and R. A. McLean. 1936. Marine bivalves collected by the
Harvard-Bahama Expedition of 1935. Mem. Soc. Cubana Hist. Nat.
10: 157-168.

Corgan, James X. 1969. Pleistocene marine molluscan faunules from the
Little Bahama Bank (abst.). Tennessee Acad. Sci. J. 44: 49.

Dall, W. H. 1894. Cruise of the steam yacht "Wild Duck" in the Bahamas,
Jan. to April 1893, in charge of Alexander Agassiz. II. Notes on
the shells collected from the shores of the Great Lagoon, Watling
Island, Bahamas. Bull. Mus. Comp. Zool. 25: 113-123.

let dtaatatates 1896. The Mollusks and Brachyiopods of the Bahama Expedition
of the State University of Iowa. Bull. Lab. Nat. Hist. State Univ.
Iowa 4: 12-21.

-------- +- 1900. A new species of Cerion (Cerion stevensonii).
Nautilus 14: 65.

---------- 1905a. Report on the land and fresh water shells collected
in the Bahamas in 1904, by Mr. Owen Bryant and others. Smithsonian
Misc. Coll. 47, part 4: 433-452 + pl. 48-49.

------- --- 1905b. Fossils of the Bahama Islands, with a list of the non-
marine mollusks. in: Shattuck, George Burbank. The Bahama Islands.
Geographical Society of Baltimore. Johns Hopkins Press. Baltimore,
Maryland. pp. 23-47.

113

Dall, W. H. 1910. List of shells collected by Dr. John I. Northrop in
the Bahamas, identified by Professor William Healey Gall, Smith-
sonian Institution. in: John I. Northrop. 1910. A naturalist in

the Bahamas. Memorial Volume. Columbia Univ. Press. New York,
Neo ¥. pp. 99-102),

D'Asaro, Charles N. 1965. Organogenesis, Development, and metamorphosis
in the queen conch, Strombus gigas, with notes on breeding habits.
Bull. Mar. Sci. 15: 359-416.

~--------- 1970. Egg capsules of prosobranch mollusks from South Florida
and the Bahamas and notes on spawning in the laboratory. Bull. Mar.
Sci. 20: 414-440.

Doran, E. B. 1958. The Caicos conch trade. Geogr. Rev. 48: 388-401.

Durrell, Zoé C. 1972. The innocent island - Abaco in the Bahamas.
Durrell Publications. Distributed by Stephen Greene Press.
Brattleboro, Vermont. 157 pp.

Emery, D. L. 1924. Collecting in southern Florida, the Bahamas, and
Cuba. Nautilus 38: 56-62.

Hebard, Edgar B. 1967. Pleistocene mollusks from New Providence
Island, Bahamas. Nautilus 81: 41-44.

Henderson, J. B. 1901. Land shells of Fortune Island, Bahamas.
Nautilus 15: 85-86.

woccccen-- and G. H. Clapp. 1913. Cerion (Strophiops) Biminiense, sp.
nov. Nautilus 27: 64-65.

Hjalmarson, J. and L. Pfeiffer. 1858. Beitrage zur Fauna von
Westindien. Malak. Blatt. 5: 135-155 + 2 pl.

Jacobson, Marris K. 1965. On some land shells of Eleuthera, Bahamas.
Nautilus 78: 120-125.

Krebs, Henry. 1867. Catalogue of marine mollusks collected in the
Bahama Islands in November 1866. Ann. Lyc. Nat. Hist. N. Y. 8:
427-431.

Little, Colin. 1965. Notes on the anatomy of the queen conch, Strombus
gigas. Bull. Mar. Sci. 15: 338-358.

McGinty, Paul L. and Thomas McGinty. 1948. The Pioneer - Bahama
- expedition--May, June 1947. Mollusca 2: 40-55.

McGinty, Thomas L. 1948. A report on the Mollusca collected by the
Pioneer Bahama expedition. Mollusca 2: 56-69.

114

McLean, R. A. 1936. Some marine bivalves from the Bahama Islands.
Nautilus 49: 116-119.

Maynard, Charles Johnson. 1889-1896. Monograph of the genus Strophia,
a group of tropical and sub-tropical land shells. Contr. Sci. 1:
129) 69-79 25-155 3 SSO 2 MO = LONI 5 AO:

---------- 1924. Supplement to fall catalogue of Cerionidae with
descriptions of new species collected in the Bahama Islands in
summer of 1924. West Newton, Massachusetts. Privately published.
6 pp.

Mayr, Ernst and Carmela Berritto Rosen. 1956. Geographic variation
and hybridization in populations of Bahama snails (Cerion). Amer.
Mus. Nov. 1806: 1-48.

Moulding, Mary Baker. 1967. Shells at our feet. An introduction to
shelling in the Bahamas. Sea Scapers. Chicago, Illinois. 102 pp.

Parodiz, J. J. 1957. New records of freshwater gastropods from the
Bahama Islands. Ann. Carnegie Mus. 35: 1-9.

Pfeiffer, L. 1867. Drei neue West Indische Pneumonospomen. Malak.
Blatt. 14: 165-166.

Pilsbry, H. A. 1898. Land shells of Gun Cay, Bahamas. Nautilus 12:
26-27.

---------- 1931.- List of landand fresh-water mollusks collected on
Andros, Bahamas. Proc. Phila. Acad. Nat. Sci. for 1930. 82:

---------- and M. Black. 1931. Cerions of Andros, Bahamas. Proc. Phila.
Acad. Nat. Sci. 82: 289-296.

---------- and C. N. Grimshaw. 1936. Oxystyla undata undata in Florida.
Nautilus 50: 19-20.

Plate, L. 1907. Die Variabilitat und die Artbildung nach dem Prinsip
geographischer Formenkelten bei den Cerion-Landschnecken der Bahama-
Inseln. Arch. fur Rassen und Geseilschafts-Biologie 4: 433-475;
581-614 + 5 pl.

Randall, John E. 1964a. Contributions to the biology of the queen
conch, Strombus gigas. Bull. Mar. Sci. Gulf Carib. 14: 246-295.

oan cc----- 1964b. The habits of the queen conch. Sea Frontiers 10:
250-239).

Richards, Horace Gardiner. 1954. Pleistocene mollusks from Andros
Island, Bahamas. Nautilus 67: 120-121.

PLES

Robertson, Robert. 1959a. Observations on the spawn and veligers of
conchs (Strombus) in the Bahamas. Proc. Malac. Soc. London 33:
164-171.

---------- 1959b. The mollusc fauna of Bahamian mangroves. Rept. Amer.
Malac. Union 1959: 22-23.

---------- 1959c. Marine mollusks of Bimini, Bahama Islands. Ph.D.
Diss. Harvard Univ. Cambridge, Massachusetts. 294 pp. + 17 pl.

Simpson, Charles Torrey. 1894. Distribution of the land and fresh-
water mollusks of the West Indian Region, and their evidence with
regard to past changes of land and sea. Proc. U. S. Natl. Mus.
17: 423-450.

Smith, Egbert T. 1952. Romance of sea shells, over 100 of the most
interesting Florida and Bahama shells and other specimens of marine
life illustrated and described. 3rd ed. Published by the author.
Bee Myers, Florida, 28 pp.

Vanatta, E. G. 1919. Polygra plana bahamensis Van., n. var. Nautilus
Soe 2

‘Warmke, G. L. and R. Tucker Abbott. 1962. Caribbean seashells.
Livingston Publ. Co. Narberth, Pennsylvania. 348 pp.

Wurtz, Charles B. 1950. Results of the Catherwood-Chaplin West Indies
Expedition, 1948. IV. Land snails of North Cat Cay (Bahamas), Cayo
Largo (Cuba), Grand Cayman, Saint Andrews and Old Providence.
Procserhila. Acad. Nat. Sei) 102? "95-110.

Z. ZOOLOGY, GENERAL AND MISCELLANEOUS INVERTEBRATE
Anonymous. 1917. Sponge fishing in the Bahamas. Geogr. Rev. 3: 324.

Agassiz, Alexander. 1888. Three cruises of the United States Coast and
Geodetic Survey Steamer ''Blake.'' Vol. 1-Chapter 5: Relations of
the American and West Indian fauna and flora. Bull. Mus. Comp.
Zool. 14: 113-124.

Andrew, Warren and Nancy V. Andrew. 1953. Some annelid and Sipunculid
worms of the Bimini region. Amer. Mus. Nov. 1617: 1-16.

Bayer, Frederick M. and Elisabeth Deichmann. 1958. Two new plexaurid
gorgonians from the Bahama Islands. Bull. Mar. Sci. Gulf Carib. 8:
224-235.

Bollman, Charles H. 1893. Notes upon a collection of Myriapods belonging
to.the U.;S. National Museum... Bull. U. S. Natl.'Mus. 46: 190-220.

116

Borror, Arthur C. 1973. Tracheloraphis haloetes, sp.n. (Ciliophora,
Gymnostomatida): Description and a key to species of the genus
Tracheloraphis. J. Protozool. 20: 554-558.

Brattegard, Torleiv. 1968. Marine biological investigations in the
Bahamas. 2. On an association between Acanthopleura granulata
(Polyplacophora) and Dynamene spp. (Isopoda). Sarsia 32: 11-20.

---------- 1969. Marine biological investigations in the Bahamas.
10. Mysidacea from shallow water in the Bahamas and southern
Florida, Part, 12)..Sarsia +3593 917-106).

---------- 1970a. Marine biological investigations in the Bahamas.
11. Mysidacea from shallow water in the Bahamas and southern
Florida.) Part, 2.) .Sarsia) 415 1-55"

---------- 1970b. Marine biological investigations in the Bahamas.
13. Leptostraca from shallow water in the Bahamas and southern
Florida. Sarsia 44: 1-7.

Brattstrom, Hans. 1968a. Marine biological investigations in the
Bahamas. 1. The cruise in 1967, general report. Sarsia 32: 1-9.

-o-------- 1968b. Marine biological investigations in the Bahamas.
6. The cruise in 1968, general report. Sarsia 35: 1-8.

Cairns; J., Jr..and.J.sA. Ruthven: 1972: A test of the cosmopolitan
distribution of fresh-water protozoans. Hydrobiologia 39: 405-427.

Carter, Howard P. 1972. Infraciliature of eleven species of the genus
Euplotes. Trans. Amer. Micro. Soc. 91: 466-492.

Chamberlain, Ralph V. 1918. The Chilopoda and Diplopoda of the West
Indies. Bull. Mus. Comp. Zool. 62: 151-262.

---------- 1952. The centipeds (Chilopoda) of South Bimini, Bahama
Islands, British West Indies. Amer. Mus. Nov. 1576: 1-8.

Chappuis, P. A. and C. D. Deboutteville. 1956. Etudes sur la faune
interstitielle des Iles Bahamas récoltée par Madame Renaud-Debyser.
I. Copépodes et Isopodes. Vie et Milieu 7: 373-396.

Clarke, William D. 1955. A new species of the genus Heteromysis
(Crustacea, Mysidacea) from the Bahama Islands, commensal with a
sea-anemone. Amer. Mus. Nov. 1716: 1-13.

Clarkson, M. J. and L. N. Owen. 1959. The parasites of domestic animals
in the Bahama Islands. Ann. Trop. Med. Parasit. 53: 341-346.

Couch, John A. 1973. Ultrastructural and protargol studies of
Lagenophrys callinectes (Ciliophora, Peritrichida). J. Protozool.
20: 638-647.

117

Craig, G. Y. 1967. Size-frequency distributions of living and dead
populations of pelecypods from Bimini, Bahamas, B.W.I. J. Geol.
75: 34-45.

Dana, J. D. 1872. Corals and coral islands. Dodd, Mead and Co.
New York, N. Y. 440 pp. (Later editions: 1874, 1890).

Darlington, Philip Jackson, Jr. 1938. The origin of the fauna of the
Greater Antilles, with discussion of dispersal of animals over
water and through the air. Quart. Rev. Biol. 13: 274-300.

Deboutteville, C. D. 1956. Etudes sur la faune interstitielle des Iles
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118

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120

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—_

x U.S, GOVERNMENT PRINTING OFFICE : 1975 O - 573-620

ix ( 5 December, 1976

ATOLL RESEARCH
BULLETIN

192. THE NATURAL HISTORY OF JOHNSTON ATOLL,
CENTRAL PACIFIC OCEAN

by A. Binion Amerson, Jr., and Philip C. Shelton

Issued by
THE SMITHSONIAN INSTITUTION
Washington, D.C., U.S.A.

Errata

In Atoll Research Bulletin #186 the names of the authors, Roger
B. Clapp and William 0. Wirtz, II, were accidently omitted from the

cover.

On the title page of #186 the publication information at the
bottom should have read as it appears below. The part of this page
below the dotted line may be cut out and pasted over the bottom of the
title page of #186.

Issued by
The Smithsonian Institution
with the assistance of
The Fish and Wildlife Service
U.S. Department of the Interior
Washington, D. C., U.S.A.
February 15, 1975

In ARB #184 a line was omitted in final typing. On page 5, in
paragraph 4, line 4, after the word nesting there should be inserted
"does occur to some extent. It was not determined if increased nest-
ing".

We regret the occurrence of these errors.--Eds.

om
sla Te :
Pk ke

a font

ATOLL RESEARCH BULLETIN
NO. 192

THE NATURAL HISTORY OF JOHNSTON ATOLL,
CENTRAL PACIFIC OCEAN

by A. Binion Amerson, Jr., and Philip C. Shelton

Issued by
THE SMITHSONIAN INSTITUTION
with the assistance of
The Fish and Wildlife Service
U. S. Department of the Interior

Washington, D. C., U.S.A.

December, 1976

ACKNOWLEDGMENT

The Atoll Research Bulletin is issued by the Smithsonian Insti-
tution as a part of its Tropical Biology Program. It is co-
sponsored by the Museum of Natural History and the Smithsonian Press.
The Press supports and handles production and distribution. The
editing is done by the Tropical Biology staff, Botany Department,
Museum of Natural History and by D. R. Stoddart.

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

The sole responsibility for all statements made by authors of
papers in the Atoll Research Bulletin rests with them, and statements
made in the Bulletin do not necessarily represent the views of the
Smithsonian nor those of the editors of the Bulletin.

Editors

F. R. Fosberg
M.-H. Sachet

Smithsonian Institution
Washington, D. C. 20560

D. R. Stoddart

Department of Geography
University of Cambridge
Downing Place
Cambridge, England

TABLE OF CONTENTS

Page

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else Om pend 15c) HL aby Cx area reese 5. ie! eieica.ca «: ae 01/6 niin) 5 (otenalerekeliecclleate) ayai'suciieueyel-s. » xvix
ITE ROGURE TOM: sc co et mertlek ah et Mery tates sive oy overlord avsyedeun tal ayaee evsueheaatetelereveie s!> « i
aris T Cua Env OMMET E <.sy as tl<! 3.5pa.Geieiw ss) a! oo 6 a0: aeyeee aucuapete: yee sjovelege\Sydtel « 3
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GEM ery a cl aca cate aiclaie fever neeika 5 Shc tees + oow' oven @ 9 bop eyaywioceliee Siestreke osye"s 7
Geomorphology....... Ms Si ate Pana aiisin m3 0.0 Sheceta eo & aeVeyeds agneiatape io ta's: 6.8 12
STN s Megat ag asks, or wish ais, ielieh OW oy cu sh'av'as 6s wie wi 'Siw Sp Sueteue Aenean ahshe @eide 15
ROMPCRAGMEC IS: srk. cSisae aslckete leis Cs clove Boerbee wie dave. 6 Se 0 6 wresdiauaenehe IS

DREN oe css PEA ERS, 0) Ore, Gach can) elimn cl axortn. @ hala soared abowayeuenateueks (appaneesdadowe!'s 15

PRTC Apt ie HO MNS el hens, MT ae CalcOe as sil & or apehchegarsy Suetauane stamakhapess ie. 0, 18
Paha ELUATE CED RE Res ct AE cy ns hci av s\ orev aviar ot leh heWauchous heeeuedate ats! ele se 22

Silt COS Ree Bae ae rata) ch clas ar sysveh oh chet sbevioncrsiensh stot hekcuskelch eget penates 6 Slee 22

SF ANH PEA PALA ert ie tad ay cu ch usin ei vl =i mosis ax si eras acai avav ov oranes eal chev arouheastohanele = eee ele 24
Major svabereMassesirand (CuGGenit:S.:.:csteyedatate sisieiaelere stage «= = 6 6 24

a Gaile Warts erscpecamiit a Taree mi oe bet cM cnyar evoveley <varey'o\'n1 a: a¥el oflovalioseteno eae 26

STS oon s neh eMC oe pth CRA ENG ce synch o: Shoncyenerehovb opis nihasNdaenadetoltovayalens te 30

SetSMae, OT PEST AM WAN ANES 55, oy syupsirsnsisnahevederesoeavalssel ave er auaeetehe ferenmie © 30
Seta TaN Ba ac =) ee area haat PERE A a ig 1s! wav ons ove ayetenecassiekuancheyatele s1c)a-s 30
Diseovery. and ‘Baris. Ba pillor atl c!6 i. .:<saerediesejeiele:yejsyerenale'<eie s\ 32
Steir LCT LOG. Aetalsi te iotetsis ves etelctalele! ola > ieneje(s isin jeieterals se fakatwuctebemsiete Tales.’ 34

Mam leiaaiay OCCUPAtsh Oils cis nintatalaisetctalclalttsie ls leieiiats to jaketsumioks sol eialatelalatatale s 43
oUPag.c sgs¢hoeobondonooda cos ond so tenons eee ae ec ee ooée Gee 47
EMU nue. cle evetebehetoisie iam teke «i letedeis sein «oh oye: laneneietobaxehabe iol acRevetenererels) oe sos 47
PMLA CNT e clin a euaMata lege umteta am ietaatieta te ise faveic valle Je Jo te Sefolsiitaal dametonetere ie se lace 47

Wits Cen ae Eanes va aindete. « wlctesaiete ts slo aKa lero 1s fa eleseParwbors gevehounvals ec 6) o/s 51
Opeslosineil Wibocsin pass 6 ca eo IIe Theron en oO mc Ocoee 31

Neeonys Men oescyculy4 Say ecco oon Ooo ocr odd me 61

ice any ene CO ara Sia at ietoiat ew pisos) ale =. 6 egeuetiapulels lakanetedeielwietele sie s\'e 62

AieriMshs LAMORE PORE A ins, cc ntlarsyiits Ranate SyayeseieReta sce soe! = 62

lymrune! IWS ibehatel one eres oo die on OO 0 0 OO COILING 63

Josie! lysine ~SVap esas oie doasotca ds othogbdooan 63

sig! ibslisiyaly has ceo goemeeo coo on ae oo or Con OnG rue 65

CreateannZuln ano: SOOO oon aoc oo So cdo CoO oeo colon 65

Meni m ad Gpeeta ceo ro Cal aes edegeyeaeneyeveyevaroraratiens (everain elas + 65

Hime eb ral ess Bynepey awl ey. tots A CUeENS. s,s ,4.cchadeuacete avauch neato, Mhelsiel os eS 6. tie 65
Gniadardal ((CoethenEera ba) sig afar, cis acaucreyeneceueie ale cre ele SS es yercha 67
MACS Ca’ ener S SANS Hay aicctets. 1 alec Sins ay casbaciels opeadteyeyegeisoers Gok s isle 70

(gu SlsiGhels alc oS oO EO COO DOU OC EP OOO es CD GO OL OOOO OT CCID CHISE 70

INE TEMIEOD OG cicyatard Weksielekalsiels ciel ctehener cielo, flacelereveqeneteietels = eels. diene. so s' sce 70

ea tesla chatted ti eWeliatte atatetlalaiieia otal Nakeliclie oy aici diel sieiel hella) «leiele snes sie 70

FREI Ge Sit relied ae yaa hee cee. oa 5. al euansieneheKe (al eyakerspeleie a os ole. 6 EL

Medically, ImportanitteSpeCLesic;«, .\c1s cic eee crejeneie «le ee ie

ICRC o Beles Cad oceoc Sano COC a oot OOOO cmos 79

GOs METIACK AC ate nse laila\lsiel sted cile e/a «ale iela starayerelers «ens 84

RMETUOMGTS Sil aero) cicle ei cllopsl wellsuscenebeilcie ehefiste s! sve olete isles 86

MBULNOMINEZI. 54 Boog ao BOO UD ONC CHO De COCO OOOO tIO OO 86

EP OHO Calle iel cites clei cheelieier slieile (clei elle (a1 slailallatiel elias el ee \e 86

tt

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Eehino@ermaitals%s\':<.si0e ovo1c) eo chaters epetete rele: <foveie ccereislansiovere Siemeie wanes 86
VERCEDT ACCS 5:5) cisic7s cic «o/s ooieie sucvorene oder enerorenstens cuchcyoncrellshavetaveverenene renee 86
EiShese te cteke iekchevon: atatal Macletctcle shovetewsyiel stoRchonele stiches ciancte’ stepeuetenenaie 89
Pelagic sFISheS 0 jenctpotscnenciels crorhovetstcte ater aicke oh a er kenae 89
Inshore (PaShes) ech dicte etevchenetenede keystones Rls eictols: sfietelic sieneper sions 90
Distribution and Abundance Within the Atoll......... SO
Effects) of (Dredging titi A tertorsilers total ls a aleve anaveve a tevane 92
Cisuater Pe, “FISHES fo elets ecle eats eNererete oleteialcvelote sels cielc eects 103
Species “Accounts /.).).\...'s 46 sie see eee Blavsrs ete sre ies ovate: sealers 110
Moray: *Be lic,’ Fatsneigietons <covenoneratet o ctatichiouevone cleto ote rata craters 110
SuUBGEONEASHSS Me ipotoherener severe erepenaio: seis ctcenetche scone, s auch erate Hail
REPETUSS La. chats Pelay hooste crober enobenene taeemsteetctey or ahctetshe lores nie arene a2
Spectes) “ACCOUTES)...5.f2lrate stevcieloreterers stelciecerctote sretoteteietcce: Sates ILIEZ,
Bhack (Sea Tur tle cael ate cea ers cro oioieisete’s Oe eS otete ete,
HOUSE GECKO. ahs telate to otel slop onetoneorena aleneranerensre late a iene iotsrane 114s}
FOX (GECKO. Fitts s ctelercters oie cre ecate oust oratoneiate anes cen ons eveus ILLS}
Moremi (GSekOr tects oxo sterolere cacieioreisioreotersrctets cts. stoners 113
Snake=eyed 4S lcimle cies erarersteretsye stopeuete tererete core crete ercrete TUS
BLP sense iret hats N hostels atc attonanetalelatal ce loteretermetano terete errs taiete ctenaie cienams 114
MMERO GUCETOM rp octenstetoasten<toneroteraretenen aiietist chalctisnsvotas staha ies atelctone 116
SEab id's! sy. jot stake stetoretsotcrohe ictenoModenotererererstenstere enetencte chore oie toremene 118
BLCOAOLS tatetatese literate tonetitanetetateretercd shateletetetelatote clelcte steiebe 118
Former, Breedensics parc cisreleaercne els ect ouotenerskotenslic onerens'e ss vets 118
VASE OTIS £2 faletalel Sa tetehetstoliohcteletcret one rel crcholcteleloletetitetelerseer che 118
Waterfowl Marsh, ‘and Gand /Birdser srt alee cl erere ere ete che EIS
Resudlar Mis ramitsi.s.*.sorscttateretstoneralet ster ohotalete tote ctetalatet chats JE)
bere Sullar Vials athorsi.tassts ohevevonetctetel ovel ohatict atel ol slcl.ctctonnenatens IES)
Strage hers «bese let cratare Motetet ctor etater shovel aleve: olatel banclanene 119
Acietidentalisi.e.i 4.000 st notch tcolcte stastehe ciaters cig seer 119
Introductions. Aware PNET Mat ately ene Ta ana tsd aes IA)
Annual Gye lies. ts... Foe) te teidichantene sates crete oe ae es oe catetens LLY)
Breeding aCy Chess tatwers siete checcbelat ets’ s oieccye selene ee eo rerere 120
Winter ‘and\*Sprameu aiecnttets soe tee teeta ee 120
Sprang “and “Sunmens eae oat eee en eae 120
Stamm Pentti tetas tet erctote Oe cree tee roe cates 22
Summer "anid: “Faull eset Sere ene eee ear ae 122
EXtend edie ty 225 OR een Meee cs ane 122
Population Cycles wee eae ss see ne Moles acta tn 122
Breeding Seabirds....... eieret sr clavere sbonenee set enciener 122
Former Breeding ‘Scabirdses. s22o. se oon. one 125
Reguillan Milerantto tee w ae ae oe eee 125

Irregular Visitors, Stragglers, and
AccidentalleBirds 2.5... -. I Depo hope beatae Pah 2)5
Introduced eBid Swen cece c octets eevee crerelere sc stevens 126
AtE-SCa *Biidsrt es herciclncietcrete iste onave oucteforsierer ce otonerenele 126
Effects of Weather Patterns..... PA DIDLIOIO COD Boat 134
Ecological ‘Distribution Within-the Atoll. sc... 53. 135
Seabed Sierra tect cot sel atetisrenel ey stot eeten ch ole rers eel seekers cere] oles 1635
Lbaerers BUI Von end os oOo abinod Boo aSbadGa 35)

Vili Hs OES it Bhat o3l0 Stheneke Pee tN Seo Mes cg ce. Siete 135

Page

Waterrowl. Marsh, and Land Birdseses ests 6 c+. cnc 135
Resullar Mierants..- os Sac eer e a ncihcae eee en 135
Irregular Visitors, Stragglers, and 135
Necidentallors, tc oe ne oe ee eee crete es wicca oe) ot ILS }5)
HERO GUC tT OMS 7s els ate rat ERS fale Ste Pace les a 610s one 135

Wotlanwd NeCOuMES...'svsc cs sts ee se ene ciate wroetate ee eels 136
icant pista rid, % "soar with eens ore ee ee olaeve aboie 136

et ccna elSMari cls, *. 7.2. tatostalstererers 6 hee i) Sa ara yeeee 136
Johnston. Vsland'."s*. at. tle och ees alatetets @ aie 6 ea. sire 136

SAMA USMAM Ae retene sw tetelotete eter cians co eter ciate eaten Gia wis tele US7

MOVE EULESS an Din GdIII Orn Petra Oo DONG Gus Gono cond 139
Bandine wand Interictand Movememtct cs ct. slo ties es ccs + 141
Banepa tie oe ne ere eee ae eee ee coats 141
imterrsland) Movenene. 1 ceed cee eee eek ences s 141
GAA als NPM INTE TOS. tot tact cle ote osche othe oe oi tialie@) a/eravereie 145
eS CMMACIUC CR eer ren cance et ten tt tate 8.8 earanne EMO A ue cat graie 148
SpeECiMeusmaeie ecm ces crore cet entane Mave ore ere cen eee nm wrepaienate 149
SPECTESTACCOUMES oc cee Pek eee Co OR eS Ce TR Tee cic ale espe
Black=footedehtbabtross. ou sae tose tc oie meets. cles etnies Sy
TaysamyALDAELOSS ite ce oe oc cree tie ee ose Se tnie one © elone ws ees)
PMOCr ome HCC teens cot we hee tert nrer waver k: relates 155
Budwer Ss) betwee si cc ste sis cieicis 2.00 o'e withe e erdis eyale se 155
Wedge—carlled "Shearwater. oo. cess sts. ve te se ss se ais 165
ChETSEMAS TONCAEWALECE. cc cc cetcce ccs os cote ate eae 178
Newelbts TSnearwabers cect s ccc ccs ee sc ce ae 8 eialare 187
Sookye Stone retrehs 2a 5 7555 SARL BEE So os tie 188
Red=palWed TLEOpTreDUEds Woe o eter cic alee ce severe « eealaie 188
ReU=taived Eroptebird: ts) sfc, 222 SOM ce es 192
White—catled tt boptebard: * S40 0y 2 os eee. ols crcl ata 205
Bilte= Laced BoObye fees erere fume oe che are er olla evcheve 206
BELOW BOODY Re coe re ne ee ene ee es slaiahale 2)
REd=footed Booby! Tees Poet kee s Ce ee ee ee eee ae 229
Greattlragatrebind. 27./). T0228. Ay Ree ARR 244
RESSCEP PE UPateD PEC. cree eto eee sete eo eters tole trotalche eamene 260
CarClevEprece es se cee se ee ee ee Ss Re 260

12s Ua) 52 ial Le ee ee aa aS ieee eR eck eek eae CERES 262
AMEV Tan WLBEONM Scat she ke oe teed one ee ee lees eis: « 265
NOGENCEM OMOVE We Te etnies ce ete cree cre setae eretal crater elo loess 266
Domestic Chickents. © +o Ae leh Ane eee AS 266
BECC LENCO. sre sc aie eerste ciel ererencrale atetenetel ole aieleRa tate 266
American Goulden PLOVETS (c's o e8t o75 che sie atetane alors cveoietere 267
Black—bewltedmePloviend, «sisie sc sia sess cieteteye aieietote tenants ts 273
Sema padmated LOVES ee ss) ele/etaia cists ls atetatele ate leteiatereteret= 273
BEsret eth ohed. CUT he Wa mic clcle eieie elelete ctekeia tel ctcre tel enetone 274
ILEASSis WOT INO ALC USES eke minrencinig:d Gicvc Oo D1a1G0 cic(o.u. 010.0 274
SPOEHeC MS anG pipes cc test leis te mieleteleteta sal aalenelaleneren sone 277
WISNER eo ierepetevchs aiveire ovrelic one leliehel cuavertoue er coshane tone oletiele ore corets 278
Wandienad nip ib atEMen tics lcleisrersatelorsseleie nie stelelereiecheie ereiey sie 279
Ritilahy Whe 6 Ao goood dOU OC NOON GodoGoUoomDOUE 281
DOwAEChercuespeCueS s elersieiciaile or sre ae oslo eiloreeevelalel exeickel'e le 286

SANGIN Se Ooo SONG Goo OU agp oodcodoGadgOlC 287

ie,

WwW

Page
Western Sandpipers cpeisccic sisieneccusic dice veisinustere 6 erenonanenene 289
Pectoral) Sandpiper= .icvlesccxele's ae Soils Sib hind Gicie - 289
Sharp=taided, Sand palperiyergerccs cise 6c occiale © see el eiee one 291
BuEf—breasted: Sandpipers cckekecleie sie ici eic ere iele eer etoiere 292
RULES. oe je re ote oie odevewelere ore eaerere ns HeleeReleioncioNolel ener encienelel enero 292
Wilsons Phallarope... 3-4. Rus assusae ohoieueteaeye verte eis 293
Glaucous-winged Gull............. sfeye! slic elie) ehevel slemaieue 293
Herr ime” Cull as cies seca nedene eee sersucrs SouonDcoKOOOb 294
awe hime Gules A6o ee perer ee a kreeina Ny oe Sah Ao ee 294
Franklin" ss Guilds 20 %c ts epee tases siecs.tue 6 oS oe nh ree 295
Cults pecLess 62% sc suc c euele mcr. oon Gunes Soudoodso 0 296
Gray =bacCKed erm it tc/discweroters: ecouecevn here ih a6 ob sue tie ones 297
DOObY) Lerner clchere oloereeneneionerene clelereieacholenecenelera ee teretre 305
Elegant Perm. 212.6% »:6veceuenenctcncue’s sie cre cue lerecs cio cue erere iene 322
Bilwe=or ay™ Noddy sists ctu. cicvarsreneeckereraiencia sichele) e cutpelereieraene B22
Brown, Noddy. ..0 acc oe odisiodid dS blaliGod diss Goddid ouc.c% 323
Black Nod dys. cuercnkes oe eesietle eos cee eee Coc cre ee eue eee 337
Whi termleriny ss bt ase ocr siereneus eterenenelnts acre se eels on eieeienone 345
ROC DOV.C's cs) ey 01s) sale econenenele Scum cbold doo dg dduidd coo 350
SHOBE—eCaried™ Owilkcs cre crenencsesveleneyeecgeveieicac ocie oper oeroeneiehe 3511)
Sey iarrle & % 35.6 eke nee anne eR eon ean ts - 353
Japanese Whitteseyecris alarcrersicrse/spele eis vere alec oreo toners 354
SOCTEEY, Hameh cs ercreye cue roncnencheterciesiel sere. cnei che ts) ol oeuenenenene 355
Mammal Sis: ./e se: 0 sealer o cves stone ofene erenerenonsten crogeiere) chee, ace1e) chepencherey creretonene 355
Species Accounts......... sicheNoneke le’ hale cre cieie sicueperclelereroiekene 359
HOUSC TMOWSE'. ca «lars ol sete chenetol clei cisie «elie /sieileic/ eles’ steer oneionens 356
ROO Rabat. ac ste cere SOU OOO U COO Oates Gibiod om os SNS
DOMES CIC DOG eiex creep stoners hetelcusrevel ores) 06 selec: sicre ols ereietenene 35)
DOMES CEC Cathe oc ieraeusvoneeicrencle’ an.5. 050 aenove als cuelie- ciel eles oemens 360
Hawattan®’ Monk Sead cr. sc)epetnieconesic «sin ieie seus avers oie ene letene 361
European Rabbit........ Buel oie! che. cletsurielaieLehowecel caste reine 361
Feological (SienitaCaneek. spersieciecherenatelens eieioickeree ienensieiete SUS esp a5 6o 56 361
Beological Values>otr Johnston AGoiM oie: .s.1- 2 cress cis oye e so eel ercvene S36:
Johnston Atoll: Some Ecological Considerations............. 362
The Place of Johnston Atoll in the Natural
BPOSPN EEC crc cre, ole ekeleierenwereicl = elare) ckersie cle oveieNel cael ete ereier ne reneeete 362
The Importance of Johnston Atoll as a Scientific
Laboratory eo%:3 5508) S sieve ts ool ovencuateie rev sitiie Sous ne ueveentioie erat aeuenens 363
Future> Potent Latico cco cus cxcce tho cuapektns Weuste) ccoms ion susie cps see, ue ebererens 364
Ecological: Sienifticance: of Johnston Atoll... cy o5 cssus toc) ce cre ous 365
Johnston Atoli:” Ecological’ Sinise CanGets:. «, seus sic oie releleseie siete 367
SUMIMAMIeY, sive gaits! slike catiecaleransusierene cua tevel omaieteroveceueanisiolepercnezehopeie ouiers ter exenshelione ee canekes 369
Acknowliedements vis:5:5 cies o's wie ao ouels ose wens sioucuehic: sbelo cuecsystoucue! s:spaoncyeie © ove eis 369
insli crap Webb of =k WOn ll 2f-\e erin Sn merrrrco o.oo Oo 1d Gide Did HO OO oS UO Udo doco S72
Addendum... 32). «. SCE Enea He iGO tic COLE OC ON bec Holy CUES U6 codicil Gc 383
385

Appendix MabwWwesreecre eoeoeoe oe eoeoeovnee eee eoeoGeooeeF2F FFF FOOL HOB SCHOLL OC LEO

LIST OF FIGURES

Figure Page
1. Map of the central Pacific Ocean. facing 1

2. Map of Johnston Atoll; adapted from Wennekens (1969) and
Ashmore (1973). 4

3. Looking northeast over Johnston and Sand Islands from 3,000
feet, 12 November 1935 (U.S. Nat. Archives, R.G. 80
photograph). 5

4. The northeast end of Johnston Island as it appeared from
the south in 1923 (B.P. Bishop Museum photograph by R.S.
Palmer). 6

5. Johnston Island, seen from the north at 3,500 feet,
10 October 1939 (U.S. Nat. Archives, R.G. 80 photograph). 6

6. Looking north over Johnston Island, 9 April 1942 (U.S.
Nat. Archives, R.G. 80 photograph). 8

7. Profile map of Johnston Island, Johnston Atoll, 1923,
1942, and 1964; adapted from Ashmore (1973) and USNOO
Chart No. 5356, 20 May 1963 and 23 December 1967. 8

8. Sand Island from the north, probably in 1939 (U.S. Nat.
Archives, R.G. 80 photograph). )

9. Sand Island from the southeast, 18 March 1940; buildings
later covered the entire island, including the northeast
peninsula (U.S. Nat. Archives, R.G. 80 photograph). 9

10. Sand Island as it appeared from the southeast about
1962, with the U.S. Coast Guard LORAN Station completed

(official U.S. Coast Guard photograph). 10
11. Sand Island as it appeared from the west, circa 1965,
“: after 1964 construction had added a few acres to the south

Side of the man-made portion of the island (official U.S.

Coast Guard photograph). 10

12. Johnston Atoll reef profiles; adapted from Emery (1956)
and Wennekens (1969). A: outer reef; B: isolated
patch reefs. 14

13. Sequence of sea level changes, Johnston Atoll (Ashmore,
IBS /E))ic 14

14. Mean monthly air and sea temperatures, Johnston Atoll,
1931-1972; air: (solid line), daily maximum and minimum
(dashes), extremes (dots); sea: means (barred). Adapted

from Seckel (1962) and U.S. Department of Commerce (1972). 16

VL

Figure
5%

16.

Lye

18.

iL)

20.

21.

Magee

2's

24.

Zs

26.

27.

28.

Percent frequency of wind from E and ENE (open bar) and
ESE and NE (stippled bar), Johnston Atoll.

Mean monthly wind speed, Johnston Atoll, 1931-1972;
means (solid line), upper extremes (dots). Adapted from
U.S. Department of Commerce (1972).

Johnston Atoll mean U components; adapted from Joint
Task Force Seven (1959).

Johnston Atoll mean V components; adapted from Joint
Task Force Seven (1959).

Mean monthly precipitation, Johnston Atoll, 1931-1972;
means (solid line), extremes (dots). Adapted from U.S.
Department of Commerce (1972).

Mean monthly relative humidity, Johnston Atoll, 1931-
1972; 0100 hours (dots), 0700 hours (dashes), 1300
hours (solid line), 1900 hours (barred). Adapted from
U.S. Department of Commerce (1972).

Mean monthly sky cover, sunrise to sunset, Johnston
Atoll, 1931-1972. Sky cover is expressed in a range
of 0 to 3 as being clear, 4 to 7 partly cloudy, and 8
to 10 cloudy. Adapted from U.S. Department of
Commerce (1972).

Mean monthly number of days of clear skies (solid line),
partly cloudy skies (dots), cloudy skies (barred), and
precipitation of 0.01 inch or more (dashes), Johnston
Atoll, 1931-1972. Adapted from U.S. Department of
Commerce (1972).

Pacific Ocean currents (King, 1967).

Seasonal and tidal flow regimes, Johnston Atoll
(Wennekens, 1969).

Johnston Atoll inferred lagoon circulation, January-
February 1965 (Kopenski and Wennekens, 1966).

Johnston Atoll inferred lagoon circulation and turbid
outflow (westerly flow), July-August 1965 (Kopenski
and Wennekens, 1966).

Johnston Atoll inferred circulation and turbid outflow
(easterly flow), July-August 1965 (Kopenski and Wenne-
kens, 1966).

Tidal wave records, Johnston Atoll (Wennekens, 1969).

Aly!

IL)

ILE)

20

Zu

(B72

23

25

27,

28

29

29

31

Figure

29%.

30.

31.

B2:.

318)6

34.

S516

36.

2 ae

38.

39.

40.

Ae

42.

43.

Algae collection stations, Johnston Atoll, 1965-1966
(Buggeln and Tsuda, 1969).

Distribution of the five most imvortant plant species
on the original portion of Sand Island, Johnston Atoll,
July-August 1963.

Distribution of dominant plant species on the original
portion of Sand Island, Johnston Atoll, May 1967.

Johnston Atoll showing invertebrate and fish collection
stations, dredged areas, and the extent of silt laden
water (Brock, et al., 1966).

Distribution of positive samples of Orntthodoros
capensts for 1965, Sand Island, Johnston Atoll.
Inserts show bi-monthly infestation rates for the
three major areas.

Bi-monthly tick infestation rates (stipples) compared
with breeding bird population fluctuations (bars),
Sand Island, Johnston Atoll, 1965.

Areas used by Sooty Terns and Wedge-tailed Shearwaters
on Sand Island, Johnston Atoll, 1965.

Fish collection stations, Johnston Atoll, 1963-1964
CBrockte hala, 1965)".

Breeding cycles of seabirds at Johnston Atoll; stippled
area represents eggs, barred area young, and black dots
non-breeding birds.

Monthly cumulative bird populations, Johnston Atoll,
1963-1969.

Monthly mean shorebird populations for Johnston Atoll,
1963-1969; Golden Plover (solid line), Ruddy Turnstone
(dots), Wandering Tattler (dashes).

Diurnal bird populations at sea 175 miles southwest of
Johnston Atoll, 1963-1967.

Monthly diurnal population fluctuations within the at-
sea grid by species group, 1963-1967.

Total nocturnal bird sightings at sea 175 miles southwest

of Johnston Atoll, 1963-1967.

Nesting areas of ground nesting birds (except Sooty Terns)

on the original portion of Sand Island, Johnston Atoll,
L963).

VIL

66

66

67

82

82

83

Oi.

iyzik

124

126

3 2

139

133

138

DELL

Figure

44,

45.

46.

47.

48.

49,

50.

See

D Lie

SS}e

54.

Dis

Nesting areas of birds which normally nest in low
vegetation on the original portion of Sand Island,
Johnston Atoll, 1963.

Bulwer's Petrel chick, about 2 months old, in front of
rock crevice nest site along causeway, Sand Island,
Johnston Atoll, 25 August 1966 (POBSP photo by P.C.
Shelton).

Concrete slabs on east shore, at base of northeast
peninsula, Sand Island, Johnston Atoll, 8 May 1967.
Christmas Shearwaters nested regularly under the large
slabs at left, and Bulwer's Petrels attempted to nest
at least once under the slabs in center of photo.
Brown Noddies nested in the edge of the Sesuviwm
growing just above the storm beach (POBSP photo by
P.¢..-Shel ton).

Distribution of Bulwer's Petrel nest sites, Sand
Island, Johnston Atoll, 1964-1969.

Means and extremes of semimonthly estimates of Bulwer's
Petrel numbers, Sand Island, Johnston Atoll, 1964-1969.

Diurnal occurrence of Bulwer's Petrels at sea 175
miles southwest of Johnston Atoll, 1963-1967.

Distribution of Wedge-tailed Shearwater nest sites on
Sand Island, Johnston Atoll, 1963-1969. Numbers indi-
cate chicks banded in 1968.

Wedge-tailed Shearwater chick in front of burrow under
Lepturus repens clumps, Sand Island, Johnston Atoll,
2 September 1966 (POBSP photo by P.C. Shelton).

Wedge-tailed Shearwaters among Tribulus, Boerhavia, and
Lepturus on southwest Sand Island, Johnston Atoll, mid-
morning July 1963 (POBSP photo by A.B. Amerson, Jr.).

Means and extremes of semimonthly estimates of Wedge-
tailed Shearwater numbers, Sand Island, Johnston Atoll,
1963-1969.

Diurnal occurrence of Wedge-tailed Shearwaters (light
phase) at sea 175 miles southwest of Johnston Atoll,
1963-1967.

Christmas Shearwater chick, 2 and 1/2 months old, in
shallow trench burrow under Lepturus repens clump on
east shore of Sand Island, Johnston Atoll, 29 August
1966 (POBSP photo by P.C. Shelton).

138

ALS)

IES)//

158

159

164

167

168

168

169

176

180

Figure

56.

5

58.

59.

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

Christmas Shearwater chick, 2 months old, and adults in
bunker on east shore, Sand Island, Johnston Atoll,
1 September 1966 (POBSP photo by P.C. Shelton).

Distribution of Christmas Shearwater nest sites, Sand
Island, Johnston Atoll, 1963-1969.

Means and extremes of semimonthly estimates of Christmas
Shearwater numbers, Sand Island, Johnston Atoll, 1964-
1969.

Distribution of Red-tailed Tropicbird nests on Sand
Island, Johnston Atoll, 1967.

Distribution of Red-tailed Tropicbird nests on the
original portion of Sand Island, Johnston Atoll, 1969.

Distribution of Red-tailed Tropicbird nests on the man-
made portion of Sand Island, Johnston Atoll, 1969.

Red-tailed Tropicbird on nest under Tournefortta bush
northeast of transmitter building, Sand Island,
Johnston Atoll, 25 February 1967 (POBSP photo by P.C.
Shelton).

Looking east from top of power building, Sand Island,
Johnston Atoll, 5 September 1969. Several Red-tailed
Tropicbirds nested under these Pluchea odorata bushes
(POBSP photo by P.C. Shelton).

Means of semimonthly estimates of Red-tailed Tropicbird
numbers, Sand Island, Johnston Atoll; 1965-1966 (solid
line) compared with 1967-1969 (dots).

Means of semimonthly estimates of Red-tailed Tropicbird
numbers, Johnston Island, Johnston Atoll; 1965-1966
(solid line) compared with 1967-1969 (dots).

Diurnal occurrence of Red-tailed Tropicbirds at sea 175
miles southwest of Johnston Atoll, 1963-1967.

Blue-faced Booby adult and chick, Johnston Atoll, July
1923 (B.P. Bishop Museum photo).

A roosting club of Blue-faced Boobies on Sand Island,

Johnston Atoll, 2 June 1941 (U.S. National Archives, R.G.

photo).

Means and extremes of semimonthly estimates of Blue-
faced Booby numbers, Sand Island, Johnston Atoll, 1963-
ISTO)

80

180

181

182

Oa

192

OZ

IU )3'

193

196

197

204

208

208

209

Wo

UD

74.

Zoe

76.

ila.

13\

Ue

80.

Sie.

Distribution of Brown Booby nests on Sand Island,
Johnston Atoll, 1969.

Brown Boobies in early stages of courtship and nest
building, east hill of Sand Island, Johnston Atoll,

25 February 1969. Vegetation is mostly Boerhavia with
scattered clumps of Trtbulus (POBSP photo by P.C.
Shelton).

Brown Booby adults and 31-day-old chick, west shore of
northeast peninsula, Sand Island, Johnston Atoll,

9 July 1966. Brown Noddies roost and nest here also;
Sooty Terns are in the background (POBSP photo by P.C.
Shelton).

Means and extremes of semimonthly estimates of Brown
Booby numbers, Sand Island, Johnston Atoll, 1963-1969.

Distribution of Red-footed Booby nests, Sand Island,
Johnston Atoll, 1963-1967.

Means and extremes of semimonthly estimates of Red-
footed Booby numbers, Sand Island, Johnston Atoll, 1964-
1969.

Four Red-footed Boobies nesting on artificial platforms
(framework of an old boat) on the southeast beach of
Sand Island, Johnston Atoll, 5 May 1969 (POBSP photo

by 2.¢. Shelton).

Red-footed Booby nest placed on ground on crest of hill
east of transmitter building, Sand Island, Johnston
Atoll, 4 May 1969. Subadult Blue-faced Booby roosting
at lower right; Sooty Tern adults and chicks and Brown
Noddy adult nearby (POBSP photo by P.C. Shelton).

Diurnal occurrence of Red-footed Boobies at sea 175
miles southwest of Johnston Atoll, 1963-1967.

Great Frigatebird nesting areas, Sand Island, Johnston
Atoll, 1963-1969.

Great Frigatebirds nesting on ground on crest of hill

east of transmitter building, Sand Island, Johnston Atoll,

May or June 1964. Sooty Tern adults and chicks surround
the colony (POBSP photo by A.B. Amerson, Jr.).

Great Frigatebird colony on south shore, Sand Island,
Johnston Atoll, 26 February 1969 (POBSP photo by P.C.
Shelton).

21D

215

216

23

239

236

236

243

246

247

247

Figure

82.

83.

84.

33.

86.

87.

88.

89.

90.

91.

92).

93.

94.

95.

Great Frigatebird roosting areas, Sand Island,
Johnston Atoll, 1963-1969.

Means and extremes of semimonthly estimates of Great
Frigatebird numbers, Sand Island, Johnston Atoll,
1963-1969.

Diurnal occurrence of Great Frigatebirds at sea 175
miles southwest of Johnston Atoll, 1963-1967.

Moving averages of semimonthly estimates of Golden
Plovers, Sand and Johnston Islands combined, 1963-
1969.

Distribution of Golden Plovers on Sand Island, Johnston
Atoll, 26 December 1965.

Moving averages of semimonthly estimates of Wandering
Tattlers, Sand and Johnston Islands combined, 1963-
1969.

Moving averages of semimonthly estimates of Ruddy Turn-
stones, Sand Island, Johnston Atoll, 1963-1969.

Dowitcher species photographed on Sand Island, Johnston
Atoll, 8 September 1969 (POBSP photo by P.C. Shelton).

Moving averages of semimonthly estimates of Sanderlings,
Sand and Johnston Islands, 1963-1969.

Distribution of Gray-backed Tern nest sites, Sand
Island, Johnston Atoll, 1966.

Distribution of Gray-backed Tern nest sites, Sand
Island, Johnston Atoll, 1967.

Gray-backed Tern on nest (center foreground) among Sooty
Terns in Ipomoea,and Lepturus on south shore of Sand
Island, Johnston Atoll, 21 April 1969 (POBSP photo by
P.C. Shelton).

Gray-backed Tern nesting area on southwest islet, Sand
Island, Johnston Atoll, 16 April 1969. Two adults, one
chick, and four eggs are shown (POBSP photo by P. C.
Shelton).

Means and extremes of semimonthly estimates of Gray-backed
Tern numbers, Sand Island, Johnston Atoll, 1964-1969.

Peay.

259

269

2EE

280

283

286

288

298

298

300

300

302

beg

LTT

Figure
96.

Oke

98.

99.

100.

Oa:

102.

103.

104.

105.

106.

107.

HOS:

Sooty Terns (tiny dots) utilize almost all available
ground nest sites on the original portion of Sand
Island, Johnston Atoll, May 1964 (mosaic by A.B.
Amerson, Jr., from POBSP photographs taken at the
620-foot tower level by R.W. Merrill).

Sooty Tern nesting areas, Sand Island, Johnston Atoll,
LOGH

Nesting Sooty Terns cover virtually the entire south-
west portion of Sand Island, Johnston Atoll, 15 April
1969. Most of the Wedge-tailed Shearwater burrows
are under the Lepturus in this area. The southwest
islet and old dock are visible at upper left (POBSP
photo by P.C. Shelton).

Nesting Sooty Terns northeast of the transmitter
building and on the northeast peninsula, Sand Island,
Johnston Atoll, 30 March 1967 (POBSP photo by P.C.
Shelton).

Means and extremes of semimonthly estimates of adult
Sooty Tern numbers, Sand Island, Johnston Atoll, 1963-
1969.

Means of semimonthly estimates of Sooty Tern eggs and
youtig, Sand Island, Johnston Atoll, 1963-1969.

Interisland movement of Sooty Terns involving the Hawaiian

Islands, Johnston Atoll, and Wake Atoll (Gould, 1974b).

Diurnal occurrence of Sooty Terns at sea 175 miles
southwest of Johnston Atoll, 1963-1967.

Orange-streamered Sooty Terns at sea 175 miles southwest

of Johnston Atoll, 1963-1967.

Nocturnal occurrence of Sooty Terns at sea 175 miles
southwest of Johnston Atoll, 1963-1967.

Brown Noddy nest distribution, Sand Island, Johnston
Atoll, April 1967.

Brown Noddies and Sooty Terns on the southwest portion
of Sand Island, Johnston Atoll, July 1923. Johnston
Island is visible in the background (B.P. Bishop Museum
photo by Chapman Grant).

Brown Noddies nesting and roosting on the southwest
islet, Sand Island, Johnston Atoll, 9 May 1967 (POBSP
photo by P.C. Shelton).

310

310

Sly

314

Sily/

320

Syl,

S21

325

328

328

LTLL

Blgure Page

109. Means and extremes of semimonthly estimates of Brown
Noddy numbers, Sand Island, Johnston Atoll, 1963-
1969. 330

110. Black Noddy nesting and roosting areas, Sand Island,
Johnston Atoll, 1963-1967. 338

111. Black Noddy nest in Amaranthus plants south of the
transmitter building, Sand Island, Johnston Atoll,
7 May 1967. Sooty Tern adults and chicks nearby (POBSP
photo by P.C. Shelton). 339

112. Black Noddy nest with 12-day-old chick in Amaranthus
plant south of the transmitter building, Sand Island,
Johnston Atoll, 4 June 1969 (POEBSP photo by P.C.
Shelton). 339

113. Means and extremes of semimonthly estimates of Black
Noddy numbers, Sand Island, Johnston Atoll, 1964-1969. 340

114. White Terns roosting on undercut beach rock on shore of
Johnston Island, Johnston Atoll, July 1923 (B.P. Bishop
Museum photo). 347

115. White Tern chick hatched on framework of old radio
telescope, Johnston Island, Johnston Atoll, 19 June
1969 (POBSP photo by P.C. Shelton). 347

116. Means and extremes of semimonthly estimates of diurnal
White Tern numbers, Johnston Atoll, 1963-1969. 349

117. Diurnal occurrence of White Terns at sea 175 miles
southwest of Johnston Atoll, 1963-1967. 349

XLV

Table

106.

1s

2

oye

14.
NS)e

16.

7
ite}
19.

20.

LIST OF TABLES

Ownership and control of Johnston Atoll (adapted from
Bauer, 1965).

Distribution of benthic marine algae at Johnston Atoll.
Vascular plants known from Johnston Atoll.
Cnidaria (Coelenterata) from Johnston Atoll.

Distribution and abundance of corals at Johnston Atoll,
1964-1965.

Distribution and abundance of Mollusca from Johnston Atoll.

Annelida from Johnston Atoll.
Marine Arthropoda from Johnston Atoll.

Insects recorded from Johnston Atoll; adapted from
Chilson (1953).

Berlese sample data--Ornithodoros capensis infestation
rates for Sand Island, Johnston Atoll, 1965.

Berlese sample data--bimonthly Ornithodoros capensis
infestation rates for Sand Island, Johnston Atoll, 1965.

Orntthodoros capensts infestation rates from Berlese
samples of hosts" nests.

Mallophaga collected at Sand Island, Johnston Atoll by
the POBSP; adapted from Amerson and Emerson (1971).

Echinodermata from Johnston Atoll.
Inshore fishes recorded from Johnston Atoll.

Toxicity of fish species tested for ciguatera at Johnston
Atolls

Birds from Johnston Atoll.
Status of birds on Johnston Atoll.

Peak breeding period of Johnston Atoll birds.

Monthly occurrence of non-resident birds at Johnston Atoll.

Page

41
49
oy

68

69
Hak
13

74

77

80

84

85

87
87

93

103
114
116
120

123

Table

Jade

22

23).

24.

Ze

26.

27's

28.

Zu

30.

Sle

32).

B3'.

Bylye

S5)F

36.

57 5

Occurrence of birds at sea in the grid 175 miles southwest
of Johnston Atoll.

Status and area of derivation of birds observed at sea
near Johnston Atoll.

Sooty Tern mortality from guywire strikes, Sand Island,
March-July 1965.

Birds banded at Johnston Atoll, 1963 to 1973.
Yearly band return totals for Johnston Atoll.

Interisland movement of banded birds involving Johnston
Atoll.

Bird specimens and summary of new records from Johnston
Atoll.

Productivity of Bulwer's Petrels on Sand Island, Johnston
Atoll, 1964-1969.

Extreme dates of significant events in the breeding cycle
of Bulwer's Petrels on Sand Island, 1964-1969.

Banding and recaptures of Bulwer's Petrels, Sand Island,
Johnston Atoll, 1964-1969.

Productivity of Wedge-tailed Shearwaters on Sand Island,
Johnston Atoll, 1963-1968.

Extreme dates of significant events in the breeding cycle
of Wedge-tailed Shearwaters, Sand Island, Johnston Atoll,
1963-1969.

Banding and recaptures of Wedge-tailed Shearwaters, Sand
Island, Johnston Atoll, 1963-1969.

Productivity of Christmas Shearwaters on Sand Island,
Johnston Atoll, 1963-1969.

Extreme dates of significant events in the breeding cycle
of Christmas Shearwaters, Sand Island, Johnston Atoll,
1963-1969.

Banding and recaptures of Christmas Shearwaters, Sand
Island, Johnston Atoll, 1963-1969.

Distribution and nesting success of Red-tailed Tropicbirds
by habitat type, Sand and Johnston Islands, 1967-1969.

128

130

140

142

143

146

150

161

162

165

Aby/ak

173

17/7)

183

185

187

195

LVL

Table

38.

Bo)

40.

41.

42.

43.

44,

45.

46.

47.

48.

49.

50.

Sh

Di2s.

Do

Productivity of Red-tailed Tropicbirds on Johnston Atoll,
1964-1969.

Incubation and fledging period of Red-tailed Tropicbird
eggs and chicks, Sand Island, Johnston Atoll, 1967-1969.

Banding and recapture of Red-tailed Tropicbirds, Sand and
Johnston Islands, 1963-1973.

Sightings of White-tailed Tropicbirds on Johnston Atoll.

Semimonthly population estimates of Blue-faced Boobies on
Sand Island, Johnston Atoll, 1963-1969.

Blue-faced Boobies banded on Sand Island, Johnston Atoll,
by the POBSP.

Productivity of Brown Boobies, Sand Island, Johnston
Atoll, 1963 through 1969.

Success rates of different clutch sizes of Brown Boobies,
Sand Island, Johnston Atoll, 1964 through 1969.

Percentages of Brown Booby chicks resulting from one, two,
and three egg clutches, Sand Island, Johnston Atoll, 1964
through 1969.

Renesting attempts, incidences of two chicks reared in

one nest, and number of chicks raised by foster parents or

by hand, Sand Island, Johnston Atoll, 1964 through 1969.

Percent of Brown Booby chicks produced and success rates
of nests by thirds of breeding seasons, Sand Island,
Johnston Atoll, 1964 through 1969.

Nesting success of known-age Brown Boobies, Sand Island,
Johnston Atoll.

Extreme dates of significant events in the Brown Booby
breeding cycle, Sand Island, Johnston Atoll, 1964-1969.

Number of Brown Booby nests established by semimonthly
periods, Sand Island, Johnston Atoll, 1964-1969.

Banding and recaptures of Brown Boobies, Sand Island,
Johnston Atoll, 1963-1969.

Productivity of Red-footed Boobies, Sand Island, Johnston
Atoll, 1963-1969.

Page

199

201

203

206

211

212

218

AAs)

220

222.

223

224

225

226

228

239

Table

54.

55.

56.

Sis

58.

59%

60.

61.

62.

63.

64.

65).

66.

Oi.

68.

69.

Known age Red-footed Booby nesting, Sand Island, Johnston
Atoll, 1963-1969.

Extreme dates of significant events in the Red-footed
Booby breeding cycles, Sand Island, Johnston Atoll, 1963-
16S:

Plumage ratios of Red-footed Boobies, Sand Island, Johnston
AeEoll, 967 .

Banding and recaptures of Red-footed Boobies, Sand Island,
Johnston Atoll, 1963-1973.

Productivity of Great Frigatebirds, Sand Island, Johnston
Atoll, 1963-1967.

Fate of young Great Frigatebirds banded on Sand Island,
Johnston Atoll, 1963-1967.

Extreme dates of significant events in the Great Frigate-
bird breeding cycles, Sand Island, Johnston Atoll, 1963-
1969).

Plumage ratios of Great Frigatebirds, Sand Island, Johnston
Atoll.

Observations of Pintails on Johnston Atoll.

Observations of Bristle-thighed Curlews on Sand Island,
Johnston Atoll, 1963-1969.

Numbers and percentages of Ruddy Turnstones sighted on
Johnston Atoll that were color-marked on St. George Island,
Brpilofs!, Kald 1966.

Observations of Pectoral Sandpipers on Sand Island, Johnston
Atoll, 1963-1968.

Extreme dates of significant events in the Gray-backed Tern
breeding cycles, Sand Island, Johnston Atoll, 1963-1969.

Mark-release-recapture calculations of Sooty Tern breeding
populations, Sand Island, Johnston Atoll, based on February-
April 1967 recaptures.

Extreme dates of significant events in the breeding cycle
of Sooty Terns, Sand Island, Johnston Atoll, 1963-1969.

Orange-streamered Sooty Tern sightings within a 50-mile-
radius of Johnston Atoll and in the at-sea grid 175 miles
southwest of Johnston Atoll, 1964-1967.

238

239

242

253

254

255

285

290

304

313

315

LVELL

Table

70.

Udke

U0

73h

74.

126

Ue

THEE

78.

79.
80.
Silks

O2e

83.

Nesting material used by Brown Noddies, Sand Island,
Johnston Atoll; observations by B.A. Harrington, about
1 May 1968.

Extreme dates of significant events in the Brown Noddy
breeding cycles, Sand Island, Johnston Atoll, 1964-1969.

Daily egg laying, hatching, and losses for Brown Noddies,
Sand Island, Johnston Atoll, July-September, 1964-1969.

Known-age Brown Noddy recaptures taken from eggs or chicks,
Sand Island, Johnston Atoll, 1967.

Primary molt scores of Brown Noddies, Sand Island, Johnston
Atoll.

Nesting success of Black Noddies on Sand Island, Johnston
Atoll, 1963-1969.

Calculations of numbers of Black Noddies using Sand Island,
Johnston Atoll, 1967.

Primary molt scores of Black Noddies caught roosting on
Sand Island, Johnston Atoll, February-April LOGT:

Extreme dates of significant events in the Black Noddy
breeding cycles, Sand Island, Johnston Atoll, 1964-1969.

Observations of Short-eared Owls on Johnston Atoll.
Records of Japanese White-eyes on Johnston Atoll.
Distribution and status of mammals at Johnston Atoll.

Breeding condition of house mice autopsied, October 1964-
March 1965.

Evidence of breeding condition in house mice live-trapped
and released, October 1966 and February 1967.

334

$35

335

336

341

342

344

344

Boz

354

356

65/7

BD

LIST OF APPENDIX TABLES

Appendix Table

ils

De

POBSP personnel participating in bird studies on
Johnston Atoll, July 1963-September 1969.

Annotated list of vascular plant species recorded
from Akau Island, Johnston Atoll.

Annotated list of vascular plant species recorded
from Hikina Island, Johnston Atoll.

Annotated list of vascular plant species recorded
from Johnston Island, Johnston Atoll.

Annotated list of vascular plant species recorded
from the man-made portion of Sand Island, Johnston
Atoll.

Annotated list of vascular plant species recorded
from the original portion of Sand Island, Johnston
Atoll.

Bird specimens collected on Johnston Atoll.

Pre-POBSP observations of Sooty Terns on Johnston
Atoll.

387

393

396

418

429

443

473

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THE NATURAL HISTORY OF JOHNSTON ATOLL,
CENTRAL PACIFIC OCEAN

by A. Binion Amerson, Jr., and Philip C. Shelton

INTRODUCTION

Johnston Atoll consists of two highly modified natural islands and
two completely man-made islands totaling about a square mile in surface
area lying on a 14 by 7 mile coral reef platform in the tropical Pacific
Ocean at 16°45'N, 169°31'W (Figure 1). The nearest land is French
Frigate Shoals in the northwestern Hawaiian Islands, about 450 nautical
miles north-northwest; Honolulu, Hawaii is 717 nm northwest; the Marshall
Islands lie almost 1,200 nm southwest, Howland Island is 1,050 nm south-
southwest, and Kingman Reef in the Line Islands is about 850 nm southeast.

Although the atoll was discovered in 1/796 and claimed by the United
States in 1858, it was uninhabited except for brief visits by guano miners,
adventurers, government scientists and military personnel until 1936,
when the U.S. Navy began developing first a seaplane base and later an
airstrip and refueling facilities which served as an important link from
the Hawaiian Islands to the western Pacific during World War II.

After the war, operations diminished until the late 1950's when
the atoll was used for atmospheric nuclear testing. Although the last
tests were executed in 1962, facilities have been maintained "...for re-
suming nuclear testing in the Pacific Ocean should the Test Ban Treaty
of 1963 be abrogated by a foreign world power" (Bauer, 1965: 1).

Most recently the atoll has become the storage site for obsolete
chemical warfare agents, including nerve gas formerly stored on Okinawa,
and herbicides used in southeast Asia.

The atoll was made a federal bird refuge by executive order of
President Calvin Coolidge in 1926. This order remains in effect, although

1/ Paper Number 80, Pacific Ocean Biological Survey Program, Smithsonian
Institution, Washington, D.C.

2/ Present address: Department of Systematics and Ecology, The University
of Kansas, Lawrence, Kansas 66045

3/ Present address: Department of Biology, Clinch Valley College, Wise,
Virginia 24293

subsequent executive orders have given primary jurisdiction over the
atoll to military agencies, first to the Navy, later to the U.S. Air
Force. Presently a Joint Task Force has jurisdiction over all the
atoll except for Sand Island, which is the site of a U.S. Coast Guard
LORAN station.

The atoll is an unincorporated territory (=possession) of the
United States. This is distinguished from an incorporated territory
(=territory) in that it is "...territory to which the constitution [of
the United States] has not been expressly and fully extended" (U.S.

Department of State, 1965: 9).

Although the oceanic region in which Johnston Atoll lies is rela-
tively unproductive, life is abundant on the atoll itself. At least
half a million seabirds use the atoll for roosting and nesting. Mest
of these are of one species, Sooty Tern (Sterna fuscata), but the
total known avifauna is 56 species, including 12 regular breeders and
half-a-dozen regular migrants. Other terrestrial forms are scarce, and
are limited mainly to introduced species, including five species each of
mammals and reptiles, all of which were introduced, at least 87 species
of insects, many of which were introduced, and well over one hundred
species of plants, only three of which are native.

The inshore marine biota is richer. To date 194 species of
inshore fishes have been recorded, sea turtles visit the atoll regularly
in small numbers, porpoises are occasionally seen outside the lagoon,
and in 1968 and 1969 Hawaiian Monk Seals visited the islands for the
first time so far as is known. Tropical marine invertebrates are
abundant in the lagoon.

Military occupation and construction during the past 40 years
have so drastically altered the physiography of the islands that little
of the original habitat type remains. Breeding populations of three of
the 15 seabird species known to have bred on the islands were eliminated,
while a few others were able to increase through adaptation to man-
made changes. For most bird species the alteration of habitat and
continued disturbance, sometimes through planned efforts to move birds
to less critical areas, sometimes through ignorant or malicious perse-
cution, has resulted in severely decreased roosting and nesting space
and consequently in reduced populations.

As part of its study of populations, breeding biology, and
movements of seabirds of the central Pacific Ocean, the Pacific Ocean
Biological Survey Program (hereafter referred to as the POBSP) of the
Smithsonian Institution, Washington, D.C., maintained one or more
biologists on Sand Island, Johnston Atoll almost continuously from
7 July 1963 until 9 September 1969 (Appendix Table 1). Former POBSP
employees have visited the atoll twice since 1969. Brian A. Harrington
studied Sooty Terns from 18 March through 3 June 1971; A. Binion Amerson,
Jr. visited Johnston 6-12 November 1973 for the Office of International
and Environmental Programs, Smithsonian Institution, to update ecological
data.

Although primary emphasis was on the 12 species of breeding
seabirds, POBSP studies also included arthropods, reptiles, mammals,
and terrestrial plants. This report is an attempt to summarize the
major results of these studies and to compile a comprehensive summary
of other scientific efforts made on Johnston Atoll. The major thrust
of the report is island-oriented rather than species-oriented, and
discussions presented are primarily descriptive rather than analytical.
Hopefully the data presented will be useful in developing a detailed
analysis of the central Pacific ecosystem, and for comparison of
population and breeding phenomena of the seabirds and other groups
with similar data from other islands where the same species occur.

This report is based primarily on the field notes and semi-

monthly reports made by the 29 POBSP personnel (Appendix Table 1) who
maintained the Sand Island station. These voluminous data are stored

in the National Museum of Natural History, Washington, D.C. This final
report is a combination of a manuscript by Shelton and a technical
report by Amerson (1973). Amerson's ecological baseline report was

used as part of a 10 May 1974 U.S. Air Force Environmental Impact
Statement concerning proposed disposal of chemical herbicide on Johnston
“Atoll.

PHYSICAL ENVIRONMENT

The physical characteristics of Johnston Atoll and its surrounding
waters are relatively well studied. The following descriptions are based
primarily on accounts published by government agencies or by scientists
under government contract.

Johnston Atoll lies between the latitudes of 16°40'26" and
16°47'25" North and the longitudes of 169°24'15" and 169°33'58" West
(USNOO Chart 5356).

Islands

The two natural islands, Johnston (16°45'N x 169°32'W) and Sand
(16°45'N x 169°30'W), originally 46 and 10 acres in extent, lie a mile
and a half apart on the southern margin of the lagoon, which is the
shallow portion of the reef platform (Figs. 2 and 3). Summit Peak,
near the northwest end of Johnston Island, originally stood 48 feet
above sea level (Christophersen, 1931: 3) (Figs. 4 and 5). The highest
point on Sand Island was and remains about 15 feet above sea level.
Both islands were formed of beachrock--sand and gravel cemented by
calcium carbonate--and loose sand, presumably wind and wave transported
material (Emery, 1956, and Ashmore, 1973).

From 1939 through 1942 construction for military operations
leveled Johnston Island and enlarged it to 211 acres (Fig. 6), and
built another islet about the size of Sand Island and connected it to
Sand Island by a narrow causeway. These additions were built mostly of
coral material dredged from the lagoon in deepening and lengthening ship
channels and seaplane landing areas. By 1944 Johnston Island had a

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Figure 4.’ The northeast end of Johnston Island as it appeared from the south
A in July 1923 (B.P. Bishop Museum photo by R. S. Palmer).

mite Civics foe

om a Se wie

Figure 5. Johnston Island, seen from the north at 3,500 feet, 10 October 1939
(U.S. Nat. Archives, R.G. 80 photograph).

6,100 foot runway, and both islands were virtually covered with
buildings, roadways, and gun emplacements. None of the original
shoreline or vegetation of Johnston Island remained.

The airstrip on Johnston was again enlarged in 1951-52, and in
1963-64 additional construction with coral dredged from the lagoon
enlarged Johnston Island to 570 acres (Fig. 7), with a 9,000 foot runway,
added a few acres to the artificial portion of Sand Island, and built
two entirely new islands, Akau (North) at 16°45'52"N x 169"31'03"W and
Hikina (East) at 16°45'26"N x 169°29'19"W, of 24 and 17 acres respect-
ively, within the lagoon (Bauer, 1965).

The present surfaces of Johnston, Akau, and Hikina Islands, and
the artificial portion of Sand Island, consist of hard packed coral
material ranging from sand to cobble size. Buildings, antennas, roads,
and (on Johnston only) runways and taxiways dominate these islands.
Vegetation is restricted to sparse lawns, scattered bushes and trees,
and thinly scattered weedy species on areas that are not frequently
disturbed.

Although buildings formerly occupied most of the original portion
of Sand Island (Figs. 8 and 9), these were removed in the late 1950's.
A few concrete foundations and gun emplacements remain. The only new
structures added since then are the LORAN-C transmitter building and
the 625-foot transmitter tower. Ground wires radiate from the base of
the tower on or just below the surface, and guywires extend to concrete
pillars, most of them beyond the shores of the island in the lagoon
(Figs. 10 and 11). A roadway leads to the transmitter building from
the causeway connecting the two portions of the island. Except for
these structures, the surface of this original island is approximately
like that of both Johnston and Sand Islands prior to their disturbance.
The interior is covered with deep, loose coral sand, bound by roots of
the grass, Lepturus repens, and perforated by burrows of Wedge-tailed
Shearwaters. A beachrock layer is exposed around most of the perimeter
of the island.

Geology

Emery (1956) described the geology of Johnston Island based on
information from 56 borings for building foundations, ranging to 36
feet below mean low water, and from six wells drilled to depths ranging
from 86 to 191 feet below mean sea level. None of these holes reached
non-calcareous rock. Two layers of beachrock were encountered, one
above sea level, and presumably correlated with the one still exposed
on Sand Island but covered or destroyed on Johnston, and one at 4.3 to
6.4 feet below sea level. This lower layer was also found 8 feet below
sea level along bases of the parallel, linear algal reefs that extend
between Johnston and Sand Islands. These reefs were thought to result
from growth of coralline algae on the outcropping edges of submerged
beachrock.

Below the beachrock the deeper parts of the wells showed alter-
nating sand, loose coral, and sand and coral, with little certainty of

Figure 6. Looking north over Johnston Island, 9 April 1942 (U.S. Nat.
Archives, R.G. 80 photograph).

16°45'

————
0.5 Nautical Miles

169°33' 169232"

Figure 7, Profile map of Johnston Island, Johnston Atoll, 1923, 1942, and
1964; adapted from Ashmore (1973) and USNOO Chart No. 5356, 20
May 1963 and 23 December 1967.

Figure 8. Sand Island from the north, probably in 1939 (U.S. Nat. Archives,
R.G. 80 photograph).

Figure 9, Sand Island from the southeast, 18 March 1940; buildings later
covered the entire island, including the northeast peninsula
(U.S. Nat. Archives, R.G. 80 photograph).

10

Figure 10.

Figure 11.

Sand Island as it appeared from the southeast about 1962, with
the U.S. Coast Guard LORAN Station completed (official U.S.
Coast Guard photograph).

|

construction had added a few acres to the south side of the |
man-made portion of the island (official U.S. Coast Guard
photograph).

Sand Island as it appeared from the west, circa 1965, after 196

|
|

correlation from well to well. Mud of unknown origin occurred in
several wells.

Several occurrences of non-calcareous rock are known from the
islands. Several hundred pounds of pumice floated ashore in 1953,
presumably from volcanic activity on San Benedicto Island off Mexico
(Emery, 1956: 1516, and Richards, 1958). Maxwell S. Doty found a small
piece of rhyolite imbedded in beachrock on Sand Island. ‘This could

have been rafted on driftwood or could have arrived ‘as ship's ballast
(Emery, 1956: 1516). Pumice also floated ashore in the 1960's.

Although Johnston Atoll lies several hundred miles from the
mearest island, it is fairly closely associated with two submerged
mountain ranges. The nearest is the Mid-Pacific Mountains, also called
the Marcus-Necker Rise, which lie in an arc extending from near the
middle of the Hawaiian Ridge (Necker Island) first southwesterly, then
westerly to Wake Island, then northwesterly to Marcus Island. Barkley
(1962: 3) considered Johnston to be a part of this range, but Kroenke
and Woollard (1965: 365) and Ashmore (1973: 3) considered it to be the
northeastern extremity of Christmas Ridge (formerly the Line Island
Ridge). Johnston lies about 100 nm southeast of the crest of the Mid-
Pacific Mountains and is separated from it by water over 5 km deep.

It is more directly aligned with the long axis of the Line Islands, but
it is separated from the nearest of these (Kingman Reef) by a distance
of about 740 nm and by depths of over 5 km, although submerged mountains
rise to much shallower depths in several places between the two (Menard,
Peers ae, Pa 14 ard 1.12,. p. 19).

Both ranges were built up from the ocean floor by repeated volcanic
flows. Subsequent isostatic readjustment and erosion at and above sea
level then lowered the mountains. Corals and coralline algae grew in
shallow waters around these peaks, and under favorable conditions
maintained islands by growing upward as rapidly as the underlying
range sank. But not all peaks subsided at a time when coral growth
was favored, or they sank too rapidly for corals to maintain the surface.
Particularly in the Mid-Pacific Mountains, a large number of flat
topped "guyots," presumably drowned islands with their tops 1 to 2 km
below the surface, were discovered during World War II (Hess, 1946).

Menard (1964: 92) discussed the relative ages of the central
Pacific mountain groups. In summary, many of the volcanoes forming
the Mid-Pacific Mountains were thought to be high islands by middle
Cretaceous time (roughly 100 million years ago) on the basis of
fossils dredged from the summits of Hess, Cape Johnson and other
guyots a few hundred miles west of Johnston (Hamilton, 1956, and
Matthews, et al., 1974). Although vulcanism and island formation con-
tinued for some time in the region, the islands were eroded and the
region gradually submerged. Some of the islands probably still existed
into late Mesozoic-early Cenozoic time, but except for Wake none has
survived to the present in the central Pacific. The Line Islands
probably were formed in late Mesozoic or early Cenozoic time, 60 to
70 million years ago.

2

Details of the geological history of Johnston Atoll itself are
lacking. When it first appeared as an island, its original size,
rate of erosion and subsidence, and rate of coral growth are unknown.
No deep drilling or seismic studies which would reveal the depth of
the coral cap have been reported.

Gravity measurements by Kroenke and Woollard (1965) show Johnston
Atoll to be intermediate between the main Hawaiian Islands and the
Line Islands in Bouguer anomaly values, which may indicate that the
base of heavy volcanic material is more massive under Johnston than
under the Lines but less massive than under the Hawaiian Islands, where
heavy basalts stand far above sea level in the southeast and lie at
most a few hundred feet below sea level at Midway at the northwest end
of the chain (Ladd, Tracy, and Gross, 1967). Bouguer anomaly gradients
on Johnston were lower by about half than in the Hawaiian Chain,
which may mean that there are no volcanic necks or plugs involved in
the upper structure of Johnston in contrast to the Hawaiian Islands,
including the northwest chain.

Geomorphology

The atoll consists of a coral platform with over 50 square miles
of area under less than 100 feet of water. A marginal reef, exposed
only at lowest tides, extends for 9 miles along the northwest margin
of the platform. A broad shallow ridge, on which lie Johnston, Sand,
and Hikina Islands, extends from the west end of the marginal reef
eastward, then northward to enclose a lagoon approximately 7 by 2
miles in extent, with depths generally between 10 and 30 feet. The
remainder of the platform lies south and east of the lagoon and is
mostly deeper (see Fig. 2).

Emery (1956) studied the composition of the reef platform by
making visual estimates of the relative percentages of sand, coral,
and coralline algae on photographs of about 4 percent of the platform,
in which the bottom material could be identified by color to a depth
of about 10 fathoms. Surfaces of coralline algae dominated along the
main outer reef, and many patch reefs less than 4 fathoms deep through-
out the lagoon were topped with coralline algae (Fig. 12). The deep
eastern part of the platform showed no evidence of algal patch reefs.
Coral-dominated areas occurred throughout the lagoon and platform, and
coral made up the substrate of many algae-topped patch reefs. Between
patches of coral and coralline algae were strips of sand, including
dune-like structures near Johnston and Sand Islands. Sand dominated
the eastern half of the platform.

The Johnston Atoll platform differs from typical atolls in that
the main outer reef extends only about one-fourth of the way around
the perimeter. In addition, neither the well-defined northwestern
marginal reef nor the poorly developed southern reef is in a clearly
defined windward or leeward position; most atolls display better
developed reefs on the windward side.

Emery (1956) thought that either the windward portion of the reef
had been removed by erosion during a time of lowered sea levels c- that
the platform had tilted southeastward. He slightly favored the t.lting
hypothesis. E.H. Bryan, Jr. (ms.) noted that the coral beds on Sand
Island dipped 4 to 5 degrees southeastward, which would indicate pos-
sible tilting in that direction.

Ashmore (1973) studied the morphology of the atoll on the basis
of data collected mainly by a team of U.S. Naval Oceanographic Office
engineers and Naval personnel from the USS MAURY during 1964. These
data showed features that were not evident to Emery, and led Ashmore
to favor a sea level change to explain the present morphology of the
atoll, although he acknowledged the possibility of a combination of
sea level change and tilting.

The major feature Ashmore found that did not fit the tilting
hypothesis was that the platform was composed of two more or less
distinct levels, neither with measurable slope (Fig. 13). These were
separated by an uneven scarp, 500 to 1,000 yards wide, with an average
gradient of 1 to 2 percent, extending across the entire platform.

The higher level is approximately 25 to 30 feet below mean low
water and, including the lagoon, covers about 24 square miles. It is
irregular in outline with significant embayments south of Johnston
Island, at the eastern tip of the platform, and at a break in the
northernmost section of the outer reef. The most striking feature of
this level is the great number of sink-like holes, commonly 55 to 60
feet deep, and some exceeding 1/2 mile in length.

The lower level has depths of 55 to 65 feet below mean low water
and extends over an area of about 30 square miles. A depression at
least 90 feet deep and about 5 square miles in area lies in the extreme
eastern portion of this level. Otherwise the lower level is somewhat
dished, with a distinct rim extending to within 33 feet of mean low
water, along much of the southern and western edge. In contrast to the
upper level, the lower level has few knolls, no sinkholes, and no
patch reefs.

Ashmore suggested that these terraces were formed as sea levels
fell at the end of the Sangamon Interglacial, about 75,000 years ago,
or with rising sea levels at the end of the Wisconsin glaciation, 10
to 12 thousand years ago. The sinkholes in the upper level were
thought to have been formed by fresh water erosion and hence were
interpreted as strong evidence that this level was older than the lower,
which would indicate formation of the terraces during stable periods
within a general period of falling sea levels. But the upper level
appeared to Ashmore to be too well preserved to have withstood 65,000
years of subaerial erosion during low waters of the Wisconsin glacial
period, which would indicate more recent formation, during rising
sea levels. Present data are insufficient to resolve the problem.

13

14

®

90°%o CORALLIN
/NGINE E ALGAL RIDGE

SO %o CORAL

TALUS OF DEAD CORAL AND ALGAE

Figure 12, Johnston Atoll reef profiles; A: outer reef, B: isolated patch
reef, Adapted from Emery (1956) and Wennekens (1969).

CROSS SECTION-NOT TO SCAN |

|
|
|
|

ay may asec eesecacecere sanseenrsfar\cancsecccsecevecennensersse

{
j
| OUTER REEF SAND ISLAND
|

#\-Present sea level.

f-Assumed to predate G because of the depth of the solution holes.

|| %3-Assumed to predate both S&& due to the absence of Solution ho!es deeper than C.

Figure 13, Sequence of sea level changes, Johnston Atoll (Ashmore, 1973).

IS)

Climate

Weather records are available for the atoll from about 1931 to
the present, but standardized data have been recorded only since 1952
(Joint Task Force Seven, 1959, and U.S. Dept. of Commerce, 1972).
These records show a tropical marine climate with little variation
in temperature and wind speed, but great variability in rainfall.

There is no evidence that climatic conditions differ from one
part of the atoll to another. The islands are too low and small to
affect their own weather significantly. The only observed influence
of the islands on their own weather was the development of well-
organized thermal updrafts generated by solar heating of air next to
the surfaces of Johnston and Sand Islands on the rare days when winds
were less than 8 to 10 knots. Attention was drawn to this updraft
from Sand Island by the column of soaring Great Frigatebirds and
Sooty Terns that developed within it. Individual birds entered this
column near the west shore of the island and rose to near the cloud
bases at over 1,000 feet altitude in 15 to 30 minutes. During this
time they drifted a third to half a mile downwind from the island.
Above the birds there appeared to be a cumulus buildup at the top of
the column of air. There appeared to be a similar but larger cumulus
buildup downwind from Johnston Island at the same time, but fewer birds
used this updraft, probably because it was farther from the bird
colony on Sand Island. There was no indication that precipitation was
generated from this phenomenon.

In terms of winds, temperature, and precipitation, two broad
seasons can be distinguished on the atoll. The 4-month "winter"
including the months December through March is characterized by
slightly lower temperatures, more variable winds, and heavier pre-
cipitation than the 8-month "summer" extending from April through
November.

Temperature

The mean annual temperature is 79.3°F (26.3°C). Daily ranges
are usually only 7 or 8 degrees (F), and the daily maximum and minimum
temperatures vary only a few degrees throughout the year (Fig. 14).
The extremes range from a low of 62°F (16.5°C) (December 1964) to a
high of 89°F (31.5°C) (October 1968, July and November 1969), which is
lower than the daily range frequently encountered in continental areas.

The constancy of temperature results from the air masses passing
over the atoll having been modified by close contact with the ocean
for thousands of miles. Thus the air temperature is near that of the
water. Sea surface temperatures (Fig. 14) vary little from day to
day and change only slowly with the seasons.

Winds

Johnston lies within the belt of strong easteriy trade winds through-
out most of the year, but these reach a maximum in d2p*h, speed, and

16

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oy e——-e-——0~
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= 70°F
Lu
ke
60RF

J eur iM A M J J A 5 0 Nee

Figure 14. Mean monthly air and sea temperatures, Johnston Atoll,
1931-1972; air: (solid line), daily maximum and minimum
(dashes), extremes (dots); sea: means (barred). Adapted
from Seckel (1962) and U.S. Department of Commerce (1972).

steadiness during the summer. Only during the winter period, when the
world wind belts shift southward to their maximum extent, do trade winds
(NE through ESE) occur less than 80 percent of the time (Fig. 15).

During this period light variable winds and westerlies occur occasionally
associated with the passage of organized disturbances--easterly waves,
troughs aloft, and weak cold fronts--characteristic of more temperate
regions.

Mean annual surface wind speed is 15.1 miles per hour, and the
monthly means range only from 13.6 to 16.0 (Fig. 16). Monthly extremes,
excluding 1972, range from 35 mph in July to 49 mph in March and
November. Mean monthly extremes are 43 mph. Until 1972 there was no

Ee Beit (C. sesNieeT)

Figure 15.

MILES PER HOUR

Figure 16.

17,

100

60 ll
50
40
30
20
10
0
Deer a pile whee

J A S 0 N D

J

Percent frequency of wind from E and ENE (open bar) and ESE and
NE (stippled bar), Johnston Atoll. on

105 =
we SZ

NX é *

Mean monthly wind speed, Johnston Atoll, 1931-1972; means (solid
line), upper extremes (dots). Adapted from U.S. Department of
Commerce (1972).

18

record of a storm of hurricane force (winds of 64 knots or more)
having passed over Johnston. On 19 August 1972 hurricane Celeste,
spawned off the southern coast of Mexico some 3,000 nm to the east
nearly 2 weeks earlier, passed 25 miles northeast of Johnston. The
wind speed on the island hit an all time high of 104 mph, and did
minor damage on the atoll. Observations from weather satellites
indicate that tropical storms in the Johnston Atoll area, though
infrequent, may not be as unusual as was once supposed.

Aloft winds are described by Figures 17 and 18 in which east-
west or "U'' components and north-south or "V" components of wind
direction and velocity are plotted against altitude in feet and
barometric pressure in millibars. Since the air flow is predominantly
east-west at all altitudes, the "U" components show by far the highest
velocities.

The comparatively shallow trade winds are shown below the lowest
O isopleth of Figure 17. Above the trades, air currents in the upper
trophosphere (20,000 to 60,000 feet) are predominantly westerly
throughout the year, with a maximum of speed, depth, and steadiness
in March and April. These winds have a considerable north-south
component in December and January, with an especially strong northerly
component in February (Fig. 18).

Above the trophosphere (above the highest O isopleth on Figure
17), stratospheric flow is dominated by the Krakatoa easterlies, but
there is considerable seasonal variability. Winter is characterized
by relatively light winds generally with east predominating in the
lower stratosphere (60,000 to 90,000 feet), west dominating the mid-
levels (90,000 to 110,000 feet), and easterlies again above 110,000
feet. A fairly rapid transition occurs during spring to strong,
steady easterlies, reaching a maximum in July and August. There is
a sharp transition back to light variable winds in the fall.

Precipitation

Mean annual precipitation is 26.11 inches, but year-to-year
variation is great (Fig. 19). For example, the total for 1968 was
42.27 inches, making it the wettest year on record, while 1969 had
only 17.11 inches, next to the lowest yearly total recorded. Lowest
was 12.86 inches in 1953. This variation is reflected strongly in
growth of vegetation on the islands. Twice during POBSP studies, in
spring 1966 and in late 1969 and again in November 1973, extended dry
periods resulted in the original portion of Sand Island becoming a
virtual desert. March, April, and May 1966 were driest on record for
these months, and May 1966, with only 0.11 inch was the driest month
ever recorded.

Atmospheric disturbances that occur during the four winter
months bring in cooler, less humid, less stable air, which results
in more cumulus build-up and heavier, more frequent precipitation than
occurs during the 8-month summer season, when light showers resulting

yeuscoe -20 +30 -40 50-60 -70
A 1
110,000
10
100,000
15
90,000 20
80,000 30
ho
70,000 50
60,000 80
110
50,000
150
40,000 200
250
300
30,000
20,000 900
10,000 700

Altitude in

Figure 17,

120,
110,000

100,
90,
80,

70,

60,

e

50,
40,
30,
20,

10,

Altitude in feet

Figure 18.

850
feet

JAN FEB APR MAY JUN JUL AUG SEP ocT NOV DEC

Seven (1959).

000
7
10
000
15
000 20
000 30
000 40
50
000 80
100
000
150
000 00
250
000 300
000 500
000 700
50

JAN

FEB MAR APR MAY JUN JUL AUG OCT NOV DEC

Johnston Atoll mean V components; adapted from Joint Task
Seven (1959).

ig)

Barometric

pressure in
millibars

Johnston Atoll mean U components; adapted from Joint Task Force

Baronetric
pressure in
millibars

Force

20

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Mean monthly precipitation, Johnston Atoll, 1931-1972; means
Adapted from U.S. Department ¢

(solid line), extremes (dots).

Commerce (1972).

Figure 19.

21

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80 (Bs
79 2%. * pus! ‘e.
vo 4 + ; : .” , 4 . a ;
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ie eT al le, net | ae a aaa a ee | le

igure 20. Mean monthly relative humidity, Johnston Atoll, 1931-1972; 0100
hours (dots), 0700 hours (dashes), 1300 hours (solid line), 1900
hours (barred). Adapted from U.S. Department of Commerce (1972).

2

from small build-ups of cumulus clouds generated by the shallow trade
winds are the usual source of precipitation. The four winter months
average 2./5 inches of rainfall and the eight summer months 1.87 inches.
However, in April and again in September-November heavy showers asso-
ciated with the passage of tropical storms or depressions are not
uncommon.

Relative Humidity

The mean relative humidity is 75 percent, being highest at 0100
hours (78 percent) and lowest at 1300 hours (69 percent). Monthly mean
values vary little throughout the year, but are definitely lower in
January and February and in June and July than during the rest of the
year (Fig. 20).

Sky Cover

Mean monthly sky cover, sunrise to sunset only, is 6.0 ona
scale of O for no clouds to 10 for complete sky cover (Fig. 21), with
little variation throughout the year. During an average year there are
75 clear days, 172 partly cloudy days, and 118 cloudy days (Fig. 22)

SKY COVER

Figure 21. Mean monthly sky cover, sunrise to sunset, Johnston Atoll,
1931-1972. Sky cover is expressed in a range of 0 to 3 as
being clear, 4 to 7 as partly cloudy, and 8 to 10 as cloudy.
Adapted from U.S. Department of Commerce (1972).

MEAN NUMBERS OF DAYS
W

23

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=
$
$
$
>
s
3
>
>
$

S

$

$

SS
>
$
S
8

.
C TO)

Mean monthly number of days of clear skies (solid line), partly
cloudy skies (dots), cloudy skies (barred), and precipitation of
0.01 inch or more (dashes), Johnston Atoll, 1931-1972. Adapted
from U,S. Department of Commerce (1972).

24

Oceanography

Characteristics of the waters surrounding Johnston Atoll are
described on the basis of studies made by the Bureau of Commercial
Fisheries, U.S. Fish and Wildlife Service (Seckel, 1962 and 1968, and
Barkley, 1972) and more detailed studies of the immediate vicinity of
the atoll and within the atoll are based on studies by the Office of
Naval Research (Kopenski and Wennekens, 1966, and Wennekens, 1969).

Major Water Masses and Currents

The major surface water mass of the Hawaiian and Johnston
region is called North Pacific Central (Fig. 23). These waters have
a salinity generally greater than 34.8 o/oo and range up to 35.3 o/oo
(Seckel, 1962 and 1968). They flow in a huge clockwise gyre, the
southern limb of which moves past the Hawaiian Islands and Johnston
Atoll as the North Equatorial Current, a belt of water several hundred
miles wide and several hundred feet deep moving at up to 1/2 knot.
South and east of this water mass is the California Current Extension,
consisting of waters transitional between North Pacific Central and
North Pacific Equatorial, a smaller mass of lower salinity, usually
less than 34.2 o/oo. The California Current Extension stretches
westerly during the summer to recirculate into the Equatorial Counter-
current west of 175° longitude, but weakens in the fall to recirculate
between 160° and 170°W (Seckel, 1962: 407). Seckel pointed out that
the spring and summer invasion of the Hawaiian Islands region by the Cal-
ifornia Current Extension correlates with the availability of skipjack
(Katsuwonus pelamis). He thought the correlation could be a direct
result of the invasion of the lower salinity waters or could result
from a dynamic effect produced when this current entered the region.
This season also correlates fairly well with the breeding seasons of
most Hawaiian and Johnston Atoll seabirds.

The Equatorial Countercurrent does not reach the immediate
vicinity of Johnston Atoll, but may be near enough to make a signifi-
cant contribution to the atoll's biota. It is a shallow easterly
flowing current, with salinity less than 34.5 o/oo, which flows at
up to 2 knots roughly between 5°N and 10°N latitude (Svedrup, Johnson,
and Fleming, 1942). During spring and summer, when it is farthest
north, it may reach to within 300 nm of Johnston Atoll and thus
probably is within feeding range of Johnston seabirds (Gould, 1974a).

This current spirals clockwise to an observer facing east, with
the lateral (north-south) component about a fifth the velocity of
the west to east velocity. Thus there is upwelling and divergence
along the boundary between the countercurrent and the westerly flow-
ing waters of the North Equatorial Current. Svedrup, et al. (1942:
712), Ashmole and Ashmole (1967), and Gould (1974a) present evidence
that this boundary and waters flowing away from it are highly productive
of marine organisms.

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26

Local Waters and Currents

Waters near the atoll were described by Wennekens (1969) as
having temperatures ranging between 25 and 27°C and salinities between
34.6 and 34.8 o/oo in the upper 100 meters. Salinity increased with
depth to a maximum slightly in excess of 35 o/oo between 100 and 200
meters, then decreased to a minimum of slightly over 34.3 o/oo at
about 400 meters, then decreased slowly to about 34.6 o/oo at 2,000
meters. The main thermocline was found between about 100 and 400
meters, with temperatures decreasing from about 25 to about 7°C.

Below 400 meters temperature decreased slowly to about 2°C at 2,000
meters. The region of steep gradients in both temperature and salinity
between 100 and 400 meters was highly stable because of rapid change
in density correlated with these changes in physical characteristics.
Intermixing of waters above and below this region was inhibited.

Although Timme (1963) reported zooplankton levels in the waters
moving past Johnston to be very low, and Ashmore (1973) reported that
fishes were seldom seen beyond the immediate atoll area, the large
submerged bases of atolls such as Johnston deflect the waters and
cause leeward eddies with local areas of nutrient turnover
and enrichment (Sette, 1955). King (1967) and Gould (1974a) show
that areas within 100 miles of these island upwellings provide most
of the food resources for large seabird colonies.

Wennekens (1969) described the interactions of the North
Equatorial Current and local tides with the atoll, based on extensive
studies with dye markers and parachute drogues designed to measure
flow at depths down to 3,000 feet. The purpose of his study was to
obtain information useful for minimizing effects of pollution of
atoll waters. His information indicated that deeper waters flowed
rather smoothly around the atoll, but that a distinct "island wake"
formed in the surface layers. He recognized three seasonal flow
regimes or patterns around and over the atoll (Fig. 24).

One regime extended from late November to early March and was
characterized by a strong offshore current setting generally south-
west, with island wake confined mainly to the southwestern quadrant
and a prominent convergence forming along the seaward margin of the
island platform during rising tide. Barkley (1972) suggests that
in February Johnston's wake extends some 600 km downstream.

The second regime lasted from early March through mid-June and
was characterized by a strong offshore current setting generally
northwest with island wake confined mainly to the northwest quadrant.
An active easterly counterflow developed along the seaward edge of
the reef during periods of falling tide.

The third regime extended from mid-June to early December and
was characterized by moderate offshore current setting generally
westward. Island wake was confined mostly to the western and north-
western quadrants.

27

CS ca eT

REGIME I

Strong Offshore Current

:
ii

Osciiiaies W-S

2
Spee
Jonnston Island 256 = ™M
Seasonal SO)
Current Regimes aran t=
nz
ee
anid
20° 4
a
=

Falling Tide

Rising Tide
Flow deflects
to right

Flow deflects
to left

Daily Tidal! Flow Regime

Seasonal and tidal flow regimes, Johnston Atoll (Wennekens, 1969).

Figure 24.

The effects of local tides were to deflect currents to the left
during rising tide and to the right during falling tides. During the
June to December period, when offshore currents were only moderate,
local tides induced a rotary or semi-rotary motion to the currents,
reversing the westerly flow during falling tides.

The flow within the lagoon was complicated by the effects of
tide and the presence of islands and channels. Some of the patterns
inferred from earlier work (Kopenski and Wennnekens, 1966) are shown
in Figures 25 through 27. Wennekens' (1969) publication contains
numerous diagrams and aerial photographs showing these flow patterns

in great detail.

Kopenski and Wennekens (1966) thought the disturbing influence
of man on the marine environment of Johnston Atoll to be minimal,
except for localized destruction of living coral and the release of
fine sediments. They suggested that the enlargement of Johnston and
Sand Islands and the creation of the two small islands affected the
general circulation only in a minor way. At the local level, however,
they found that circulation had been greatly affected. The dredging
of boat channels, turning basins, etc., especially between Akau Island,
Sand Island, and the southern tip of the main reef had created new and
artificial flow channels. The geometry of the shoreline of the man-
made islands also contributed to the creation of vortexes, stagnation
points, and venturic effects, which shift in location and vary in

"ISLAND WAKE"
EDDY

RGENC F

Sel |
os SONVE

NORTH EQUATORIAL CURRENT

169° 30'

Johnston Atoll inferred lagoon circulation, January-

Figure 25,
February 1965 (Kopenski and Wennekens, 1966).

29

OURING WESTERLY FLOW, TURBID WATER
1S TRANSPORTED TO THE WEST AND
SOUTHWEST AS INDICATED )

16° 45'

TURBULENT

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FIELD ot %
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mo

CURRENT ROTATES
AND SETS WESTWARD

169° 30°

Figure 26. Johnston Atoll inferred lagoon circulation and turbid Surtion
(westerly flow), July-August 1965 (Kopenski and Wennekens, 1966).

DURING EASTERLY FLOW, TURBID ; TURBID "CLOUD"
WATER BECOMES "SQUEEZED" BROKEN OFF.
ALONG THE WESTERN SEAWARD

MARGIN OF THE ISLAND PLATFORM
AS INDICATED. 7,

16° 45'

SS Whitecaps
Ye

TURBID "CLOUD" NE
BREAKING OFF CURRENT ROTATES
AND SETS EASTWARD

169° 30'

Figure 27. Johnston Atoll inferred lagoon circulation and turbid outflow
(easterly flow), July-August 1965 (Kopenski and Wennekens, 1966).

30

intensity as the regional circulation changes. In addition, the
westerly extension of Johnston Island created a backwash immediately
south of the island during the summer, resulting in very limited
mixing of waters in that area.

Tides

Wennekens (1969) described the local tides as "...mixed type,
predominantly semi-diurnal, usually with two daily high and low waters
exhibiting strong inequalities in the amplitudes of the high waters.
The inequalities in the high reach a maximum at the time of maximum
moon declination....'' His tide graphs show the highest high tides to
have amplitudes of a little under 3 feet and the lowest high tides with
amplitudes of under 2 feet.

Seismic or Tsunami Waves

The characteristics of seismic or Tsunami waves--long gravity
waves caused by submarine earthquakes, landslides, or plutonic activity--
at Johnston Atoll were described by Wennekens (1969). From available
historical data (Fig. 28), he found the largest amplitude wave recorded
to be about 3.4 feet, resulting from the Chilean earthquake of 22 May
1960. The crest of the wave at Johnston Island appeared as a pro-
gressive rise in sea level, with maximum elevation being reached in
25 to 30 minutes. The period of seismic waves at the island appeared
to be between 45 and 60 minutes.

Wennekens (1969: 6-7) further suggested that historical records
---indicate the effects of a Tsunami at Johnston Island should be
minimal, consisting of transient rise in water levels, occasional
breakers to higher than normal levels, and local flooding of low areas
.--[and] that no large breaking wave or bore is likely to occur at the
island."

Ww

HISTORY

Johnston Atoll has had a varied history. From two small, in-
significant islands, the atoll has grown into a large Department of
Defense complex. Previous historical accounts have been written by
Bryan (1942), Thorp (1960), and Bauer (1965).

Major sources of information for this history of Johnston Atoll
were the Bernice P. Bishop Museum Library, Hawaii State Archives,
Hawaii State Library, Hawaiian Mission Children's Society Library,
Hawaii Historical Society Library, Library of Congress, Library of
the National Museum of History and Technology of the Smithsonian
Institution, U.S. National Archives, U.S..Naval Archives, and the
University of Hawaii Library. The staffs of all these institutions
were most helpful, and their contributions are hereby acknowledged.
Although these sources were carefully searched, none of them was
exhausted for information. What is presented here should, however,

Kamtchatka

Tide Gage Record Showing Tsunemi
JOHNSTON ISLAND
November 45, 1952

Hours G.C.T.
0 1 2 3 4 5

Tide Gage Record Showing Tsunami

JOHNSTON ISLANO, HAWAII
A \ i May 23-24, 1960 pron whe
Ap
pcan Chile

Approx. Hours G.MT.
ee NG a7 as Se 20) 2k 22 23 0 1 2 3
acnyeheercunnstecesuer rsd: és

| Tide Gog Record Showing Taunami
JOHNSTON ISLAND, KAWAL
hing March 28.29, 1964 Ab
t

Approa. Hours GAT.

NS ONSha TS nies eee, AOL gee gan 2a 0 ] 2 3 4 3 é ?
a 5

Figure 28. Tidal wave records, Johnston Atoll (Wennekens, 1969).

S}ib

provide a reasonable starting point for anyone preparing a definitive
history of the atoll. A fruitful source of information for recent
history, which we scarcely tapped, is the few thousands of military
and civilian personnel who have spent time on Johnston Atoll during
the last 40 years.

Discovery and Early Exploration

Nowhere have we found any evidence or suggestion that Johnston
Atoll was inhabited or even visited by Polynesians. Habitation would
hardly have been possible, and any visits that may have occurred
left no mark upon the islands that was discerned by later visitors.

The Spanish sailed Central Pacific waters for two and a half
centuries before any sightings of Johnston Atoll are known to have
been recorded. Annual trips in the trade winds from Acapulco to
Manila (between 13° and 14°N latitude) took them less than 200 miles
south of the Atoll. The return route, in the prevailing westerlies,
took them north, not only of Johnston but of the entire Hawaiian
Archipelago. Dahlgren (1916) and Stokes (1939) thoroughly explored
the possibility of Spanish discovery of the Hawaiian Archipelago, a
myth that grew and spread from several sources but is negated by both
these authors. We have not seen Dahlgren's paper, but Stokes contains
no hint of anything that might have been Johnston Atoll. Stokes
does not entirely discount the possibility of a sighting of the
Hawaiian Islands by other than Spanish, since they were not tied to
the trading routes as were the Spanish. The junior author perused
a number of maps in the Map and Cartographic Division, Library of
Congress, finding only two possible islands that could have been
Johnston. These were the Basse de Villa Lobos, which appears on
Arrowsmith Chart of 1798, and Izle Solitaire, on a 1706 map by Godallett.
Positions of these were too crudely determined to give any certainty
as to exact locations, and no descriptions could be found of either of
these. It is possible that a sighting of Johnston could have been
responsible for the placing of these islands on the charts, but there
is no evidence that such was the case, other than that they appear in
the general area.

The discovery of Johnston is usually attributed to HMS CORNWALLIS,
Captain Charles James Johnston, on 14 December 1807. At least one
account of the island, by Captain Joseph Pierpont of the SALLY, out of
Boston, predated the CORNWALLIS sighting by 11 years, and Krusenstern
(1811) declared that the Spaniard Don Jose Comisares saw the atoll in
1786, 21 years before the CORNWALLIS sighting. The Comisares' sighting
is poorly documented, but the 1796 grounding of the SALLY and the
PRINCE WILLIAM HENRY was reported in several newspapers and is here
regarded as the earliest account of the atoll on record.

Captain Joseph Pierpont of the SALLY published the following note
in several newspapers, including the Columbian Centinel of Boston on
24 June and 13 September 1797. It is here copied from Ward (1967: 417):

33

In lat. 16,45 N. long. 169,38 W.from London, on my
passage from the Sandwich Islands to China; the 2d of
Sept. 1796, at midnight, in company with the schooner
Prince William Henry, William Wake, Master, of London,
we both ran ashore on the south side of a reef of
coral rocks and sand, where we continued until next
day noon -- at which time the weather being very clear.
We saw two small islands of sand, bearing W. by N.

4 or 5 miles distant, and from our topgallant-mast-
head. We saw the shoal extending E.S.E. southerly
round to W.S.W. -- but how far we were not able to
determine.

Keep the Lat. 17 N. and this shoal will not be seen.
Joseph Pierpont

N.B. It is hoped that the printers of America, will
give the above a place in their papers.

Krusenstern himself (1811: 288) suspected the presence of land
in the vicinity of Johnston Atoll in 1804:

On the 15th of June we saw in lat. 17° and long. 169°30'

an extraordinary number birds that hovered round the ship
in flocks up upwards of a hundred; this raised our hopes

of meeting with land very considerably; but although the
night was perfectly clear and we kept a good lookout, there
was none to be percieved. I cannot; however, but think,
that during the night, we must have passed near some island
or rock, standing above water, that serves as a resting
place for these birds, for we again saw several the next
morning, nor did we lose sight of them until noon.

The CORNWALLIS sighting was announced by Lt. Wm. Henry Smyth, an
officer on the ship. He placed the islands at 16°53'20" N, 169°31'30" W.
The ship's log described the islands as "Two very low islands having a
dangerous reef to the eastward of them, and the whole not exceeding four
miles in extent" (Marshall, 1827: 173).

Otto von Kotzebue (1821: 258-259), another Russian explorer, de-
scribed the islands but it is doubtful if he actually saw them:

The islands which Mr. Johnstone [sic] discovered in 1807,

on board the frigate Cornwallis, in the W,S.W. of the
Sandwich Islands, and which we looked for in the spring

of 1817, are, like the island of Sala y Gomez, perfectly
naked rocks which do not seem to belong to the formation of
low islands. The reefs which are united to them form shoals
extending to great distances from them, which are very
dangerous to vessels that navigate these seas.

No further sightings of the islands are known between the
possible Kotzebue sighting of 1817 and that of one of the ships of
the Wilkes Expedition in 1841 (Wilkes, 1844, 5: 288):

Mr. Knox [S.R. Knox, Acting Master of the U.S. Schooner
FLYING FISH] reported to me that after his separation,

on the 30th of November [1841], he stood for the position
of Cornwallis Island, as laid down by Arrowsmith in
longitude 169°31'W., latitude 16°50'N., without seeing
any indication of land. Twenty-two miles to the south-by-
east of this position, he discovered a reef, which
surrounded an extensive lagoon, extending north-east and
southwest ten miles, and in the opposite direction five
miles. On the northwest side of this reef there are two
low islets: the one to the westward was covered with
bushes, but no trees; the other was no more than a sand
bank. This reef lies deep. The longitude of the
westernmost islet was found to be 169°45'36" W., and

the latitude 16°48' N.

Guano Period

This section is taken primarily from sources in the U.S. National
Archives, Record Group 59, U.S. Department of State, Guano letters.

The next known possible sighting of the island was by William H.
Parker, captain of the REINDEER, who passed through the area in
January 1852 on a trading voyage to the Ladrone Islands and China,
and later claimed to have seen several islands at Johnston Atoll.
What he actually saw is difficult to determine, because his descrip-
tions fit nothing actually present. It is possible he saw nothing,
but knowing islands had been described in the area, made up a good-
sounding story.

After the U.S. Congress passed the Guano Act 18 August 1856,
Parker attempted to claim the islands under the act. On 26 August
1857 he formed a partnership with attorney Richard F. Ryan, of San
Francisco, giving Ryan one half of all profits from discovery. On
29 August Ryan wrote Secretary of State Lewis Cass, giving notice of
Parker's discovery and asking prompt action on their claim to the
islands. The petition was refused on grounds that Parker and Ryan
were not occupying the islands and could not finance their exploitation.
On 19 November Ryan submitted an analysis of the guano to the State
Department. This states: '"'The guano on Cornwallis Island is of
extraordinary good quality, consists of 82.53 of Azote [nitrogenous
material] in 100 parts and 17.47 of Hydrogen, the deposits are large
and from its dryness is most valuable as an article of commerce."
The source of this guano, or its report, sound suspicious, for there
was no mention that Parker actually landed on these islands in 1852,
and no one had been back. On 19 December 1857 Parker and Ryan made
an agreement with Robert G. Byxbee and Asa B. Stoddard, owners of the
100-ton schooner PALASTINE, giving the ship owners five-eights interest
in the venture, they in turn agreeing to send the ship to the islands

35

to claim them and to bring back a load of guano. On 20 December

Ryan dispatched another letter to Cass, enclosing an amended declara-
tion of Parker's claim, stating that a vessel was to be sent for a
load of guano in 2 weeks. It also pointed out the potential
importance of the islands to the United States as a coaling station.

The PALASTINE departed San Francisco 8 January, under Captain
William R. Perriman, and with Parker on board. They arrived on the
island 9 March 1858, and left 16 March, arriving back in San Francisco
22 April. Subsequent statements, most of them sworn and notarized,
varied greatly as to exact procedures on landing. Parker's statement,
dated 28 April 1858, stated that he and Rich, the mate of the PALASTINE,
and two seamen went ashore on Johnston Island and erected a flagstaff
with a cross on top with the inscription "this island was taken
possession of by W.R. Perriman, master of the schooner Palastine on
behalf of the owners and charterers of the schooner Palastine in the
name of the United States of America, James Buchanan, President." He
took latitude and longitude 16°46'N, 169"28'W, and with the mate
explored the island and took 13 "gunny bags" of guano (about 1,200
pounds). They also landed on Sand Island (called by him Agnes, for
his daughter), and erected a flagstaff with a similar inscription as
the one placed on Johnston Island. Parker described the islands quite
accurately in his statement and gave the position as 16°46'N, 169°28'W.

The signed statement dated 1 May 1858 by Perriman and Rich did
not differ significantly from that of Parker but added that there
was no sign of previous occupation. It too gave a good description
(neither description, however, mentioned birds) and the position given
differs only slightly from that given by Parker. Position given in
the various letters and statements all tended toward minor variations,
some of which probably resulted from actual variations in measurements,
and some from mistakes in copying. The most significant part of the
statement, for later questions of ownership, was: '"...that possession
was immediately taken of the said islands by the deponents Perriman
and Rich in company with the said Parker, and same were occupied by
them during this time."

In May, Ryan and Louis Blanding (an attorney for the group) cor-
responded with Secretary of State Cass, calling attention to Parker's
claim and giving notice that a ship had been sent, guano removed, etc.,
and including the sworn statements of Parker and of Perriman and Rich.

On 8 June, Byxbee, Stoddard, and Ryan, together with William
Thompson and Cornelius L. Place, formed the Pacific Guano Company,
incorporated under California law. Parker was not included, but
apparently he owned stock in the company, for records dated between
26 June and 6 November of 1858 show that he sold a total of 80 shares
of stock in it to five different people.

Meanwhile, a second attempt was made to claim the island, this
time for the Government of the Kingdom of Hawaii. Samuel C. Allen,

recorded as being from Honolulu and from San Francisco, obtained from
King Kamehameha IV a royal commission dated 31 May 1858 to take pos-
session of unclaimed islands in the name of the King. Aboard the
KALAMA, under Capt. Watson, he arrived at Johnston Atoll 14 June, and
according to a sworn statement by Edward Watts, a cook and steward on
the KALAMA, Allen and the Captain removed the flagstaff and cross put
up by the PALASTINE crew, and presumably destroyed these and the in-
scriptions. They claimed the islands, Cornwallis (Johnston) on the
14th and Kalama (Sand) on the 19th; after erecting a Hawaiian flag on
them and taking a small sample of guano, they returned to Honolulu.
Kamehameha IV formally claimed the islands for his Kingdom in a
Proclamation 27 July 1858, in which it was stated that the islands
had been found "derelict and abandoned," by the KALAMA crew.

The PALASTINE returned to Johnston, arriving 22 July and the crew
found their flags, etc., replaced by those of the KALAMA. They remained
on the islands 13 days, loading about 70 tons of guano, before leaving
two men on the island and returning to San Francisco, arriving 31 August.
Meanwhile, Louis Blanding, attorney for the Pacific Guano Company,
wrote on 13 August to James W. Borden, U.S. Commissioner in Honolulu,
asking Borden to inform the Hawaiian Government of his company's
prior claim to the island, and requesting "some positive disapproval
of the act." The company had most likely heard of the trespass via
the King's proclamation in Honolulu, since their ship had not yet re-
turned when the letter was written.

On the return of the ship, Blanding wrote Lewis Cass, U.S. Secretary
of State, informing him of the details of the trespass by the KALAMA, and
placing the blame on the crew of the KALAMA, stating that the King
probably did not know of the prior claim at the time he issued his
proclamation. When the Hawaiian Government was informed of the facts,
via dispatch from Commissioner Border, 8 September, Allen's royal com-
mission was rescinded as far as Johnston Islands were concerned, and it
was amended to the effect that the commission was void in cases of
prior claim by other powers. This had been the intent of the original
commission, but it was not specifically worded. The King then solicited
from Borden an opinion as to whether the flag, cross, and inscriptions
left by the PALASTINE's crew 9 March fulfilled the requirements for
claims under the Guano Act. This act required that the islands be
occupied by the claimant or their agents. Byxbee, President of the
Pacific Guano Company, had requested similar information from Cass
in a letter dated 19 August 1858 requesting a statement as to the
validity of Parker's claims and the means by which all requirements of
the Guano Act could be met.

On 9 November the Hawaiian Government requested Borden to ask the
Pacific Guano Company not to send vessels to the islands to load guano,
adding that the Hawaiian Government was asking the same of Allen, until
ownership had been settled, to avoid a confrontation by the two partici-
pants on the island. Newspaper articles at the time indicate the King
relinquished claim to the island, but our search has not produced the
actual documents.

37

A confrontation did occur, however, for Allen sent Captain
Borland in the ship GAUNTLET to the island for a load of guano. On
arrival, he found the two Pacific Guano Company men in possession of
the island and told them he intended to load the GAUNTLET, using force,
if necessary. The Pacific Guano Company had in the meantime sent
A.D. Piper in the ship RADIANT, under Captain Hallett, to take command
of the island and to prepare for shipping guano. The GAUNTLET was
at anchor when Piper arrived and he immediately protested its presence
to Borland, but offered to sell him a load of guano at $4.00 per ton,
or to charter his ship for his own firm--the Pacific Guano Company.
Borland apparently was more intimidated by the difficulties in loading
from the islands than by threats or deals, so he accepted neither offer.
Captain Hallett concurred that loading was impossible, and refused to
load the RADIANT. Both Borland and Hallett then left the island,
leaving Piper and 11 men in armed possession of the island. Piper
set his men to work preparing a wharf, railway, etc., for shipping guano.
The FENIMORE COOPER, under Captain John M. Brooke, was ordered to the
island in March 1859, to survey the islands and make soundings. He
arrived 14 March and learned the above story from Piper (Brooke, ms;
see also, Brooke, 1955).

Meanwhile, it appeared that Parker was being squeezed out of
the Partnership which had become the Pacific Guano Company. The company
had pursued the claim, through protests over the intrusion by Allen's
ships from Honolulu, and by letters to the State Department requesting
information as to proper means of complying with and fulfilling the
requirements of the Guano Act. In support of the contention that
Parker himself had no claim to the islands, Perriman and Rich again on
25 and 26 October 1858 made sworn statements as to the happenings on
the first landing on the islands in March. They stated that Parker
had said there was but one large island with trees on it, which was
visible on the horizon for 20 miles, and which had no reefs around it.
What they found instead after beating about for 7 days, was the two
small islets, visible only 8 miles, with no trees, and with extensive
dangerous reefs all around. There is suspicion that this story was
at least in part concocted, because Parker's original claims described
no less than 7 islands and islets, some said to be no more than cays;
it is therefore not too likely that he would have told Perriman there
was only one island while they were searching for them.

Perriman further claimed that he and two others went ashore on
Johnston at 0900 on the first day, and that Parker did not land until
the second boat in the afternoon. In his first statement, Perriman
said he and Rich "...in company with...Parker..." had taken possession
of the island. Here he asserts that Parker was not along on the first
landing, although he does not say what part Parker had in the claiming
ceremonies. Interestingly, in a newspaper interview in 1892, 34 years
after the first landing, Perriman asserted that Parker did not go
ashore at all.

Who was trying to pull what on whom in the hassle that took
place over the claim to the island is not clear from the available

38

documents. Parker may not have even seen the islands before the
first PALASTINE trip--his descriptions up to that time certainly
did not fit any existing islands. Whether the organizers of the
Pacific Guano Company were trying to get rid of him as a partner
in the business, or whether he was trying to use the company as a
means for claiming the islands for himself, or both, is not clear.
But that both were occurring simultaneously is not out of the
question.

In any case, by 6 November 1858 Parker had sold all his ownings
in the company--80 shares. On 22 February 1859 Parker died in a
hotel in Washington D.C., supposedly there to perfect his claim to
the islands.

Parker's supposed widow, Harriet B. Parker, pursued his claim
to the islands, but a judicial ruling from Jeremiah Black, later in
1859, stated that since Parker had not actually occupied the islands,
and the Pacific Guano Company had, the company was the proper possessor
of the islands. Consequently the company posted bond for $100,000.00
for rights to exploit the island. In 1859 attorneys for Parker's heirs
and for the Pacific Guano Company wrote several letters to the Secre-
tary of State and to President Buchanan attempting to verify their
claims.

The amount of guano being taken off during 1859 and subsequent
years must have been fairly substantial, if the number of ships and the
statements of their captains can be trusted. There is, however, some
question as to the value of the product shipped. It was reputed by
at least two ship captains to be of high quality, easily obtained,
etc., but Hague, in 1862, wrote that: "...from Johnston's Islands

one or two cargoes have been brought to this country [the United States],

the greater part of which proved, I believe, to be sand. These are
described as three small islands (probably islets of one atoll) con-
taining but little guano and that mixed with coral sand." Whether the
companies continued in contention is not known, but they probably did,
for ships of the American Guano Company were reported in and out of
Honolulu to and from Johnston Island in 1860 and 1861.

The Civil War caused an interruption in both guano exploitation
and in attempts to pursue claims, but shortly after it was over,
Parker's only heirs, his widow Harriet B. Parker and their daughter
Agnes Parker Burtt (for whom Agnes Island, now Sand Island, was named
by Parker) again took up the claim of the guano islands. William H.
Parker, Jr., Parker's only son, had died of wounds suffered while
serving with the Union Army at Antietam.

In 1872, Parker's widow was successful in getting Congress to
amend the Guano Act to allow legal heirs to claim islands of deceased
original claimants. She had no success, as far as is known, in ob-
taining clear title to the islands, or any compensation for whatever
guano was removed. Their principal claims were that a document
awarding the islands to the Pacific Guano Company bore the forged

39

signature of Secretary of State Cass; that Parker was acting alone
in claiming the islands; that he had not sold stock in the Pacific
Guano Company; and that Perriman and Rich made statements in October
which conflicted with those made in May in order to help the Company
beat Parker out of his claim.

About 1879 a third heir appeared. Mrs. Malvina H. Parker, of
San Francisco, who claimed to have been married to W.H. Parker 16 July
1858 in San Francisco, and furnished proof in the form of sworn state-
ments by the Justice of the Peace who performed the ceremony. She,
together with Agnes Parker Burtt, disclaimed the assertions by Harriet
B. Parker, whom they now referred to as Harriet B. Fisher, of her rights
to the guano islands claims. Apparently Parker had divorced Harriet
B. Parker sometime before 1858. Records of this were not searched
for, but probably could be found. Malvina H. Parker actually sold her
claims to the island to a man named Hueston, of Michigan, who in turn
sold it back to her.

Ships known to stop at Johnston Island after the Civil War were
few. Presumably most of these stopped for guano, but as far as is
known, there was no permanent, continuous occupancy during this time.
The only major shipwreck known to have occurred on the atoll happened
in 1889. This is remarkable in view of the difficulty in seeing the
island, and the number of whalers, guano ships, etc., plying these
waters throughout the 18th and 19th centuries. The whaler J.B. HOWLAND
wrecked on the island 26 December 1889. One or two men were lost in
getting the men ashore over the coral heads of the lagoon. All sur-
vivors were subsequently picked up and returned to Hawaii.

In 1892 Great Britain attempted to claim Johnston as a potential
cable station. HMS CHAMPION was dispatched from Honolulu with orders
to leave for the island without telling anyone where they were headed.
This they did, and finding the island totally abandoned, claimed it
for Great Britain. The CHAMPION's captain described the island and
made a small sketch map of it--quite accurate for johnston, but crude
for Sand which they saw only from a distance. They also found remains
of the wreckage of the whaler.

On learning of the British claim, an agent, Lomberg, for the
Pacific Guano Company claimed the company had not given up possession
of the islands, and expected to return there to resume operations, and
also expected protection by the United States Government. There are
no records indicating that the company did return to the islands.

The State Department, however, still listed Johnston as an island
bonded to the U.S. as late as 1893.

In 1905 there was renewed interest by Parker's heirs. Mrs. Agnes
Parker Burtt York, Parker's daughter, in agreement with William H.
Underwood and LeGrande Brown, visited the islands aboard the Steamer
IWILANI 14 February 1905 and brought off a small cargo of guano.
Shortly after this visit, Mrs. York sold the islands to Underwood for
$250,000.00, most of which was to be paid from a 10 percent royalty on

40

all guano sold. On 9 June 1905 Underwood sold the islands to Phineas
Coleman for $1.00 plus unnamed "considerations." On 3 August Coleman
sold the islands to the United States Guano Company for $10,000,000.00.
Whether Mrs. York had a clear title to the islands before she sold to
Underwood, or if any money actually changed hands in these deals is
not known. Furthermore, as far as is known, no further ships were
sent out by any of these parties.

In 1909 the Territory of Hawaii leased the islands to Max
Schlemmer, of Honolulu, for 15 years (Govt. lease #661, 20 September
1909, filed with office of Commissioner of Public Lands, Territory of
Hawaii). Johnston had not, however, been included in the Territory of
Hawaii when the latter was annexed by the United States on 30 April
1900. Schlemmer never removed guano nor did he fulfill the requirements
of his lease, which included planting 500 coconut trees per annum and
maintaining them throughout the lease. During June 1914 Captain George
Pliltz, in company with Edward M. May, visited the atoll and found it
to be uninhabited and unimproved with no sign of coconut trees. Con-
sequently, in 1918 the lease was voided, and another lease was given
to C.K. Ai, of Honolulu, whose Japanese firm, C.K. Ai and Company,
proposed to use the island as a fishing station. His party of Chinese
stayed on the island only one day, however, before they mutinied and
returned to Honolulu.

The first scientific expedition to visit the islands, excepting
the possible visit by the Wilkes ships in 1841, and the surveying done
from the FENIMORE COOPER in 1859, was done by the party visiting the
islands aboard the minesweepers TANAGER and WHIPPOORWILL in July 1923.
This party was headed by Alexander Wetmore, then of the Bureau of
Biological Survey of the Department of Agriculture, later to become
Secretary of the Smithsonian Institution. Other members of the ex-
pedition represented the Bernice P. Bishop Museum of Honolulu. Their
special interests and the published results of their trip are discussed
under the Biota section in this report. Commander John Rodgers, later
to become famous for seaplane flight from California to Hawaii, and
two other aviators were along and made aerial photographs of the islands
and reefs with the aid of a small seaplane. The party was on the
islands from 10 through 20 July 1923.

Because of this visit, the atoll was made a federal bird refuge
on 29 July 1926 by Executive Order No. 4467 of President Calvin Coolidge
and placed under control and jurisdiction of the Department of Agricul-
ture. In 1940 this responsibility was transferred to the U.S. Department
of Interior. This Order still remains in effect, although subsequent
executive orders have given jurisdiction over the atoll to military
agencies (Table 1).

In 1924 the TANAGER and WHIPPOORWILL returned to the islands as
part of Mine Squadron Two, which included the LUDLOW and the BURNS.
These four ships, together with the tender PELICAN, with one seaplane
on board, left Honolulu 3 June 1924 and all but the PELICAN which had

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43

mechanical difficultues and was delayed a day, arrived off Johnston

on the 6th. On the 7th work began in setting up signals on the islands
for surveying the lagoon, which was done from 10 through 20 June.
Accurate position determinants were made, currents were studied, a tide
guage was operated, and extensive soundings were made throughout the
lagoon. Although there was no record of any scientific personnel on
board, collections of coral, marine growth, land shells, etc., were
made for the Bishop Museum. Eight coconut and 12 ironwood trees were
planted "...in favorable locations" on Johnston Island. Recommendations
were made for use of the atoll for seaplanes, as it was thought that
seaplane runways could be developed easily by blasting coral heads.

The signals erected on the island were removed so subsequent visitors,
notably the Japanese, would not know the survey had been made. The ex-
pedition left the atoll 22 June and arrived back at Pearl Harbor on the
26th (U.S. Nat. Archives, R.G. 37, Letter from G.V. Stewart, Commander,
Mine Squadron Two, to Chief of Naval Operations).

During the late 1920's the island was visited by at least one
fishing vessel out of Honolulu, the LANIKAI; Captain Bill Anderson
(who was along on the 1923 expedition with Wetmore, and who had been
on the island, again from a fishing vessel, in November 1922) and
Lorrin A. Thurston, a Honolulu reporter, were the principal members
of the crew. Thurston reported his 1927 experiences, including the
finding of the wreckage of the J.B. HOWLAND, in the Honolulu Advertiser,
5 and 26 August 1928.

Military Occupation

No further military activity occurred on the atoll until 1933,
when Johnston was included in an exercise in extended reconnaissance
flights by Navy patrol planes. Data obtained from the 1924 survey
were to be rechecked "...with view to actually determining the prac-
ticability of utilizing Johnston Island as a temporary aircraft base"
(U.S. Nat. Archives, R.G. 45, Chief of Naval Operations to Commander
in Chief, U.S. Fleet, 26 January 1933).

The USS PELICAN left Pearl Harbor 17 April 1933 and arrived off
Johnston 20 April. A small boat attempted to enter the lagoon through
a reef blasted through the reef west of Johnston Island by the 1924
expedition, but was unsuccessful because of high surf. Entry was
made around the southwest end of the reef, and landing and mooring
areas for seaplanes were searched for in vain in the west and north-
west parts of the lagoon. On the 21st a desperate attempt was made to
find suitable mooring areas, and finally the area between Johnston and
Sand Islands was picked and a message sent out to that effect.

Meanwhile, a squadron of seaplanes had left Hawaii and flown to
French Frigate Shoals, following a line of ships spaced at intervals
along the route. After the planes arrived at French Frigate Shoals,
the ships realigned themselves between there and Johnston Atoll, and
on 22 April four planes left for Johnston, arriving at noon. The only
adverse incident was the striking of a large bird by one of the planes

Ad

as it landed. The damage was repaired successfully on the island.
All planes were fueled and serviced and departed for French Frigate
Shoals on the morning of 25 April (see Amerson, 1971),

A general description was made, including the information that
there was no vegetation except for "long grass." Apparently the trees
planted in 1924 did not survive. Johnston Island was considered long
enough for landing small land planes, with considerable grading
necessary beforehand. The presence of large numbers of birds on both
islands and the islands being riddled with holes made by the birds were
considered as major hazards to aircraft operations, especially if land
planes were to be used. It was concluded that the atoll offered
",..only fair possibilities for making a good advanced base" (U.S. Nat.
Archives, R.G. 45, letter, Lt. F.M. Hughes, USN, to Commander, Aircraft
Squadrons and attending craft, Fleet Air Base, Pearl Harbor, T.H.).
Several photographs were made during this operation, and these are on
file in the National Archives.

On 18 December 1934 the atoll was visited by the Coast Guard
ship ITASCA in a vain search for the lost aircraft STAR OF AUSTRALIA.
Both islands were briefly explored by Captain C.T.P. Ulm and two com-
panions and described very generally by Jan Jabulka (1934), a reporter
on board the ship.

Because of the atoll's strategic military location, President
Franklin D. Roosevelt, on 29 December 1934, by Executive Order No.
6935, placed the atoll under the Department of the Navy. Provision was
made within the new Order to keep Executive Order No. 4467, thus pro-
tecting the breeding birds and habitat.

In 1935 the ITASCA again visited the atoll on 23 June and 23
September. The general description of the islands noted that an old
guano shed was present, but lying flat, and that narrow guage railroad
wheels were still present. On 12 November 1935 a squadron of 6 Navy
planes flew to the islands from French Frigate Shoals and they returned on
the 13th.

In 1936 the Navy began developing the atoll. By 19 October
1936 a 210-foot pier had been built out from the south side of Sand
Island and a sinuous, narrow boat channel had been opened through the
reefs from the south into the seaplane landing area northwest of Sand
Island (see Fig. 8). Reports indicate that a reconnaissance flight
to the atoll had occurred earlier in the year by VP Squadron Ten.
First blasting operations were carried out by the advance party, and
in October the USS WRIGHT arrived with more explosives (U.S. Nat.
Archives, R.G. 45, letters of 15 October and 23 November 1936, from
Commander K. Whiting, Aircraft Squadrons and attending craft, Fleet
Air Base, Pearl Harbor, T.H., to Hydrographer).

The PELICAN and TANAGER arrived on 30 March 1937, and the SWAN
on 31 March, all carrying explosives for enlarging the channel. As a

45

result of this last effort, the area for seaplane landing and mooring
was about doubled and 24 planes could be accommodated. Blasting was
also done around the pier. Recommendations were made for further im-
provements to allow tenders to get closer to the anchorage area ot the
planes. A map of the lagoon dated 14 April 1937 showed a straight

boat channel from the south side of the lagoon into the seaplane landing
area, and on Sand Island a mess hall was marked at the northwest side

of the base of the pier, and on the southeast side a signal tower

(teow Nat... Archives, R:G. 45, letter” from Lt. Commander J.L. Cotton;
USN, Johnston Island Expedition, to Commander, Minecraft, Battle Force).

On 8 April 1937 two VP-6's made a round trip from Pearl Harbor to
Johnston and back in 10 and a half hours to rescue a sick seaman (Bryan,
1939).

In late 1939 the Navy awarded a contract for construction of a small
base and in October or November a force of 60 men were sent to the atoll
to begin work on a@1,150,000.00 warplane, minecraft, and submarine base
for the U.S. Navy, to be completed in a year (Anon., 1939). At first a
lagoon seaplane landing area with headquarters on Sand Island was built.
But this tiny islet soon was overcrowded (see Fig. 9), and expansion in
1940 was made to Johnston Island itself (see Fig. 10). The work, details
of which can be found in Woodbury (1946), was rushed and the Naval Air
Station was commissioned ahead of schedule on 15 August 1941 (Bauer,
1965). On February 14 of 1941 Presidential Executive Order 8682 set
up the Johnston Island Naval Defensive Sea Area and Johnston Island
Naval Airspace Reservation to include respectively the "waters between
extreme high water marks in the three-mile marine boundaries surround-
ing the island, and the airspaces over these waters and islands."

On 15 December 1941, four months after being commissioned, Johnston
Island was shelled for 10 minutes by Japanese surface craft. Again on
21, 22, and 29 December both Johnston and Sand Islands were shelled
from offshore ships; one Japanese submarine was reported sunk by land
guns. There were no injuries to personnel but civilian workers were
demoralized. The shelling caused considerable damage to various
facilities (see Bauer, 1965).

Construction continued until April 1942; projects included dredging
channel approaches and seaplane landing areas, construction of bomb
shelters, living quarters, landplane runways, storage sheds, and gun
emplacements.

In addition to being an air station during early World War II,
patrol submarines used the atoll as a refuel base. By 1944 and
throughout the rest of the war, it became one of the busiest air trans-
port terminals in the Pacific, servicing planes going to and from the
Pacific front some 4,000 miles to the west.

At the end of World War II, the Navy continued the Naval Air
Station at reduced strength but as activity decreased, the status was
degraded to that of a Naval Air Facility. Sand Island was abandoned

46

in 1946. By order of the Secretary of the Navy, operational control
of Johnston Atoll was transferred on 1 July 1948 from the U.S. Navy
to the U.S. Air Force; Navy, however, retained technical jurisdiction.

During the Korean airlift in 1951 and 1952, Johnston Island again
assumed military importance. The airstrip was enlarged by dredging and
new buildings and improved utilities were added. For a short time,
military personnel were permitted to have their dependents accompany
them to Johnston; this ended in October 1956.

On 25 January 1957, the Treasury Department was granted a permit
for the U.S. Coast Guard (now under the Department of Transportation)
to operate a LORAN transmitting station on Johnston Island. Also on
13 September 1957, the Department of Commerce was permitted to operate
a Weather Bureau Station on the Island.

On 22 April 1958 operational control of Johnston was assumed by
the commander of Joint Task Force Seven. As part of Operation Hardtack,
two missiles carrying thermonuclear devices were fired from Johnston
Island into the stratosphere to obtain information on the effects of
nuclear detonations at high altitudes. Teak, the first of these shots,
was detonated at 2150 hours on 1 August 1958, at an altitude of 252,000
feet. The second shot, Orange, was detonated at 2130 hours on 12
August at approximately 100,000 feet. These were the first megaton
devices detonated in the stratosphere by the United States (Hines, 1962).
The Pacific phase of Operation Hardtack lasted until 19 August 1958.

Operational control of Johnston was again assumed by Joint Task
Force Eight (JTF-8) and the Atomic Energy Commission (AEC) on 17 January
1962 for the purpose of conducting additional high-altitude nuclear
tests. Shot Starfish, an explosion of about 1.5 megatons, was conducted
at an altitude of 200 kilometers on 9 July 1962. Another test, scheduled
for 25 July 1962 was aborted after the test missile was destroyed.

During these nuclear tests, an elaborate water sprinkler system
was installed on the original portion of Sand Island to protect the
birds living there. In addition, other protective devices were used,
including smoke pots placed upwind as a shade screen and aerial flares
to divert the birds' attention from the flash of the blast itself.

The Secretary of Defense granted permission on 10 December 1959
for transferring the U.S. Coast Guard LORAN-A and -C Station to Sand
Island. The Coast Guard facility on Sand Island was completed in 1961
and presently maintains a complement of approximately 25 men. A U.S.
Air Force Satellite Tracking Camera Station was established on Sand in
1964.

As of 1965, Johnston Atoll was under joint operational control of
JIF-8 and AEC. It serves as headquarters and base of operations for
resuming U.S. atmospheric nuclear testing in the Pacific should the
Test Ban Treaty of 1963 be nullified by a foreign world power. Tenants
on Johnston Island included: U.S. Air Force and Navy units, and

47

personnel of the Air Defense Command, Space Systems Command, and
Pacific Missile Range.

In the early 1970's various chemical agents, including herbicides
left over from the Vietnam Conflict, were brought from the western
Pacific and stored on Johnston Island. These were placed downwind in
a security area on the southwest end of the island. As of 10 May 1974
measures were being considered by the Environmental Protection Agency
to safely dispose of these agents.

In the spring of 1973, operational control of Johnston Atoll came
under the Defense Nuclear Agency. The atoll is still a Naval Defensive
Sea Area and Air Space Reservation, and access is restricted. The
atoll is a bird sanctuary under the Department of Interior.

BIOTA

The following treatment of Johnston's biota does not treat taxa
evenly. Some are entirely deleted because they are so far outside the
knowledge of the authors and others who worked on the atoll, and
outside the objectives of the POBSP, as to make their discussion mean-
ingless. References to all pertinent papers are included where they
are known, but there is implied no pretense that the references included
comprise a comprehensive bibliography.

Emphasis of the POBSP was on analysis of the natural history of
the terrestrial vertebrates and the terrestrial vascular plants, with
by far the greatest emphasis on the birds. In the following treatment
the only groups listed in detail besides these two categories are the
inshore fishes and insects.

Plants

The marine and terrestrial plants of Johnston Atoll are well known.
Both have been heavily disturbed by man.

Algae

Only one marine algal species was known from Johnston Atoll prior
to 1965 (Moul, 1964). As part of a study of the effects of dredging on
Johnston's marine environment, 67 species of benthic marine algae were
collected by the Hawaii Institute of Marine Biology, University of
Hawaii, 17-22 August 1965 and 17-21 December 1965 (Buggeln and Tsuda,
1966). Collections were made in 14 habitat areas (Fig. 29). Additional
collections from 16 other habitat areas were made 14-24 June 1966 by
the Department of Radiation Biology, University of Washington; from
these collections 26 more species of algae were added to the known
species list (Buggeln and Tsuda, 1969).

The distribution of the 93 species of benthic marine algae known
from Johnston Atoll (Table 2) shows that 12 species were found only from
the marginal reef, while 33 species were found only in the lagoon waters.

48

OAs
169 33 30

Johnston Atoll

r 8)

East Island

0 Tos

rs aA
10 2% “Sand

% .8sIsland

Johnston Island F

Redrawo from Brock gtaj., (1965).

Figure 29. Algae collection stations, Johnston Atoll, 1965-1966 (Buggeln
and Tsuda, 1969).

Of these 33 lagoon species, 11 were found only in open water, 11
occurred only in the inshore Johnston Island area, and 2 were taken
in the inshore Sand Island area.

Buggeln and Tsuda (1966) suggest a vertical distribution of the
Johnston algae--especially Chlorophyta and Rhodophyta--in the sub-
littoral zone, such that as light intensity decreases with depth, the
number of algal species also declines. These authors, as well as
Brock, Henkelem, and Helfnich (1966), found a correlation between in-
creased silt in dredge-affected areas and decreased number of algae
species. The silt caused a reduction in light intensity at all depths;
lack of light reduced photosynthesis and thus eliminated plants.

They also found that in the newly dredged and widened ship channel
(areas 10 and 11), the new surface had been invaded by sparse Cyanophyta
and Rhodophyta species. In some of the dredged areas rather thick
algal mats developed over the whole bottom. The blue-green alga
Lyngbya majescula was the dominant species; others included Sehtzgothrix
caleteola and Phormtdtum submembranaceum.

49

Table 2. Distribution* of benthic marine algae at Johnston Atoll

Lagoon

Division Marginal Open Inshore Inshore
Species Reef Water Johnston Sand
Cyanophyta
Anacystts dimtdtata 1 il
Entophysalts deusta 3
Sehtzothrtx caletcola 3 6 5) 2
Hydrocoleum lyngbyaceum it 1 it
Mterocoleus chthonoplastes 4 i!
Microcoleus tenerrtmus 1 1
Microcoleus vaginatus 1
Lyngbta aestuartt i il 1
Lyngbta confervoides 1
Lyngbia lutea i
Lyngbya majuscula il 3 1 2
Sptrulina tenerrima 1 il
Symploca atlantica iL 1
Osetltatorta nigroviridts 1
Phormtdium submembranaceum 3 3 1
Hormothamton enteromorphotdes 2 i
Calothrtx crustacea 1 1
Calothrix scopulorum 3 3 4 il
Isactis plana 2 > iL
Chlorophyta
Palmogloea protuberans iL
Enteromorpha kyltnit 1
Cladophora crystallina 2 2 1
Cladophoropsts sp. a
Valonta ventricosa it
Dietyosphaerta versluystt 6 6 4 1
Broodlea compostta 1
Mterodtetyon setchelltanum 5 2 1
Dervesta martina il
Derbesta sp. 1
Caulerpa ambtgua i al 2 1
Caulerpa racemosa macrophysa il
Caulerpa urvilltana 3 1 1
Bryopsts pennata 5 4 4 al
Pseudochlorodesmts parva 2 3
Codtum arabteum it
Codium sp. 3 2 4
Halimeda discoidea 3 2 il
Halimeda tuna 3
Acetabularta clavata 2
Acetabularta mopbit 3 2 2
Acetabularta tsengtana il

Acetabularta sp. iL il

Table 2. (continued)

Lagoon

Division Marginal Open Inshore Inshore
Species a2 0 ee Reet eater. Johuston eer
Chrysophyta
Ostreobium retnecket 2
Phaeophyta
Ectocarpus breviarticulatus 3
Eetocarpus tndtcus 3 3 4 1
Eetocarpus trregularts 1
Eetocarpus sp. 1
Sphacelarta furetgera 1 il
Sphacelaria novaehollandiae 5 ih 5 2
Sphacelaria trtbulotdes 1
Dtetyota sp. 2 al it
Pocoektella vartegata 40 5 3
Rhodophyta
Asterocystts ornata 1
Gontotrichum alstdit 1
Erythrotrichta sp. if
Gelidtum ecrinale perpustllum 3 3 iL
Geltdium pustllum pustllum 4 3 2
Wurdemanta sp. 3 1 2
Janta captllacea 3 6 2 2
Janta decussato-ditchotoma 4 2 2 ih
Amphtroa sp. 1 i
Hypnea espert 4 4 4 il
Lomentarta hakodatensts il HH ah il
Champta parvula 2
Anttthamnton anttllarum 2 uh i 1
Callithamnton marshal lensis 2 2
Callithamton sp. 1
Centroceras aptculatum 5 5) 4 2
Centroceras clavulatum Th 3 Hl iL
Crouanta minuttsstma if
Ceramtum affine 3 5 1 1
Ceramium fimbriatum at
Ceramium gractllimum byssotdeum 4 4 4 if
Ceramium huysmanstt 3 3) ik
Ceramtum maryae 1
Ceramtum vagabunde 2 2
Ceramtum zacae il 2 aL
Ceramtum sp. 2
Crouanta mtnuttssima 1
Grifftthsta metcal fit 2

Grtffithsta ovalts 1

Syl

Table 2. (continued)

Lagoon
Division Marginal Open Inshore Inshore
Species Reef Water Johnston Sand
Rhodophyta (cont.)
Grtffithsta tenuts 4 al 2
Grtffithsta sp. ib
Dasya adherens iL al
Dasya stnicola 3 if
Dasya sp. iL 2
Taentoma macrourum it 3
Caloglossa leprteurit al
Heterostphonta wurdemantt laxa 2 2
Herpostphonta SPP- 4 3 1 ib
Polystphonta SPP- 5 7 5 1
Laureneta sp. 4 5 1
Chrondria repens 4 5

*Figures indicate total number of collection stations from which samples
were taken. See Figure 29 for collection station localities; Marginal
Reet mlocalities: 1,2,4,12,27,28,29; Lagoon Open Water: 3,5,6,8,9,10,11,17;
Lagoon Inshore Johnston: 13,14,15,16,18,19,20,21,22,23,24; Lagoon Inshore
Sands, 7,25,26,30.

Vascular Plants

Table 3 lists the 51 families, 109 genera, and 127 species of vascu-
lar plants that have been identified from the four islands at Johnston
Atoll. This number is remarkable, for in 1859 Brooke (ms.) noted only two
plants and in 1923 the TANAGER-WHIPPOORWILL expedition found only three
species of vascular plants (Christophersen, 1931). By 1946, 27 species
were recorded from Johnston Island (Fosberg, 1949), and by 1954, 43
species occurred there (Newhouse, 1955). By 1963, 77 species were known
from both Johnston and Sand Islands (POBSP, 1964).

The three plant species recorded by 1923 probably reached the
atoll through natural means either by water currents, air, or birds.
A few species found since possibly also arrived via these same ways,
but the majority of the remaining 124 species has been man-made intro-
ductions. Some of these introductions were intentional, others came as
stowaways or adventives.

Original Flora
Lt. John M. Brooke (ms.) noted the following in his journal for 16

March 1859: "There is a downy creeping plant bearing a yellow flower,
and grass, these are the only representatives of the vegetable kingdom."

52

Table 3.

Family
Species
Common Name

Polypodiaceae Ferns
Polypodtum scolopendrta
Nephrolepsts sp.

Araucariaceae
Araucarta heterophylla
Norfolk Island pine

Pandancaceae
Pandanus tectortus?
Screw-pine, hala

Graminea Grasses
Cenchrus echinatus
Sandbur

Chlorts barbata
Fingergrass

Cynodon dactytlon
Bermuda grass

Dactyloetentum aegypttum
Crowfoot grass

Digttarta sanguinalts
Crabgrass

Echtnochloa erus-gallt
Barnyard grass

Eleustne tndica
Goose grass

Eragrostts tenella
(incl. amabilts)
Lovegrass

Lepturus repens
Bunch grass

A = Adventive; N = Native;

Planted;

Vascular plants known from Johnston Atoll

Islands

John-
Akau Hikina ston

A
P
iP.
A
A
A y
A A
A
A A A
A A
A N
S = Seed only

Sand

Orig.

Sand
Man-
made

Table 3. (continued)

Family
Species
Common Name

Graminea (cont.)
Paspalum dtlatatum
Dallas grass

Saccharum of ftetnarum
Sugarcane

Setarta vertictllata
Bristlegrass

Sporobolus virgintcus
Dropseed

Zea mays
Corn

Cyperaceae Sedges
Cyperus rotundus
Fimbristylts cymosa?

Palmae Palms
Cocos nuctfera
Coconut palm

Araceae
Anthurtum andraeanum
Anthurium

Liliaceae
Allium fistulosum
Welsh onion

Alltum sp.
Chives

Aloe sp.
Aloe

Cordyline fruticosa
Cordyline

Sansevterta trtfactata

Bowstring Hemp

Akau

Hikina

Islands

John-
ston

Sand
Orig.

53

Sand
Man-
made

54

Table 3. (continued)

Family
Species
Common Name

Amaryllidaceae
Crinum astatteum
Crinum sp.

Hymenocallts itttoralis
Spider lily

Bromeliaceae
Ananas comosus

Zingiberaceae
Alpina sp.
Ginger

Musaceae
Helteonta humtlts

Strelttazta reginae
Bird of Paradise

Orchidaceae Orchids
Eptdendrum sp.
Vanda sp.

Casuarinaceae
Casuarina equitsettfolta
Ironwood

Moraceae
Ficus mtcrocarpa
Banyan

Urticaceae
Pilea microphylla
Artillery plant

Polygonaceae
Coceoloba uvtfera
Sea-grape

Chenopodiaceae
Chenopodium murale
Goosefoot, Pigweed

Amaranthaceae Pigweeds
Amaranthus dubtius

Akau

Hikina

Islands

John-
ston

Sand
Orig.

Sand
Man-
made

3)P)

Table 3. (continued)

Islands
Family Sand
Species John- Sand Man-
Common Name Akau Hikina ston Orig. made
Amaranthaceae (cont.)
A. sptnosus A
A. vtridis A A A A
Nyctaginaceae
Boerhavia sp. A N N A
Bougainvillea sp. P P
Aizoaceae
Tetragonta tetragontotdes P
New Zealand Spinach
Sesuvtum portulacastrum A A A A A
Portulacaceae
Portulaca oleracea A A A A A
Purslane
Caryophyllaceae
Spergularta marina A A A A A
Lauraceae
Persea americana P P
Avocado
Cruciferae
Lobularta maritima P A
Sweet Alyssum
Rosaceae
Ertobotrya japonica P
Loquat
Leguminosae
Acaeta farnestana A A
Sweet Acacia
Crotalaria ineana A
Rattlebox
Leueaena latistltque A A
Phaseolus sp. P

Bean

Table 3. (continued)

Family
Species
Common Name

Leguminosae (cont.)
Ptsum sativum
Pea

Mucuna sp.

Pithecellobtum dulee
Manila Tamarind

Prosopts pallida
Algarobe, Kiawe

Vigna martina
Beach pea

Zygophyllaceae
Tribulus cetstotdes
Puncture Vine

Ruthaceae

Citrus auranttfolta
Lime

Citrus stnensts
Orange

Euphorbiaceae

Aleurttes moluecana
Candlenut, Kukui

Codtaeum vartegatum var.

ptetum
Croton

Euphorbta atoto?
Spurge

EZ. prostrata
Spurge

E. prob. heterophylla
Spurge

Islands

John-
Akau Hikina ston

Sand
Man-
made

Table 3. (continued)

Family
Species
Common name

Euphorbiaceae (cont.)

E. glomertfera
Spurge

E. htrta
Spurge

E. puleherrima
Poinsettia

Pedilanthus tithymelotdes
Slipper flower

Ricinus communts
Castor bean

Anacardiaceae

Mangtfera indica
Mango

Sehtnus terebinthtfoltus
Christmas berry tree

Tiliaceae

Triumfetta procumbens

Malvaceae

Htbtseus ttltaceus
Hau

Htbtscus sp.

Thespesta populnea
Milo tree, Portia tree

Stda sp.

Sterculiaceae

Waltherta tndtca

Guttiferae
Calophyllum tnophyl lum

False Kamani

Akau

Hikina

Islands

John-
ston

SY,

58

Table 3. (continued)

Family
Species
Common Name

Combretaceae
Termitnalta catappa

Indian almond, Kamani

Myrtaceae
Eucalyptus sp.

Araliaceae

Brassata actinophylla
Octopus tree

Polysetas gutlfoylet
Wild coffee

Caricaceae
Carica papaya
Papaya
Plumbaginaceae
Plumbago auriculata
Plumbago, Leadwort

Apocynaceae

Catharanthus roseus
Madagascar Periwinkle

Nerium oleander
Oleander

Plumerta acuminata
Frangipani

Plumerta rubra
Frangipani

Thevetta peruvtana var.
auranttaca

T. peruvtana(=neretfolta)
Yellow Oleander

Convolvulaceae

Ipomoea indtea

Akau

Hikina

Islands

John-
ston

Sand
Orig.

Sand
Man-
made

59

Table 3. (continued)

Islands
Family Sand
Species John- Sand Man-
Common Name Akau Hikina ston Orig. made
Convolvulaceae (cont.)
I. pes-caprae A A A
Beach Morning Glory
I. macrantha 2
Merremta tuberosa P
Wood Rose
Hydrophyllaceae
Nama sandwtcensts A
Boraginaceae
Cordia sebestena P P
Kou, Geiger-Tree
Heltotroptum curassavtcum A A A
Tournefortta argentea B P P P
Tree Heliotrope
Ver benaceae
Staechytarpheta jamaicensts A
Vitex ovata P P
Solanaceae
Capstcum frutescens P P
Papper
Nteottana glauca A
Solanum lycoperstcum P? P? P P?
Tomato
Solanum melogena P
Eggplant
Bignoniaceae
Tabebuta pentophylla Pp
West Indian Boxwood
Rubiaceae
Gardenia sp. Pp

Coprosma Sp- P

60

Table 3. (continued)

Islands
Family
Species John- Sand
Common Name Akau —_ Hikina ston Orig.
Cucurbitaceae
Citrullus lanatus var. é s
vulgaris
Watermelon
Cuecumts melo P
Muskmelon
Goodeniaceae
Seaevola taccada P P
Compositae
Btdens ptlosa : A
Burmarigold
Conyza bonartensts A A A
Emilta sonehtfolta A
Heltanthus annuus P
Sunflower
Pluchea indtea A A A
Pluchea carolinensis A A A A
Pluchea * Fosbergit A

Sonehus sp. (oleraceus x
asper)? A A
Sow-thistle

Tagetes sp. i? P
Marigold

Vernonta etnerea A
Ironweed

Zinnia elegans 12 P

Zinnia

Sand
Man-
made

61

"The nearest approach to the plant I could find in my elementary
Botany is the portulaca, the same plant grows on Laysan. I gave a
specimen to Dr. Hilebrand at Honolulu."

Brooke (Zoe. cit.) also mentioned that Capt. A.D. Piper, in charge
of the island, had "...planted potatoes and pumpkins, but they withered
away.'' Brooke gave Piper seeds of the following: watermelons, muskmelons,
squash, egg plant, pepper, asparagus, sorrel, lettuce, and leek. If Piper
planted these, their ultimate fate was the same as that of the potatoes and
pumpkins, for none survived until the next known description of the atoll's
vegetation, made 64 years later.

Members of the TANAGER Expedition found only three species of vas-—
cular plants growing on Johnston Atoll in 1923: Lepturus Perey e: Tribulus

etstotdes, and Boerhavia sp. [usually referred to as diffusa (Christo-
phersen, 1931: 4).

The grass Brooke found growing in 1859 almost certainly was Lepturus,
and the "downy creeping plant bearing a yellow flower" was most likely
Tribulus. Brooke's tentative identification of it as a Portulaca seems
unlikely in view of the lack of downiness of Portulaca and of its absence
from the atoll in 1923. It is curious, however, that Brooke did not men-
tion the armed fruits, if indeed the plant was Trtbulus. Although these
fruits are not so conscicuous in March as later in the season, old fruits
remain sharp enough to penetrate thin shoes for many months, and certainly
should have been present and noticeable to Brooke.

If Boerhavia were present in 1859, it apparently was not abundant
enough for Brooke to notice it readily. While it seems unlikely that
such a plant could be missed if it were present in anything near the
abundance it was found in 1923, it seems equally unlikely that such an
easily distributed species would have to await the help of man to reach
Johnston Atoll.

The ambiguity of this situation leaves us in doubt as to whether the
original vegetation of the atoll consisted of two or three species. In
either case, it is, as Fosberg (1949) pointed out, one of the smallest
floras known. All three plants in question have propagules that could be
transported easily by birds by sticking to or in the plumage (Boerhavia
and Lepturus) or by sticking into the feet (%rtbulus).

It is not surprising that the cultivated plants Brooke and Piper
introduced to the atoll died out, but it is remarkable that none of the
common weedy species from other parts of the world became established
from ship's cargoes. It would be most interesting to know if any of
these were indeed introduced prior to 1923, and if so, how long they per-
sisted. Perhaps this situation is evidence that many common weeds require
continued disturbance by man to assure their persistence.

Recent Vegetation

In the years since 1923, the atoll's flora has been enriched by
numerous introductions, both intentional and otherwise. Wetmore (ms.b)

planted eight "clips" of Hau (Hibiscus ttliaceus) 18 July 1923. These

may not have survived for the species was not included on the next list

of the flora, compiled by Bryan (ms.b) in 1944. This list showed 29

species of native and introduced plants. The next published list was that of
F.R. Fosberg (1949) based on collections made 1 November 1946 by his brother,
K.P. Fosberg. This showed 27 species, including eight not listed by Bryan
in 1944, bringing the total known flora to about 35. Doty and Newhouse

(ms.) summarized previous findings, including additional data from Fosberg,
together with results of a study of the vegetation fostered by Lt. Col.

Jack Bently, Base Commander, in 1953 in which 44 additional species were
found. Of these 44, 20 were grown only in yards under care of residents.
About 51 species apparently grew untended then, with an additional 18

tended, for a total flora of 69 species.

POBSP studies, principally by A. Binion Amerson, Jr., Charles H.
Lamoureux, and Philip C. Shelton, added 48 species to the Johnston Atoll
flora. Included were the first collections from the two new islands, Akau
and Hikina.

Island Accounts

Appendix Tables 2 through 6 present annotated lists of all vascular
plant species known to have occurred on each island at Johnston Atoll,
with the exception of introductions that have failed, such as some of the
cultivated species introduced by Brooke and Piper, and certainly many
others.

Land clearing and dredge fill have increased the size of the two
original islands and have made two new islands, thus bettering the op-
portunity for more plant species. Disturbed soil together with freedom
from competition by an established flora have provided conditions suitable
for many kinds of plants. But some plants have not been able to survive
the poor soil and climate conditions of Johnston. Thus, many of the
ornamental species intentionally introduced by man would not survive if
not frequently cared for by man.

Because of human disturbance, the plants occurring on the four
islands are different in species composition.

Akau Island: There are 38 species of vascular plants known from
Akau Island. This island has a much larger flora than Hikina Island,
both of which were built at the same time, partly because of its closer
position to Johnston and Sand Islands, but more importantly because of
more interest in plants by workers on Akau than on Hikina. Most of
Akau's flora consists of species transplanted from Johnston Island. Six
species recorded from Akau, however, are not known from the other three
islands.

This man-made island was completed in 1964; by September 1967, 31
species were growing there. Sixteen of these 31 were obvious introductions;
the remaining 15 were considered to be adventive and not intentionally
planted. Only a few of each of the introduced species existed, but at

63

least five of the adventive species were common. Fimbristylts sp. grew
over most of the island and was the most dominant species. Spergularis
marina was widespread, and small plants of Sesuvium portulacastrum were
scattered about. FEleustne tndtca was fairly abundant in a few small areas.
Two patches of Cynodon dactylon, one about 30 feet in diameter, grew in a
slight depression on the north-central portion of the island (POBSP, 1967b).
Similar plant distribution existed in November 1973.

Hikina Island: Only 14 species of vascular plants have been recorded
from Hikina Island (Table 3). All are known from at least one of the other
three islands; three are known from seed only.

The island was completed in 1964 and by September 1967 five species
of growing plants and seeds of three species were recorded. Two of the
five growing species were represented by one specimen each; both were
obvious introductions. Fimbristylts sp. was the dominant plant and oc-
curred scattered on almost the entire island. Sesuvitum portulacastrum was
the second most abundant species and grew in clumps, the largest of which
was 6 by 9 feet. Spergularta marina, the third ranking species, was widely
scattered over the island, but most plants appeared to be dead. The three
seeds were all on the beach (POBSP, 1967b).

The island was revisited in February 1969 and two additional species
of grass were recorded. The flora looked about as it did in September
1967, except that the Fimbritstylts was more evenly spread, although little
changed in density (POBSP, 1969). The flora was the same in November 1973,
except for a large Pluchea bush growing in the north-central portion.

Johnston Island: In 1923, as mentioned earlier, only three plant
species were known (Christophersen, 1931). Wetmore's 1923 photographs
show Lepturus repens as the dominant species. U.S. Navy aerial photographs
taken in 1935 and 1939 (see Figs. 3 and 5) reveal vegetation growing over
the entire island; another taken in May 1941 shows original vegetation
everywhere except around the buildings on the northeast corner and in an area
north to south across the center. By 1942, another aerial photograph (see
Fig. 6) shows no vegetation at all and numerous buildings.

Twenty-seven plant species were recorded in 1946; these consisted
mainly of obvious introductions and accidental weeds. No original vegeta-
tion remained; the whole island was occupied by runways and buildings with
disturbed ground in the open places and along paths and roadsides (Fosberg,
1949).

In January 1954, Newhouse (1955) found 43 plant species; 20 were not
under cultivation and 23 were under cultivation in dependents’ homes and
gardens. Three plant species recorded in 1949 were not observed in 1954.

By 1967, 111 plant species had been recorded from Johnston Island
(Table 3). Of these 111, 3 were native, 61 were planted, 44 were adventive,
and 3 of questionable status. In addition, 56 of the 111 were not known
from the other three islands.

64

Continued disturbance on Johnston Island in 1967 prevented any
meaningful development of vegetation except in a few scattered locations
near the runway. Pluchea carolinensis, which at times has been kept
trimmed, had developed conspicuous stands on the south side of the runway,
especially northeast of the Weather Bureau building; nesting Red-tailed
Tropicbirds were especially attracted to these stands. Only one other
species, Cenchrus echtnatus or sandbur, was considered to be fairly
abundant on open disturbed areas. Other species ranged in number from
individual plants to widely scattered, but not abundant plants (POBSP,
1967b).

In November 1968, most of the Casuarina equtsettfolta, or ironwood
trees, were removed from atop the underground hospital, thus destroying
some of the prime White Tern roosting and nesting habitat (POBSP, 1968).
By November 1973, Casuarina trees near the tennis courts were supporting
nests of both White Terns and Black Noddies.

Sand Island: A total of 54 species of vascular plants has been
recorded from Sand Island (Table 3): 3 were native species, 23 were
planted, 26 were adventive, and 2 occur only as seeds. Nine species
found here are not known from the other three islands.

Original: Three species--Lepturus repens, Boerhavia repens,
and Trtbulus ctstotdes--were recorded in 1923 (Christophersen, 1931).
Ground-level photographs taken in 1923 by Wetmore reveal L. repens to be
the dominant species. U.S. Navy aerial photographs taken during the mid-
1930's (see Fig. 3) show the island completely covered by vegetation.
The arrival of a permanent military station in 1939 (see Fig. 8) signaled
destruction of the original flora; by March 1940 (see Fig. 9) vegetation
remained only on the north peninsula.

POBSP personnel recorded 11 species of plants here in 1963. Some 80
percent, or approximately 400,000 square feet, of the original portion, was
covered by vegetation. The five most dominant species (Fig. 30) were, in
order of importance, Lepturus repens, Trtbulus ctstoitdes, Sesuvtum
portulacastrum, Boerhavia repens, and Amaranthus virtdts. The first three
species covered 84 percent of the vegetated area (Lepturus 50 percent,
Tribulus 30 percent, and Sesuvtum 4 percent). The eight other species
filled the remaining 16 percent and were primarily scattered over bare
soil and rock (POBSP, 1964).

Seasonal climatic conditions and human disturbance often affect
the islands' flora. For example, in early February 1967 the vegetation
appeared drier than it had in October 1966, although several species had
spread noticeably during the winter. At least two species, Boerhavia
repens and Tribulus etstoides, were reduced in numbers. Heavy rains in
February and March brought the growth up to the October 1966 level; there
was only a slight decline in greenness by early May. By late August and
September, when the flora was usually at its lowest growth, the vegetation
appeared at least as vigorous and green as in May, and much more so than
at the same time in 1966.

65

By 1967, 25 plant species were known from the original portion of
Sand Island (Table 3). Of these 25, 3 were native, 2 were planted, 18
were adventive, and 2 occurred as seeds. Plant distribution as of late
1967 (Fig. 31) and late 1968 was similar to that in late 1964 except that
Boerhavia was more widespread on the eastern half (POBSP, 1967b, 1968).
Sparse plants were observed in November 1973, most likely resulting from
low spring rainfall coupled with large numbers of Sooty Terns.

Man-made: The man-made portion was started in late 1939 and by
May 1941 was nearly completed. A U.S. Navy aerial photograph taken in
February 1942 from a height of 600 feet reveals no vegetation.

POBSP personnel recorded 19 species of plants here in 1963; 9 of
these 19 were not growing on the original portion (POBSP, 1964).

By 1967, 50 plant species had been recorded (Table 3). Of these 50,
23 were obviously introduced, and 27 were probably adventive. In addition,
29 of the 50 species were not growing on the original portion, and seven
were not growing on the other three islands in the atoll.

In 1967, pioneer adventive species including Fimbristylis, Conyza,
Sonehus, Cenchrus, and Pluchea were increasing on the newest coral,
dredged in 1964. Other species, including Cynodon, Sesuvtum, and
Euphorbia htrta, were replacing the above first four on the older dredged
portion. Sceaevola continued to grow well around the buildings in 1968
and 1969, thus providing nesting habitat for Red-tailed Tropicbirds
(POBSP, 1967b, 1968, and 1969). A similar distribution was found in
November 1973.

Invertebrates

The invertebrate fauna of Johnston Atoll is not well-known and while
scattered collections have been made, no extensive systematic sampling
programs have been conducted. Most of the published lists appear to be
spotty and the ecological aspects of the zoogeography have been largely
neglected.

Published reports and museum collections center around the macro-
invertebrates of the major phyla Cnidaria, Mollusca, Annelida, Arthro-
poda, and Echinodermata. Of these phyla, even the corals--the most
conspicuous group of the atoll marine fauna--are not well studied.

There are little or no data available on other groups such as the
sponges (Porifera), the acoelomate bilateria (e.g., Platyhelminthes),
the pseudocoelomates (e.g., Aschelminthes), the minor coelomate proto-
stomes (@.g., Priapulida, Sipunculida, Echiurida), the lophophorate
coelomates (e.g., Phoronida, Ectoprocta, Brachiopoda), and the minor
deuterstome phyla (e.g., Pogonophora, Hemichorda).

The invertebrate fauna (especially the Mollusca) known from Johnston
Atoll, however, indicates that Johnston may serve as a unique filter
bridge between the south-central (Polynesian) and western (Micronesian)
Pacific fauna and the Hawaiian fauna.

66

: Lepturus repens

© Tribulus cistoides

e Sesuvium portulacastrum

Boerhavia sp. N
Amaranthus viridis
0 100 200
== |
Feet

Figure 30, Distribution of the five most important plant species on the
original portion of Sand Island, Johnston Atoll, July—August 1963.

== Lepturus repens ; e Scaveola taccada

oO Tribulus cistoides @ Ipomoea indica

e Sesuvium portulacastrum © Vigna marina

© Boerhavia sp. Sonchus sp.

* Amaranthus viridis @ Pluchea carolinensis

ew Ipomoea pes-caprae @© Tournefortia argentea N

vw Portulaca oleracea

4 Cynodon dactylon

||| Cenchrus echinatus, Eleusine indica, & Setaria veticellata
0 100 200
eed

Feet

Figure 31. Distribution of dominant plant species on the original portion
of Sand Island, Johnston Atoll, May 1967.

67

Cnidaria (Coelenterata)

Eighteen species in 11 genera of Cnidaria (hydras, jellyfish, sea
anemones, and corals) are known from Johnston Atoll (Table 4).

The distribution and abundance of the seven genera of coral reported
by Brock, et al. (1965, 1966) are presented in Table 5. Collection sta-
tions are shown in Figure 32. Two genera are quite restricted, occurring
only in areas 1 and 2 near the marginal reef. The other five genera are
distributed widely throughout the lagoon.

Dredging operations in 1964 destroyed over 1,100 acres of reef;
dredging itself destroyed 700 acres of living coral while newly deposited
"coral aggregate'' destroyed more than 400 acres. In addition, over 7,000
acres of reef and lagoon were more or less seriously affected by silty
water. The reduction in the percentage of living coral in these silty
water areas varied from none to 40 percent, with 10 percent being roughly
an average figure. A parallel reduction in the number of associated
invertebrate species and fish species also occurred.

JOHNSTON ISLAND AND REEF
|

oss"

1645'

.
oe
.

Legend: Black - recently formed land
areas. Cross hatch - areas of reef
altered by recent dredging. Dots -
areas of lagoon often covered by
turbid water. Dense dots - area of
lagoon and sea covered by very turbid
water. Numbered dots - previous
stations. Numbered arrows — new
stations (transects) studied.

6
| CARLA tata.

30°

Figure 32. Johnston Atoll showing invertebrate and fish collection stations,
dredged areas, and the extent of silt laden water (Brock, et alc,
1966).

68

Table 4. Cnidaria (Coelenterata) from Johnston Atoll*

Class Wells Brock Present
Family 1934 eb al, Paper
Species 1965

Hydrozoa
Milleporidae
Mtillepora tenera x
Mtllepora sp. X
Stylasterinidae
Dtsttehopora sp. X
Stylaster sp. x

Anthozoa

Pocilloporidae
Poetllopora damicornis x
Poetllopora eydouxt X
Poetllopora meandrina X

Acroporidae
Acropora humtilts
Acropora hyacinthus
Acropora retusa
Aeropora tumtda
Monttpora verrucosa
Monttpora sp.

Agariciidae
Leptastrea sp.
Pavona vartens X
Pavona sp.

Fungiidae
Fungta scutarta x

Poritidae
Portites lutea X

Isopheliidae
Telmatactts decora 2 X

PIP PS PS Pt Pd

P<

P<

*Taxonomic order follows Bayer, et al. (1956).

69

Table 5. Distribution and abundance of corals at Johnston Atoll,

1964-1965*
3
Cd) —
0) r= 3 3 9
Se an Oe ir ee
Qa xo ° v © hag autos
QO 4 2) DS we » is) Sy aS)
oe ae oe ke
Areas <x SSS re ee Percent of living coral
1 (1964) a Gina 4 DED, NaS Cag 6 Mound tops 70 to 100%
Lower sides 40 to 50Z
2 (1964) XE fo XO ue ee Ka OK Ridge tops 99%
Channel floor 17%
3 (1964) ED, SER 6 x > i oe 80 to 90%
4 (1964) Krak >; ui ied) Cane ¢ 20% or less
4 (1965) Ke Ke eum aXe 10 Eo” 207
Se Clo 64) x x DL RS 6 90 to 95%
5. anGl965)) Xo ox x oe ok 50. t0 757
6 (1964) 5 eau Coige 5 cea pilin 50 to 75%
6 (1965) X Xx X X X xX SOE 07
7 (1964) ek ONT ae Ky uP KE th OR 20%
7 (1965) 5 Se ee haa eae 15%
8 (1964) Gi aX Sie eK 20 to 30%
8 (1965) eek Tage io: aI 15 to 20%
9 (1964) xX Sia x 50 to 60%
10 Reef **¥ oe > aaa 10%
TO MGITEE 0%
10 Bottom 0%
11 Reef X xX xX xX Xx 60 to 70%
PA Glae £ 0%
11 Bottom xX SOR,
12 Reef xX X 110) jee) LOY,
RASS CE 0%
12 Bottom 0%
13 Reef X X X xX X 110) {2@) IS YZ
ils) (Gulatiene 0%
13 Bottom 0%
Number of X's 15eel6ers 1 LEwOUOY \6ae2 Oz

*Adapted from Brock, et al. (1965, 1966).
**Areas 10 through 13, 1965 only.

70

Mollusca (by Harald A. Rehder)

The 58 species listed in Table 6 comprise only those mollusk species
found in the collection of the National Museum of Natural History and by
no means represent the complete fauna of this atoll. A thorough survey of
the lagoon and a search of the reef and shores of both Sand and Johnston
Islands for the smaller species would increase the list materially.

As might be expected, most of the species are widely distributed
throughout the Indo-Pacific region. Six, however, are otherwise restricted
to the Hawaiian Islands; these are Trochus intextus, Turbo articulatus,
Nertta ptcea, Cypraea granulata, Maculotriton species, and Pertsternta
cerocea. Two others, Strombus maculatus and Cypraea schitlderorum, are found
in Micronesia and Polynesia, as well as in the Hawaiian Islands, but not
elsewhere.

On the other hand, 12 of the species listed are uncommon to rare in
the Hawaiian Islands, and three--Planaxis zonatus, Mitra columbelltformis,
Parvtperna dentifera--have not as yet been recorded from Hawaii.

In brief, although the molluscan fauna of Johnston Atoll shows an
affinity with that of the Hawaiian Islands, it is also closely related to
the fauna of Micronesia and Polynesia. A more thorough survey of the
mollusks of Johnston Atoll should help to clarify its biogeographical
position.

Annelida

At present only 12 species belonging to eight families of Polychaeta
are known from the lagoon waters at Johnston Atoll (Table 7). Edmondson,
et al. (1925) listed six species collected during the 1923 TANAGER Expe-
dition. Brock, et al. (1965) reported three species from 1963-1965
collections. The present paper lists six species found in the collections
of the National Museum of Natural History and the Bernice P. Bishop Museum.
Additional collecting will most likely result in an increased number of
known species from the atoll.

Brock, et al. (1965, 1966) showed that dredging at Johnston slightly
increased the number of polychaetes, but reduced their weight.

Arthropoda

Marine

The lagoon waters at Johnston Atoll support a total of 75 species
belonging to 20 families of Crustacea (Table 8). Edmondson, et al. (1925)
reported 59 species collected by the 1923 TANAGER Expedition, while Brock,
et al. (1965, 1966) reported 12 species collected from 1963 through 1965.
The present paper lists 61 species found in the collections of the National
Museum of Natural History and the Bernice P. Bishop Museum. Further col-
lecting at Johnston Atoll possibly would result in additional known species.

Al

Table 6. Distribution and abundance* of Mollusca from Johnston Atoll

Marginal Johnston Sand Lagoon fill
Gastropoda: N.W. Reef Island Island Sand Island

Trochidae
Trochus tntextux Reeve U

Turbinidae
Turbo arttculatus Reeve M

Neritidae
Wertta plitcata Linnaeus M
Nerita poltta Linnaeus
Werita albtetlla Linnaeus
Nerita ptcea Recluz V

<aC8 8

Littorinidae
Littortna pintado Wood
Littorina undulata Gray
Littortna cocetnea (Gmelin)

(=) (=

Planaxidae
Planaxts zonatus A. Adams U

Modulidae
Modulus tectum (Lamarck) U

Cerithiidae
Rhinoelavits sinensts (Gmelin) U
Rhtnoelavts arttculatus Adams U
& Reeve
Certthtum mutatum Sowerby U
Certthium nestotteum Pilsbry U
& Vanatta

Hipponicidae
Sabita conica (Schumacher) U U

Strombidae
Strombus maculatus Sowerby U U

Cypraeidae

Cypraea granulata Pease U
Cypraea helvola Linnaeus U M
Cypraea poraria Linnaeus U
Cypraea caputserpentts Linnaeus

Cypraea moneta Linnaeus U
Cypraea macultfer (Schilder)

Se} eS

*V = Very abundant; M = Moderately abundant; U = uncommon.

Table 6. (continued)

Marginal Johnston Sand Lagoon fill
Gastropoda (cont.): N.W. Reef Island Island Sand Island

Cypraeidae (cont.)
Cypraea tsabella Linnaeus
Cypraea carneola Linnaeus
Cypraea schilderorum (Iredale) U

sas

Naticidae
Polintces (Mamtlla) melano- U
stoma (Gmelin)

Cymatidae

Cymattum (Septa) ntcobarteun U
Roding

Cymattum (Septa) aquattle (Reeve)

Cymatium (Septa) gemmatum (Reeve)

Cymattum (Ranularta) murtcinum
Roding

Distorsto anus Linnaeus ; U

SVE eS

Tonnidae
Tonna (Quitmalea) pomum (Linnaeus) U

Muricidae
Maculotriton species
Drupa morum Roding
Drupa ricinus (Linnaeus)
Morula uva Roding
Morula granulata (Duclos)
Drupella ochrostoma (Blainville)
Nassa sertum Bruguiere M

Sans
os.

asad

Coralliophilidae
Coralltophtla vtolacea Kiener M
Quoyula madreporarum (Sowerby) M

Buccinidae
Ptsanta ignea (Gmelin) U U

Nassariidae
Nassartus (Retitcunassa) der- U
mestina (Gould)

Fasciolariidae M
Peristernia crocea (Gray)

Mitridae
Mitra (Strtgatella) colum- U
betltformts Kiener

13

Table 6. (continued)

Marginal Johnston Sand Lagoon fill
Gastropoda (cont.): N.W. Reef Island Island Sand Island

Turbinellidae
Vasum turbinellus (Linnaeus) U U U

Conidae
Conus pulteartus Hwass
Conus nanus Sowerby
Conus rattus Hwass
Conus vttultnus Hwass
Conus miles Linnaeus
Conus flavidus Lamarck U

Seq ee

Terebridae
Terebra crenulata Linnaeus U

Bivalvia:

Isognomonidae
Tsognomon perna (Linnaeus) M
Parviperna denttfera (Krauss) U

Trapezidae
Trapezium oblongum (Linnaeus) U

Tellinidae
Arcopagta (Seutarcopagia) U
scobtnata (Linnaeus)

Table 7. Annelida from Johnston Atoll*

Class Edmondson Brock
Family CEuat. Cbrals Present

Species 1925 1965 Paper

Polychaeta
Amphinomedae
Eurythoe complanata (Pallus) x
Eurythoe pactfitca Kinberg X
Hermodtce pinnata Treadwell X
Cirratulidae
Ctrratulus sp. X X
Eunicidae
Eunice sp. x
Polynoidae
Hololeptdella ntgropunctata (Horst) X
Phyllodocidae
Phyllodoce stigmata Treadwell X X

74

Table 7. (continued)

Class Edmondson Brock
Family Be Cubs et al. Present
Species 1925 1965 Paper
Nereidae
Nerets kobtensts xX
Pertnerets hellert (Grube) X xX
Leodicidae
Lystdtce fusca Treadwell X
Lystdice sp. x
Leodocidae
Leodtce sp. X

*Taxonomic order follows that in the Annelida collection of the National
Museum of Natural History.

Table 8. Marine Arthropoda from Johnston Atoll*

Class Brock
Subclass Edmondson Cb Gls
Family ebrak. 1965), Present
Species 1925 1966 Paper
Crustacea
Cirripedia
Lepadidae
Lepas anatifera Linnaeus X
Malacostraca

Squillidae [=Chloridelidae?]
Pseudosquilla oculata (Brullé)
Palaemonidae
Coralltoecarts graminea (Dana)
Harptltopsts depressus (Stimpson)
Jocaste lucina (Nobili)
Palaemonella tenutpes Dana
Peridemenaeus trtdentalus (Miers)
Gnathophyllidae
Gnathophyllum americanum Guerin
Alpheidae
Alpheus brevipes Stimpson X
Alpheus bucephalus Coutiére
Alpheus elypeatus Coutiere
Alpheus collumianus Stimpson
Alpheus crasstmanus Heller
Alpheus dtadema Dana
Alpheus gracilts Heller
incl. subsp. stmplex (Banner)
Alpheus levtuseulus Dana

PP OPS PS PS PS PS PS PS PX PS PS ri
a P<

PS PS PS PS PS OPS

a
rs

75

Appendix Table 8. (continued)

Class Brock
Subclass Edmondson Cink
Family et al. 1965, Present
Species 1925 1966 Paper

Alpheidae (cont.)
Alpheus lottint Guérin
Alpheus paracrinitus Miers
Alpheus paragracilis Coutiére
Synalpheus paraneomeris Coutiére
Hippolytidae
Lysmata paucidens (Rathbun) x X
Saron marmoratus (Olivier) X
Painuridae
Panultrus marginatus (Quoy & Gaimard) X
Panulirus penctllatus (Olivier) a8 x
Scyllaridae
Parrtbacus antareticus (Lund) X Ki
Axiidae
Axtopsts johnstont Edmondson x x
Galatheidae
Galathea spinosorostris Dana 4
Diogenidae
Aniculus aniculus (Fabricius)
Caleinus elegans (N. Milne-Edwards)
Caletnus herbstit de Man
Caleinus latens (Randall)
Dardanus haanit Rathbun
Dardanus megistos (Herbst)
Dardanus punctulatus X
Dynomenidae
Dynomene hisptda Desmarest X X
Calappidae
Calappa hepatica (Linnaeus) X
Leucosiidae
Nyeta spectosa Dana x
Majidae
Perinea tumtda Dana Xx
Sehtzophrys hilensts Rathbun Xx
Portunidae
Cataptrus tnaequalts (Rathbun)
Portunus longtspinosus (Dana)
Thalamita admete (Herbst)
Thalamitotdes qutdridens A. Milne-
Edwards
Xanthidae
Carpiltus convexus (Forskal)
Chlorodtella asper Edmondson
Domeeta hispida Eydoux & Souleyet
Etisus electra (Herbst)

Pi Ps PS PX
rd Pt Pd

Pd PS PS PS

PPS PS Pd Pd

Pd PS PS Pd
Pd PS Pd Pd

PI Pd PS OPS
ms
PS PS PS PS

Table 8. (continued)

Class Brock
Subclass Edmondson (HE Oilbn
Family Ctl. 1965, Present

Subspecies 925 1966 Paper

Xanthidae (cont.)
Leptodius sanguineus (H. Milne- X x
Edwards)
Leptodtus wataluanus Rathbun
Ltocarptlodes btunguts (Rathbun)
Liocarpilodes integerrtmus (Dana)
Ltomere bella (Dana) X
Lophozozymus dodone (Herbst)
Phymodius laysant Rathbun x
Phymodtus ntttdus (Dana) X
Pilodius aberrans (Rathbun) xX
Pilodius areclata (H. Milne-Edwards)
Platypodta eydouxt (A. Milne-Edwards) X
Pseudoltomera spectosa (Dana) x 2
Tetralta glaberrima (Herbst) - Xx
Tetralia spp.
Trapezta cymodoce (Herbst)
Trapezta digttalts Latreille
Trapezta ferruginea Latreille
Trapezta tntermedia Miers
Trapezta maculata (MacLeay) X
Trapezta rufopunctuta (Herbst) x
Trapezta spectosa X x
Ocypodidae
Ocypode laevis Dana Xx
Grapsidae
Grapsus strigosus (Herbst)
Grapsus tenutcrtstatus (Herbst)
Pachygrapsus minutus A. Milne-
Edwards
Pachygrapsus pltcatus (H. Milne- X
Edwards)
Hapalocarcinidae
Hapalocaretnus marsuptalts X X
Stimpson
Pseudoeryptochtirus crescentus X X
(Edmundson)

DM PS PS PS PS OS OO OP OX

PM PS PS
P< PS Pd
mM PS PS OPS

rr PX
MP PX

*Taxonomic order follows Chase (pers. corres.).

Tih

Very little is known about the distribution and abundance of
Johnston's marine arthropods. Brock, et al. (1965, 1966), in studying
the effects of dredging, did reveal that dredging reduced the number and
weight of xanthid crabs and alpheid shrimp inhabiting living cor: .
(vasiform Acropora) heads, but showed little change in crab and shrimp
populations inhabiting living coralline (Poroltithion) algae.

Terrestrial

Insects totaling 68 species of 35 families were known from the islands
aeedonnston Atoll by 1952 (Bryan, et al... 1926; Chillson,, 1953). These are
listed in Table 9. POBSP personnel collected insects at Johnston from
1963 through 1969; these collections have not, as yet, been fully identified.

Table 9. Insects recorded from Johnston Atoll; adapted from Chilson (1953)

Orthoptera
Blattidae
Blattela ltturtcollts (Walker)
Cuttlta soror (Brunner)
Pertplaneta amertcana (Linnaeus)
Pyenoscelus surtnamensts (Linnaeus)
Dermaptera
Labiduridae
Antsolabts maritima (Gene)
Euborellia annultpes (Lucas)
Mallophaga
Menoponidae
Austromenopon sternophtlum (Ferris); on tern.
Thysanoptera
Aeolothripidae
Franklintella sulfurea Schmutz
Hemiptera
Lygaeidae
Nystus terrestris Usinger
Geocorts puncttpes (Say)
Reduviidae
Zelus renardit Kolenati
Nabidae
Nabts capstformts Germar
Gerridae
Halobates sertceus Eschscholtz
Homoptera
Aphididae
Aphts gossyppt Glover
Aphts medteagtnts Koch
Margarodidae
Iecerya purchast Maskell
Pseudococcidae
Pseudococeus (cttrt complex)
Pseudococcus sp. perhaps eitrt (Risso)
Ferrtstana virgata (Cockerell)

78

Table 9. (continued)

Homoptera (cont.)
Coccidae
Coceus sp.
Coceus hespertdum Linnaeus
Satssetta nigra (Nietner )
Satssetia oleae (Bernard)
Diaspididae
Asptdtotus lataniae Signoret
Chrysomphalus dtetyospermt (Morgan)
Pinnaspts sp.
Pinnaspts strachant (Cooley) (of Ferris and Rao)
Neuroptera
Hemerobiidae
Sympherobius sp. may be barbert Banks
Lepidoptera
Tineidae
Tineola uterella Walsingham
Ereunetts incerta Swezey
Pterophoridae
Trichopttlus oxydactylus (Walker)
Phalaenidae
Achaea janata (Linnaeus)
Laphygma exempta (Walker)
Coleoptera
Dermestidae
Dermestes ater Degeer
Histeridae
Caretnops quattuordecimstrtata (Stephens)
Anobiidae
Lastoderma serricorne (Fabricius)
Tenebrionidae
Alphitobtus ptceus (Oliver)
Coccinellidae
Coelophora tnaequalts (Fabricius)
Seymnus loewit Mulsant
Seymnus notescens Blackburn
Curculionidae
Dryotrtbus mimettcus Horn
Macraneylus tmmigrans (Perkins)
Hymenoptera
Encyrtidae
Aenastus advena Compere
Leptomasttx dactylopit Howard
Formicidae
Solenopsts geminata rufq (Jerdon)
Monomorium pharaonts (Linnaeus)
Cardtocondyla sp.
Tetramortum guineense (Fabricius)
Paratrechina (Nylanderta) sp.
Paratrechina longicornts (Latreille)

19

Table 9. (continued)

Hymenoptera (cont.)
Sphecidae
Chalybion bengalense (Dahlbom)
Vespidae
Polistes fuscatus aurifer Saussure
Megachilidae
Megachile fullawayt Cockerell
Diptera
Syrphidae
Simosyrphus (Xanthogramma) grandicornis (Macquart)
Xanthogramma scutellaris (Fabricius)
Syrphus sp.
Sarcophagidae
Gontophyto bryant Lopes
Sarecophaga sp.
Sareophaga dux Thomson
Sareophaga barbata Thomson
Calliphoridae
Phaenteta sp.
Muscidae
Musca domestica Linnaeus
Musea domestica vicina Macquart
Athertgona exctsa (Thomson)
Milichiidae
Desmometopa sp.
Agromyzidae
Agromyze pustlla Meigen
Hippoboscidae
Olfersta spintfera (Leach); from frigate birds.

Medically Important Species: Medically important arthropods have,
however, been studied (POBSP). These include 2 ticks, 5 chiggers, 2 nasal

mites, 23 biting lice, and 2 louse flies. No fleas or mosquitoes are known
from Johnston Atoll.

Ixodides: Two ticks, Ornithodoros capensts and Orntthodoros denmarki
(family Argasidae), are known from Johnston Atoll. A third tick, [yodes

Laysanensts (family Ixodidae), occurs on islands in the north-central
Pacific, but has not been recorded from Johnston Atoll. It is expected,
however, to eventually reach Johnston Atoll because of bird movement between
islands in the north-central Pacific. Another tick, [rodes amersont, which
occurs on islands in the south-central Pacific, is not expected to reach
Johnston Atoll because of a lack of bird movement between islands in the
north-central and south-central Pacific (Amerson, 1968).

Philip (1965) reported that two viral isolates were made from
Orntthodoros capensis Neumann taken from Sooty Tern and Brown Noddy nests
at Sand Island. He stated that this arbovirus may be related to ''Hughes
virus" which was isolated by Hughes, et al. (1964) from Orntthodoros

80

denmarkt taken in tern nests on Bush Key, Dry Tortugas, Florida. The
arbovirus was later named "Johnston Atoll virus" by Clifford, et al.
(1968), who noted that in addition there were also two unnamed viruses
from seabird ticks taken at Sand Island, Johnston Atoll.

In order to obtain a better understanding of the relationship
between Orntthodorns capensts and its hosts, a program was started in
January of 1965 by the POBSP to monitor the tick population on Sand
Island, Johnston Atoll, each month. Berlese samples were taken (using
a 5" x 5" x 1" metal sampler) at random over the entire island which had
been divided into sections (Fig. 33).

Berlese samples for 1965 totaled 515 for the entire island and 159
of these were positive for 0. capensts; the infestation rate was 31 percent.
Since the original part of Sand Island is the most important to the avi-
fauna, 479 of the Berlese samples were taken from this part. The investa-
tion rate for the original part during 1965 was 33 percent and Table 10
presents the variation in each of the eight areas. The infestation rate was
definitely highest (51 percent) on the northeast peninsula (Areas H and K)
and diminishes gradually toward the western part of the island. This is
shown in Figure 33 by the distribution of the positive samples in the
different areas. :

Table 10. Berlese sample data--Ornithodoros capensis infestation rates
for Sand island, Johnston Atoll, 1965.

Areas
A B C D E F H K Total
No. samples negative if Zee 58 57 48 44 43 32
No. samples positive 4 4 4 5) 6: p35 46 44 158
Total samples ie 2 a7, 637 = 773.) 165 90° . 87 479
Percent positive 364 167 Show 04 220, V42- 5 ol 33%

This variation in infestation rates in the different areas could be
due to several reasons: (1) ecological differences in the various areas,
(2) bird distribution over the island, and (3) man's disturbance of the
island. Ecological differences in the various areas are few since the
vegetation and soil is similar throughout the island and the island itself
is small. As will be shown later, the bird distribution, especially at the
peak population, covers the entire original portion of the island. The
USCG LORAN Station was built in 1960 and during construction all old
buildings, etc., left from earlier activities, were bulldozed into the
surrounding lagoon. This activity undoubtedly disrupted the soil and
thus in turn may have destroyed any ticks which were present. The penin-
sula, being further away from the center of the disturbance, was not severely
disrupted. This was verified when photographs taken in 1923 were compared
with 1963 photographs. It is known that ticks were very numerous on Sand
Island as early as 1923 (Wetmore, ms.a).

Bi-monthly infestation rates for Sand-Johnston (Table 11) reveal that
there was an increase in the rate from January-February (32 percent) through

81

May-June (43 percent) and then a decrease through the fall months to a

low of 17 percent in November-December. This seasonal fluctuation is

shown in Figure 34 to be correlated with the seasonal population of all

the breeding birds, mainly Sooty Terns, on the island. Briefly, the 1965
breeding cycle of the Sooty Terns was as follows: late October (1964)--

few observed; November (1964)--sharp increase in night calls; December
(1964)--increase in numbers, first roosting on 14th; late January (1965)
--170,000 birds at night, only a few during day; February--thousands sitting
on the island by the 10th, 25,000 incubating by the 15th, 40,000 by end of
the month; mid-March--peak egg count of 140,000; April--82,000 chicks hatched
by the 9th, but numbers dropped to 65,000 due to mortality; May--20,000
young were flying; June--well over 50 percent of young were flying and large
numbers left the island, only 50 eggs remained; July--most of the young left
the island; August--17,000 first half, 2,700 by end of month; September--
few birds remained; October--one chick present, not over 10 adults a day;
November--night calling increasing; December--by mid-month birds began
setting down on the island, late month roosting totaled 50,000 birds. The
area occupied by the Sooty Terns is shown bi-monthly in Figure 35.

Increase in the population and breeding activity of the Sooty Terns
during the spring months directly influenced the tick population. The
number of ticks per positive sample also increased during this period
(Table 11) as follows: January, February--3.5; March, April--9.4; May,
June--8.0. During the summer and fall months the number of ticks per
positive sample showed a decrease and then an increase as follows: July,
August--4.0; September, October--18.0; November, December--6.7. The in-
crease in numbers during September and October is possibly due to low
sample size (5) and the fact that samples were purposefully taken in
places where ticks were plentiful (i.e., under rocks). The overall
average for 1965 was seven ticks per positive sample.

The bi-monthly infestation rates for the three major areas (north-
east peninsula, east half, and west half) of Sand Island are shown in
Figure 33 and Table 11. These rates show that the tick populations in
the different areas fluctuate similarly to that for the entire island
(Fig. 34), as all three areas reach a peak in the May-June, July-August
period. There is, however, a variation in the infestation rates in each
area for each bi-monthly period. This again shows that the northeast
peninsula area not only has a higher rate overall, but the rate was higher
each bi-monthly period than the other areas. For instance, during May-
June, the rate on the peninsula was 89 percent, while on the east portion
it was 49 percent, and on the west portion it was 18 percent. As has already
been pointed out (and shown in Figure 35), the Sooty Terns and other species
utilize almost all of the available land areas during the spring and summer
months, so the bi-monthly infestation rate variation for the different areas
was not due completely to the number of birds using any particular area of
the island.

The 515 Berlese samples taken on Sand-Johnston in 1965 are shown by
host-source in Table 12 and demonstrate various nest infestation rates for
Orntthodoros capensis. On Sand Island the highest nest infestation rates
occur in the Brown Booby at 50 percent and the Brown Noddy at 49 percent.

82

0
JF MAMJ JA SO ND
Areas H & K

100 200

Figure 33. Distribution of positive samples of Ornithodoros capensis for
1965, Sand Island, Johnston Atoll. Inserts show bi-monthly
infogt ation rates for the three major areas.

0 as
JFMA MJ JA SOND
Areas E & F

EL

Hee

MJ $30

Figure 34. Bi-monthly tick infestation rates st eh compared with breeding
bird population fluctuations (bars), Sand Island, Johnston Atoll,
1965);

83

February

= =
October Gund yp) December Feet

Figure 35. Areas used by Sooty Terns (stippled) and Wedge-tailed Shearwaters
(barred) on Sand Island, Johnston Atoll, 1965.

84

The Sooty Tern, by comparison, has an infestation rate of only 26 percent;
however, many of the samples labeled Miscellaneous Litter (rate 33 percent)
probably should be listed under Sooty Tern nests since the species is so
widespread over the island. The remaining infestation rates are as follows:
Red-footed Booby 20 percent, Great Frigatebird 20 percent, Red-tailed
Tropicbird 7 percent, and Wedge-tailed Shearwater 5 percent, with the rest
being 0 percent. Comparative data from Green Island, Kure Atoll, north-
western Hawaiian Islands, are also shown in Table 12.

Table 11. Berlese sample data--bi-monthly Ornithodoros capensts
infestation rates for Sand Island, Johnston Atoll, 1965.

Areas A, B, C, and D

JF MA MJ JA 50 ND Total
No. samples negative if 20 42 7 10 49 129
No. samples positive 0 2, 9 2 0 4 17
Total samples iL 22 Sil 9 10 5p) 146
Percent positive 0% 9% 18% Ing 0% 8% 27,

Areas E and F

No. samples negative 4 iti 33 10S) 3 35 LOS
No. samples positive 2 7 32 8 0 2 Syl
Total samples 6 18 65 27 3 37 156
Percent positive 33% 39% 49% 30% 0% 5% 33%
Areas H and K
No. samples negative Pad 8 2 43 i 12 87
No. samples positive 10 da uy) 33 5) 14 90
Total samples Syl 19 189 76 6 26 176
Percent positive 327 582 89% 44% 83% 504% SIU
All Areas
No. samples negative 26 39 Th. 69 14 96 Sal
No. samples positive a2 20 58 43 5 20 158
Total samples 38 59 135 nee E9 116 479
Percent positive 327 34% 43% 38% 267, 17Z 337,
No. of ticks per
positive sample Sig 5) 9.4 8.0 AOr ~ L650 G7 7.0

Trombiculidae: Seven genera of 16 species of larval trombiculid mites,
commonly referred to as chiggers, are known from POBSP collections in the
central Pacific (Brennan, 1965; Brennan and Amerson, 1971). Five species
taken off six host species have been recorded from Sand Island. Neoschoengastta
ewingt was taken off Golden Plover and Brown Noddy, as well as EromELtter
taken in the Sooty Tern nesting area. Blankaartia amersoni was recorded
from the nasal cavities of Bulwer's Petrel, Red-tailed Tropicbird, Sooty
Tern, and Brown Noddy. Guntherana domrowi was taken from Sooty Tern.
Guntherana sp. was taken from a house mouse. Womersta strandtmannt was
recorded off Sooty Terns. The first three species listed above are found
both in the north- and south-central Pacific. Guntherana sp. is known only
from Sand Island, Johnston Atoll. Womersta strandtmannt at Sand-Johnston is
a first and only record from the Pacific; it is also known from the North
American Gulf and Atlantic coasts.

Table 12. Orntthodoros capensis infestation rates from Berlese
samples of hosts' nests

Host - Source

Black-footed Albatross

Laysan Albatross

Wedge-tailed Shearwater

Christmas Shearwater

Bonin Petrel

Bulwer's Petrel

Red-tailed Tropicbird

Blue-faced Booby
Brown Booby
Red-footed Booby
Great Frigatebird
Sooty Tern

Noddy Tern

Black Noddy
Hawaiian Monk Seal
Polynesian Rat

Misc. Litter

Totals

# Do not occur.
* Resting area.

Green I., Kure Atoll
1964-1965

Total Nests
Examined

ZS

28

12

167

398

Total Nests
Infested

ho

66

Infestation

85

Sand I., Johnston Atoll

Total Nests
Examined

267

Sys)

1965

Total Nests
Infested

ILD)

Infestation

Rate

33%

86

Rhinonyssidae: Amerson (1967) examined the nasal cavities of 460
adult and immature Sooty Terns taken throughout their 1965 breeding
cycle on Sand Island, Johnston Atoll, for Rhinonyssidae, or nasal mites.
Incidence of rhinonyssids of two genera, Lartnyssus and Sternostoma,
in adult terns increased during the egg-laying period and decreased
(possibly due to feeding of young) after the eggs had hatched.
Incidence in young Sooty Terns was low (2 percent) during their first
4 months, suggesting that transference of mites from adult tern to
their young is not very efficient. During the 5th month, however, the
incidence rose to 29 percent, suggesting that the mites were elsewhere
in the host and they require time to travel to the nasal cavity.

Mallophaga: Chilson (1953) listed one species of Mallophaga, or
biting lice, from terns at Johnston Atoll. Amerson and Emerson (1971)
recorded 23 species of 11 genera and two families from POBSP collections
(Table 13). Of the 12 bird hosts, all but two species were seabirds;
the two were both shore birds. Five Mallophaga species were taken from
the Wedge-tailed Shearwater, four species were taken from the Sooty
Tern, and three species were from the Brown Noddy.

Hippoboscidae: Two species of hippoboscids, or louse flies, were
recorded by POBSP personnel off five species of seabirds at Sand
Island (Maa, 1968). Olfersta aenescens was taken from the Great Frigate-
bird, Sooty Tern, as well as from a Berlese sample containing litter.
Olfersta sptnifera was recorded from the Wedge-tailed Shearwater, Red-
tailed Tropicbird, Great Frigatebird, and Brown Noddy. These two
hippoboscid species are pantropical in the Pacific, Indian, and Atlantic
Oceans.

Echinodermata

A total of 37 species belonging to 13 families of Echinodermata
are presently known from the lagoon waters of Johnston Atoll (Table
14). Edmondson, et al. (1925) and Clark (1949), reporting primarily on
material collected by the 1923 TANAGER Expedition, each recorded 24
species. Brock, et al. (1965, 1966) recorded six species from 1963-
1965 collections. The present paper lists 26 species found in the
collections of the National Museum of Natural History and the Bernice
P. Bishop Museum.

Brock, et al. (1965, 1966) studied the distribution and abundance
of conspicuous echinoderms at Johnston Atoll. Of nine species studied

in 13 areas, Trtpneusten gratilla was found in 12 areas and Heterocentrotus

mamnillatus was found in 10 areas. They concluded that dredging
affected both distribution and abundance of echinoderms.

Vertebrates

The vertebrates of Johnston Atoll are well known. Extensive
ecological studies have been made on the fishes, birds, and mammals.
Although the species composition of reptiles is well known, very little
is known about their populations.

Table 13. Mallophaga collected at Sand Island, Johnston Atoll by
the POBSP; adapted from Amerson and Emerson (1971)

Suborder
Family

Species Host

Ischnocera
Philopteridae

Halipeurus heraldtcus
Haltpeurus pactftcus
Naubates harrisont
Peettnopygus graciltcornis
Peetinopygus sulae
Quadraceps btrostris

Quadraceps charadrit hospes

Quadraceps tmpar

Phoenix Petrel
Wedge-tailed Shearwater
Wedge-tailed Shearwater
Great Frigatebird
Red-footed Booby

Sooty Tern
Black-bellied Plover
Wandering Tattler

Quadraceps separatus Brown Noddy
Quadraceps sp. Gray-backed Tern
Saemundssonta albemarlensts Sooty Tern

Saemundssonta hexagona
Saemundssonta puellula
Trabeculus hexakon

Red-tailed Tropicbird
Wedge-tailed Shearwater
Wedge-tailed Shearwater

Ablycera
Menoponidae
Actorntthophilus tnetsus Brown Noddy
Aetorntthophtlus ptceus piceus Sooty Tern
Actorntthophylus sp. Brown Noddy
Austromenopon atrofulvum Sooty Tern

Austromenopon squatarolae

Colpoecephalum angulaticeps

Etdmanntella albescens
Procellartphaga patlula
Procellartphaga sp.

Black-bellied Plover
Great Frigatebird
Blue-faced Booby
Wedge-tailed Shearwater
Phoenix Petrel

Table 14. Echinodermata from Johnston Atoll*

Class Brock
Subclass Edmondson et al.
Family Chal. Cilank | 71965, ~ Present
Species 1925 1949 1966 Paper
Echinoidea
Diadematidae
Dtadema sp.. Ke
Echinothrtx calamaris (Pallas) x
Echinothrix dtadema (Linnaeus) X x
Echinometridae
Echinometra mathaet (de Blain- xX x xX
ville)
Echntometra oblonga (de Blain- X X X

ville)

88

Table 14. (continued)

Class

Subclass Edmondson

Family
Species

Echinometridae (cont.)
Echtnostrephus actculatus
A. Agassiz
Echtnostrephus molarts
(de Blainville)
Heterocentrotus mammnillatus
(Linnaeus)
Heterocentrotus trigonartus
(Lamarck)
Toxopneustidae
Tripneustes grattlla (Linnaeus)
Brissidae
Brissus latecarinatus (Leske)

Holothuroidea
Holothuriidae
Acttnopyga maurtttana (Quoy &
Gaimard)
Acttnopyga obesa (Selenka)
Aettnopyga sp.
Holothurta atra Jaeger
Holothurta dtffictlts Semper
Holuthurta hilla (Lesson)
Holothuria impattens (Forskal)
Holothuria pardalis Selenka
Hotuthurta parvula (Selenka)
Synaptidae
Chtridota rtgida Semper
Opheodesoma spectabilis Fisher
Polyplectana keferstetntt
(Selenka)
Stelleroidea
Asteroidea
Oreasteridae
Culetta arenosa Perrier
Culetta novaeguineae Muller &
Torschel
Ophidiasteridae
Linekta multtfora (Lamarck)
Acanthaster idae
Acanthaster planet (Linnaeus)
Mithrodiidae
Mithrodta bracleyt Verrill
Mithrodia fishert Holly

Cu Gils
1925

a
2.

Clark
1949

MP PM PX

Brock
Be Che
1965,
1966

Present
Paper

Pi PS PM OPS PS PS PS

Appendix Table 14. (continued)

Class Brock
Subclass Edmondson et al.
Family et al. Cllank, Wel9GS; Present
Species 925 1949 1966 Paper
Stelleroidea (cont.)
Ophiuroidea
Ophiactidae
Ophtaetts savtgnyt X X x
(Muller & Troschel)
Ophiocomidae
Ophtocoma ertnaceus
Muller & Troschel Xx xX X
Ophtocoma ptea (Muller & : =
Troschel)
Ophtocoma sp. X
Ophtocomella eltippertont xi
Ophtocomella sexradia (Duncan) x
Ophiodermatidae
Distichophts elarkt 1K
Ophtopeza spinosa (Ljungman) x

*Taxodermic order follows Hyman (1940), Durham, et al. (1966a, 1966b),
and Beaver, et al. (1967).

Fishes

The fish fauna of Johnston Atoll is well known and is herein
separated into two categories: pelagic fishes and inshore fishes.

In the interest of consistency, taxonomic order follows that used
by Gosline (1955) and Brock, et al. (1965, 1966), whose identification
and nomenclature follow Schultz, et al. (1953).

Pelagic Fishes

Numerous large pelagic fishes occur around Johnston Atoll. No
extensive species list exists for this area, but fishes here include
tuna (wahoo, skipjack, and yellowfin), barracuda, and sharks (Chapman,
1946; Halstead and Bunker, 1954a). Two sharks, not included in the
inshore reef fauna but known from there, include the white-tipped
shark, Trtaenodon obesus (family Triakidae) and the grey sand shark,
Carcharhinus mentsorrah (family Carcharhinidae). Additional species of
sharks are most likely to occur in the lagoon. Barracuda and tuna also
are known from the lagoon waters (Brock, et al., 1965, 1966).

89

90

Inshore Fishes

A total of 194 species of inshore fishes are known from Johnston
Atoll (Table 15). Of these 194 species, 27 were reported by Smith and
Swain (1882), of which five were described as new. Fowler and Ball
(1925) listed 72 species collected in 1923 by the TANAGER Expedition;
one of these was described as new. Nine species, three of them new,
were reported by Schultz, et al. (1953). From collections taken in
1951, Halstead and Bunker (1954a) listed 60 species and Gosline (1955)
recorded 111 species. Brock, et al. (1965, 1966) reported 143 species
from collections taken during August 1963 through August 1965. Asa
matter of further historical interest, 109 species were known by 1954,
49 species not previously reported were added by 1955, and an additional
36 species were added by 1966.

Gosline (1955) suggested that the Johnston Atoll fish fauna is
made up of four components: (1) endemics, (2) fish that have made
Johnston a stopping point on their migrations north, (3) fish that
have made Johnston a stopping point in their southward travels, and
(4) the pelagic fish to whom Johnston is of little or no significance.

Only two species of Johnston fishes have not been taken elsewhere.
These are Centropyge ntgrtocellus and C. flammeus, both butterfly-
fishes; neither are abundant at Johnston (Gosline, 1955).

Since only two species are endemic, it would appear that Johnston
is not isolated in the evolutionary sense. In contrast, according to
Gosline, nearly one half of the Line Island inshore fishes do not occur
in Johnston or Hawaiian waters. Because of the wide stretches of deep
water between the northernmost Line Islands and Johnston, Johnston Atoll
is comparatively isolated from the south. Johnston's fish fauna is much
more closely allied to that of the Hawaiian Islands. This is seen in
the fact that the majority of the tropical species present at Johnston
are also found in Hawaii.

The present-day Hawaiian fish fauna is thought to have been derived
from more tropical waters. Yet presence of many tropical species at
Johnston that are now unknown from Hawaiian waters would seem to indicate
that there is a limit to how many warm water species Hawaii can absorb.
Eventually it is expected that more tropical species will establish
themselves in Hawaiian waters. Gosline has suggested that Johnston
Atoll was probably the original port of entry of Hawaii's inshore fish
fauna.

Distribution and Abundance Within the Atoll

Halstead and Bunker (1954a) generalized the fish distribution at
Johnston and divided the atoll into three zones: (1) northern (periphe-
ral) reef area, (2) southern (shoal) reef area, and (3) bank (lagoon)
shoals. The northern reef area was characterized by scombroid fishes,
barracuda, sharks, and other pelagic species living on the seaward

91

(open-water) side of the reef and eagle-rays, moray eels, butterfly,
squirrel, surgeon, damsel, surmullet, puffer, and file fishes on the
lagoon side. The southern reef area was not examined but a large
shark population was thought to exist in the open water; fish species
were believed to be fewer in number and variety here, compared to the
northern reef area. The bank shoals or lagoon area was characterized
by: moray eels, wrasse, parrot, coronet, surgeon, butterfly, squirrel,
damsel, and puffer fishes in the patch reef portion; flounders,
blennies, surmullet, pompano, puffers, pomacentrids, stingrays, and
triggerfishes in the sandy areas; and trumpet, parrot, damsel, pompano,
trigger, and butterfly fishes in the wreckage or dump area to the
southwest of Johnston Island.

As part of an extensive study on the effects of dredging at
Johnston, Brock, et al. (1965) studied the distribution and abundance
of 75 species at nine localities within the lagoon during August 1963
to June 1964 (Fig. 36). Areas 1, 2, and 4 were near the marginal reef

JOHNSTON ISLAND AND REEF

16°45

Black - recently formed land areas.
Cross hatch - areas of reef altered by
recent dredging. Stipling - land area
prior to recent dredging. Dots - area
often covered by turbid water during
the dredging operation (incomplete in
southwest area where turbidity extended
far out to sea). Hatching - trapping
stations.

Numbered dots - stations studied. A - John-
ston Island, B - Sand Island, C - East Island,

Does D - North Island, E - North Island Trap Area,

eS onouR = nar F — Small Boat Channel Trap Area, G - Sand
Island Trap Area, H - Marginal Reef.

Oan-
169°33 30°

Figure 36. Fish collection stations, Johnston Atoll, 1963-1964 (Brock, et
UNS IL TS) A

92

and each contained higher numbers of species, respectively 59, 45, and
43 species (mean = 49 species), than areas 3, 5, 6, 7, 8, and 9 in the
lagoon proper. Numbers of species in the lagoon areas ranged from 28
to 39 species, with the mean being 33 species. Actual numbers of
individuals were also highest in the marginal reef zone; however,
ratios of numbers of species to numbers of individuals appear to be
higher in general in the lagoon zone.

Commonness of species, based on the number of areas each were
found in, show that of the 75 species, six species were found in all
nine areas. The most abundant species was Acanthus sandvicensis; it
was found in all areas and was dominant in five areas. The next three
more abundant species, in order, were Searus sordidus, Ctenochaetus
strigosus, and Acanthurus achtlles; each was dominant in three areas
as well as found in all areas. Twelve species were found only in one
area; these were considered rare in abundance.

Brock, et al. (1966) continued their distribution and abundance
studies during August 1964 to August 1965. They added four new study
stations (see areas 10, 11, 12, and 13, Fig. 32) and examined effects
of recently completed dredging operations. They collected fishes in
the bottom and sides (cliffs) of dredged areas, as well as non-disturbed
reef areas. In general, the cliffs were barren of fish: two areas had
no fish and two had three or nine species. Bottoms of three areas
were also quite desolate: one had no species, one had three species,
and the other had eight species. The remaining bottom area had 25
species, one more than the 24 species found in the adjacent reef. In
general, the remaining non-dredged reef portion had fair numbers of
species: 23, 19, and 14.

Effects of Dredging

Brock, et al. (1965) found no effects of dredging on fishes in
their work through June 1964. But Brock, et al. (1966) did find the
fish fauna adversely affected by dredging from August 1964 through
August 1965.

By resurveying areas 4 through 8 they were able to compare numbers
and abundance of species. The number of species of fish observed
during 1965 was about 50 percent less than that observed during 1964
for stations 4 and 5, and about 10 percent less than stations 6, 7,
and 8. From 2 to 13 species (mean = 7) new to each area were also found.
If these were subtracted from the 1965 totals, then the difference
between years would even be higher.

Brock, et al. (1966) also noted that the number of species rated
as common or dominant was less for all stations (except one changing
from common to dominant in station 8) for 1965 as compared to 1964.
Their data suggest that while the total number of species observed for
the two years changed very little for stations 6, 7, and 8, their
abundance changed significantly.

93

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103

Ciguateric Fishes

The first identified cases of ciguatera-type fish poisoning occurred
at Johnston Atoll in 1950. According to Halstead and Bunker (1954a),
there were about 20 cases of ciguatera poisoning between May 1950 and
May 1951. Fishes thought by island workers to be dangerous to eat
included: puffers (Tetraocontidae), triggerfish (Balistidae), saboti
(Kuhliidae), ulua (Carangidae), "tuna" (probably Euthynnus yaito, a
skipjack), "red or blue toothless snapper" (?), moray eels (Muraenidae),
and surmullet (Mullidae).

Following this 1950 poison outbreak, Halstead and Bunker (1954a)
studied the ciguatera problem at Johnston Atoll. They collected fishes
in May, October and November 1951. They found that of 60 species (211
specimens) of 21 families tested, 47 species (78 percent), and 98
specimens (46 percent) were toxic (Table 16). Both the musculature
and viscera of specimens were found to be toxic. Although the bioassay
method used in this 1954 study was later found to be of questionable
value (Banner, et al., 1960), numerous species of inshore fishes were
toxic as evidenced by the numerous cases of ciguatera poisoning re-
ported (Banner and Helfrich, 1964; Brock, et al., 1965).

By 1963, however, the poison problem appeared to have subsided
(Banner and Helfrich, 1964). In August 1963, 21 species (224 specimens)
of 11 families were examined for poison (Table 16) by Brock, et al.,
(1965). Of these 21 species, only two (10 percent) were toxic: the
white-tipped reef shark Triaenodon obesus (11 percent) and the large
moray eel Gymnothorax javantcus (80 percent). By interviewing island
construction workers (mostly from Hawaii), they also discovered that
Many varieties of lagoon fishes were eaten locally. Most “expressed a
willingness to eat any of the Johnston...fishes except black ulua,
moray eels, and sharks."

Brock, et al. (1965: 23) described the situation at Johnston Atoll
as "...what might be expected near the termination of a period of
occurrence of ciguatera poisoning in fishes. Such a period would become
first evident by the herbivorous species becoming toxic. Following the
initial stage both herbivorous and carnivorous species would be toxic,
the latter through predation of the herbivores. This period may last
for a number of years. Its termination would be marked first by the
absence of young toxic herbivores, followed by the absence of young
toxic carnivores. The residual effects would be evident through the
occurrence of ciguatera toxin only in the large, long lived predators.
Once a fish becomes toxic it appears to remain so for long periods of
time even though it does not continue to consume toxic prey. However,
the toxin locked up in the bodies of the larger predators can recycle
back into other predators either large or small upon the occasion of
their death from old age or disease or some other cause."

Randall (1958) suggested that the toxin responsible for ciguatera
was produced by algae, possibly a blue-green algae, which occurred
early and abundantly during the successional invasion of a newly

104

denuded substrate by algae. The algae would be eaten by herbivorous
animals and these, in turn, by carnivorous animals. Brock, et al.
(1965) noted that if Randall's hypothesis was valid, a substantial
increase in the number and species of toxic fish, beginning with the
herbivorous species, should follow the dredging operations at Johnston
which began in August 1963. At the beginning of this dredging, none
of these herbivorous species was found to be toxic; only the larger
predacious sharks and eels were toxic.

Ciguatera poison cases, in fact, did continue (Brock, et al.,
1966). Several poisonings (one serious, the others mild) were reported
in May 1964 involving a predacious black ulua Caranx melampygus. In
addition, two rather serious cases occurred in August 1965, with several
species of reef fishes involved.

Likewise, a slight increase in numbers of toxic fishes occurred
(Brock, et al., 1966). Of 38 species examined during August 1964 to
August 1965 (number of specimens unknown), six species of five families
(16 percent) were toxic. As in the 1963 study, the moray eel Gymnothorax
javantecus was most toxic at 48 percent, with the white-tipped reef
shark Trtaenodon obesus second at 19 percent toxic (Table 16). Another
shark species--the grey sand shark Carcharhinus mentsorrah--was also
found to be toxic (11 percent); this species was not toxic in the 1963
tests. Three other species--all herbivorous--not toxic in 1963 were
also toxic in the 1964-1965 tests. They were Ctenochaetus strigosus,

a surgeon fish, and Melitchthys buniva and M. vidua, both triggerfishes;
all three fed mainly on algae. Furthermore, in some of the dredged
areas thick algal mats developed over the whole bottom, with the blue-
green algae Lyngbya majescula being the dominant species.

From these data, Brock, et al. (1966) suggested that a new cycle in
the production of ciguateric fishes at Johnston may be in an initial
stage. These data also support Randall's 1958 hypothesis discussed
earlier for indeed following dredging (1) the blue-green algae increased,
(2) the number of toxic herbivorous fish species increased, and (3) the
number of ciguatera poisonings increased.

Published Johnston Atoll toxin data from 1966 through 1973 has been
very limited, but does suggest that little change in toxicity levels has
occurred. Brock (1972) found that ciguatera toxin in the moray eel
Gymnothorax javantcus has not statistically changed in level from 1963
to 1970; these data are presented in a later section.

It is not known whether seabirds are affected by ciguatera poisoning;
however, Halstead and Bunker (1954b) discovered that experimental chickens
are apparently unaffected by the less virulent fish poisons. Most seabirds
at Johnston and elsewhere in the central Pacific, moreover, do not feed on
inshore lagoon fishes. At Johnston, only Brown Boobies have been observed
feeding in the lagoon and this species, White Terns, and Black Noddies
normally feed offshore near the atoll; the remaining species usually feed
well at sea (King, 1967).

105

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110

Species Accounts

Although the fish fauna of Johnston Atoll is well known, very little
is known about the biology of individual species occurring there. Only
one species, a moray eel, and one family have been studied extensively.

MORAY EEL Gymnothorax javantcus

Status

Abundant breeding species; present year-round throughout the lagoon
and reef waters. Breeds during the summer months. Highest of all
fishes toxic to man at Johnston; up to 80 percent are toxic.

Ecological Distribution

Halstead and Bunker (1954a) first collected two specimens of
Gymnothorax javanteus from Johnston Atoll in 1951. Additional obser-
vations and collections were reported by Brock, et al. (1965, 1966)
during 1963 through 1965. Brock (1972) found it to be very abundant
from 1963 to 1971. In all, 1,165 specimens were taken at Johnston by
the Marine Toxins Program of the Hawaii Institute of Marine Biology,
University of Hawaii.

This species is a coral reef dweller, is very cryptic in color, and
hides in reef crevices. At Johnston, it has been found throughout the
lagoon and reef waters. It has been found under rubble in extensively
dredged areas, but is more commonly encountered in areas of dense coral
growth. Its vertical range is unknown, but has been collected at
Johnston at depths ranging from 0.3 to 17 meters.

Populations

Although Brock, et al. (1965, 1966) found G. javanicus rare in
their studies, Brock (1972) found fair numbers (6 to 164, mean = 50) in
a sample of 648 eels collected at 13 stations throughout the lagoon.
Additional population estimates or studies are lacking.

This species--perhaps the largest of the Indo-Pacific morays-—-
probably lives to be older than 20 years. The only eel predators of
significance at Johnston appear to be sharks (Brock, 1972).

Annual Cycle

Gymnothorax javanitcus occurs at Johnston Atoll throughout the year,
although breeding occurs only in June, July, and August. Females re-
lease about 188,000 to 291,000 eggs per spawning. Males mature at 3.4
kg and females at 4.6 kg in weight. Males, however, attain a larger
size than females.

This species feeds chiefly crepuscularly and nocturnally; however,
attempted diurnal feeding has been observed. The variety of food items

ILL

indicates it is an opportunistic predator; 89 percent consisted of
fishes. Gut content analysis revealed, however, that this species
feeds infrequently (Brock, 1972).

Ciguatera

Halstead and Bunker (1954a) first documented ciguatera at Johnston
in 1950 and noted that toxin occurred in one of two Gymnothorax javanicus
specimens examined. Brock (1972) examined 887 eels of this species
taken at Johnston and found no statistically significant differences
between the levels of toxicity from 1963 through 1970.

Analysis of the distribution of toxicity showed no significant
differences in the levels of toxicity among the 14 habitat areas;
however, there was a significant correlation of toxicity with depth.
This further suggests that there is more toxin in the food of G.
javantcus at depths of 6 to 12 meters, compared to depths of 0 to
6 meters. This species also displayed a significant increase in
toxicity with increasing body size. No significant difference in
the toxicity of the flesh from males and females existed, however.
Up to 80 percent of all specimens examined were toxic (Brock, 1972).

Movement

Mark and recapture methods, as well as indirect observations,
suggested that on a short-term basis (less than one year) Gymnothorax
javanteus has a definite home range and its migration is restricted
(Brock, 1972) .

SURGEONF LSHES Acanthuridae

Jones (1968) studied ecological relationships between 13 species of
surgeonfishes, family Acanthuridae, found on Johnston Atoll in an attempt
to determine habitat separation and feeding habits.

He concluded that although there appeared to be unlimited food
available for all species, the fishes were separable on ecological
grounds. He recognized four species groups by habitat (mid-water, sand
patch, subsurge reef, and surge zones); three different ways of feeding
(zooplankton feeders, grazers, and browsers); and at least four dif-
ferent categories of foods eaten (zooplankton, interstitial material,
coarse algae, and filamentous algae). He also described numerous
modifications of the alimentary canal which suggested interspecific
differences in food handling. He hypothesized that there must be in-
frequent periods of food shortage during which selection pressure was
high enough to favor adaptations that would tend to reduce interspecific
competition.

a2

Reptiles

Two reptile species--"'one long brown species and one shorter grey
species''--were recorded by E.H. Bryan, Jr. (ms.) in July 1923 when the
first scientific visit was made to Johnston Atoll. He indicated that
these were then abundant. Five species of reptiles are now known from
the Atoll. All but one species, the Black Sea Turtle, were probably
introduced by man. They are all known from various islands throughout
the central Pacific.

Of the five species, two species occur only on Johnston Island and
two species occur on Sand Island. Only one species occurs on both
islands. As of 1969, none was known from Akau and Hikina Islands.

Species Accounts

There are no general references that illustrate the reptiles of
Johnston Atoll. Taxonomy of the turtles follows Carr (1972) and
Amerson (1971). Order of listing for lizards follows Brown (1957).

BLACK SEA TURTLE Chelonta agasst2t

Status

Regular uncommon visitor; known from the lagoon, offshore Johnston
Island, and Sand Island.

Observations

Brooke (ms.), who visited Johnston Atoll in March 1859, commented
about the lack of turtles: "The reefs are covered with fish of various
kinds. Mullet abound, but there are no turtles.'' Wetmore (ms. a and b),
likewise, recorded no turtles at Johnston Atoll in July 1923.

POBSP personnel recorded sea turtles in the shallow marginal reef
area west of Johnston Island in July 1963. An adult (USNM 163581) was
collected 20 November 1966 on the beach of Sand Island. Island personnel
in 1973 reported seeing 10 to 12 turtles offshore of Johnston Island
throughout the year. A longtime resident estimated harvesting 12 to 15
per year.

Annual Cycle

The Black Sea Turtle apparently visits Johnston Atoll year-round.
No records exist of it breeding on the atoll, although perhaps it did
in small numbers prior to inhabitation by man. This species breeds
during the summer in the northwestern Hawaiian Islands, especially at
French Frigate Shoals (Amerson, 1971).

ses

HOUSE GECKO Hemtdaetylus frenatus
Status

Common introduced breeder; occurs only on Sand Island.
Observations

It is not known when House Geckos were introduced to Johnston Atoll.
POBSP personnel collected two specimens from Sand Island in May 1964
(USNM 154207-208) and two more in May and March 1966 (USNM 163579-580).
FOX GECKO Hemtdactylus garnottt
Status

Uncommon introduced breeder; occurs on Johnston and Sand Islands.
Observations

POBSP personnel collected a Fox Gecko from Johnston Island 2 July
1966 (USNM 166825). Another was collected at Sand Island 22 August 1968
(USNM 166863).
MOURNING GECKO Leptdodactylus lugubrts
Status

Uncommon introduced breeder; occurs only on Johnston Island.

Observations

A Mourning Gecko was collected (USNM 156990) from Johnston Island
28 October 1964 by POBSP personnel.

SNAKE-EYED SKINK Ablepharus boutontt poectlopleurus
Status

Uncommon introduced breeder; occurs only on Johnston Island; present
status unknown.

Observations
Thirteen specimens of this species were collected by E.H. Bryan,

Jr., on Johnston Island in July 1923 (USNM 66810-22). None has been
recorded since.

114

Birds

Several sources were used in assembling the common and scientific
names of the birds occurring at Johnston Atoll (Table 17). The names
used in the American Ornithologists' Union's Check-list of North American
Birds (1957), as amended by Eisenmann, et al. (1973), were followed for
species occurring in North America. In the interest of consistency,
seabird names agree with those which appear in Watson's Smtthsontan
Identtficatton Manual: Seabirds of the Troptcal Atlantic Ocean, and
King's Smithsontan Identification Manual: Seabtrds of the Tropical
Paetfie Ocean. Taxonomic order follows that of Peter's Check-ltst
of Btrds of the World, volumes I, II, and III, with the exception of
the Procellariiformes, which follow Alexander, et al. (1965), the
Anseriformes, which follow Delacour (1954, 1959), and the Charadrii-
formes, which follow Bock (1958) and Jehl (1968).

Table 17. Birds from Johnston Atoll**

Order Procellariiformes
Family Diomedeidae E
Diomedea nigripes* Black-footed Albatross
Diomedea tmmutabtlis* Laysan Albatross
Family Procellariidae

Pterodroma atba*
Bulwerta bulwertt
Puffinus pactficus

Puffinus nattvttatis
Puffinus puffinus newellt*

Family Hydrobatidae

Oceanodroma tritstramt*

Order Pelecaniformes

Family Phaethontidae
Phaethon aethereus*
Phaethon rubrtcauda
Phaethon lepturus*

Family Sulidae
Sula dactylatra*
Sula leucogaster
Sula sula

Family Fregatidae
Fregata minor
Fregata artel*

Order Ciconiiformes
Family Ardeidae
Bubulcus tbits*
4d

Order Anseriformes
Family Anatidae
Anas acuta*

Anas [=Mareca] americana*
Anas [=Spatula] elypeata*

Phoenix Petrel

Bulwer's Petrel
Wedge-tailed Shearwater
Christmas Shearwater
Newell's Shearwater

Sooty Storm Petrel

Red-billed Tropicbird
Red-tailed Tropicbird
White-tailed Tropicbird

Blue-faced Booby
Brown Booby
Red-footed Booby

Great Frigatebird
Lesser Frigatebird

Cattle Egret

Pintail
American Wigeon
Northern Shoveler

Table 17. (continued)

Order Galliformes
Family Phasianidae
Gallus gallus

Order Falconiformes
Family Falconidae
Faleo peregrinus tundrius*

Order Charadriiformes

Family Charadriidae
Pluvialis domintea*
Pluvialts [=Squatarola] squatarola*
Charadrius semtpalmatus*

Family Scolopacidae
Numentus tahtttensis*
Tringa [=Totanus] flavtpes*
Actttts macularta*
Catoptrophorus semtpalmatus*
Heteroscelus tneanus [=ineanum]*
Arenarta tnterpres*
Limnodromus sp.*
Caltdrts [=Crocethia] alba*
Caltdris [=Ereunetes] mauri*
Caltdrts [=Erolita] melanotos*
Caltdrts [=Erolia] acuminata*
Tryngttes subruficollis*
Philomachus pugnasx*

Family Phalaropodidae
Steganopus tricolor*

Family Laridae
Larus glaucescens*
Larus argentatus*
Larus atrictlla*
Larus ptptxean*
Larus spp.*
Sterna lunata
Sterna fuscata
Thalasseus elegans*
Procelsterna cerulea*
Anous stolidus
Anous tenutrostrits
Gygts alba

Order Columbiformes
Family Columbidae
Columba livia

Order Strigiformes
Family Strigidae
Asto flammeus*

a5

Domestic Chicken

Peregrine Falcon

American Golden Plover
Black-bellied Plover
Semipalmated Plover

Bristle-thighed Curlew
Lesser Yellowlegs
Spotted Sandpiper
Willet

Wandering Tattler
Ruddy Turnstone
Dowitcher species
Sanderling

Western Sandpiper
Pectoral Sandpiper
Sharp-tailed Sandpiper
Buff-breasted Sandpiper
Ruff

Wilson's Phalarope

Glaucous-winged Gull
Herring Gull
Laughing Gull
Franklin's Gull
Gull species
Gray-backed Tern
Sooty Tern
Elegant Tern
Blue-gray Noddy
Brown Noddy
Black Noddy
White Tern

Rock Dove

Short-eared Owl

116

Table 17. (continued)

Order Passeriformes

Family Alaudae

Alauda arvensts*

Family Zosteropidae

Zosterops japontca*

Family Estrildidae

Lonehura striata

Skylark

Japanese White-eye

Society Finch

**Resident birds are unmarked; non-resident birds are marked with an *.

Table 18.

Introduction

The 56 bird species recorded belong to 10 orders, 19 families,
and 38 genera; 22 species are classes as seabirds, and 34 species are
waterfowl, marsh, and land birds (Table 18).

Akau

Seabirds:

Breeders

Bulwer's Petrel

Wedge-tailed Shearwater
Christmas Shearwater
Red-tailed Tropicbird

Brown Booby 2
Red-footed Booby

Great Frigatebird R
Gray-backed Tern B*
Sooty Tern

Brown Noddy 2
Black Noddy

White Tern

Former Breeders
Black-footed Albatross

Laysan Albatross
Blue-faced Booby

Visitors

Phoenix Petrel

Newell's Shearwater

Sooty Storm Petrel
Red-billed Tropicbird
White-tailed Tropicbird O
Lesser Frigatebird

Blue-gray Noddy

Hikina

Status of birds on Johnston Atoll

Sand

Johnston Original Man-made

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Table 18. (continued)

Akau Hikina Johnston Original Man-made

Waterfowl, Marsh, and Land
Birds:
Regular Migrants
Pintail
American Golden Plover
Bristle-thighed Curlew
Wandering Tattler
Ruddy Turnstone
Sander ling
Pectoral Sandpiper
Irregular Visitors
American Wigeon
Northern Shoveler
Glaucous-winged Gull
Herring Gull
Laughing Gull
Short-eared Owl
Stragglers
Cattle Egret
Franklin's Gull
Accidentals
Peregrine Falcon
Black—bellied Plover
Semipalmated Plover
Lesser Yellowlegs
Spotted Sandpiper
Willet
Dowitcher species
Western Sandpiper
Sharp-tailed Sandpiper
Buff-breasted Sandpiper
Ruff
Wilson's Phalarope
Gull species
Elegant Tern
Skylark
Japanese White-eye
Introductions
Domestic Chicken
Rock Dove
Society Finch

Present Breeders
Former Breeders
Total species

AAD DA

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118

B = Breeder; R = Recorded; O = Overflier. Capital letters indicate
status 1963-1969; lower case letters indicate status 1923-1962, if
different than at present.

* bred only in 1964
xk bred only in 1973.

Seabirds

Of the 22 seabird species recorded at Johnston Atoll, 12 are
breeders, three are former breeders, and seven are visitors (Table 18).

Breeders: All 12 resident breeding species nest in the Hawaiian
Islands and other parts of the tropical Pacific. These 12 appear to
have closer affinities with their respective northwestern Hawaiian
populations than with those in the Line and Phoenix Islands. See
also the Banding and Movement section.

Former Breeders: Three species formerly bred at Johnston Atoll.
The Black-footed Albatross presently breeds only in the Hawaiian Islands
and the Bonin-Volcano Islands area, while the Laysan Albatross presently
breeds exclusively in the Hawaiian Islands. Both species normally range
north of 10°N. The Blue-faced Booby breeds throughout the tropical
Pacttac.:

Visitors: In the surrounding ocean one may find both seabird
species that use or breed at Johnston Atoll and those that seldom
occur there. These latter species (Table 18) include both breeders
from the Hawaiian Islands and Line-Phoenix Islands, as well as
breeders from other areas of the Pacific who migrate into or through
the surrounding waters during their non-breeding season (POBSP, 1967a).
Normally most of these 28 bird species stay at sea, but, because of
sickness or bad weather, any could alight on the islands at Johnston
Atoll.

The seven seabird visitors to Johnston Atoll come from the north,
south, and east Pacific.

The Newell's Shearwater breeds exclusively in the main Hawaiian
Islands, while the Sooty Storm Petrel breeds only in the Hawaiian
Islands and the Bonin-Volcano Islands area. Although the White-tailed
Tropicbird and Blue-gray Noddy breed throughout the tropical Pacific,
those that visit Johnston are thought to breed primarily in the Hawaiian
Islands.

Visitor seabirds from the south Pacific are the Phoenix Petrel
and Lesser Frigatebird. Both are equatorial, south-central Pacific
breeders. The Lesser Frigatebird has been recorded from the western

EY)

Pacific, Wake, and Kure, and may have a clockwise migration pattern
that occasionally includes Johnston on the return to their southern
Pacific breeding islands.

From the east Pacific comes the Red-billed Tropicbird which
breeds on islands along the coast of Central and South America.

Waterfowl, Marsh, and Land Birds

The 34 species of waterfowl, marsh, and land birds recorded at
Johnston Atoll (Table 18) are herein divided into regular migrants
(seven species), irregular visitors (six species), stragglers (two
species), accidentals (16 species), and introductions (three species).

Regular Migrants: One of the regular migrants is the Pintail
which breeds in temperate North America, Europe, and Asia. The re-
maining six species are shore birds which are all arctic breeders.
Of the seven species, three--the Pintail, Bristle-thighed Curlew,
Pectoral Sandpiper, and Sanderling--are regular, but uncommon. The
remainder are common.

These regular migrant shorebirds breed during the summer in the
Northern Hemisphere and migrate south for the winter. Some use
Johnston Atoll as a "wintering ground"; others, particularly first-
year birds, spend the summer here.

Irregular Visitors: Whereas regular migrants have a set pattern
to their movements, the six irregular visitors have no set pattern in
their visits to Johnston. This group contains two ducks, both
temperate breeders; three gulls, all subarctic and temperate breeders;
and an owl, which breeds in the north-central Pacific and various
temperate areas.

Stragglers: The two species in this group rarely visit Johnston,
but the atoll is within their range. The Franklin's Gull breeds in
northwestern North America, and the Cattle Egret has its closest
breeding population in the main Hawaiian Islands.

Accidentals: The accidental species at Johnston are so classified
because they are out of their normal range of occurrence. Fourteen
species are arctic and temperate breeders. Two land bird species are
introduced north-central Pacific breeders. The rain-water puddles on
Johnston may attract accidental birds, especially shorebirds.

Introductions: Three species of birds have been introduced by
man. The Rock Dove breeds and the Domestic Chicken probably breeds;
both species are semi-tame. The Society Finch is caged.

Annual Cycles

Among the bird species at Johnston Atoll, annual breeding and
population cycles vary.

120

Breeding Cycles: The seabirds breed during all seasons of the
year (Fig. 37). Most have short, distinct breeding periods; some
have extended breeding cycles. Based on peak breeding periods, the
12 present breeding seabird species are grouped as follows (Table 19):
winter and spring breeders (one species), spring and summer breeders
(nine species), summer breeders (one species), summer and fall
breeders (one species); none is a fall and winter breeder. Two
former breeding species were probably winter and spring breeders;
another was most likely a spring and summer breeder.

Table 19. Peak breeding period of Johnston Atoll birds

Winter-Spring Spring-Summer Summer Summer—-Fall
Gray-backed Tern Christmas Shearwater Bulwer's Petrel Wedge-tailed
Red-tailed Tropicbird Shearwater

Brown Booby
Red-footed Booby
Great Frigatebird
Sooty Tern

Brown Noddy
Black Noddy
White Tern

Winter and Spring: The single winter and spring breeder is the
Gray-backed Tern. In the northwestern Hawaiian Islands, where space
and nest site competition is no problem, this species is a spring and
summer breeder. Apparently because of nest site competition with Sooty
Terns on the reduced space available at Sand Island, the Gray-backed
Terns have shifted their nesting cycle so that first eggs are laid by
early or mid-winter. Last chicks fledge by late summer. The two
former breeders, Black-footed Albatross and Laysan Albatross, were
probably winter and spring breeders.

Spring and Summer: Of the nine species with a spring and summer
peak breeding period, one is a procellariiform, four are pelecaniforms,
and four are charadriiforms. Egg laying and fledging within this
group varies. Only two species actually start laying eggs during the
spring months: Christmas Shearwater and Black Noddy. Individual birds
of the remaining seven species commence laying during the winter months--
the Brown Noddy as early as December, the Red-tailed Tropicbird, Brown
Booby, Great Frigatebird in January, and the Red-footed Booby, Sooty
Tern, and White Tern in February. Of the nine spring-summer breeders,
only the young of two species (Black Noddy and White Tern) completely
fledge by the end of the summer months. The young of two species
(Christmas Shearwater and Sooty Tern) normally commence fledging in
the summer and extend into early fall. The Brown Noddy extends its
fledging period to late fall. Fledging of the remaining four species
(Red-tailed Tropicbird, Brown Booby, Red-footed Booby, and Great
Frigatebird) extends into early winter. The Blue-faced Booby, a
former breeder, probably had a spring and summer peak breeding season.

12d

Bulwer's Petrel e202 000 00 EEE ITT IDX

Wedge-tai led
Shearwater 00000000 9000

Christmas

Shearwater e00 00000 — pps [|| | PT

Red-tailed
Tropicbird

Brown Booby

Red-footed Booby

Great Frigatebird

Gray-backed Tern S @00 000000

Sooty Tern

Brown Noddy

Black Noddy

White Tera

J F M A M J J A S 0 N D

Figure 37. Breeding cycles of seabirds at Johnston Atoll; stippled area
represents eggs, barred area young, and black dots non-breeding
birds.

22

Summer: Only one species, Bulwer's Petrel, a procellariid, has
a peak breeding. season that is contained in a single three-month
period. Actually this species could be considered as a spring and
summer breeder since courtship begins in late March and first eggs
are laid the last of May. Or it could be considered a summer and fall
breeder because a few fledglings do not leave until early September.
The cycle of this species in the northwestern Hawaiian Islands is in
fact slightly later and is there considered to be a summer and fall
breeder.

Summer and Fall: The Wedge-tailed Shearwater, a procellariiforn,
is the only species at Johnston Atoll that has a summer and fall peak
breeding period. Eggs are laid in early June and young fledge by late
fall or early winter.

Extended: Although all 12 breeding species have a peak breeding
period, five species are considered to have an extended breeding
season. These are the Red-tailed Tropicbird, Brown Booby, Red-footed
Booby, Great Frigatebird, and Brown Noddy. In all of these species,
the egg laying period and fledging period each extend at least over
a three-season span.

Population Cycles: Many of the bird species using Johnston Atoll
leave the Atoll during part of each year; others stay throughout the
year (Fig. 37 and Table 20). Even so, all have a population buildup
sometime during the year.

Breeding Seabirds: Although the breeding population consists
of 12 species, five species are dominant both in total numbers and in
their importance to the surrounding ocean (Fig. 38). The Sooty Tern,
with a mean population of 300,000 to 310,000 breeding birds in March,
April and May, makes up 95 percent or more of the total island
population between February and July; perhaps as many as 600,000
Sooty Terns use Johnston Atoll annually.

Red-footed Boobies, whose mean population ranges up to 3,/50
birds, ranks second in seabird numbers in winter and spring; most of
these birds are transients for only a few young are produced each year.
The Brown Noddy ranks third in mean population numbers, but ranks
second in number of young (about 1,000) produced each year. The
Wedge-tailed Shearwater ranks fourth in numbers of adults using the
atoll and is present only from March to early December; it ranks
third in numbers of young (up to 250) produced annually. The Great
Frigatebird ranks fifth in population numbers (a mean peak of 750 in
March and April), but only produces about 130 fledglings yearly. Mean
monthly populations for all the other species, combined, never totals
more than 600, nor lower than 300.

As is evident from Figure 38, bird population numbers are extremely
high (up to 310,000) from mid-winter through summer. This coincides
with the peak breeding periods. During the fall and early winter the
total population usually is below 9,000 birds.

£23

Table 20. Monthly occurrence of non-resident birds at Johnston Atoll

Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Black-footed Albatross xX xX
Laysan Albatross Xx Me 5
Phoenix Petrel xX

Newell's Shearwater x x

Sooty Storm Petrel a6
Red—billed Tropicbird ee

White-tailed Tropicbird Re kere, ke ee ONE PEN xX
Blue-faced Booby se eee Kk OK OX ORL UK CX OUX:COUX
Lesser Frigatebird xX xX

Cattle Egret nol) Week” XX RU RR Re eK
Pintail x) eee” * * * * ae ON a
American Wigeon Re ae
Northern Shoveler xx

Peregrine Falcon

American Golden Plover XEN ARG ce ON ee
Black-bellied Plover xX xX
Semipalmated Plover

Bristle-thighed Curlew EV I RS RR KOO 1
Lesser Yellowlegs

Spotted Sandpiper

Willet

Wandering Tattler De Peer see CER RE oe re
Ruddy Turnstone OR EY Rie eR ee PAG eee
Dowitcher sp.

Sanderling Mee a OR ee, eK
Western Sandpiper

Pectoral Sandpiper ie > FOE
Sharp-tailed Sandpiper

Buff-breasted Sandpiper

Ruff X

Wilson's Phalarope x
Glaucous-winged Gull X
Herring Gull

Laughing Gull

Franklin's Gull x

Gull sp. X hee eK. X
Elegant Tern Rm

Blue-gray Noddy 7 ne xX

Short-eared Owl XG Kar emg RK

Skyiark

Japanese White-eye xX X X

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124

320,000

270,000

ect Sooty Tern

220,000 Red-footed Booby

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Wedge-tailed Shearwater
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J iP M A M J J A S 0 N D
Figure 38. Monthly cummulative bird populations, Johnston Atoll 1963-1969.

25

Of the 12 breeding species, seven are present on the atoll through-
out the year (Fig. 37). The remaining five species spend from 1 1/2
months to six months away from Johnston. Most of these probably leave
the central Pacific and migrate to distant areas.

Former Breeding Seabirds: The Blue-faced Booby is present, though
uncommon, throughout the year (Table 20). Black-footed Albatross have
been recorded on Johnston Atoll in November, and March, while Laysan
Albatross have been observed during November, December, and July; both
albatrosses may occur offshore during late winter and early spring.

Regular Migrants: Of the seven regular migrants, three--American
Golden Plover, Wandering Tattler, and Ruddy Turnstone--are known in all
months (Fig. 39 and Table 20). Although the Wandering Tattler and
Sanderling are found in low numbers throughout the year, American Golden
Plovers and Ruddy Turnstones show peak population periods in fall and
mid-winter and early spring; these peaks correspond to migrations from
Arctic breeding areas in fall and migrations north in mid-winter and
early spring. A slight population decline occurs in early winter;
lowest populations occur during the summer months when only immatures
remain on the atoll.

The Bristle-thighed Curlew is known from all months except July.
The Pintail has been observed in most years from September through March;
captive birds have been kept from April through August. The Pectoral
Sandpiper has been recorded in most years in April and May and in
September, October and November.

Irregular Visitors, Stragglers, and Accidental Birds: The monthly
occurrence of the irregular visitors, stragglers, and accidental species
is presented in Table 20.

As for the seven visiting seabirds, the White-tailed Tropicbird
has been recorded from all months except for June, and October and
November. The Red-billed Tropicbird, however, is known only from
April, May, and June. Similarly, the Blue-gray Noddy is known from
April, May, and July. The Lesser Frigatebird has been recorded during
March and August. The Newell's Shearwater also has only occurred during
two months--June and August. The Phoenix Petrel and Sooty Storm Petrel
have been seen only one month, respectively September and December.

Of the six irregular gull visitors, the Laughing Gull has been
recorded from four months, the Herring Gull from three months, and
Glaucous-winged Gull from two months. The American Wigeon and
Northern Shoveler are each known from two months. The Short-eared Owl,
however, has been observed in all months except August, September, and
October.

One of the two stragglers, the Cattle Egret, is known from all
months except June and July, while the other, the Franklin's Gull, is
known from only March.

130

120

110

100

NUMBERS

J F M A Maw J A S 0 N D

Figure 39, Monthly mean shorebird populations for Johnston Atoll, 1963-1969;
Golden Plover (solid line), Ruddy Turnstone (dots), Wandering
Tattler (dashes).

Of the 15 accidental waterfowl, marsh, and land birds, the
Japanese White-eye is known from five months, the Black-bellied Plover
from four months, the Semipalmated Plover and gull species from three
months each, the Peregrine Falcon, Spotted Sandpiper, Dowitcher
species and Sharp-tailed Sandpiper from two months each; the remain-
ing eight species are known from one month each.

Introduced Birds: The three introduced bird species are known
from all months. Furthermore, both breeding species probably breed
year-round.

At-Sea Birds: In August 1963, a 50,000 square mile, rectangular,
pelagic grid was established by the POBSP, centered approximately 175
miles southwest of Johnston Atoll. In all, 42 monthly survey cruises
were conducted in this grid through February 1967. Birds, primarily
seabirds, were observed along 22,898 miles of daytime travel and along

27,

10,819 miles of night travel. Pelagic observations maintained for
more than 2,500 hours in daytime and more than 1,150 hour at night
recorded 33,261 birds of 41 species. Tables 21 and 22 show, respect-
ively, monthly occurrences and area derivation of these species.

Figure 40 presents the monthly fluctuation in the number of birds
within this grid. A definite 12-month diurnal cycle exists which has
four basic features.

There is a population low from late December through early March
and is primarily a reflection of the absence of Sooty Terns and Wedge-
tailed Shearwaters. The major elements of the avifauna at this time
are probably Johnston Atoll non-breeding species, although at least
one north Pacific non-breeding species and two northwestern Hawaiian
Islands breeding species regularly occur in small numbers.

A spring peak exists from late March through early May and is
primarily the result of the arrival of Johnston Atoll breeding species
in the area. In addition, the number of birds is augmented by the
passage of migrants moving between distant areas, primarily from the
SouEn Pacific to the north Pacific.

There is a summer plateau from late May through August. Johnston
Atoll breeding species predominate at this time. The numbers are,
however, supplemented by sojourners from the south Pacific.

The cycle shows a fall migration peak-—-higher even than the spring
peak--from September through early December. Migrating seabirds, moving
from the north Pacific to the south Pacific, make up the bulk of the
pelagic population at this time. Sooty Terns and Wedge-tailed Shear-
waters, perhaps from the northwestern Hawaiian Islands and passing
through on migration, also contribute to the September peak.

Figure 41 presents the monthly diurnal population fluctuation
within the grid by species group. Two groups, the shearwater-petrel
group and the tern group, account for over 90 percent of all birds
present. The shearwater-petrel group accounts for 54 percent of all
birds throughout the year; it is the most abundant of all groups from
July through December. The tern group comprises 39 percent of all birds
present throughout the year; it is the most abundant group from March
through June. The tropicbird group accounts for two percent of all
birds present year-round. The booby group accounts for only one percent
of all birds present throughout the year, but is the most abundant group
from January through February.

In all, 21 seabird species were recorded during the nocturnal ob-
servations. Because of the difficulty in observing most species, only
Sooty Terns were counted with any degree of accuracy. The nocturnal
annual cycle (Fig. 42), therefore, primarily reflects the Sooty Tern
cycle rather than the total population. Nevertheless, two population
peaks were prominent, one in February, and another in March.

128

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SATIW JYVNOS OOL Yad SYIEWAN

133

O JOHNSTON ATOLL

Degrees North
q

i Tern

8q. miles

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(excluding S-SB)

S-SB Sooty-Siender-bilied

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Figure 41. Monthly diurnal seabird population fluctuations by species group

within the at-sea area 175 miles southwest of Johnston Atoll,
1963-1967.

BIRDS PER 10 HOURS

Figure 42. Total nocturnal bird sightings at sea 175 miles southwest of

Johnston Atoll, 1963-1967.

134

Effects of Weather Patterns: The annual breeding and population
cycles presented in the two preceding sections were based on data
from normal weather patterns from 1963 through 1968. The 1969
weather pattern, however, was abnormal.

The 1969 season advanced normally until early February, when
the weather pattern was disrupted by a series of low pressure dis-—
turbances moving across the area from the west. During this period
winds shifted to the south, occasionally southwest, and for one brief
period of strong winds and heavy rain, to the north. Heavy rains
fell on several days during this time, particularly 3 and 8 February.
The normal trade wind pattern was re-established in the last half of
February, and no further weather disturbances occurred through the
end of April.

Concurrent with these weather changes came a decline in numbers
and changes in behavior of nearly all bird species on the Atoll. Most
drastically influenced were the three major terns: Sooty Terns, Gray-
backed Terns, and Brown Noddies, but Christmas Shearwaters, Red-tailed
Tropicbirds, Brown Boobies, and Great Frigatebirds were noticeably
affected.

The connecting factor between the weather pattern and changes in
bird numbers remains obscure. Food availability may have decreased,
making it necessary for the birds to stay at sea in order to fulfill
their nutritional requirements, or perhaps the combination of changed
wind patterns and heavy cloud cover affected the navigational pro-
cedures of the birds such that they were unable to find or return to
the island.

The restoration of normal weather in late February was followed
by increases in numbers and nesting activity in all breeding birds,
but most were delayed in comparison with other years. Sooty Terns
were about ten days behind their usual cycle, and Brown Noddies did
not commence laying until over a month later than in former years.

There was evidence that unusually favorable feeding conditions
prevailed in March and early April. Sooty Terns spread to occupy a
greater area than ever before, and two-egg clutches, many of them
identical in proportions and color pattern, were more common than
in any other year. Brown Noddies accelerated laying so that, although
the first eggs were not laid until 20 March, the total number laid by
mid-April was three times the total produced by that date in 1968.
Gray-backed Terns re-nested beginning in late March, and laid several
times more eggs than they normally lay in the re-nesting attempt.

Four Brown Booby pairs laid three-egg clutches, the first ever re-
corded for Johnston Atoll. The only outstandingly conflicting
evidence comes from Christmas Shearwaters, which did not begin laying
until early April, and produced only three eggs during April. Perhaps
the shortage of food in February had a more drastic effect on these
birds because of the larger size of their egg and consequent greater

35

need for heavy feeding during its development. Shearwaters are
thought to lay no replacement eggs, and the forced resorption of a
partially developed egg might block further egg production in that
year.

Ecological Distribution Within the Atoll

The four islands at Johnston Atoll differ in size, height,
soil, vegetation, fresh-water supply, and degree of human disturbance.
Major differences in avifaunal distribution, especially breeders, are
found between disturbed and non-distrubed islands (Table 18).

Of the 56 bird species known from the atoll, 52 are known from
Sand Island. Furthermore, 44 species are known from the original
portion, while 35 are from the man-made part, In all, 35 species
are known from Johnston Island, while eight are from Akau Island
and five are recorded from Hikina Island.

Seabirds:

Breeding: Thirteen breeding seabird species are known from
Sand Island; 11 species presently nest there (Table 18), Likewise
14 breeding species are known from Johnston Island; however, only
three species currently nest there. One species has been known to
nest in recent years on man-made Akau Island; two have nested on
man-made Hikina Island.

Visitors: All seven seabird visitors are known from Sand
Island, possibly attracted there by other seabird species; six are
recorded from the original portion, while only two are from the man-
made part (Table 18). Three species are known from Johnston Island.
One species is known from Akau Island, none from Hikina Island.

Waterfowl, Marsh, and Land Birds:

Regular Migrants: All seven regular migrants are known from
both original and man-made portions of Sand Island (Table 18).
Likewise, all but one species-—-the Pectoral Sandpiper--are known
from Johnston Island. Four species have been recorded from Akau
Island; three species are from Hikina Island.

Irregular Visitors, Stragglers, and Accidentals: In all, 22

irregular visitor, straggler and accidental waterfowl, marsh and
land bird species are known from Sand Island: 16 from the original
portion and 12 from the man-made part. Ten species have been
recorded from Johnston Island. Only one species is known from Akau
Island; one is known from Hikina Island.

Introductions: Two of the introduced species are known from
Johnston Island. One species has been recorded from Sand Island.

136
Island Accounts:

Avifaunal components of each island, listed alphabetically,
are discussed in the following section.

Akau Island: Eight bird species have been recorded (Table 18).
The island was built in 1964 and the low population and breeding
numbers recorded since are due primarily to human disturbance.

Total daytime populations are extremely small; populations,
especially of roosting shorebirds, probably increase at night.

Only one species--Gray-backed Tern--has nested on Akau. That
occurred only in 1964 after the island was completed, but before
building construction started. The nests were placed on bare sand
but most were destroyed by construction; none has nested since.
Great Frigatebirds roosted on various objects on the island, and
Brown Boobies probably roost here occasionally. White-tailed
Tropicbirds have also been seen flying over the island.

Four regular migrant shorebirds--American Golden Plover, Ruddy
Turnstone, Bristle-thighed Curlew, and Wandering Tattler--have been
observed on the beaches of Akau.

Only one irregular visitor--a land bird--has been recorded.
Short-eared Owls sometimes roosted here when present on the atoll.

Because of human inhabitation (buildings, etc.), small size,
and lack of nesting habitat, Akau Island attracted very few bird
species. However, with the proposed reduced human activity after
fall 1973, the island will no doubt attract additional bird species
and perhaps even breeding seabird species.

Hikina Island: Since Hikina Island was built in 1964 only six
species of birds have been recorded, primarily because of human
disturbance (Table 18).

From 1964 through 1969 one seabird occasionally roosted here
and three of the regular migrant shorebirds commonly feed on the
beaches. Roosting shorebirds frequented the island at night.

Hikina Island has continued to attract small numbers of bird
species because of human disturbance, the island's small size, and
a lack of suitable nesting habitat. Two species of seabirds, however,
did nest in 1973 because of reduced human disturbance. These two
species and most likely others will continue to breed here as long
as human disturbance is kept at a minimum.

Johnston Island: Birds totaling 35 species have been recorded
from Johnston Island (Table 18). Of these 35 species, 17 were sea-
birds and 18 were waterfowl, marsh, and land birds.

Of the 17 species of seabirds, four presently breed and ten
formerly bred on the island. Since 1963, only one of these ten

137

former breeders has ever been recorded from Johnston Island. This
decline in breeding species, as well as a correspondingly low bird
population, can be directly attributed to human disturbance beginning
in the late 1930's.

Presently White Terns utilize the tallest Casuarina trees and
various man-made objects on which to roost and lay their eggs. Like-
wise, Black Noddies roost and nest in the tallest Casuarina trees.
Red-tailed Tropicbirds nest under the larger bushes--particularly
Pluehea--scattered about the island. Wedge-tailed Shearwaters nest
under the denser bushes and have been known to nest under various
buildings.

Sand Island: Fifty-two species of birds have been recorded
from Sand Island (Table 18). Of these 52 species, 44 are known
from the original portion and 35 are known from the man-made portion.

Of the 35 species known from the man-made portion, three species
of seabirds presently breed. An additional six species of seabirds
nested here while the island was uninhabited during the late 1940's
and 1950's. Bulwer's Petrels presently nest in cavities formed by
the rocks along the causeway. Red-tailed Tropicbirds and at times
Wedge-tailed Shearwaters nest under the dense bushes, especially
Seaevola and Tournefortia, growing around the buildings.

Of the 44 species recorded from the original portion, 11
species of seabirds presently breed. In addition, two breeding
species of seabirds no longer nest either here or elsewhere on the
atoll. The population cycles shown in Figure 38 are predominantly
of birds on Sand Island. During the spring and summer, Sooty Terns
nest on the bare ground over most of the island (Fig. 35) and are
the most prominent species. The 1963 nesting areas for other species
are shown in Figures 43 and 44. Nesting areas are also illustrated
under individual species accounts. Brown Noddies nest on the ground
around the periphery of the island. Black Noddies nest in the low
Amaranthus bushes, when present. Gray-backed Terns nest on the
ground on the northeast peninsula and the southwest islet. Red-
tailed Tropicbirds nest under low vegetation, and various cement
slabs and other objects, about the island. Wedge-tailed Shearwaters
nest in burrows over much of the island; most burrows are placed so
that vegetation roots help support the surrounding soil. Christmas
Shearwaters, and at times Bulwer's Petrels, nest under cement slabs
and various wooden timbers lying about on the surface of the island.
Brown Boobies nest on the ground on the southeast hill, the south
edge, the northeast peninsula, and the southwest islet. Red-footed
Boobies build their nests on the east hill, on the Tournefortta bush
northeast of the transmitter buildings, and on the various pilings
around the island. Great Frigatebirds nest along the east hill and
the south edge.

138

© Wedge-tailed Shearwater

e Brown Noddy

ee ese,

@ Gray-backed Tern

N
* Christmas Shearwater
© Brown Booby
@ Red-tailed Tropicbird
NST OS 62005 90
Ge A HS
rome)
0 100 200
|
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Figure 43. Nesting areas of ground nesting birds (except Sooty Terns) on
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%* Black Noddy

@ Great Frigatebird N
0
@ Red-footed Booby
0 100 200
(= = |
Feet

Figure 44, Nesting areas of birds which normally nest in low vegetation on
the original portion of Sand Island, Johnston Atoll, 1963,

139

Mortality

The largest cause of mortality during the POBSP studies was
the guywire system of the LORAN-C antenna on Sand Island. This
system contained 24 top-loaded guywires which stretched from the
top of the 625-foot tower to concrete anchors located in the lagoon
in a circle around the island. In addition, three other sets of
guywires stretched from part way up the tower to two sets of concrete
anchors located on or near the island itself. One of these latter
sets, stretched low across the southeast corner of the island,
accounted for an abnormally high amount of mortality because it
crossed a heavily-used bird area and was closer to the ground than
the other guywires.

Table 23 shows monthly mortality of Sooty Terns from this
guywire system recorded during five months of the breeding season
in 1965. Since roughly 600,000 adult Sooty Terns use the island
during the breeding season and about 60,000 young fledge annually,
the 5,062 dead birds represent about 0.8 percent of the total
population. Adult mortality is about 0.5 percent, while immature
mortality is about 3.7 percent. Adult Sooty Tern losses were
greatest just as egg laying began (March) when highest numbers
swirled low over the island; chick losses were highest when they
began to fly. Of the 5,062 dead birds 14.4 percent were banded
(adults 9.2 percent, immatures 21.1 percent). There appeared to be
a decline in numbers of adult Sooty Terns lost to the guywires
during the last two years (1968 and 1969) of POBSP studies.
Possibly the remaining birds were becoming wary of the wires as
undoubtedly most adults had had close brushes with the wires and
it is possible they could learn to avoid them.

Mortality records were also kept for other species during the
five-month study. Although not as spectacular number-wise as for
Sooty Terns, two other species actually had higher mortality rates.
Surprisingly, shorebirds had the highest rate at 5.8 percent (nine
dead). Great Frigatebirds were next at 1.8 percent (24 dead, 8.3
percent banded). Although no figures are available, a high percentage
of the Great Frigatebirds that fledged each year broke their wings
on these wires and died. The mortality rates for Brown Noddies
(33 dead, 18.2 percent banded) and Wedge-tailed Shearwaters (25
dead, 44 percent banded) were the same--0.8 percent--as for Sooty
Terns. Red-footed Boobies (nine dead, 11.1 percent banded) had
0.3 percent mortality, whereas Red-tailed Tropicbirds (one dead)
had 0.5 percent mortality.

Dead birds from the guywires presented little disposal problem
because of the presence of dermestid beetles (Dermestes ater).
These insects were at their lowest numbers in mid-winter when re-
turning Sooty Terns first began hitting the wires. The population
quickly responded to the increase in food and by the time the terns
reached their peak abundance, the dermestids could eliminate

146

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carcasses, even of frigatebirds and boobies, in little more than a
day. This made it unnecessary, and even undesirable, to initiate
any kind of dead bird removal program except where carcasses ap-
peared near buildings.

Of lesser importance in cleaning carcasses were cockroaches,
ants, and flies. Fly populations generally were low because
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Banding and Interisland Movement

The use of U.S. Fish and Wildlife Service bands is a widely
accepted method for marking birds. By using these serially numbered,
metal (usually aluminum), leg bands at Johnston Atoll, and throughout
the central Pacific, the POBSP was able to (1) study bird migration,
(2) obtain species longevity, (3) recognize individuals from their
neighbors, and (4) estimate populations by the mark-and-recapture
method. To facilitate sighting Johnston-banded birds at sea and on
distant islands, most banded adults and young were also tagged with
a blaze-orange, plastic, leg streamer.

Banding: N.P. Ashmole, then of the B.P. Bishop Museum, Honolulu,
Hawaii, banded the first birds at Johnston Atoll in February 1963;
he banded 72 seabirds of five species. Birds totaling 303,732 of
21 species were banded at Johnston Atoll by POBSP personnel from
July 1963 through September 1969 (Table 24). The POBSP banded more
birds here than anywhere else in the central Pacific. Amerson, then
of the Smithsonian Institution, banded 97 birds of five species in
November 1973.

From 1967 through 1969, primary emphasis turned to recapture
of banded birds, but chicks of most species were banded in as large
numbers as possible in order to continue to mark known-age birds
for future studies.

By far the majority of these 303,901 birds was banded at Sand
Island. Only a few of two species--Red-tailed Tropicbird and White
Tern--were banded at Johnston Island. None was banded at Akau and
Hikina Islands.

Of the total banded (Table 24) there were: 285,526 Sooty Terns,
7,979 Brown Noddies, and 6,517 Wedge-tailed Shearwaters, 1,047 Red-
footed Boobies, and 1,005 Great Frigatebirds; the remaining 16
species only accounted for 1,827 banded birds.

Interisland Movement: In all, 18 species totaling 60,932 birds
have been recaptured at Johnston Atoll since 1958 (Table 25). Of
this total, 60,526 birds were originally banded on the atoll, while
406 were banded elsewhere.

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The 406 captures of birds from other localities involve 13
species, but represent only 388 birds as some were captured more than
once. These birds were originally banded at 15 localities (Table 26).
The majority came from the northwestern Hawaiian Islands with the most
from French Frigate Shoals (127 birds, or 33 percent). Laysan Island
was second at 51 birds (13 percent) and Kure Atoll third at 40 birds
(10 percent). Wake Atoll, however, was second overall with 60 birds
(15 percent) moving to Johnston. Only five birds, three being
questionable records, possibly moved to Johnston Atoll from islands
to the south.

In addition to birds coming to the Atoll, 345 birds of 12 species
originally banded at Johnston Atoll were captured on 20 other atolls or
island groups and from 13 at-sea localities (Table 26). Of these 345
banded birds, 273 (79 percent) went to the Hawaiian Islands and Wake,
while 50 (17 percent) moved to the west and southwest Pacific. Twelve
banded birds (most questionable records) possibly moved to the Phoenix
Islands.

The number of individual banded birds involved in interisland
movement, both to and from the atoll, totals 733. French Frigate Shoals
(232 birds), Laysan Island (92 birds), Kure Atoll (75 birds) and Wake
Atoll (73 birds), in that order, are the etolls most frequently involved.

The overall POBSP banding and recapture program in the central
Pacific has shown that bird movement between the Line and Phoenix
Islands area and the Hawaiian Islands and Johnston area was virtually
nil (Amerson, 1968). Movements between islands within each of these
areas was much larger, and the number of birds returning to their island
of banding was still greater. Banded birds from both areas, however,
were recaptured in the far western Pacific.

Banding data seem to indicate that Johnston Atoll is a major focal
point for interisland movements in the north-central Pacific. Movements
to and from Johnston Atoll are highest among all the islands in this
area. This may simply result from full-time POBSP personnel coverage
at Johnston, or it may actually reflect a high degree of movement to
and from the atoll (POBSP, 1967b).

Avifaunal Affinities

The Johnston Atoll avifauna shows clear affinities with the
Hawaiian Islands, both taxonomically and in movement patterns. These
are, of course, the closest islands to Johnston.

Particularly significant are movements of breeding species:
Red-footed Boobies, Great Frigatebirds, Blue-faced Boobies, Brown
Noddies, Wedge-tailed Shearwaters, and Black Noddies (list is in descend-
ing order of numbers of interisland movements from Table 26), in which
most of the recorded movements involving Johnston are with the Hawaiian
Islands. The major exceptions are Sooty Terns and Red-tailed Tropicbirds,

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148

in which a disproportionately high number of movements are from Wake
Island, which lies west of Johnston at a greater distance than any of
the Hawaiian Islands.

Furthermore, those breeding species for which subspecies or
color phases can be used to determine affinities, are allied with the
northern islands: Wedge-tailed Shearwater, Red-footed Booby and Blue-
gray Noddy are of the typical northern color phases (or subspecies in
the case of the Blue-gray Noddy).

As for the non-breeding species, movements of birds not banded but
collected, however, remain mostly from the Hawaiian Islands: Newell's
Shearwater, Sooty Storm Petrel, White-tailed Tropicbird, Cattle Egret,
Blue-gray Noddy, Short-eared Owl, and Japanese White-eye. There are
a few recorded movements from the south and east. The Phoenix Petrel
and Lesser Frigatebirds are equatorial breeders, although Lesser
Frigates have been recorded from Wake and the northwestern Hawaiians,
and may have a circular migration pattern that includes (occasionally)
Johnston on the return to the equatorial breeding islands. From the
eastern Pacific comes the Red-billed Tropicbird, and from the northeast,
possibly via Hawaii, have come a number of accidentals: Glaucous-winged
Gull, Laughing Gull, Franklin's Gull, Elegant Tern, and possibly
Herring and Western Gulls.

Techniques

Although most of the study techniques used by the POBSP at
Johnston Atoll were straight-forward and self-explanatory, a few
comments on the accuracy and relative values of the results are
necessary to clarify the meaning of the data, and to emphasize that in
some cases, tables and discussion are carefully labelled to state what
the figures actually represent, rather than what they ideally should
represent.

For example, reported numbers of birds in the semimonthly reports
ideally were to be the total number of birds using the atoll during the
period in question. Clearly, this figure is unattainable for even the
most conspicuous and easily enumerated species. It would require
constant surveillance on the entire atoll for the entire period, with
all new birds detectable. Thus the figure can mean no more than the
number of individuals of a given species known to use the atoll during
the period. In many cases figures reported really are not estimates of
total numbers using the islands, but of maximum seen at one time, which
is probably far lower in nearly every instance than the total number
using the island during the period, because it is extremely unlikely
that all would be present at any one time, even at night for roosting.
Reported counts, then, should be regarded as indices to the numbers
using the atoll, rather than accurate estimates, although in a few
cases they may be fairly accurate.

In general, the fewer individuals and to a lesser extent, the
larger and more conspicuous the bird, the more likely is the count to

149

accurately represent the true number of individuals using the island.
An extreme example is the Cattle Egret, which occurred no more than
two at once, were comparatively large and conspicuous and were highly
predictable in habits and habitat.

It is recognized that the capture-mark-recapture (or observe)
calculations for populations do not meet the assumption of no turnover
in the population between marking and observation. However, it is the
only technique available in most cases and the estimates are presented
in a few cases where no other objective estimates are available. They
may even be used for estimating the rate of turnover, as for example
in Sooty Terns, Black Noddies, and White Terns. Some indication of
populations are thus possible.

Rapid turnover of personnel necessarily had a deleterious effect
on data gathering, through studies being dropped or changed when per-
sonnel changed. However, a large body of reliable information was
assembled in the field notes and semimonthly reports. Basically these
species accounts will summarize in detail those aspects of the life
histories and populations for which comparable data were obtained for
all species of ecological equivalence.

Specimens

The first scientific bird specimens were collected by Kern in
March 1859 when the U.S. Schooner FENIMORE COOPER visited Johnston
Atoll (Brooke, ms.). What species were collected are not known;
however, they were lost when the ship later burned and sank in Japan
(Brooke, 1955).

Prior to the first POBSP in July 1963, 93 specimens of 16 bird
species--all collected by Alexander Wetmore in July 1923--were known
fiom Johnston Atoll. POBSP personnel collected 616 specimens of 41
<pecies from 1963 through 1969. These 709 specimens of 41 species
are all housed in the bird collection of the National Museum of Natural
History (USNM), Washington, D.C.

Fifty-one new species distributional records--two first specimen
records of species previously unknown for the central Pacific Ocean,
one first specimen record of a species previously known only from a
sight record in the north-central Pacific Ocean, 38 first specimen
records for the atoll, and ten first sight records--and four new
species breeding records are reported herein. Table 27 summarizes
these specimen and distributional records. These records fall into
two categories, one composed of species that regularly occur on
islands in the north-central Pacific Ocean but which cannot be con-
sidered unusual, and the other group composed of species of uncommon
or seldom-documented occurrence on islands in the north-central
Pacific Ocean.

150

Table 27. Bird specimens and summary of new records from Johnston

Atoll
Collected Specimens

Species Wetmore POBSP Total New Records
Phoenix Petrel i i sr
Bulwer's Petrel 9 1 10 sr br
Wedge-tailed Shearwater 7 95 100 sr
Christmas Shearwater 10 il dail sr, br
Newell's Shearwater 2 2 sr
Sooty Storm Petrel i i sr
Red-tailed Tropicbird 5) 24 29 sr
Blue-faced Booby 7 3 10 sr, br*
Brown Booby 5 8) 14 sr
Red-footed Booby 8 2 29 sr
Great Frigatebird 6 61 67 sr
Lesser Frigatebird oq
Cattle Egret ie
Pintail 4 4 sr
American Wigeon 3 2 Z sr
Northern Shoveler 1 1 sr
Domestic Chicken 58
Peregrine Falcon I il sr
American Golden Plover 1 18 IW) sr
Black-bellied Plover UL i sr
Semipalmated Plover 2 2 sr
Bristle-thighed Curlew 3} 5 Sr
Lesser Yellowlegs i! i sr
Spotted Sandpiper i il sr, SFR
Willet ye
Wandering Tattler 2 2 4 sr
Ruddy Turnstone 1 9 10 sr
Dowitcher species rE
Sanderling a i} sr
Western Sandpiper 18
Pectoral Sandpiper 8 8 sr
Sharp-tailed Sandpiper 2 2 sr
Buff-breasted Sandpiper 1 1 sr
Ruff 1 1 sr
Wilson's Phalarope 1 iL Sr, SR
Glaucous-winged Gull 2 2 sr
Herring Gull Ts
Laughing Gull il iL
Franklin's Gull il 1 sr
Gull species 1** 1** E
Gray-backed Tern 5 14 19 sr
Sooty Tern 2 224 236 sr
Elegant Tern ‘ it 1 sr, SFR
Blue-gray Noddy 2 it 3 sr
Brown Noddy 6 78 84 sr

Black Noddy 6 6 sr

ils)

Table 27 srt (eont2)

Collected Specimens

Species Wetmore POBSP Total New Records
White Tern i! 7 14 sr
Rock Dove tbe
Skylark r
Japanese White-eye 2 2
Society Finch r

Totals 93 616 709

br=first breeding record; r=first sight record; sr=first specimen
record; SR=first specimen confirmation of a species previously known
only from a sight record in the north-central Pacific Ocean; SFR=first
specimen record for a species previously unknown from the central
Pacific Ocean.

*extirpated; **specimen lost.

Species Accounts

For descriptions and illustration of the 56 species of birds
recorded herein, the reader is referred to the ornithological sources
cited previously, especially King (1967).

BLACK-FOOTED ALBATROSS Dtomedea ntgrtpes
Status

Former uncommon breeding species; recent regular but scarce
winter visitor at sea near Johnston Atoll; two questionable records
of birds landing on Johnston and Sand Islands during the 1960's.
Small numbers probably nested on the sandy portions of Sand Island,
as well as on Johnston Island, prior to military occupancy.

Ecological Distribution

Johnston Island: One possible record exists. Sundell, a POBSP
employee, wrote in his field notes 22 November 1963: ''While on
Johnston in the evening I talked with an engineer who claims to have
seen a Black-footed Albatross over the island on November 17." No
further confirmation of this record was obtained.

Sand Island: The only breeding record for the atoll was a sight
record made by a member of Wetmore's 1923 party: '"'W.G. Anderson, of

152

our party, who had come in November 1922, as navigator for a fishing
sampan, informed me he saw one, with its egg, on the beach of Sand
Island. None were present during our work in July 1923" (Wetmore,
TIS!s¢b)he

One recent sight record is questionable. On 11 March 1964 a
Black-footed Albatross followed the USS ENERGY to within about 2.5
miles of Sand Island and departed just after a harbor tug came to assist
the ship to port. Bratley, a POBSP employee who was on Sand at the
time, reported in his field notes: "I could see the Energy at this
time from Sand, but I could not see if there was any bird following it."
On 12 March, Coast Guardsmen described to Bratley a bird seen sitting
on the causeway of Sand and flying around over Johnston that may have
been the Black-footed Albatross that had followed the ship in the day
before. No further confirmation could be made of this record.

Populations

The occurrence of albatross on Johnston Atoll in the past is
poorly recorded. During the 1800's there were no scientific visits and
no known observations reliable enough to determine the species composi-
tion of the atoll's avifauna. Brooke (ms.) reported "an albatross"
as being collected on 15 March 1859. A guano prospector, Le Grand
Brown, listed "the Gooney" as being present in 1905 (U.S. Nat. Archives,
R.G. 59, guano letters), but no quantitative significance can be
attached to this record. In addition to Anderson's breeding sight
record in November 1922 (Wetmore, ms. b), he (Anderson, 1954) reported
seeing a Black-footed Albatross there in 1929, but this may have been
an erroneous listing of the bird he saw in 1922.

Rice and Kenyon (1962) interpreted Wetmore's observations, including
the absence of albatross bones in the piles of bones left by feather
hunters shortly before Wetmore's visit, as evidence that albatross were
extirpated [prior to the time of said feather hunters]. It is equally
likely that the albatross population never exceeded a few birds, even
before the first feather poachers arrived. Johnston Atoll is the
southernmost known breeding island for Black-footed Albatross, unless
the record (Rice and Kenyon, 1962) for Taongi Atoll (14°35'N) in the
northern Marshalls is valid. Amerson (1969), however, found no
evidence to substantiate this Taongi breeding record.

An elderly Polynesian construction worker on the islands in the
late 1930's told Amerson of the POBSP in 1964 that albatrosses of both
species were on the island when construction work began, and that
they were shot by marines. Whether they were shot for lack of better
entertainment, or whether there was an official effort to rid the
islands of these birds because of the hazard to aircraft, is not known.

An article in the New Yorker magazine (Anon., 1945) listed the
Black-footed Albatross as occurring on Johnston Atoll, but this listing
was "based in part on the British Admiralty Sailing Directions, and

153

in part on information supplied by U.S. Navy personnel who had been
stationed on the island....'' (Hutchinson, 1950: 186), and is of little
value in determining the status of the species on the atoll.

Even if man left Johnston Atoll it is doubtful if Black-footed
Albatross would return to its islands to breed because of their strong
tendency to nest on their place of hatching. Those that may have
survived the feather hunters and early military construction have
probably long since gone elsewhere, or most likely have died.

Annual Cycle

The sight record of an egg in November 1922 by Anderson (Wetmore,
ms. b) falls within this species’ breeding cycle in the northwestern
Hawaiian Islands. There, adults arrive during the last part of
October, eggs are laid about mid-November, young hatch beginning late
January, and adults and young leave by the end of July (Richardson,
ICIS

Banding and Interisland Movement

No Black-footed Albatross were banded and no specimens were
collected from the island by POBSP personnel. No interisland movements
are known.

At-Sea Distribution

Brooke (ms.) saw "a black albatross" on 19 March 1859 while
sailing west two days from Johnston Atoll.

Black-footed Albatross do occur at sea in the grid southwest of
Johnston Atoll, where they have been found in small numbers from
December through April (POBSP, 1967a). The POBSP collected three
specimens at sea near Johnston Atoll (Appendix Table 7). They are
seldom found below 10°N in the central Pacific (King, 1967).

LAYSAN ALBATROSS Dtomedta tmmutabilts
Status

Former uncommon breeding species, presently a rare visitor; a
regular but rare winter visitor at sea near Johnston Atoll. Small
numbers nested on Johnston Island prior to mitlitary occupancy.

Ecological Distribution.

Johnston Island: Wetmore (ms. b) recorded one young bird on the
high barren areas of Johnston Island in July 1923. None was observed
in late April or May 1928 by Thurston (1928), although this species
may have been present in the late 1930's (see Black-footed Albatross
account). A few visited here in 1949, probably November or December
(Jensen, 1949).

154

Three were present in early winter 1957 (Manandic, pers. comm.).
Manandic, former SSgt., USAF, stationed on Johnston in 1957, knew
the birds of the atoll quite well and used the term "albatross" not
"sooney'' to describe the birds in question. He described them as
large white birds, with hooked beaks, a description that could hardly
have fit any other species.

Sand Island: One adult with a bare brood patch landed on the
road to the transmitter building on 10 December 1965. It was banded,
photographed, and ectoparasites collected before being released by
POBSP personnel. Another adult was seen soaring over the western
portion of Sand for at least ten minutes on 26 December 1966 by POBSP
personnel.

Populations

For the Laysan Albatross, as for the Black-footed Albatross,
Johnston Atoll was previously the southernmost extreme of the breeding
range. Small numbers bred on the atoll prior to military occupancy,
but now the species is a rare visitor.

The discussion of the occurrence of albatrosses before 1923, and
of Rice and Kenyon's (1962) interpretations of Wetmore's observations,
are included in the Black-footed Albatross account and apply equally
to the Laysan Albatross.

It is also unlikely that this species will return to Johnston
Atoll to breed, even if man decides to abandon the atoll in the future.

Annual Cycle

Wetmore (ms. b) wrote of the one young bird, nearly ready to fly
in July 1923, as: "This should not be taken as indicative of the
rarity of the species, since by July--most members of this species had
left their breeding grounds [in the northwestern Hawaiian Islands], and
were at sea.'"' The annual cycle of the Laysan Albatross in the Hawaiians
is similar to that described for the Black-footed Albatross, but the
cycle is about two weeks later (Richardson, 1957).

Banding and Interisland Movement

The only Laysan Albatross banded by POBSP personnel on the atoll,
an adult banded on Sand 10 December 1965, was seen and identified by
its orange, plastic, leg-streamer over Kure Atoll, nearly 1,000 miles
to the northwest, three days later, 13 December, by POBSP personnel
(Woodward, 1972).

At-Sea Distribution

Laysan Albatross have been observed only during one month--
February--in the grid southwest of Johnston Atoll (POBSP, 1967a).
This species seldom is found below 15°N in the central Pacific (King,
1967)

155

PHOENIX PETREL Pterodroma alba
Status

Accidental; one specimen record from Sand Island.
Observations

The only Phoenix Petrel recorded from Johnston Atoll was an adult
female Kepler and Lehner collected after it landed "in a Tribulus bush
in the Wedge-tailed Shearwater colony..." on 18 September 1964. The
bird appeared to be in good health and was kept alive for a few days
before it was killed and preserved.

Phoenix Petrels breed in the summer in the Line and Phoenix
Islands (Clapp, pers. comm.) and occur south to about 24°S latitude
in the central Pacific (Murphy and Pennoyer, 1952). Small numbers
occur at sea north to the Hawaiian Islands (King, 1967) but may only
rarely alight on islands in this area. POBSP (1967a) personnel
recorded Phoenix Petrels at sea in low numbers in the grid southwest
of Johnston Atoll only during October, November, January, and February.

Specimens

The Phoenix Petrel noted above is a new specimen record for
Johnston Atoll. Furthermore, it has never been recorded on land in
the Hawaiian Islands.

BULWER'S PETREL Bulwerta bulwertt
Status

Uncommon breeding species, present from March to early September.
Nests in rock crevices along the causeway, on the east and west shores,
and on the southwest islet of Sand Island; previously nested on
Johnston Island. About half of the 40 to 50 adults that return each
year attempt to breed, and at least five to ten young are produced
yearly. Adults return in March, lay in May or June, and depart just
ahead of the chicks which hatch in late June or early July and fledge
in late August or early September. Nests are in rock crevices mainly
along the man-made causeway of Sand Island.

Ecological Distribution

Johnston Island: -Wetmore (ms. b) in July 1923 found about 400
nesting in crevices in the rock ledges of Johnston Island with young
ranging from a week old to those partly covered with contour feathers.
Wetmore's field notes (ms. a) reveal something of their habits and
habitats:

156

Common, nesting in the rock ledges along the beach where
I hear their barking calls day and night. They are more
advanced in their breeding here than farther north as they
have young from a week old to those growing contour
feathers. The small young are as usual in petrels very
fluffy with heavy down about the head that almost hides
the eyes. Young beginning to molt into first plumage
appear much larger than the adult and in reality are
heavier because of the heavy layer of greasy fat that
covers the entire body. They remain hidden in the ledges
during the day, but at night come out into the open
shuffling about on the breast. At day they seek shelter
when exposed to light.

The larger ones resent handling and attempt to bite as do
adults when handled at this season. The young utter a
low whistling wheeze somewhat similar to the note of a
squab.

Adults delight to rest a few inches apart and with ex-
tended head and swelling throat utter their comical little
barking call, a note common to both sexes. When quiet they
rest prostrate on the breast and moving merely shuffle along
barely raising the body free of the sand and dropping back
at once.

About 400 on Johnston Island.

None has been recorded on Johnston Island since, most likely because
the natural rock crevices where Wetmore found them in 1923 have all been
destroyed by military activities.

Sand Island: Wetmore (ms. b) saw none on Sand in July 1923.
Because of their nocturnal and secretive habits, Bulwer's Petrels were
completely overlooked by POBSP personnel in 1963 but were, however, re-
corded as nesting from 1964 to 1969.

Bulwer's Petrels were seldom found except in the rock crevices where
they nested (Figs. 45 and 46). The distribution of these sites is shown
in Figure 47. Most nests were along the causeway, where angular rocks
and blocks of concrete form deep crevices in which the birds lay their
eges on a thin mat of grass stems. There were also a few sites under
chunks of concrete on the east and west shores of the original portion of
the island, some of the same ones used by Christmas Shearwaters. The
sizes of cavities required by these two species appear to be sufficiently
different to prevent competition between them for sites. On the south-
west islet a bird was heard calling in 1967 but could not be reached or
seen in the crevices, several feet deep, between concrete chunks used to
build the bunker. In 1969 a chick almost ready to fledge was caught there.
Nesting probably occurred there in other years as well. In 1967 one bird
was found among rocks along the south shore, but no nests were ever
discovered there.

57,

Bulwer's Petrel chick, about 2 months old, in front of rock
crevice nest site along causeway, Sand Island, Johnston Atoll,
25 August 1966 (POBSP photo by P, C, Shelton).

Figure 46. Concrete slabs on east shore, at base of northeast peninsula, |
Sand Island, Johnston Atoll, 8 May 1967, Christmas Shearwaters i
nested regularly under the large slabs at left. and Bulwer's
Petrels attempted to nest at least once under the slabs in center |
of photo. Brown Noddies nested in the edge of the Sesuvium i
growing just above the storm beach (POBSP photo by P. C. Shelton). .

158

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159

Rarely were Bulwer's Petrels found in the open, away from rocky
cover. One of the 1965 chicks that returned in 1969 was first found
sitting in the road on the causeway but it was found later in 1969 ina
cavity on the causeway. On 3 August 1966 a lone calling bird with a
bare brood patch was found on open ground north of the transmitter build-
ing among Sooty Terns. Almost exactly three years later, 2 August 1969,
the same bird appeared in the same place, again sitting alone among Sooty
Terns and calling. Possibly this bird was hatched, or had formerly nested,
at a now destroyed site under a rock on concrete slab near where it ap-

peared in these two years.

Populations

These nocturnal and secretive petrels are among the most difficult
to census of the less common birds using the island. They were completely
overlooked in 1963. Semimonthly estimates by different observers varied,
resulting in a wide spread in the ranges for estimates shown in Figure 48.
The mean of these estimates for each period probably represents the true
numbers fairly well, although the spring buildup may be steeper than shown.

80
Adults

70 @--e
Canes Means
60 Samnene .
° esoee Extremes
50

40

NUMBERS

30

20

10

20 ee:

NUMBERS
S
&
je)
=
a)

J Bact A M J J A S OPS N D

Figure 48. Means and extremes of semimonthly estimates of Bulwer's Petrel
numbers, Sand Island, Johnston Atoll, 1964-1969.

160

The number of adults handled each year probably represents 75 to 90
percent of the returning birds on average, although it may have been no
more than 50 percent in 1966. The upward trend in numbers handled each
year at least partly resulted from an increase in birds present and was
not entirely due to increased effort to catch them.

If one makes the reasonable assumption that Wetmore's 1923 visual
estimate of 400 Bulwer's Petrels was slightly low, this species has been
reduced by perhaps 90 percent since 1923. The natural rock crevices on
Johnston Island where Wetmore found these birds in 1923 have all been
destroyed. The birds using the atoll now depend completely upon the
crevices on Sand Island between rocks and chunks of concrete used to build
the causeway, and on space under slabs of concrete scattered around the
island.

As in the case of Christmas Shearwaters, it is difficult to see where
any Bulwer's Petrels would have been able to nest during the years of in-
tensive military use of the two islands, and it is likely that during
several years there was no successful nesting.

Under present conditions there probably are more available nest
sites than are being used, and barring further disturbance the population
should be able to increase to some extent.

Production of chicks was relatively high for this species and ap-
peared to be increasing from 1964 through 1969 (Table 28). Losses were
mainly of eggs rather than chicks, and most losses were attributed to
flooding by high tides. If the adult population was about 50 in 1968
and 1969, then the number of chicks produced was about 20 percent of this
number. Mortality rates for adults are unknown, but the apparent increase
in total numbers indicates that there must be a net gain from year to year
--more chicks are produced than adults lost. In 1968 the first known-age
chicks returned to the island--the single chick banded in 1964 and one
banded in 1965. Neither of these was recaptured in 1969, but three more
of the six 1965 birds returned in 1969. Thus of seven chicks banded in
1964 and 1965, five survived long enough to return to the island, one at
age three years and the others at four. None of these known-age birds
was known to breed, but two of the 1965 chicks that returned in 1969
developed bare brood patches and engaged in courtship calling. All re-
turned in late May or later, at least two months after established breeders
return, and all returned to approximately the same section of the island
where they had hatched.

Annual Cycle

Table 29 shows the available dates of major events in the annual cycle
of Bulwer's Petrels on Sand Island during POBSP studies. Because the birds
are secretive, returning to the island at dusk or after dark, and also re-
turning at the time Sooty Terns are most abundant and most demanding of
research time, observations were not as complete as for the more conspicu-
ous species. Exact date of laying was not determined for any eggs,
although dates of hatching and fledging were known for several.

67

Table 28. Productivity of Bulwer's Petrels on Sand Island, Johnston
Atoll, 1964-1969*

Chicks
Hatched Fledged

Percent Percent of

Year Eggs No. Percent No. of eggs hatching
1964 6 3 50 3 50 100
1965 12 9 15 6 50 67
1966 10 6 60 5 50 83
1967 iL} 14 93 10 67 7Ai
1968 13 ot 85 1 85 100
1969 5 10 67 10 67 100
Totals, 7. 53 67 45 64 85

*A11 egg and chick figures are minima, considered to be about 90 percent
complete [and the number of adults handled represents about 90 percent
of the total].

Bulwer's Petrels were completely absent from the atoll for about six
months, from late September to late March. The first birds probably re-
turned to the island each year in the last week of March (see Figs. 37
and 48), although in some years none was seen until nearly mid-April.

On arrival they come in at about dark and court in the nest sites through
the night, leaving sometime in the early morning. They are most easily
found at this time, for their courting calls can be easily heard. Many
nest cavities are too deep for an observer to reach the birds, or in

some cases even to see them, but at this season they can be called out
readily by imitating their courting calls.

First eggs were found usually by the end of May, but it is likely
that laying began by mid-May in most years. The latest known laying was
about 5 July 1969. This egg, and the latest laid in other years, did
not hatch, and probably was produced by young adults, laying for the first
time. Exact incubation periods were not determined, but two 1968 eggs
required at least 44 and 47 days. Earliest chicks appeared in late
June and none was known to hatch after mid-July. The data are insuf-
ficient to draw definite conclusions, but apparently the successful
nesters are closely synchronized. Egg laying may extend over six or
seven weeks, but the latest eggs never survive; thus all the chicks hatch
within about three weeks of each other.

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Fledging took place from the last week of August through mid-
September. The period from hatching to fledging, determined for eight
1968 and 1969 birds, ranged from 57 to 67 days (mean = 62 days).

After the eggs were laid only one adult was found at a time at the
nest, and after hatching the chicks were guarded only for the first few
days. Nearly all five-day-old birds measured in 1969 were unattended,
even at night. Apparently the deep, well-protected nest sites make
parental protection unnecessary and the adults stay only long enough to
feed their chicks, then immediately return to sea.

Pairs and single birds without eggs continued to return to the nest
sites and court through early August when other birds were incubating or
feeding chicks. Latest calling was heard 14 August in 1969. By the time
the chicks are ready to fledge all adults are very scarce; not only do
the non-breeders cease to come back but those with chicks abandon their
offspring for several days before they fledge. No adults have been
recorded after mid-September, and in most years none was found after the
end of August.

Molt probably occurs entirely while the birds are away from the
island. Only a few birds were examined carefully for molt, but all those
examined in early spring had completely new plumage and none examined
late in the breeding season had begun to molt.

Specimens

Appendix Table 7 lists the ten specimens of Bulwer's Petrel col-
lected from Johnston Atoll. Wetmore collected nine (skins: 5 male, 4
female) and POBSP personnel collected one (skin: 1 male) in 1964; all
are housed in the USNM. This is considered as a new specimen record,
as well as a new breeding record.

Banding and Movements

Tables 24 and 30 show banding and recapture efforts on Bulwer's
Petrels. These birds do not dig and consequently band wear was much less
than on shearwaters. Thirteen of the 22 adults banded in 1964 were re-
captured in 1969 and all bands were legible and in no immediate danger
of being lost. None was rebanded.

A few birds may have changed mates or taken new ones when the former
mate was lost, and a few birds changed nest sites, but generally pair
and site bonds appeared to be very strong. Members of at least two
pairs stayed together from 1964 through 1969, one pair definitely using
the same nest site each year and the other using the same or another
close by.

No interisland movements of Bulwer's Petrels to or from Johnston
Atoll are known.

164

At-Sea Distribution

Bulwer's Petrels were found in the grid in all months except
December, January, February, and June (Table 21 and Fig. 49). Since
the grid population is too high to be accounted for by Johnston birds,
and the peaks occur just before and after the breeding season, it is
unlikely that Johnston based birds make more than a minor contribution
to the grid population. The September peak in grid birds is certainly

concurrent with the major southward migration from the Hawaiian chain
(POBSP, 19672).

2.0

lo)

1.0

O85

BIRDS PER 100 SQUARE MILES

J F M OA M J J A S 0 N D

Figure 49. Diurnal occurrence of Bulwer's Petrels at sea 175 miles southwest
of Johnston Atoll, 1963-1967.

165

Table 30. Banding and recaptures of Bulwer's Petrels, Sand Island,
Johnston Atoll, 1964-1969

Adults handled

Nie 2 ae Young Total new

Year bandings Returns* Total banded bandings
1964 22 0 22 i 23
1965 15 iS: 30 6 21
1966 3 9 22 3) 6
1967 3, Zo 30 6 et
1968 ita 29 40 10 2a
1969 6 40 46 8 14
62 128 190 34 96

*Each bird counted only once each year recaptured.

WEDGE-TAILED SHEARWATER Puffinus pactftcus
Status

Abundant breeding species, present from March to early December.
Nests in shallow to deep burrows, mainly in dense Lepturus on the west
half of the original portion of Sand Island; a few nest on Johnston
Island. Ranks third behind Sooty Terns and Brown Noddies in number of young
produced each year (300 to 450), and fourth behind these two species and
Red-footed Boobies in number of adults using the island (3,000). After an
absence of about three months, adults return in March and immediately begin
courtship and burrow digging, mainly in dense Lepturus on the west half of
the original portion of Sand Island. Eggs are laid in late June and early
July and most chicks hatch during the last three weeks of August. Fledg-
ing begins in November and the last chicks and adults usually leave in
early December.

Ecological Distribution

Johnston Island: Brooke (ms.) described what most likely were
Wedge-tailed Shearwaters from Johnston Island on 15 March 1859, as follows.

The ground is undermined by petrels who live two together,
and in walking over the island I came across many pairs sit-
ting or lying cozily together at the entrance of their
retreats, basking in the Sun and screened from the wind by

166

the grass. They were quite noisy and many were billing
and cooing. The people onshore call them diggers from
the fact of their living in holes:

If these were Wedge-tailed Shearwaters--there are no records of
Bonin Petrels from Johnston Atoll--their breeding cycle then was not in
the same season as today.

Wetmore (ms. a and b) noted large numbers of Wedge-tailed Shearwaters
on Johnston Island in July 1923. The island was honey-combed with burrows.
Most were incubating in burrows up to four feet long; others were under
little mats of vegetation above ground. Various other visitors noted this
species on the island, including Thurston (1928) in April 1928, Clark
(1945 a and b) in May 1945, and Fennell (1948) in 1948. POBSP personnel
recorded a few wedge-tails each year during 1963-1969.

Johnston Island, where most of the original birds nested, probably
became less suited to them throughout the war years. From 1963 to 1969
a few wedge-tails were atrracted to the fairly dense stand of Vitex
trtfolta, growing in loose gravelly sand on a point between the main street
(Arthur Avenue) and the east taxiway. They dug shallow burrows there, or
simply sat on the ground under the bushes; a few eggs were laid some years
but no young were known to fledge. A few birds also used the crawl space
under at least one building and burrowed under concrete slabs on top of
an underground building.

Sand Island: Wetmore (ms. a and b) recorded this species on Sand in
July 1923. He, as did Thurston (1928) in April 1928, noted the soil on
the island was honey-combed with shearwater burrows. Moynihan (1957)
found at least a few on the original part of Sand sitting on the ground
in the open or just inside the mouths of their burrows during the daytime
in April 1957. POBSP personnel recorded this species as very numerous
on the original portion from 1963 through 1969. Amerson recorded them
with almost fledged young in November 1973.

Figures 35, 43 and 50 show the distribution of Wedge-tailed
Shearwaters on Sand Island. The preferred habitat from 1963 to 1973 was
deep sand, stabilized by dense, undisturbed Lepturus, which was prominent
only on the west half of the original portion (Figs. 51 and 52). Many
burrows were dug in more open sandy areas, but without grass roots to hold
the loose sand, heavy rains caused many of these burrows to cave in. They
also burrowed under sidewalks around the transmitter building and the
abandoned gun emplacement. The man-made end of Sand Island was too
gravelly and hard-packed for the birds to burrow, but Pluchea carolinensis
bushes growing southeast of the barracks were thick enough by 1969 to
attract at least one bird for long enough to lay an egg.

The necessary element in all these habitats was shelter from the sun.
Where this was obtained without digging, as in the gun emplacement or under
buildings, the birds made no attempt to dig, but simply laid their eggs on
the ground in a shallow cup formed of bits of grass, bones, and feathers.
Burrows were lined similarly.

167

Figure 50. Distribution of Wedge-tailed Shearwater nest sites on Sand Island,
Johnston Atoll, 1963-1969. Numbers indicate chicks banded in
1968.

Populations

The Wedge-tailed Shearwater population at Johnston Atoll consists
predominantly of light phase individuals. Light phase populations also
breed in the Hawaiian Islands and the northern Marshall Islands.

Figure 53 shows means and extremes of semimonthly estimates made
by POBSP personnel from 1964 through 1969 of Wedge-tailed Shearwater
numbers on Sand Island. These estimates were usually based on sample
counts of birds above ground, plus a liberal allowance for birds in
burrows and birds not returning to the island each night. Year to year
variations in these estimates were small and more likely resulted from
differences in estimating techniques than real variations in shearwater
numbers. Hence, there is no evidence that the number of Wedge-tailed
Shearwaters using the atoll varied during the years of POBSP studies.

On 30 September 1966 Shelton made a sample count on all parts of
Sand Island and found that 57 percent of 150 Wedge-tails had fresh orange
streamers put on some time that year. About 1,/20 birds had been streamered
in 1966, most of them in August and September. Although there was no way
to determine an exact figure, streamer losses were thought to be less than

168

Figure 51. Wedge-tailed Shearwater chick in front of burrow under Lepturus
repens clumps, Sand Island, Johnston Atoll, 2 September 1966
(POBSP photo by P. C. Shelton).

Ss ne sites

Ss

Figure 52. Wedge-tailed Shearwaters among Trtbulus, Boerhavia, and
Lepturus on southwest Sand Island, Johnston Atoll, mid-
morning July 1963 (POBSP photo by A.B. Amerson, Jr.).

NUMBERS

4,000

169

Adults
340007 *) 7 Means ry s
eee. Extremes POE» LET
2,000 al
° a eae 2
e 7: e oe "e--@
76 “. es oe
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# -
0 -
1,000
= nEggs
33 eeeee YOUNG
uu
ca 500
=
=>)
=
0
J F M A M J J A F 0 N D
Figure 53. Means and extremes of semimonthly estimates of Wedge-tailed
Shearwater numbers, Sand Island, Johnston Atoll, 1963-1969.

ten percent by 30 September. If there had been no streamer loss, then
the 1,720 birds would represent approximately 57 percent of the total
population using the island during that period:

ea. 3,000-

1,720/0.57 =
If ten percent of the streamers had been lost by 30 September, then the
total population would have been:

550/057 8="Ca. 2,700).

170

It is acknowledged that the assumptions underlying these calcula-
tions may not be fully met, but the data are presented as the best
available estimate of wedge-tail numbers on Sand Island during this
period. These figures are only about one-fourth larger than the mean
of the semi-monthly estimates, which suggests that these estimates are
a fairly good approximation of the numbers of wedge-tails using the
island during the periods involved.

The only comparative data from pre-POBSP sources are from Wetmore's
1923 observations in which he estimated 2,500 wedge-tails on Johnston
Island and 500 on Sand. Superficial comparison of these figures with
POBSP estimates and calculations would indicate that the population of
the atoll had not changed appreciably, but that the former population on
Johnston Island had moved to Sand. But Wetmore's figure is a visual
estimate, made over only a ten-day stay, in which Sand Island was not
visited at night when maximum populations would have occurred. Visual
estimates by the POBSP in mid-July averaged about 2,000 birds, or two-
thirds Wetmore's estimate. Wetmore thought that a quarter of the several
thousand birds he found killed by feather poachers were Wedge-tailed
Shearwaters. Thus, his estimate of 3,000 must have been at least a
thousand less than the original number.

Both Wetmore and Thurston stated that the entire surface of both
islands was burrowed by wedge-tails. At that time at least five times
as much dense Lepturus habitat was available as during the 1960's. If
the pre-war density of burrows (and birds) was as great as the density
during the 1960's, then the former population may have been several
times that of the present. No comparative density data exist, but it is
likely that many birds from Johnston have crowded on to Sand Island, re-
sulting in greater density than occurred on the undistrubed islands.

In summary, then, it would appear that the original population of
wedge-tails was at least 1.5 times that of the present, and may have
been two or three times the 3,000 now thought to use the atoll.

Whatever may have been the magnitude of the decrease in population
between Wetmore's visit and the 1960's, the cause is not difficult to
determine. The original portion of Sand Island--less than ten acres-—-
now has the only significant amount of natural habitat for this species.
There probably were about 50 acres available to them on the two islands
before military construction began. The ability of these birds to crowd
together to adapt to artificial habitats, such as abandoned bunkers and
crawl space under buildings, has not been great enough to allow them to
maintain their original numbers.

It is not likely that enough wedge-tails were killed, either by
feather poachers or military personnel, to lower permanently the popula-
tion. Reduced nesting success during early years of military occupancy
of the atoll probably gradually lowered the population to its present
level. Johnston Island, where most of the birds originally nested,
probably became less suited to them throughout the war years, but Clark's

(1945) and Fennell's (1948) notes indicate that more than the few dozen
largely unsuccessful nesters found there in the 1960's were using
Johnston until at least the late 1940's.

During POBSP studies the number of chicks produced each year
averaged about ten percent of the adult population (Table 31). Figures
in this table probably represent at least 90 percent of the chicks
fledged in each year. Egg numbers were less reliable because many bur-
rows were too deep to inspect. The two egg figures given are based on
counts made over the entire island, with the ratio of eggs to burrows
assumed to be the same for those that could not be inspected as for those
that were examined. These counts were made in July or August, after an
undetermined number of eggs had already been lost.

Table 31. Productivity of Wedge-tailed Shearwaters on Sand Island,
Johnston Atoll, 1963-1968

Percent of
eggs resulting in

Year Eggs laid Chicks banded* fledged chicks
1963 153

1964 304

1965 389

1966 600 150 25
1967 (300) **

1968 750 447 60
1969 Q**x%

*Number banded represents at least 90 percent of number fledged.
**Early September count. None banded.

***No counts were made in 1969. Numbers of eggs and chicks were estimated
to be approximately equal to 1967 numbers.

Egg and chick losses resulted from several factors. Each year
several dozen eggs were laid on top of the ground where there was no
protection for the incubating birds. Some of these eggs were incubated
for several weeks, but all were abandoned before they hatched. Eggs
sometimes were broken in the burrows, possibly as a result of disputes
over ownership of the burrows. Both these egg losses could be attributed
to crowded conditions. Wedge-tails normally burrow under dense Lepturus
and the burrows are fairly stable, even in heavy rains, because of the
holding power of the grass roots, but much of Sand Island is now well
vegetated and burrows in bare sand frequently cave in during heavy late
summer rains. In 1966 a dry spring and summer coupled with disturbance
from machinery in May when the LORAN-C transmitter tower was re-guyed
and painted left little healthy vegetation and wedge-tail chick losses

172

during August and September rains were greater that year than any other.
Dogs killed at least 35 chicks in 1963 and at least 11 in 1964. The
main culprit, a Coast Guard dog named "Jeffroe," was too aged to have
much impact after those years although he survived until late 1968.

Only a few chicks were killed by other dogs.

Adult mortality was very low and must not have been more than ten
percent per year both on and away from the island, since an annual chick
production of about this magnitude appeared to maintain the population.

Annual Cycle

Table 32 and Figure 37 shows inclusive dates for the major events
in the annual cycle of Wedge-tailed Shearwaters on Johnston Atoll. The
approximate numbers of adults, eggs, and chicks are shown on Figure 53.

Striking features of the wedge-tail's cycle are the uniformity from
year to year and the shortness of the laying period. Among Johnston
birds, this species holds the record on both counts. Wetmore's comment
that most birds were incubating in mid-July 1923 indicates that the breed-
ing cycle was the same then as now. In 1969, when egg laying was delayed
in several other species, wedge-tails arrived on schedule. The unusual
weather pattern thought to be responsible for the disruption of breeding
in the other species occurred in early to mid-February and ended about
three weeks before the wedge-tails returned. Other species did not re-
gain normal status until after wedge-tails returned but wedge-tails,
being the latest breeders on the atoll, apparently were not at a vulnerable
stage in their cycle in the few weeks prior to returning to the islands.

The synchrony of egg laying in this species is remarkable. Virtually
all the eggs appear to be laid within a month, and most in the last two
weeks of June. Accurate dates for last egg laying are not available for
most years, but in 1969 the southwest islet bunker was checked at least
every other day through June and early July and all the 16 eggs were laid
between 12 June and 1 July. Also, about three dozen eggs laid on top of
the ground where they could be seen readily were laid between these dates.
Close synchrony of laying in this species results partly from lack of re-
laying, which is also true of the other two breeding procellariiforms
(Christmas Shearwaters and Bulwer's Petrels). ‘Lack of chose synchrony
in these two species may be related to their occurring in much smaller
numbers than wedge-tails.

Wedge-tails began courtship and burrow digging almost immediately on
arriving back on the island each year. For the first few days after
they returned, they were present at night only, but soon small groups
stayed through at least part of the day. Night numbers were always at
least twice the daytime numbers. Numbers increased through late March and
April, reaching a maximum in May when courtship was most intense and
copulation occurred. The dip in early June (Fig. 49) resulted from
females staying at sea to feed during development of the egg. From this
time until after hatching began, observed numbers remained low. During
incubation adults relieved one another at intervals of about eight days

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(mean = 7.75, range = 4-12, n = 12, POBSP 1963 data). Few birds except
those incubating remained through the day, but large numbers returned
at night. Most of these appeared to be unoccupied birds--ones that did
not lay or that lost their eggs.

During the first week after hatching chicks were usually tended
by both parents at night but by only one during the day. Attendance
became less regular as the chick's food demaids increased and both
parents spent more time feeding at sea, but returned frequently, pos-
sibly every night, to feed the chick.

Maximum numbers appeared in late August through September. Daytime
numbers remained lower than during the spring courtship period, but at
dusk hundreds streamed in to the island, mostly from the southwest, and
remained through the night. Counts of 900 to 1,000 birds on the surface
were made in August and September 1966. A few hundred more were under
ground and not counted. Thus, each night approximately half the total
number of birds using the island during the year returned during this
period.

Those with chicks more than one or two weeks old apparently fed
their offspring early in the evening and spent the reaminder of the night
outside the burrows in large, loose roosting clubs, probably composed of
these and unoccupied birds. Low intensity courtship accompanied by out-
bursts of moaning as enthusiastic as those of the pre-laying courtship
period, continued through the early fall nights. By dawn, most birds had
returned to sea to feed.

Numbers declined in October, probably because birds without chicks
ceased returning. In November, adults with chicks began leaving and
numbers declined sharply. The last adults ceased returning in early
December. Chicks began leaving their burrows for short periods at night
by October, and began to fledge in November shortly after their parents
abandoned them. All were gone by mid-December.

Except for one bird found 23 February 1967, no Wedge-tailed
Shearwaters were known to have been on Johnston Atoll between mid-December
and mid-March--a period of four months. Presumably the birds spent this
time at sea, feeding and molting.

The beginning of the post-nuptial molt has not been observed, but
few birds were handled in late fall when it might have started. Adults
have nearly completed their molt by the time they return in the spring.
In five of 12 birds examined 13 April 1967 the distal primary was only
about three-fourths grown on each wing. Body feathers of the returning
birds have a frosted appearance which is soon lost by wear.

Specimens

Specimens of Wedge-tailed Shearwaters from Johnston Atoll are listed
in Appendix Table 7. Wetmore collected seven specimens (4 males, 3

Ab7()

females) in July 1923. POBSP personnel collected 93 specimens from
1963 through 1969. All specimens are housed in the National Museum of
Natural History.

This is a new published specimen record.

Banding and Interisland Movement

POBSP personnel and Amerson banded 6,517 individual Wedge-tailed
Shearwaters (4,985 adults and 1,532 young) at Sand Island during 1963-
1969, 1973 (Tables 24 and 33). Rapid wear and loss of aluminum bands
made analysis difficult because 961 birds had to be rebanded. Beginning
in 1968 most adults and all chicks were banded with Monel metal bands.
Orange, plastic, leg-streamers were also put on most adults handled
each year but these too were lost fairly rapidly; virtually none was
found on returning birds at the beginning of each new breeding season.

Of the 6,517 birds originally banded on the island, 3,635 were recaptured
back there indicating strong ties to their breeding ground. Six birds,
banded as young in 1968, were recaptured back on Sand Island in November
1973; these probably had returned to the island for the first time as
they appeared to be going through pre-breeding courtship behavior similar
to that observed in other Procellariformes. The Monel bands on these

six showed little wear.

Nine Wedge-tailed Shearwaters banded as adults at Johnston Atoll
were recaptured elsewhere: six at French Frigate Shoals and three at
sea. One banded 19 September 1973 was recaptured 8 November 1965 at
sea at 41°0'S and 174°20'E, which is west of Cook Strait, New Zealand.
In addition, eight wedge-tails banded as adults elsewhere were captured
at Johnston Atoll, six from French Frigate Shoals and two from Oahu.

At-Sea Distribution

Light phase Wedge-tailed Shearwaters--the phase most prevalent at
Johnston Atoll and in the Hawaiian Islands--occurred at sea in the grid
southwest of Johnston Atoll during all months but were at their highest
in September and lowest during January, February, and March (Table 21
and Fig. 54). Year-to-year variations in density and rhythm were fairly
small.

With the exception of an anomalously low grid density in June,
the annual population cycles on Johnston Atoll and in the grid were
similar from January through July (see Fig. 53). Major differences,
however, existed between these two areas from August through December.
In the grid, numbers continued to increase through September, decreased
in October, and increased again through November and December. These
were probably post-breeding and non-breeding birds concentrating prior
to, or during migration. By October most, if not all, these birds had
left the area. Increased numbers in the grid in November and December
probably represent young birds leaving the islands and perhaps remaining
at sea for a month or more, feeding and gathering strength for dispersal

176

to the wintering area. Most of the birds observed in the grid in
December and January were in apparent first year plumage.

No orange-streamered wedge-tails from Johnston were recorded within
the grid. At least 25 orange-streamered birds were sighted, however, in
the pelagic waters around Johnston Atoll. It is possible that a large
part of the grid population consists of non-breeding birds that belong to

the larger north-central Pacific population of which Johnston birds are
only one group (POBSP, 1967a; King, 1974).

Figure 54. Diurnal occurrence of Wedge-tailed Shearwaters (light phase)
at sea 175 miles southwest of Johnston Atoll, 1963-1967.

50

40

30

20

BIRDS PER 100 SQUARE MILES

10

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178

CHRISTMAS SHEARWATER : Pufftnus nattvitatts
Status

Uncommon breeding species, present from late January to late
September. Nests on Sand Island; previously nested on Johnston Island.
About 30 birds, half of which attempt to breed, returned each year
during POBSP studies; three to ten eggs were laid and one to six
chicks fledged each year. Adults returned to the island in January,
laid from late March to early May, and left the island in August or
September. Chicks usually hatched in May or June and fledged in August
or September. Most successful nests were near the shore under boards,
logs, and concrete slabs, or in an abandoned bunker. Attempts to nest in
shallow trenches under Lepturus or Boerhavia, in the manner of birds nesting
on undisturbed islands, usually were unsuccessful, probably because the
best sites of this type were occupied by the larger and more aggressive
Wedge-tailed Shearwaters.

Ecological Distribution

Johnston Island: Wetmore (ms. a) recorded about 400 Christmas
Shearwaters here in July 1923 and described the colony as follows.

There are two small colonies on the island, where these
birds congregate amid the wedge-tailed shearwaters and
have nests in holes dug to a shallow depth in the sandy
soil. During the day all are hidden underground but at
dark they creep out and remain active all night. Adults
are calling and caressing one another though all have
large young molting into first plumage. These squirm

and bite when handled and utter a curious call cO0O

(given with expiration of breath) oo-oo (made as breath
is. inhated). The call is; like that of the adult P.
cuneatus. At dark the young creep out of their holes and
remain in the open but seek shelter either in heavy grass
or underground when the sun is high.

Adults show some wear in plumage.

The note of the adults is a curious crowing call coo-coo-
oo-00-Koo Koo Kaw, the first part drawn out and the last
given rapidly. They are especially vociferous at day-
break in the morning and for an hour after.

Clark (1945b) again observed a few in May 1945. None has been
recorded since.

Sand Island: » Wetmore (ms. a) found about 100 adults with large
young on Sand Island in July 1923. POBSP personnel recorded this species
here in small numbers from 1963 to 1969. Kridler (BSFW, 1973) recorded
four nests with eggs in May 1973.

79

Nesting habitat is discussed in detail in the Population section.
In summary, they normally nest in shallow trenches under vegetation,
usually Lepturus, but competition from Wedge-tailed Shearwaters ap-
pears to have driven them to the periphery of the island, where they
use a variety of cavities under logs, boards, concrete slabs, and an
abandoned bunker (Figs. 55 and 56). Attempts to nest under vegetation
are less successful because the only vegetation available to this species
is too sparse to provide adequate protection. Figures 43 and 57 show
the distribution of the nesting sites used during POBSP studies.

Birds without nests were found in the same general area as the
mesters. The east slope, where most of the unsuccessful nests were
located, was a favored roosting place for birds without nests. A few
were found at other sites scattered over the island, including the
Tournefortta bush northeast of the transmitter, the unoccupied grass
along the northwest shore, and the southwest islet bunker.

Populations

The mean and ranges of POBSP semimonthly population estimates
during the years1964-1969 are shown in Figure 58. Year-to-year
variations appear to be more likely due to variations in estimating
techniques than to variations in actual numbers.

Nearly all Christmas Shearwaters returning to the island each year
were handled at least once. Thus the number of new bandings, plus the
number of previously banded birds caught, is a good estimate of the total
number of birds using the island each year. This varied from 18 to 28
during the years 1963-1969 (see Tables 24 and 36). In all years ex-
cept 1963, when observations by the POBSP did not begin until July, and
1966, when incomplete records were obtained, these figures probably
represent at least 90 percent of the returning birds for the respective
years. Loss of bands may have inflated the number of new bandings,
which would compensate for birds missed. The total number of Christmas
Shearwaters returning each year was 25 to 30. The mean of semimonthly
estimates during the period of maximum abundance was about three-fourths
this number, which probably is a good estimate of the number using the
island during each semimonthly period.

Wetmore's estimate of 300 birds on Johnston and 100 on Sand in
mid-July 1923 is over 20 times the number obtained from mid-July visual
estimates during the 1960's. Loss of nesting habitat and crowding by
Wedge-tailed Shearwaters appear to be the major causes of the decrease.
Both shearwater species originally used similar habitat--sandy soil
vegetated with Lepturus. Because both species are now crowded onto
Sand Island and wedge-tails almost totally dominate available Lepturus
stands, the smaller Christmas Shearwaters have declined to a much smaller
fraction of their original numbers than have wedge-tails. During most
intensive military use of Sand Island, there probably were years in
which Christmas Shearwaters were unable to reproduce at all and the
population may have dropped even lower than during the 1960's.

180

ot

Figure 55. Christmas Shearwater chick, 2 and 1/2 months old, in shallow
trench burrow under Lepturus repens clump on east shore of
Sand Island, Johnston Atoll, 29 August 1966 (POBSP photo by
P. C. Shelton).

Figure 56. Christmas Shearwater chick, 2 months old, and adults in bunker
on east shore, Sand Island, Johnston Atoll, 1 September 1966
(POBSP photo by P. C. Shelton).

181

@ Lepturus
@ Log or Board

© Concrete Slab
o East Bunker

4

Figure 57. Distribution of Christmas Shearwater nest sites, Sand Island,
Johnston Atoll, 1963-1969.

Fortunately, Christmas Shearwaters will nest closer to water than
will wedge-tails. Cavities under concrete slabs, boards, logs, and an
abandoned bunker, all close to the shoreline of Sand Island, were the
s.tes of most successful breeding attempts during the 1960's (Table 34).
Ccmpetition from Wedge-tailed Shearwaters restricted the Christmas
Shearwaters that attempted to nest under vegetation to marginal sites
under sparse cover where nests were exposed to rain, washing sand, and
direct sunlight, as a result of which these nests were frequently
abandoned.

Nesting success (Table 34) was low for all years except 1967 and
1968 when six and five young fledged. Low egg production in 1969
probably resulted from the weather pattern disturbance in February.
Although the birds were only beginning courtship at the time, physio-
logical changes must have been induced that prevented egg production in
several birds.

Nearly all losses were of eggs rather than chicks, for only two
chicks that hatched failed to fledge, both in 1965. One of these was
killed by a dog just before it would have fledged; the other died much
younger but the cause of its death was not recorded. Egg losses
amounted to nearly half those laid; in a few cases the cause of loss
was recorded. The two lost in 1964 were inundated by high tides; two

30

25 Adults

—_—_ Means

eoooe Fxtremes

No UU MmBe Ee Rees

5

h Eggs
”n Yn
ce 3 Fa B20 eG Oey eoeee YOUNG
a ? ‘ e ‘e
= O i
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J F M A M J J A S 0 N D
Figure 58,

Means and extremes of semimonthly estimates of Christmas

Shearwater numbers, Sand Island, Johnston Atoll, 1964-1969.

lost in 1968 were broken by Wedge-tailed Shearwaters; one 1968 egg was
infertile; and one 1969 egg was abandoned when sand washed into the
shallow nest cavity dug under loose Boerhavta.

Barring further disturbance of Sand Island, or further increase in
wedge-tail numbers, Christmas Shearwaters should at least be able to
hold their own, and might be able to increase slightly. On the north-
west corner of Sand Island is a thin fringe of dense Lepturus that is
not completely overrun by wedge-tails and this conceivably could be
used by Christmas Shearwaters.

Table 34. Productivity of Christmas Shearwaters on Sand Island,
Johnston Atoll, 1963-1969

Number of Chicks Percent Percent
Number of Hatched Fledged of of

Year Eggs Laid No. for aN Eggs Laid Chicks Hatched
1963 (2)*
1964 4 2% 50 2 50 100
1965 7 By 1143 1 14 53
1966 6 Zio ere 2 33 100
1967 6 6 100 6 100 100
1968 10 Say 729.0 5 50 100
1969 3 2) bi 2 67 100

36 20% 56 18 50 90
All years:

Cavities**
20 Muh? Os 13 65 93

Natural Vegetation***

16 6 38 5 él 84
*Not included in totals.
**kUnder concrete slabs, logs, boards, or in abandoned bunker.

*kkTepturus, Boerhavia, or Tribulus.

184

Annual Cycle

From 1963 to 1969, after an absence of nearly four months, adults
usually returned to the island in late January (Table 35 and Figs. 37
and 58). The first arrivals were usually seen soaring along the east
shore at dusk; by dark they settled near the nest sites where they sat
quietly most of the night, leaving by dawn. Numbers increased fairly
rapidly during February as more birds returned each night and court-
ship began. Maximum numbers occurred in March when courtship was most
intense. A few were sometimes present during the day at this time.
Apparent numbers then declined slightly as those birds that would produce
eggs spent more time feeding at sea. In 1969, pairs were observed
copulating on 11 and 15 March. Eggs were laid by these birds 3 and 22
April, respectively. In both cases one bird returned to the nest less
than half the nights during the two weeks before the eggs were laid.
Other nights neither returned.

First eggs were laid in late March every year except 1969, when they
were delayed until April. (In 1966 the first egg was discovered in
April, but it had been laid sometime before that.) The delay in 1969
probably resulted from the disruption of the normal weather pattern in
February; no Christmas Shearwaters appeared from 5 through 9 February,
after at least three had returned regularly for several days in late
January and early February.

Laying extended over a period of nearly seven weeks, slightly
longer than for Wedge-tailed Shearwaters. If close synchrony in laying
depended to any great extent upon social facilitation, then it would be
expected that Christmas Shearwaters would be less synchronous than
wedge-tails. The small numbers of Christmas Shearwaters nesting in
scattered sites around the periphery of the island would have much less
opportunity for social contact between pairs than would be the case with
their more numerous relatives.

Exact dates of egg laying and hatching were obtained for only a
few eggs; incubation averages about 52 days (range 51-54, n = 5) and
fledging periods average about 96 days (range 88-103, n = 6).

One adult stayed with the chick during the day for the first week
or so after hatching, and both parents usually returned at night for
the first four to six weeks, after which only one usually returned,
Chicks began to venture out of their nests at night when they are two-
thirds grown, but they were quick to seek refuge if distrubed. Adults
abandoned the chicks a few days before they fledged.

During the period of incubation and chick feeding, adults without
eggs or chicks continued to return to the island at night. They arrived
at dusk, soared along the east shore until about dark, then spent the
night sitting in pairs or sometimes in groups of four or five near the
beach on the east slope. Those that had lost eggs or had nested in
former years usually returned to the site of their former nest or near
the nesting birds. Low intensity courtship continued as long as these

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birds were present. At dawn they again soared over the east shore,
occasionally calling as they flew, before heading out to sea for the
day's feeding. These birds ceased returning to the island about the
time the first chicks fledged.

Few birds were examined late in the breeding season, but two caught
tending chicks in late August 1968 had started to replace their pri-
maries. Thus the molt apparently begins shortly before the chicks fledge.
Birds returning in January had completely new plumage.

Specimens

Eleven specimens are housed in the USNM (Appendix Table 7). This
is considered to be a new published specimen record.

Banding and Interisland Movement

POBSP personnel banded 83 Christmas Shearwaters on Sand Island
during 1963-1969 (Tables 24 and 36). Rapid band wear complicated analysis
of the banding data. The aluminum bands used during the first five years
of the study seldom lasted more than two years, so the birds had to be
rebanded frequently. By 1969 several birds were wearing their fourth
bands. Beginning in summer 1968 Monel metal bands were used to reband
adults and to band all chicks. Recaptures of 1968 banded birds in 1969
showed little or no wear of the Monel metal bands.

Unfortunately, an undetermined, but hopefully small, number of
birds lost bands without being rebanded, resulting in difficulties in
tracing pair and site bonds and in inflation of the figure for total
number of birds handled. The fragmentary data indicate that both pair
and site bonds were strong--breeding birds almost always returned to the
same site with the same mate year after year.

No birds banded as chicks were recaptured. Since these birds should
not lose their bands until they return to the island and begin digging,
it is unlikely that they returned and were not discovered. Each year a
few unbanded birds appeared late in the breeding season and it is sus-
pected that these were young birds, hatched before POBSP studies began,
who were returning to the island for the first time.

No interisland movements of Christmas Shearwaters to or from
Johnston Atoll were recorded.

At-sea Distribution

Christmas Shearwaters occur casually in the grid. They were ob-
served there in every month except January and May (POBSP, 1967a).

Table 36. Banding and recaptures of Christmas Shearwaters, Sand
Island, Johnston Atoll, 1963-1969%*

Adults Young
New Returns** Total Total, new

Year bandings (rebanded) Handled Bandings bandings
1963 20 20 2 De,
1964 12 14(1) 26 2 14
1965 6 22 (13) 28 1 if
1966 6 13'\(2) 19 2 8
1967 3 17 (4) 20 6 9
1968 9 14(9) 23 5 14
1969 7 19(13) 26 2 9

63 99 (42) 162 20 83

*Includes three adults banded by N.P. Ashmole in February 1963; all
other banding was by POBSP.

**Each bird counted only once each year.

NEWELL'S SHEARWATER Puffinus puffinus newellt
Status

Irregular summer visitor; two specimen records and one sighting
on Sand Island.

Observations

The POBSP obtained three records of Newell's race of the Manx
Shearwater from Sand Island. Amerson collected the first, an adult
female, on 21 August 1963 from a group of 100 to 200 Wedge-tailed Shear-
waters on the southeast slope. Harrington collected the second, an
adult female, as it rested among Sooty Terns and Wedge-tailed Shearwaters
in the southwest grass on 13 June 1967. The next night he saw another
beside the transmitter building but did not collect it. All three were
found at night.

Newell's Shearwaters are common at sea in the north-central Pacific
from March to November (King and Gould, 1967), and a nesting ground was
discovered on Kauai in 1967 (Sincock and Swedberg, 1969).

188

Specimens

The two specimens noted above and in Appendix Table 7 constitute
a new specimen record for Johnston Atoll.

SOOTY STORM PETREL Oceanodroma tristramt
Status

Accidental; one specimen from Sand Island.
Observations

A Sooty Storm Petrel flew into the U.S. Air Force camera building
on Sand Island shortly after midnight 21 December 1968, a night of high
winds and intermittent rain. Brownell collected it. The bird was a
female with a recently ruptured ovarian follicle, recently used oviduct,
and a slightly vascularized bare brood patch, all indicating that it had
laid shortly before arriving on Sand.

Sooty Storm Petrels breed from December through May throughout the
northwestern Hawaiian Islands (Richardson, 1957; Clapp and Woodward,
1968; Amerson, Clapp, and Wirtz, in press). They normally occur at sea
in the vicinity of their breeding grounds and probably disperse north-
ward (King, 1967). Small numbers have been recorded only during February,
March, and April in the grid southwest of Johnston Atoll (POBSP, 1967a).

Specimens
The specimen noted above and in Appendix Table 7 is a new specimen

record for Johnston Atoll.

RED-BILLED TROPICBIRD Phaethon athereus mesonauta

Status

Rare visitor; three sight records from Johnston Island.

Observations

Three Red-billed Tropicbirds have been observed on Johnston Atoll,
two of which were caught and released. Some sightings attributed to
White-tailed Tropicbirds may have been Red-bills for the two are dif-
dicult to distinguish if seen only from below.

Moynihan (1957: 36) made the following comments on his survey of
67 tO LOAD EI E9570:

One individual of this species was present during every
day of my stay. It flew back and forth in company with
the Red-tailed Tropicbirds; but did not participate in

189

their displays. It did, however, come down to land with
one of the Red-tails several times. A second Red-billed
Tropicbird appeared on April 9. It also flew back and
forth with the Red-tails; but the two Red-bills appeared
to ignore one another.

This record seems to be a considerable extension of the |
known range of P. athereus (Joseph E. King, personal com- |
munication); but it is quite definite. I did not collect
the two birds; but I got excellent views of both, even
picking one of them up in my hands, and heard their shrill
screaming calls (quite different from the rather hoarse
croaking of the Red-tails).

On 11 April 1969 Shelton caught, banded, photographed, measured,
and released an adult Red-billed Tropicbird on Johnston Island; he found
it sitting under a dense Pluchea clump with nesting Red-tailed Tropic-
birds. On 21 May Sgt. Browning, USAF, saw this same bird (the only
tropicbird with a white tail and orange plastic leg streamer known to
exist) flying over Johnston. On 7 June, Huber heard the bird calling
and found it resting beside a Red-tailed Tropicbird chick under Vitex
bushes about 200 yards from the site of its banding. On six of ten
visits to this site from 7 through 28 June the Red-billed Tropicbird
was found resting a few inches to a few feet from the red-tail chick.

It was not found after 28 June. Only once was one of the parents of

the chick present, and at this time all three birds rested quietly within
a few feet of one another until the adult red-tail became restless, dis-
turbing the red-bill, which then left the area. No time was available
for detailed observations of this bird and its interactions with the
red-tails. Most observations were during the middle of the day, but
twice it was present in the evening and probably spent the night on the
island.

Measurements of this bird ( Bill: 62 mm; Wing: 303 mm; and Tarsus:
31 mm) best fit those of the eastern Pacific race, Phaethon athereus
mesonauta, as described by Murphy (1936). This race breeds in the
Caribbean, Atlantic, and in the eastern Pacific from the Gulf of Cali-
fornia to Venezuela and on the Galapagos; it ranges from Southern
California to Chile (AOU, 1957).

Another possible sighting of a Red-billed Tropicbird occurred 11
May 1969 when Sgt. Lewis, USAF, saw two tropicbirds with white tails,
both without orange streamers, one nearly twice the size of the other,
over Johnston Island. He thought the larger of these two birds may have
landed on the roof of a building.

Specimens
No specimens exist for Johnston Atoll.
There are only two specimen records of the species from the central

Pacific, one from Nihoa and one from French Frigate Shoals (Clapp and
Woodward, 1968; Amerson, 1971).

190

RED-TAILED TROPICBIRD Phaethon rubricauda
Status

Common breeding species, present throughout the year. Nests on
the ground usually under Secaevola, Pluchea, and Tournefortia bushes
on both Johnston and Sand Islands. Red-tailed Tropicbirds were rela-
tively scarce in 1923, but are now common breeders on both Sand and
Johnston Islands. Thus they are among the few bird species that have
benefitted from man's activities on Johnston Atoll. During POBSP
studies numbers using the atoll each year increased from about 150 to
about 200. Chick production increased more dramatically-—-from about 15
in 1964 to about 75 in 1969--as bushes growing on Sand Island provided
a steadily increasing number of nest sites. Eggs have been laid in
every month except November, but maximum populations and breeding occur
during the spring.

Ecological Distribution

Johnston Island: Brooke (ms.) on 15 March 1859 while at Johnston
Atoll wrote: "Kern and Thorburn returned at Sunset, bring off...a
phaeton [sic.] with an egg...." The following day he noted: "One of
the men onshore gave me a bunch of tropic bird feathers, red and very
pretty.'' These observations no doubt refer to Red-tailed Tropicbirds.

Wetmore (ms. b) in July 1923 wrote: "A few pairs were found about
the rock ledges of the shoreline. An egg ready to lay was taken from a
female that I collected. Also secured one young bird not quite able to
fly. Its notes resembled those of the adult....[There were] 8 on
Johnston Island.'' Clark (1945a) observed a pair flying in May 1945,
and Davis (1962) mentioned their presence in 1959-1960.

POBSP personnel found them nesting here from 1963 through 1969.
Kridler (BSFW, 1973) and Amerson found them nesting in 1973. Nests were
common under artificial edifices, but Seaevola, Pluchea, and Tournefortta
bushes were also commonly used. Normal human activity disturbed them
little; most nests were within a few feet of frequently traveled roads
and walkways.

Sand Island: Wetmore (ms. b) reported four here in July 1923,
and Moynihan (1957) observed it nesting under Seaevola in April 1957.
POBSP personnel observed Red-tailed Tropicbirds nesting here during
1963-1969. Kridler (BSFW, 1973) and Amerson also found them nesting
Amt UOT Si

Figures 59, 60 and 61 show the locations of active nests of Red-
tailed Tropicbirds on.Sand Island in ‘196/ and 1969, Most of the inerease
from 1967 to 1969 was under bushes on the man-made portion of the island
and on the northwest corner of the original portion where most new bush
growth had occurred (Figs. 62 and 63).

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Figure 60. Distribution of Red-tailed Tropicbird nests on the original
portion of Sand Island, Johnston Atoll, 1969.

Fuel Tanks

Camera
Site

Seaplane
Ramp

Causeway

Figure 61. Distribution of Red-tailed Tropicbird nests on the man-made
portion of Sand Island, Johnston Atoll, 1969.

193

Figure 62. Red-tailed Tropicbird on nest under fournefortia bush northeast
of transmitter building, Sand Island, Johnston Atoll, 25
February 1967 (POBSP photo by P. C. Shelton).

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Figure 63. Looking east from top of power building, Sand Island, Johnston
Atoll, 5 September 1969. Several Red-tailed Tropicbirds nested
under these Pluchea odorata bushes (POBSP photo by P. C. Shelton).

194

Table 37 lists the distribution by plant species or other cover
under which eggs were laid in 1967 to 1969. Seaevola taccada, Pluchea
odorata, and Tournefortta argentea account for about 75 percent of the
nest cover on Sand Island. On Johnston where there is a larger variety
of habitats available, these three species still make up the bulk of
the sites.

Shelter from the sun appears to be the most necessary feature for
a tropicbird nest. Seaevola is denser than the other plants which may
account for the higher rate of success in nests under it. JLepturus and
Amaranthus provide the least cover among the plants used, because they
are not only smaller and less dense than the woody species, but they are
more ephemeral. An egg laid under Amaranthus in December 1969 was well-
shaded at the time of laying, but by the time the chick was half grown
the plant had dried away to nearly nothing, leaving the chick almost
completely exposed. Had this happened earlier in the chick's development,
it might not have survived.

Nests under artificial edifices were more common on Johnston than
on Sand because there were more edifices and fewer bushes on Johnston.
Normal human activity disturbed them little; most of the nests on both
islands were within a few feet of frequently traveled roads and walkways.
Birds returning to nests frequently startled workers by dropping
Silently out of the sky and shuffling awkwardly under a nearby bush
within arm's reach of the onlooker.

No nesting material was used by Red-tailed Tropicbirds, but they
usually formed a shallow depression for the egg by scraping away loose
leaves or sand.

Populations

Figures 64 and 65 show POBSP semimonthly estimates of Red-tailed
Tropicbirds on Sand and Johnston Islands.

The total number of adults handled each year (new bandings plus
recaptures of previously banded birds) was about 130 in 1964 through
1966, but increased steadily thereafter to 174 in 1969. These figures
probably represent 75 to 90 percent of the number returning to the islands
each year. If this is true, then the total number returning was about
150 in 1964 through 1966, but increased to at least 200 by 1969.

Maximum semimonthly estimates usually ran about half this calculated
annual total, and probably fairly accurately represented the actual popu-
lation.

Wetmore's estimate of only 12 birds on the atoll in mid-July 1923
is only about a tenth the numbers estimated in mid-July during the late
1960's. Wetmore’ found only one chick and one female about to lay. In
mid-July 1969 there were 11 eggs and 49 unfledged young on the atoll.
Wetmore's figures may have been low if a few undiscovered tropicbirds
nested in dense Lepturus as they did in the 1960's, but it is unlikely

95

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196

100

15

© e Adults

NUMBERS

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NUMBERS
a

M Ohi ag A S Ove N D

Figure 64. Means of semimonthly estimates of Red-tailed Tropicbird numbers
Sand Island, Johnston Atoll; 1964-1966 (solid line) compared
with 1967-1969 (dots).

100
75
7)
ac
Lu
ca
=
=
=
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Lu
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Figure 65.

197

Adults

Jat abllnersA”\ wiMGagdh mil aor gneSud oer Not op

Means of semimonthly estimates of Red-tailed Tropicbird numbers,
Johnston Island, Johnston Atoll; 1964-1966 (solid line) compared
with 1967-1969 (dots).

198

that several biologists roaming the tiny islets for ten days could have
missed more than a few tropicbirds. The rock ledges in which the birds
nested in 1923 must have been marginal nesting habitat, sufficient for
only a few pairs.

In early April 1957 Moynihan estimated 12 birds and found three
eggs on Sand Island. He saw no tropicbirds on Johnston Island. By
contrast, at the same time in 1969 there were 23 eggs, and an adult
population estimated at 75 on Sand Island, and nine eggs and 25 birds
on Johnston Island. This suggests that most of the increase in tropic-—
birds using Johnston Atoll has occurred in very recent years. Continued
disturbance during the war years and for some time thereafter must have
made it difficult for these birds to nest.

Table 38 shows productivity of eggs and young during POBSP studies.
The 500 percent increase in fledged young--from about 15 in 1964 to
about 75 in 1969--is considerably greater than the increase in adult
birds returning to the island. This may indicate an excess of potential
breeders which are prevented from breeding for lack of suitable habitat.
As of 1969 there was no sign of this upward trend leveling off.

The survival rate of eggs and chicks is relatively high for a sea-
bird. Productivity, figured as a function of the number of pairs nesting,
is further increased by renesting. About a third of the pairs that lost
eggs early in the season relaid a month or so later. In 1967 one pair
laid three times before finally successfully rearing a chick. In 1968
at least ten pairs renested, resulting in four chicks, and in 1969 at
least 11 pairs renested, rearing eight chicks.

Most nest losses occurred in the first week after laying and often
appeared to result simply from abandonment of the eggs by the adults. In
several cases only one adult was ever found at the nest. Presumably this
was the female who deserted the egg after several days of vainly waiting
to be relieved by her mate. A few presumably infertile eggs did not
hatch even though well attended.

On Johnston Island, human disturbance, much of it unnecessary
vandalism, accounted for a fairly large percentage of egg, chick, and
even adult loss. On Sand Island a few chicks may have been killed by
the always present dogs or cats, but these animals generally left tropic-
birds alone. A 60-day old chick died after being drenched with insecticide
in 1968. Wedge-tailed Shearwaters disrupted a few nests. Ants probably
drove the adults away from at least one nest in 1967; and one egg laid
under a concrete slab near shore was inundated by a high tide in 1967.

A few eggs on Sand were lost because of disturbance by other tropic-—
birds, probably in disputes over nest sites. In 1967 two eggs were so
destroyed. In 1769 an adult was observed vigorously, and successfully,
defending its nest and egg from an intruder. Also in 1969, an adult
killed another in what appeared to be a struggle for a nest site, although
neither bird had an egg or chick.

99

Table 38. Productivity of Red-tailed Tropicbirds on Johnston Atoll,
1964-1969

Pairs
known Eggs Chicks fledged

Year to nest Laid Hatch Percent Number % of eggs % of hatch % of pairs

Sand Island:

1964 ds 14 4 29 is 14 50 ?
1965 a 19 14 74 12 63 86 2
1966 2 22 14 64 13 59 93 2
1967 36 42 26 62 22 52 85 61
1968 55 61 38 62 36 59 95 66
1969** 69 75 (53) C71) (49) (65) (92) (71)
Mean 53 39 25 64 22 57 90 66

Johnston Island:

1964 2? 36 ? ? 172 Sl ? %
1965 ty 30 ? uy 19 63 ty ?
1966 uy 705; ? 1 15 60 ? ?
1967 2 38 ? ? iat 29 ts te
1968 2a 725) 19 76 19 76 100 91
1969** 31 36 (26) (73) ((728))) (70) (96) (81)
Mean 26 32 23 75 17 53 98 86
Totals for both islands:

1964 ts 50 ? ? 14 28 2? 2?
1965 ? 49 B ? Shik 63 is ?
1966 2 47 2 2? 28 60 ts uy
1967 ? 80 z te 343) 42 ? ?
1968 76 86 57 66 by) 64 97 YL
1969** 100 iL bil (79) (W/4b)) (74) (67) (94) 74
Mean 88 7/il 68 69 39 56 96 7/3

*Figures are about 90 percent complete, except for 1967, 1968, and 1969
figures for Sand Island which are at least 98 percent complete.

*xParenthetical figures for 1969 are projected totals, assuming that the 29
large chicks (17 on Sand, 12 on Johnston) remaining 9 September fledged,
and that the three eggs (two on Sand, one on Johnston) remaining 9 Sep-
tember hatched and the chicks fledged. Any losses that might have
occurred in these eggs or chicks probably were offset by additional
laying after 9 September.

200

This competition indicates that there may be a shortage of nest
sites on Sand Island. Further evidence for such competition is the
increase in nesting tropicbirds as the abundance of introduced bushes,
mainly Seaevola taecada, Pluchea odorata, and Tournefortia argentea,
increased. During the late 1960's there was a particularly rapid
proliferation of Pluchea odorata on both the original and man-made
portions of Sand Island. On Johnston Island, vegetation is seldom left
undisturbed long enough for dense thickets to develop. Lower limbs of
bushes frequently are trimmed, leaving little cover for tropicbirds.
Pluchea odorata bushes growing along the south side of the runway on
Johnston were removed in 1967 at the suggestion of POBSP personnel to
prevent a possible hazard to aircraft if tropicbirds were attracted
to these bushes.

The tropicbird population on Johnston Island remained stable during
POBSP studies, but the increased numbers on Sand consisted partly of
birds that had formerly used Johnston. In 1968 and 1969, 38 percent of
the banded birds caught on Sand had been banded on Johnston. Only 11
percent of those caught on Johnston had been banded on Sand.

Annual Cycle

Although a few Red-tailed Tropicbirds may be found on the atoll at
any time of year (Figs. 37, 64, and 65), the largest number appears
during spring and early summer. Eggs were laid in every month except
November, but 90 percent of the annual total was laid in the six months
February through July.

In late December or early January there is an increase in the number
of displaying birds over the islands as returning pre-breeders begin
courtship. Maximum numbers are reached at midday; all leave well before
dark, usually by 1500 hours. Shortly after returning to the island, pairs
begin to spend time under bushes during the day. They stay over night
only after they have eggs except for an occasional broody bird that in-
cubates a rock for a few days. These may be males whose mates are at
sea feeding while the egg develops.

Laying increases sharply beginning in February and reaches a maximum
in late March, after which there is a steady decline through July. In-
cubation requires about 43 days (Table 39), during which time the adults
relieve each other at intervals of about a week. When the chick hatches,
the adults change off at least once every two or three days. After a
week or ten days the chick is left alone most of the time but is fed
regularly by both parents.

Young remain close to nest sites until they are nearly ready to fly.
When they are fully feathered and over two months old they emerge from
the bushes at dawn to exercise for an hour or so, return to sit quietly
for the remainder of the day and night, and emerge again only the next
morning. When chicks are about 75 days old adults cease to feed them.
Fledging occurs at 80 to 90 days (Table 39). With one or two exceptions,
chicks never returned to their nest sites after their first sustained flight.

201

Table 39. Incubation and fledging periods of Red-tailed Tropicbird
eggs and chicks, Sand Island, Johnston Atoll, 1967-1969

Mean Range Number
Incubation period 42.7 39-45 63
Fledging period 83.7 78-92% 65

*Not including one of 96 days, raised by one parent, and one of 103
days, probably raised by one parent.

Throughout the breeding season displaying birds were common over
the islands during midday. Daily counts ranged from five to at least
15 on clear days during March and April, and still occasionally reached
ten to 12 in July and August. Tradewind rain showers had no obvious
effect on displaying tropicbirds, but on rare days of heavy overcast
and steady rain none appeared.

In 1967 and 1969 virtually all nesting birds were fitted with orange
plastic leg streamers as soon as they were found on eggs. Displaying
birds seldom had streamers, indicating that birds with chicks or eggs did
not usually take part in the display flights. The few displaying birds
with streamers could have been accounted for by birds whose eggs had been
lost. On the few occasions when incubating birds were observed relieving
each other, the arriving bird flew directly to the nest and took over the
egg after a noisy ceremony lasting ten to 15 minutes. The departing bird
usually made a circle over the island, gaining altitude, then flew
directly out to sea.

Unbanded adults continued to appear in low numbers in spring and
higher numbers in summer throughout the years of POBSP studies. Many
caught in spring were later found nesting, while most caught in summer
failed to nest despite frequent observations of display in the air and
under bushes. Many, however, returned to nest in subsequent years,
nearly always in the later part of the breeding season. These observa-
tions suggest that many of these unbanded birds were young adults, returning
for the first time a year or so before breeding. Perhaps they had hatched
before POBSP studies began.

Nearly ail chicks fledged from 1963 through 1969 were banded, and
ten of these were subsequently recaptured:

Age Number Returning Remarks

2 years 2 3 No breeding attempted; one struck guywire.

3 years 4 One bred successfully; one struck guywire.

4 years 4 Two bred successfully; another attempted but

failed (see next entry).

202

Age Number Returning Remarks

5 years 1 Bred successfully after an unsuccessful
attempt as a 4 year old.

Three of these birds returned to the islands on which they had
hatched--one to Johnston, two to Sand; however, five which hatched on
Johnston returned to Sand, while only two which hatched on Sand returned
to Johnston. The two birds that struck guywires probably were return-
ing to the islands for the first time and were unfamiliar with these
obstacles.

After they abandon their chicks, adults apparently do not revisit
the islands for about six months until they return to begin a new breeding
cycle. Molt probably occurs during this time.

Specimens

Appendix Table 7 presents data on the 29 Red-tailed Tropicbirds
collected from Johnston Atoll. This is a new specimen record.

Banding and Interisland Movement

From 1963 through 1969 POBSP personnel banded 622 Red-tailed
Tropicbirds at Johnston and Sand Islands; in 1973 Amerson banded three
(Tables 29 and 40). Of these, 500 have been recaptured back on the atoll.
None banded on the atoll has been recaptured on other islands; however,
an orange-streamered adult from Johnston was seen, but not caught, at
Kure Atoll. Four birds banded on other atolls--two from Midway and one
each from Wake and Pearl and Hermes Reef--were captured on Johnston Atoll.
All of these islands are west or northwest of Johnston, suggesting that
east to west, or northwest to southwest (Wake, northwestern Hawaiians to
Johnston) movements may be more frequent for this species than north to
south (Johnston to Line, Phoenix) movements.

During the seven years of POBSP studies band wear was not a serious
problem, but a few birds banded in 1963 and 1964 needed new bands by 1969.
None was known to be lost during this time but some probably were. Great-
est wear probably occurs when the birds return to their nest sites and
scrape out shallow nest depressions with their feet.

At-Sea Distribution

The at-sea grid density cycle for Red-tailed Tropicbirds (Fig. 66
and Table 21) closely resembles the population cycle on Johnston Atoll
(see Figs. 64 and 65) and dt other breeding localities in the north-
western Hawaiian Islands.

Breeding birds apparently remain close to the nesting area during
the breeding season and may then disperse at sea for the remaining months.
The lack of orange-streamered sightings in the grid supports this, but may
also indicate that breeding birds never use the grid area. The grid

203

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204

population might then be composed of pre- or non-breeders which visit
the Atoll at irregular intervals and then only for short periods.

Year to year fluctuations in density appear to be very small and
random (POBSP, 1967a; Gould, King and Sanger, 1974).

3.0

BIRDS PER 100 SQUARE MILES
on

Je iE CO aM ea ene SOON Ni aaa)

Figure 66. Diurnal occurrence of Red-tailed Tropicbirds at sea 175 miles
southwest of Johnston Atoll, 1963-1967.

205

WHITE-TAILED TROPICBIRD Phaethon lepturus
Status

Regular visitor; most common at midday during winter, spring, and
summer on Akau, Johnston, and Sand Islands.

Observations

Moynihan (1957) saw a White-tailed Tropicbird fly across Sand
Island near midday on 7 and 8 April 1957.

White-tailed Tropicbirds are fairly regular midday visitors to
Johnston Atoll during winter, spring, and summer, when Red-tailed Tropic-
birds are displaying and nesting. None was known to land during POBSP
studies, although several hovered low over the bushes on Sand Island,
and a few hovered repeatedly in front of crevice-like openings between
raised window shutters and the overhanging roof. In August 1973,

Kridler (pers. comm.) recorded one on the ground under low vegetation
near the garbage dump on Johnston Island.

From 1963 through 1969, POBSP observers listed 47 sightings (Table
41): 28 from Sand, 18 from Johnston, and 1 from Akau. Since the amount
of time spent observing on Johnston was far less than on Sand, there must
have been proportionately more birds visiting Johnston than Sand. In
1969, however, 15 of 19 sightings were on Sand, possibly reflecting a
shift in activity to Sand similar to that described for Red-tailed
Tropicbirds, and possibly related to the attraction of white-tails to
displaying red-tails.

Most sightings were of single birds, but there were observations
of two birds in early January 1964, on 18 and 22 July 1968, and on 13
April 1969. Three birds were seen on 28 July 1964 and 8 April 1969.

Reasons for the yearly population fluctuations shown in Table 41
are not known. No information is available from the nesting areas of
these birds that could be correlated with these figures. The 1969
figures may be biased because after the first few sightings had been
made, more notice was taken of displaying birds. The three birds seen
8 April 1969 stayed at least 400 feet high, and would not have been
noticed had not their unusual calls been faintly heard. Before the fall
of 1968, these calls would have been inaudible because of noise from
the Coast Guard generators, now silenced as a result of the procurement
of electric power from Johnston Island. There is no reason to suspect
bias for any other year, especially 1966 when no birds were reported
despite the presence of well-qualified observers during most of the
season.

The seasonal pattern of occurrence shows maximum numbers from
January through April and again from July through August. The early
period corresponds with the early part of the breeding season on the
northwestern Hawaiian Islands; these birds may be fishing in the Johnston

206

area and are attracted to the island because of incipient broodiness.
The July-August increase may result from wandering pre-breeders which
are attracted to land areas slightly after the normal northern breeding
season. The late fall low probably corresponds with the general decline
in breeding activity of these birds at that time throughout their range
when all stay at sea and are not attracted to land.

Table 41. Sightings* of White-tailed Tropicbirds on Johnston Atoll

Month
Year Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. Total
L963 a7 = = = = = = Ory, 2. 0 0 0 0 2
EGA ee 2 0 0 0 0 3 2 i il 0 0 WZ
1965 2 0 0 il 0 0 (0) es 10) 0 0 0 0 3
1966 0 0 0 0 0 0 OP axa O) 0 0 0 0 0
1967 0 0 0 i 0 0 Of yal 0 = = = 2
1968. - 0 i: 0 1 0 4 2 0 1h 0 0 9
1969 0 0 3 8 4 0 es) 0 = = = 19
Totaks *5 2 4 10 5 0 S10 i 2 0 0 47

*Numbers indicate number of birds seen.

Banding and Interisland Movement

Only one bird was handled at Johnston Atoll. At 1300 on 17 August
1969 one repeatedly circled the barracks and fenced antennas on Sand
Island. It was netted on a low pass, banded, measured, photographed,
and released. This was likely an immature bird, judging from its dark
yellow bill. (Bill color was noted on three 1969 birds: one seen
9 March had an orange-red bill while one seen 28 July and the one caught
17 August had yellow bills.) There was no wing molt, but one central
tail feather was less than one-fourth grown (75 mm). This new rectrix
and parts of the body were pale pink, possibly indicating that body molt
was in progress. Measurements were: Bill: 44 mm; Wing: 272 mm; Tail:
506 mm; Tarsus: 24 mm.

At-Sea Observations

White-tailed Tropicbirds were recorded casually in the grid
southwest of Johnston Atoll in all months except March (POBSP, 1967a).

BLUE-FACED BOOBY Sula dactylatra
Status
Former common breeding species; recent regular but uncommon visitor

to Sand Island through the year. About 200 previously nested on the sandy
beaches of Johnston and Sand Islands. About seven birds visited Sand

207

Island each month during late winter and spring throughout POBSP studies;
an average of one bird visited each month during the remainder of the year.

Ecological Distribution
Akau Island: This species was suspected of roosting at night on
this man-made island during 1964-1969.

Hikina Island: Blue-faced Boobies probably roosted here at night
during the late 1960's.

Johnston Island: Brooke (ms.) while on Johnston Island on 14 March
1859 referred to "a gannet" and on the 15th noted: "Some white boobies
sat in the grass and put their snake like heads above it." These ob-
servations probably were of Blue-faced Boobies.

Wetmore (ms. a and b) found many on the beaches of Johnston Island
in July 1923. He noted that the rock ledge on the eastern shore was
used as a roosting spot with the birds standing flat-footed on level or
rounded stones and not perching on sharp projections. None has been re-
ported from the island since.

Sand Island: Wetmore (ms. a and b) noted small numbers on Sand
Island in July 1923 (Figs. 67 and 68). Non-nesting birds were observed
roosting during the day on the shore near the Brown Booby colony on the
southern tip of the man-made end of Sand Island in April 1957 by
Moynihan (1957). POBSP personnel found small numbers on the island
from 1963 through 1969.

Blue-faced Boobies visiting Sand Island in recent years most
commonly roosted on the LORAN-C antenna guywire bases. They also
frequented the south shore (near or among nesting Great Frigatebirds) ,
the east hill (among nesting Brown Boobies), the northeast peninsula
and the southwest islet. Birds were seen a few times on the large
anchor buoys between Johnston and Sand Islands.

Populations

Figure 69 shows semimonthly population estimates of Blue-faced
Boobies on the atoll during POBSP observations. These estimates and
Moynihan's (1957) figures show the present numbers are but a small
fragment of the earlier 1923 population. Johnston Island is now never
used by this species, and a few use Sand, particularly in mid-July, the
time of Wetmore's 1923 observations.

The most likely cause of this decline was killing of birds and
disturbance of nesting areas by military and construction personnel
before and during World War II. It is probable that most, if not all,
the nesting birds were killed, and none has successfully recolonized the
atoll.

Moynihan's (1957) observations of up to a dozen birds present during
the day indicates a larger population in 1957 than during the 1960's.

208

Figure 67. Blue-faced Booby adult and chick, Johnston Atoll, July 1923
(B. P. Bishop Museum photo).

Figure 68. A roosting club of Blue-faced Boobies on Sand Island, Johnston
Atoll, 2 June 1941. Two immatures are in the center foreground

(U. S. Nat. Archives, R. G. 80 photo).

209

30 e
ae Adults
25 : : —— Means
: : see. Extremes
20 : Bie.
vn ®
a : a
a : :
ca : .
a 15 : d--0--@
=
=
é

J F M A M J J A S OF IN D

Figure 69. Means and extremes of semimonthly estimates of Blue-faced Booby
numbers, Sand Island, Johnston Atoll, 1963-1969.

This many were sometimes present at night during POBSP studies, but there
were never a dozen birds on the island during the day. Birds usually ar-
rived within an hour before sunset and left by dawn. If present ratios

of day populations to night populations were the same in 1957, then the
1957 population may have been several times larger than any recorded during

the 1960's.

POBSP observations indicate a continuing decline in numbers using
the atoll (Table 42). Maximum numbers in 1966 were only about half the

210

maximum recorded in 1965, and the 1967 and 1968 maxima were less than a
quarter of the 1966 maximum. There was only a slight increase in 1969
over 1968.

The reasons for this post-World War II decline are not definitely
known. There are indications of a slight decline in young birds pro-
duced in the northwestern Hawaiian Islands, but the decline is not as
great as the decline in birds coming to Johnston. No changes in habitat
on Sand are known which would adversely affect the birds, and no changes
in oceanographic or meterologic phenomena are known. Continued disturbance
by people, including POBSP personnel, however, may have discouraged the
birds from visiting the islands, or from staying if they do visit. In
1969 care was taken not to disturb roosting Blue-faced Boobies unless
absolutely necessary for identification. After being handled once they
were not flushed or otherwise disturbed. Several of these birds returned
to the island nightly for over two months--much longer than any stayed in
years when mass Sooty Tern banding greatly disturbed all ground roosting
birds.

Barring further alteration of Sand itself or further disturbance
by man, this species should be able to hold their own. If not, Blue-
faced Boobies will become very rare in the Johnston Atoll avifauna.
Thus, even if man left Johnston Atoll it would be many years, if not
an impossibility, before this species would nest there again because
of their strong tendency to breed on their place of hatching. There are
probably no birds remaining that belong to what could be called a
Johnston Atoll breeding population.

Annual Cycle

Wetmore's (ms. a and b) observations in mid-July 1923 that "A few
(one or two pairs) had eggs and some downy young seen, but most [chicks]
were well grown" indicated a spring-summer nesting cycle similar to that
in the northwestern Hawaiian Islands. There adults are present all year,
with eggs usually present from late January through mid-August, and
fledglings from late June to November (Richardson, 1957; Amerson, 1971).

In recent years (see Fig. 69) Blue-faced Boobies have been present
throughout the year, a similarity also of the Hawaiian population.

Specimens

Appendix Table 7 presents data on the ten Blue-faced Booby specimens
known from Johnston Atoll. All are housed in the USNM. These constitute
a new published specimen record, as well as an extirpated breeding record.

Banding and Interisland Movement

in eaalas ‘4h Blue-faced Boobies (two adult males, eight adult females,
14 unknown sex adults, 19 subadults and one immature) were banded by the
POBSP on Sand Island (Tables 24 and 43). Of these 44 birds, only four
were later recaptured on Sand, while five were taken as interisland

211

Table 42. Semimonthly population estimates of Blue-faced Boobies on
Sand Island, Johnston Atoll, 1963-1969

Period 1963 1964 1965 1966 1967 1968 1969 Mean
January 1-15 = 3 5 0 2 - 0 2
16-31 - 4 7 7 3} - 0 4
February 1-15 = 7 7 17 2 1 0 6
16-28 - 7! 30 6 3) 1 0 8
March 1-15 - 7 25 9 2 1 D 8
16-31 - 7 20 1b 3 D: 2 8
April 1-15 - 1u6) 20 abt ik 2 3 9
16-30 - 19 16 7 1 0 4 8
May 1-15 - 15 8 2 1 0 4 5
16-31 - 15 5 1 0) 0 3 4
June 1-15 - 15 5 0 0) 1 3 4
16-30 ~ 172 i 0 0 at 3 3
July 1-15 - 8 0 iL 0 if 2 2
16-31 it 5 0 0 0 il 3 1
August 1-15 = 5 0 0 0 1 3 1
16-31 2 2 0 0 0 0 2 1
September 1-15 2 iL 0 0 1 0 1 il
16-30 0 I 0 uf - i il
October 1-15 0 2 0) 0 - 0) - 0
16-31 0 0 0 0 ~ 0 - 0
November 1-15 0 0? 1 0 - 0 - 0
16-30 0 0? il 0 - 0 - 0
December 1-15 0 0? 2 0 ~ 0 = 0
16-31 0 0? 0 il - 1 - 0
recaptures on the northwestern Hawaiian Islands (Table 26). In addition

21 birds banded on other islands were caught on Sand through 1969.

The ages of the birds visiting the atoll indicate they were young
birds from the northwestern Hawaiian Islands which visit the Johnston
area during their pre-breeding-age wanderings. An unusually high 31
percent of the 65 different birds handled on the atoll was banded on
other islands. None of these 21 interisland birds had been banded as
adults, although in most cases as many adults or young were banded on
the islands of origin. Although 24 of the 44 birds banded on Sand were
listed as adults at the time of banding, it is likely that most were young

212

adults which had not yet begun breeding. That only two of the 24 had
acquired the distinctive adult male voice supports this hypothesis.

French Frigate Shoals was the island from which most of the inter-
island recaptures came and to which most Johnston-banded birds went.
This is the closest island to Johnston and has one of the largest nesting
populations of Blue-faced Boobies in the Hawaiian chain.

At-Sea Distribution
Blue-faced Boobies were of casual occurrence at sea in the grid
southeast of Johnston Atoll. They were observed in all months (see Table

21), except June through September (POBSP, 1967a).

Table 43. Blue-faced Boobies banded on Sand Island, Johnston Atoll,
by the POBSP

Adult Adult Adult, sex

Date of Banding males females unknown Subadults Immatures' Total
19635 July 0 0 - 0 2 0 2
1964 January 0 1 0 0 0 alk
February 1 il il 1 0 4
March 0 il: 0 2 0 3}
April 0 2 0 2 0 4
June 0 0 1 0 0 ill
1965 February 0 0 S) 0 0 9
March 0) 0 3 0 0 3
April 0 0 0 1 all 2
May 0 iL 0 0 0) il
June ali 0 0 2 0 3}
1966 February 0 a 0 0 0 1
March 0 0 0 4 4
1967 February 0 ili 0 0 0 AL
1968 none banded
1969 March 0 0 0 2 0) 2
April 0 0 0 1 0 ill
July 0 0 0 Z 0 2

2 8 14 IS) Al 44

Pail)

BROWN BOOBY Sula leucogaster
Status

Common breeding species, present year-round. Nests are built on
the ground on sites exposed to easterly winds at Sand Island; previously
bred on Johnston Island. From 1964 through 1969 the breeding population
increased from about 70 to about 140, and the chick productivity in-
creased from 30 to 50. Thus the population returned to nearly its
original numbers after a decline of unknown extent during the war years.
Most adults probably stay close to the atoll year round but roost offshore
on pilings and navigational aids except during the spring breeding season.

Ecological Distribution

Johnston Island: Wetmore (ms. a and b) observed many here in
July 1923; a few nests--all on the ground--held chicks from two weeks
old to nearly able to fly. Wetmore reported Brown Boobies nesting on the
ground, and only on Johnston Island, in 1923. The specific location was
not mentioned, but his description of the young climbing around on the
rock ledges indicates that they were on well exposed sites. None has
been recorded since.

Sand Island: In April 1957 Moynihan (1957) noted a small breeding
colony with eggs to young of various ages in low grass and shrubbery on
the southern tip of the western or man-made portion of Sand Island. In
view of their preference for exposed slopes, it is difficult to see why
the birds would have nested on this low-lying, somewhat protected area.
Whatever the reason for this choice, the occupancy of the western end
of Sand Island by the Coast Guard in the early 1960's necessitated the
move to the present locations on the original, eastern portion of Sand
Island.

Davis (1962) also found a small number of nesting pairs on Sand in
1959-1960. POBSP personnel recorded this species as nesting here from
1963 through 1969.

Figures 43 and 70 show locations of Brown Booby nests on Sand Island
during POBSP studies. Little variation occurred during this period.
Nests appeared to be placed more or less at random within the area, ex-
cept that they were never closer together than about three feet. Individuals
usually nested in the same general area, sometimes on almost exactly the
same spot, in successive years. Ground cover apparently had little effect
on nest site selection. Sites usually were situated to allow down-slope
take-offs into the trade winds.

Nests were built on the ground (Figs. 71 and 72), and consisted of
a deep cup formed of Boerhavia, Lepturus, Sesuvium, or Tribulus, relative
amounts depending at least as much on availability as on preference. At
least two nests were built on man-made objects in the lagoon. One in
1967 was located on a concrete antenna base and another in 1968 was located
on one of the wooden navigation aids; military officials, however, dis-
couraged such attempts.

214

1967

4

Figure 70. Distribution of Brown Booby nests on Sand Island, Johnston
Atoll, 1969.

Except when they were courting, incubating, or tending chicks,
nearly all adult Brown Boobies roosted on guywire bases and navigational
aids in the lagoon, rather than on the island. A few did, however, roost
on the rock ledge along the west side of the northeast peninsula.

Populations

Figure 73 shows the means of POBSP semimonthly population estimates
for Brown Boobies at Sand Island based on 1964 to 1969 data. Estimates
varied more between observers than between years; hence the mean is the
best available representation of numbers present during any given semi-
monthly period.

The number of different birds handled each year doubled from 65 in
1964 to 129 in 1969 (see Table 24). Nearly all nesting birds were
handled in all years, at least on the main east hill colony, and several
one-year-old or older subadults were also caught each year. These
figures probably represent 90 percent of the total number of birds using
the atoll each year. If this is true, then the total number using the
atoll increased from about 70 to about 140 from 1964 to 1969.

215

Figure 71. Brown Boobies in early stages of courtship and nest building,
east hill of Sand Island, Johnston Atoll, 25 February 1969.

Vegetation is mostly Boerhavia with scattered clumps of Tribulus
(POBSP photo by P. C. Shelton).

“ eS a ite

Figure 72. Brown Booby adults and 31-day-old chick, west shore of northeast
peninsula, Sand Island, Johnston Atoll, 9 July 1966. Brown
Noddies roost and nest here also; Sooty Terns are in the
background (POBSP photo by P. C. Shelton).

216

200

oa) Adults
: 5 —— Means
150 Pa : 0 @, 3 oes (Exthemes

e::

Nou IM Be E Res
©

50

a 40
= 30 aes ee Baye aAgs Young
faa) eo °
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= 10 s M85;
0 °° Se—e—e0., @--@.,

J iz M A M J J A S 0 N D
Means and extremes of semimonthly estimates of Brown Booby
numbers, Sand Island, Johnston Atoll, 1963-1969.

Figure 73.

217

Wetmore's observation of 75 birds on Johnston in July 1923
indicates a population nearly the same size as that found during POBSP
studies, but the 18 pairs observed by Moynihan in 1957 were only about
half the number of breeders observed during the 1960's. Thus, there
appears to have been a decline in numbers of Brown Boobies during the
war years and a subsequent increase which has not yet leveled off.

Production of young each year increased in almost exactly the same
proportion as did the number of subadults handled (see Table 52).
Except for the late nesters, hens normally laid two eggs each year which
increased their chances of raising a chick. This, plus re-nesting, made
the number of successful pairs fairly high, although the percentage of
egg losses resulted from abandonment or infertility, rather than from
outside disturbance. Chick losses were mostly of second chicks in two-
egg clutches, discussed in more detail below.

Tables 44 through 48 show productivity in terms of clutch size,
first, second, or third eggs in two- and three-egg clutches, re-nesting
attempts, and the portion of the breeding season in which the nests were
initiated. Most chicks resulted from the first egg of a two-egg clutch
laid during the early part of the breeding season. But second eggs and
re-nesting accounted for enough chicks to make the difference between a
stable and an increasing population. Re-nesting would have been more
frequent in some years if pairs where nests failed had not been given
abandoned chicks from other nests. One-egg clutches had low success,
but these were almost entirely laid by young birds nesting for the first
time. Likewise late season nesting success was low, mainly because it
was during this time that young birds made their first, usually un-
successful, breeding attempts. Late season re-nesting by mature birds
had a high success rate.

It would appear that since first eggs accounted for over 80 percent
of chicks, and re-nesting success was unusually high, a more efficient
strategy for mature birds would be to lay one egg, and to re-nest, again
laying only one egg, if the first egg failed to produce a chick. It
would be advantageous to lay two eggs in the first nest only if food
conditions deteriorated before the second clutch could be laid. But the
high rate of success of re-nesters, many of them beginning quite late in
the season, indicates that food must be easily available for several
months after maximum nesting occurs.

That the few three-egg clutches laid had a higher success rate than
either one- or two-egg clutches indicates that pairs capable of laying
three eggs were easily capable of rearing a chick. None of these clutches
produced more than one chick, and in no known case did the chick come from
the third egg.

The rare cases in which both chicks of a two-egg clutch survived
appeared to result from lack of aggression toward the second chick by the
first. Normally the first chicks hatches several days ahead of the
second (the first egg is incubated immediately after being laid), and is
therefore considerably larger than the second when the second hatches.

218

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221

Within a few days the first chick becomes antagonistic toward the second
chick and pushes it out of the nest. Occasionally the parents retrieve
the expelled chick, only to have it ejected again, or one parent may brood
it outside the nest briefly. More commonly it is ignored and left to die,
and in one case an adult male ate an ejected second chick. In no case was
an expelled chick known to survive. In those cases where the second chick
did survive with the first, there was no antagonistic behavior between the
two. The parents appeared to make no distinction, and fed both regularly.
In 1969, both chicks of the pair raised from the same nest were measured
regularly. The older grew at about the average rate for single chicks and
the younger was slightly below average. Both fledged in under 90 days
which is at least as young as most single birds fledge. Thus it would
appear that the parents had no difficulty feeding both chicks.

In summary, it appears that the reproductive strategy of Brown
Boobies on Johnston Atoll retains characteristics developed under condi-
tions significantly different from those existing at present around
Johnston Atoll. The optimum brood size probably was one, whereas two
could be reared more frequently on Johnston now if first chicks were
less aggressive toward their younger brood-mates. Re-laying after
failure of the first clutch must have been less efficient insurance than
laying both eggs at approximately the same time. Thus it would appear
that food must have been more scarce and available over a shorter period
of time than under present Johnston conditions.

But at present the Johnston population is increasing, which indicates
understocking. As the population becomes greater, food may be depleted
more quickly, bringing about a return to conditions similar to those
hypothesized to have existed some time earlier in the evolutionary his-
tory of Brown Boobies. Should this happen, the rate of success of re-
nesters, and the incidence of two chicks being reared by one pair, will
decline.

Table 49 summarizes breeding information on young birds returning
to enter the breeding population. There is a steady increase in the
number of birds attempting to nest through their fourth year, and there
is also a steady increase in the rate of success of those that do at-
tempt breeding. All breeding two-year-old birds were females, but by
the fourth year there were as many males breeding as females. Birds up
to four years old generally nested later than the older established
breeders, but five- and six-year-old birds began nesting during or
slightly ahead of the period of maximum nesting for the total population.

Annual Cycle

Brown Boobies use Johnston Atoll year round (Figs. 37 and 73), but
apparent numbers fluctuate from a maximum of over 100 during the spring
breeding season to a low of about 50 during the fall. Most of the
breeders probably stay near the atoll year round, but do not return to
roost on the island as regularly during the fall, which may account for
the decline in numbers at that time.

222

Table 47. Renesting attempts, incidence of two chicks reared in one
nest, and numbers of chicks raised by foster parents or by
hand, Sand Island, Johnston Atoll, 1964 through 1969

Renesting
% of Total Number of nests Number chicks

Number % chicks from where two chicks reared by fos-
Year of nests Success renests were reared ter parents*
1964 2 50 4 it 0
1965 9 100 30 0 3
1966 5 ? 2 il 1
1967 2 100 5 i 0
1968 6 100 15 0 8
1969 12 25 6 1 3
Totals 36 68** 12%* : 4 15

*Al11 were second chicks ejected from nests by older chicks. AI11l
reared by Brown Booby pairs that had lost all their own eggs or
chicks, except the three from 1969, which were hand-fed on
Johnston Island.

**Not including 1966.

Table 50 presents the extreme dates for significant events in the
Brown Booby breeding cycle during 1964 through 1969. Courtship has been
observed by late November, but usually does not begin until January.
During courtship, pairs spend most of the day on the nesting area, but
leave in the evening to roost offshore. Egg laying has begun as early
as late January, but normally began mid-February; eggs have been recorded
as late as the end of September. Incubation required about 43 days
(1969 data: range = 41 to 47 days; n = 80) and maximum hatching occurred
in April. Chicks are closely guarded by the adults for about a month
after hatching, although the chicks by this time are over half grown.

The young fledge when about 90 days old (1969 data: range = ca. 80 to
ca. 100 days; n = 20).

Table 51 shows the number of new nests established by semimonthly
periods during 1964 through 1969. Although in 1965 and 1968 maximum
laying occurred slightly earlier than in the immediately previous years,
there was a general trend toward later nesting, discernible both in the
date of first egg and the period of maximum laying. The variation was
no more than a month over all years.

223

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225

Table 50. Extreme dates of significant events in the Brown Booby breeding
cycle, Sand Island, Johnston Atoll, 1964-1969

Year Laying Hatching Fledging

1964 23 Jan.-21 Apr. 10 Mar.-? ca. 10 June-?

1965 ca. 15 Feb.-29 May 29 Mar.-10 July 30-June—21) Oct.

1966 11 Feb.-8 July ca. 1 Apr.-l Aug. Ca. 5 July-20 Oct.

1967 12-13 Feb.-12 July 25 Mar.-23 Aug. ca. 25 June-ca. 25 Nov.
1968 20 Feb.-21 Aug. 4 Apr.-30 Sept. 27 June-ca. 10 Dec.

1969 19 Feb.-ca. 30 Sept. 2 Apr.-ca. 30 Sept. 3 July-ca. 5 Nov.

There was evidence that human disturbance may have delayed laying
slightly. In 1967 and 1969, the first eggs were laid on the southwest
islet, which is relatively rarely visited by people. Courting birds
flush readily when approached to within several yards, and disturbance
as little as two or three times a day by passing fishermen or biologists
may prevent these birds from progressing as rapidly to laying as non-
disturbed birds. Birds with eggs or young, by contrast, are docile and
usually can be caught by hand with little difficulty.

The weather disruption in February 1969 had little effect on Brown
Boobies. Nesting occurred later in 1969 than 1968, but not later than
in 1967. Courting birds decreased from about 50 birds each day in
late January to about 10 by the second week in February. By mid-
February, numbers returned to the late January level, and laying began
by the 19th, almost exactly the same time as in 1968. Laying progressed
more Slowly, however, and maximum laying occurred two weeks later in
1969 than in 1968. This delay is much less than that for several other
species in 1969. Perhaps the tendency of Brown Boobies to feed closer
to the island and in shallower water than any other breeding species
accounts for their being less affected by the changed weather pattern
than birds that feed farther at sea where food production may have been
more drastically affected.

Another significant change in the pattern of nesting, evident on
both Tables 50 and 51, is the increased variability to be expected from
a larger population. However, if social facilitation is a significant
factor in inducing breeding behavior in Brown Boobies, then the larger
number of birds present in the later years may have stimulated more
potential late nesters to lay.

Wetmore's description of the colony in July 1923 fits the stage
found in July during the 1960's. Moynihan, however, found near-fledging
chicks in early April 1957 about two months earlier than the earliest
recorded fledging during POBSP studies. Perhaps the trend toward later
nesting observed during the 1960's was a continuation of readjustment in
the breeding cycle induced by disturbance during the war years.

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226

7/2) |

During POBSP studies the maximum number of birds occurred in
April, shortly after the laying peak, when most birds were incubating
or tending small chicks. Incubation shifts occurred at least once
and probably twice each day. In March 1968, Harrington found more
females than males on nests in the early morning, with the percentage
of males increasing throughout the day. Almost all incubation at
night was done by males, and females apparently returned about dawn.
Fairly frequently both members of a pair were at the nest during the
day, but at night only those incubating remained on the island, while
the mate roosted offshore on navigational aids or guywire bases,
or possibly fed at sea. The frequent incubation shifts, and the
occasional occurrence of both members of pairs at the nest, indicate
that these birds feed fairly close to the island, perhaps within 25
miles most of the time, and that food is relatively easy for them to
catch.

Soon atter fledging the immatures begin roosting at night offshore
on the guywire bases and navigational aids, but spend much time back
on the island during the day, usually near their nest sites. On
nights of unusually high winds they frequently roost on the ground.
Some of these are still fed fairly regularly by their parents until
at least December. About half the immatures from the previous season
are on hand when the adults begin courting for the new breeding
season. These young birds vigorously beg from the adults, who
usually ignore the youngsters, and only rarely regurgitate food for
them. The yearlings acquire their subadult plumage--typical adult
pattern, except for scattered dark speckles on the breast and belly--
from about 11 to 14 months of age. Facial colors begin to differen-
tiate at this time also.

Breeding adults begin to molt before their chicks fledge. Four
birds with large chicks or second nests examined in August 1969 had
replaced most of their primaries. Replacement appeared to have begun
simultaneously with primaries 1 and 6 and proceeded distally therefrom.

Specimens

Fourteen specimens of Brown Boobies have been collected at Johnston
Atoll (Appendix Table 7); all are located in the USNM bird collection.
These constitute a new published specimen record for the atoll.

Banding and Interisland Movement

In all, 375 Brown Boobies (114 adults and subadults, 261 young)
have been banded at Sand Island (Tables 24 and 52). Nearly all nesting
birds were banded and/or handled each year. The same birds returned
each year, showing a strong attraction for the island. Individuals
usually nested in the same general area, sometimes on almost exactly
the same spot, in successive years.

Interisland movements of Brown Boobies have been few (Table 26).
Four Johnston banded birds were recaptured or observed (identified by

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228

orange streamer) on other islands. An immature banded 12 July 1966
was captured alive and uninjured on Nauru Island, 0°20'S, 167°10'E,
2,000 statute miles from Johnston on 29 December 1966. An adult
male, banded 26 April 1967, mated with a previously banded female
and raised a chick which fledged about 9 August 1967 on Sand Island;
on 1 September 1967, the male was caught on Lisianski Island. This
probably was a Lisianski native, which happened to be on Johnston
during the breeding season, but returned to Lisianski immediately
thereafter. Brown Boobies with orange streamers were sighted on La
Perouse Pinnacle, French Frigate Shoals, on 12 June 1967, and on
Taka Atoll in the Marshall Islands (about 1,200 nm southwest of
Johnston) 21 October 1964 (Amerson, 1969: 47).

There are two recorded movements of Brown Boobies to Johnston
from other islands. On 30 June 1964 POBSP personnel found in the
tunnel on the southwest islet of Sand Island a band which was put on
a young Brown Booby on Eastern Island, Midway Atoll, by Dale Rice of
the U.S. Fish and Wildlife Service 2 April 1958. This bird apparently
died on Sand some time before POBSP studies began. An adult female
banded 19 June 1966 on Wilkes Island, Wake Atoll, appeared in the
east hill colony on Sand 20 May 1969, and laid an egg. No mate was
ever seen with her or with the egg, and she abandoned the egg and
disappeared after a few days.

At-Sea Distribution

Brown Boobies were observed in low numbers in the grid southwest
of Johnston Atoll from October through April (Table 21). They were
absent from May through September, except for July (POBSP, 1967a).

RED-FOOTED BOOBY Sula sula
Status

Uncommon breeding species, as well as an abundant, transient
visitor; breeders present from February through August and probably
are year-round residents, transients present throughout the year.
Nests built primarily of Tribulus stems placed on the ground at Sand
Island and on pilings, etc., around the periphery of the island;
previously nested on Johnston Island. Transient visitors roost on
antenna guywires, rock piles, pilings, and concrete blocks around
Sand Island. During POBSP studies about a dozen pairs attempted to
nest yearly, with one to six chicks fledging; transients numbered up
to 3,000 or more at night.

Ecological Distribution

Johnston Island: About 250 were using Johnston Island in July
1923, but only six nests with eggs and a few with young from two weeks
old to those nearly grown were there. Roosting birds used the low rock
ledges along the beaches (Wetmore, ms. a and b). None has been recorded
from the island since 1923.

229

230

Sand Island: Wetmore (ms. a) in July 1923 noted that "after three
or four nights disturbance [on Johnston Island] however the majority
moved to Sand Island." During April 1957, Moynihan (1957) saw approxi-
mately 100 on the western man-made portion. Some were immature birds;
but most were adults, sitting on nests with eggs or fairly small chicks,
on the same towers as the frigatebirds. POBSP personnel found them on
Sand Island from 1963 to 1969.

The distribution of nesting sites used by Red-footed Boobies are
shown in Figures 44 and 74. Changes in sites used during years of
POBSP studies and in earlier years are discussed under the Population
section. This species normally nests on bushes or trees, but these
were completely lacking from the original flora of Johnston Atoll.
Wetmore did not give details of nesting in 1923, but did mention one
nest on the ground. Perhaps the comparatively small population of
breeders--apparently not more than 50 or 60 in 1923--is a result of this
lack of optimum habitat. Best conditions for red-foots may have occurred
immediately after World War II, when abandoned towers and antennas on
Sand Island provided raised nesting sites for perhaps 100 birds. The
only raised sites available after the Coast Guard occupied Sand Island
were pilings, concrete blocks, and rock piles around the periphery of
the island, which the birds use willingly, but with little success
because most of these are within reach of high tides and most eggs and
nests are destroyed. Only the birds nesting on the east hill had even
a fair degree of success.

There were a few bushes growing on the original end of Sand that
were large enough to support nests by 1969, but they were not in areas
frequently used by red-foots. The Tournefortta northwest of the trans-
mitter building occasionally attracted roosting birds; by 1973 it
supported at least one nester.

The favored nest building material during our studies was Trtbulus
stems which the birds gathered from growing plants. Red-footed Boobies
are a major factor in the ecology of this plant species, for in years
when its growth is not luxuriant, as in early 1967, the birds almost
entirely strip the island of it. In 1967, the red-foot nests built in
March were almost entirely of Trtbulus, but by the end of the month,
there was practically none left. The birds had pulled up all available
stems, leaving only frayed and broken stubs above ground. When the second
round of nests was built in late March and April, Boerhavia was the major
material used, with a little Sesuvitum in some cases. These materials are
thinner and less durable than Tribulus and definitely were less desired
by the birds. Vigorous sprouting from root crowns later in the year re-
established Tribulus to its normal status. Its susceptibility to
destruction in early 1967 may have resulted from the dry year of 1966,
which most plants survived with reduced vigor. In 1969, when all plant
species were unusually lush, following a year of above average rainfall,
red-foots did little damage to Tribulus, and did not significantly
decrease its abundance or vigor.

234

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The primary roosting habitat for Red-footed Boobies on Sand Island
is the guywires of the LORAN-C antenna. Most preferred are the lower
portions of those wires lying approximately perpendicular to the wind
direction. Both Red-footed Boobies and Great Frigatebirds use the wires,
but there is little mixing of the two species. In late 1966 Schreiber
observed that Great Frigatebirds arrived earlier in the evening than
Red-footed Boobies, and often landed on the lower parts of the wires.
Later in the evening as red-foots arrived in increasing numbers, the
boobies drove the frigates farther up the wires. In all cases of conflict
observed, the frigatebird moved. Huber noted in June 1969 that Great
Frigatebirds left the island earlier in the morning than red-foots,
Schreiber also found some evidence that adult red-foots usurped the
lower parts of the wires, forcing the younger birds to move higher.

Besides the guywires, roosting red-foots used rock piles, pilings
and concrete blocks around the margins of the island. After the Coast
Guard burned the planking from the old dock in August 1967, red-foots
began roosting in numbers up to 200 to 300 on the exposed iron beams
that remained.

Nesting birds attracted young non-breeders, and in the years 1963
through 1965 and again in 1969, subadults roosted on the east hill among
the nesting birds. These numbered up to 100 or more in the early years,
but no more than a few dozen in 1969, when only one pair nested on the
hill. The south shore where Great Frigatebirds and a few red-foots
nested was another favored roosting place for non-breeding red-foots.

Populations

Figure 75 shows the means and extremes of semimonthly population
estimates of Red-footed Boobies from 1964 through 1969. The population
is composed of two distinct elements: (1) resident breeding birds
numbering no more than 2 or 3 dozen, which are the remnant of a formerly
larger breeding population, and (2) transient birds, numbering upwards
of 3,000 and mostly young pre-breeders from the northwestern Hawaiian
Islands, that use Sand Island only as a roosting place while they are
feeding or traveling in the Johnston area.

In contrast to the other two booby species, red-foots occur in
relatively large numbers and nowhere near the total number visiting the
island can be caught, or even accurately counted. Even the relatively
few breederswere not handled in sufficiently large numbers to give more
than a rough idea of their numbers. Maximum numbers of roosting birds
occur only after dark and roost on the outer guywires, where they cannot
be counted with less than perhaps a 25 percent margin of error, even by
the most meticulous and experienced observers. Reported figures often
fluctuated wildly from period to period, especially if personnel changed,
and the new man made independent estimates based on his own interpretations
of apparent numbers. For this reason, no purpose would be served by
showing all semimonthly estimates. The graphed means probably represent
a fair approximation of actual numbers present, and clearly shows the
seasonal trends in numbers.

NUMBERS

233

6,000

Adults

5,000 e —— Means

coors Extremes

4,000

NUMBERS

OFM W FO O

CaN a M A M J J ARES 0) N D

Figure 75. Means and extremes of semimonthly estimates of Red-footed Booby

numbers, Sand Island, Johnston Atoll, 1964-1969.

234

The size of the original breeding population is unknown, but
Wetmore's and Moynihan's observations indicated that it was sub-
stantially larger than that found during the 1960's. Wetmore's count of
six nests with eggs, as well as young (number not given) from two weeks
old to flying indicates at least twice as many breeders as during our
studies, when no more than three eggs were found in July, and six was
the maximum number of young produced in one year.

Moynihan (1957) did not count eggs or chicks, but his statement
that most of the 100 birds he saw were on nests with eggs or small chicks
indicates a population of breeders much larger than any noted since then,
and possibly it was larger than it was before human occupation of the
atoll. The brief period between Navy abandonment of Sand Island and its
occupation by the Coast Guard may have been more favorable for Red-
footed Boobies than any period before or since. The wrecked and abandoned
towers and buildings may have provided enough elevated nest sites to
attract a larger breeding population than was present before the war, and
elimination of these sites by the Coast Guard no doubt decreased suita-
bility of the island for red-foots considerably.

If the number of breeders in 1923 was as many as 50 pairs, then a
substantial portion, perhaps half, of the first and second year birds
Wetmore observed roosting on the island could have been young, Johnston-
hatched birds. The others were most likely from other islands.
Moynihan's observations do not indicate the presence of a substantial
number of non-breeding birds. He did not, however, visit Sand Island
except during the day, when roosting birds are at their lowest numbers.
It does not seem likely, however, that anywhere near the number of
roosting birds present during Aprils of POBSP studies (2,000 to 3,000)
could have been using the atoll without his noticing them arriving at
dusk. It therefore seems likely that the large number of roosting,
non-breeding birds now using the island are attracted primarily to the
guywires of the LORAN-C antenna and that much smaller numbers of these
roosting birds used the island before the antenna was built.

Table 53 shows productivity figures for Red-footed Boobies for
1963 through 1969. These birds have the lowest rate of productivity,
in terms of chicks produced per reproductive effort (eggs laid) and
pairs breeding of any species now breeding on Johnston Atoll. Persistent
renesting does not contribute significantly to productivity, for most
successful eggs were the first laid by the pair in that year. In 1969,
four pairs laid three eggs each without producing a chick.

The poor nesting success results primarily from lack of suitable
nest sites (Figs. 76 and 77). In 1963, 1964, and 1965, most Red-footed
Boobies nested in Trtbulus clumps on the east hill, near the Brown Booby
and Great Frigate colonies. A few nests were found in these years on
concrete blocks and pilings around the island, but all the young came
from the east hill. In 1966, 1967, and 1968, all nests were around the
margins of the island, on pilings, rock piles, and the remains of an old
boat. Increased activity of POBSP personnel was the most likely cause

Table 53. Productivity of Red-footed Boobies, Sand Island, Johnston
Atoll, 1963-1969

Eggs Chicks
Percent Percent Percent

No. No. No. Percent of of of
Year Pairs Laid Hatched Hatched Fledged Hatch Eggs Pairs
1963 c4 (8)* (3) ? (3) 4 2 ?
1964 ? (12) (6) (50) 6 2 y ?
1965 9 15 6 40 4 67 27 44
1966 2 (9) 2 2 1 50 2 ?
1967 i 21 3 14 if 33 > 9
1968 (17) 27 6 22 4 67 5 ?
1969 At 22 7 32 3 43 14 27
Mean 2 16.3 4.7 32 Shad 52 abs 27

*Figures in parentheses are approximate.

of the birds failing to nest on the east hill. In 1966 particularly,
Sooty Tern banding operations probably disturbed any pre-nesting birds
on the east hill up to several times each night, which apparently was
enough to discourage them from nesting there. In 1969 a pair that began
courting on the east hill was carefully avoided until after they laid
their eggs; they were successful in raising a chick.

Losses from the peripheral nest sites most commonly resulted from
nests being washed away by high tides, usually the first spring tide
after the nest was established. The nests frequently were precariously
placed and high winds sometimes swept them off. The shallow cups formed
by the nests did not insure the egg's safety, and flushed birds often
kicked the egg off the nest platform as they flew. In 1969 a 19-day-old
chick apparently fell off its nest on the old boat and drowned. Two
infertile eggs, one each in 1968 and 1969, were incubated well beyond
the normal incubation period.

Kleen constructed several platforms of old buckets and gratings
on the pilings and derelict boat in late 1968, and one of the 1969
chicks fledged from one of these sites.

Red-footed Boobies mature relatively early, and may breed as three-
year-olds (Table 54). Interestingly, one four-year-old bird successfully

235

236

Figure 76. Four Red-footed Boobies nesting on artificial platforms (framewor
of an old boat) on the southwest beach of Sand Island, Johnston §
Atoll, 5 May 1969 (POBSP photo by P. C. Shelton).

Figure 7/7. Red-footed Booby nest placed on ground on crest of hill east of
transmitter building, Sand Island, Johnston Atoll, 4 May 1969.
Sub-adult Blue-faced Booby roosting at lower right; Sooty Tern
adults and chicks and Brown Noddy adult nearby (POBSP photo by
PP. Ge. Shelton).

231

raised a chick in 1969, which gave the known-age birds as high a rate

of success as the general population. More known-age birds may have
nested but were not caught, for only a little more than half the nesting
birds were handled each year.

Table 54. Known-age Red-footed Booby nesting, Sand Island, Johnston
Atoll, 1963-1969

Found breeding as:

Year of No. 3-year-old 4-—year-old 5-year old
Hatching Fledged* No. hs No. vs No. th
1963 3 1 33
1964 6 2 33 iL 17 7 33
1965 3 1 33 1**k 33

Total 12 3 3)8\ 2 25 3 33

*Less those known dead before maturity.

**kOnly known age bird known to raise a chick.

Annual Cycle

Breeders: The breeding birds probably are year-round residents on
Johnston Atoll, and courtship has been observed in all months. But
intensive courtship usually begins in January, when displaying adults
are seen daily, and nest building begins. Laying normally begins in
February and continues through the following four to six months (Table
55 and Figs. 37 and 75). Most eggs laid in April and later were by
renesting birds that lost their first eggs. Relaying usually occurred
about a month after the previous egg was lost.

The incubation period ranges from 44 to 46 days (mean = 45.4 days;
n= 7). Chicks usually begin hatching in early April and may continue
until late September. The fledging period ranges from 95 to 101 days
(n = 4). Young birds are known to have fledged from early July to early
December. These young birds remain near their nest sites for at least
a few weeks after fledging, but soon begin roosting farther away.

Incubation shifts probably occurred fairly often--at least once a
day--but may not have been as frequent as shifts in Brown Boobies.
After hatching, chicks were attended closely for the first few weeks,
but not as closely or as long as Brown Booby chicks. Unlike Brown
Boobies, both Red-footed Booby adults rarely appeared together at the
nest or with the chick. These observations possibly indicate that red-
foots have more difficulty finding food than Brown Boobies, or at least
that they travel farther from the island in search of food. The

238

Table 55. Extreme dates of significant events in the Red-footed
Booby breeding cycle, Sand Island, Johnston Atoll, 1963-

1969

1963 7 —s bor Auge Pate On 2S. Ulsy; ? (to. 23 Sept.

1964 23. Feb. toubyss July @ dil t4s Apr. tou? ? -tota Sept.

1965 by 17 Feb.) to 16) July737 wApr. tom2) Sept. late July to 10 Oct -

1966 5 Feb. to late Apr. Early May to early June eq. 15 Aug. (only
one bird fledged)

1967 + 8 Feb. to 1-4 Sept. mid-May 15-25 Aug. (only
one bird fledged)

1968 by 14 Beb. (to) 28"Aug. 3cas7 97 Apr. ‘to 23 Sept. 9 July to early Dec.

1969 19 Mar. to ca. 21 Aug. 4 May to ca. 25 Sept. 12 Auge toy ly, Aug.

comparatively large number of non-nesting red-foots--at least a few
hundred at any time of year--might provide more competition for food
than occurs in the smaller Brown Booby population. This comparison
is not wholly pertinent if, as is suspected, Brown Boobies feed
mostly inshore and red-foots over the open ocean, for the areas
available to the two species would not then be of comparable size.

Throughout the breeding season, adult birds commonly roost on
the pilings and rock piles where they nest. A few of these birds
probably use these perches year-round. Whether or not there is a
decline in numbers of breeding birds during the non-breeding months
is not known, for these birds are not entirely separable from non-
breeders, and during this season they are not easily caught for
identification.

Non-breeders: The annual cycle in numbers shown on the population
graph (Fig. 75) represents non-breeding birds almost entirely. Highest
numbers occur in mid-winter months at about the beginning of the
Johnston breeding season, followed by a slow decline through spring,
summer, and early autumn, reaching a low in October. The sudden rise
in numbers in late fall comes after the end of the breeding season on
the northern islands.

In late fall and early winter, most of the birds are adults or
older subadults,,at least 1.5 years old, with white breasts and heads,
but dark speckling on their backs and wings, typical of birds in their
second and third years. Until late February, birds of the past
breeding season--dark headed birds, with dark underparts or at least
a dark breast band--are rare. Apparently these birds do not leave the

239

nesting islands until after the adults and older subadults. But in
late February the proportion of these dark young in the Johnston
population increases and by late March they are the most numerous
age-class present (Table 56).

Table 56. Plumage ratios of Red-footed Boobies, Sand Island,
Johnston Atoll, 1967

Adults and Subadults Subadults less Roteaue
Date over 1 year old* than 1 year old** Counted
Zarda. 95% by, 704
18 Feb. 95% Dy, 54
21 Feb. 947% 6% ks
28 Feb. 887 WA 69
73) Wize 36% 63% 180
24 Mar. (night) 742 26% al
RADE 497 Sylyé 67
12-28 June 1969 20% 80% ALAA

(hand caught and
banded)

*Plumage white or mostly white.

**Plumage essentially gray-brown.

About the time the youngest birds reach maximum numbers on Sand
Island, total numbers of roosting birds begin to decline, apparently
as the adults return northward for their breeding season, and many of
the older subadults follow. Throughout the remainder of the year,
the approximately l-year-old birds make up most of the non—-breeding
population, becoming, in late fall, the second year, white-breasted
birds that make up the bulk of the population until the next crop of
young arrives in late winter.

Where these l-year-old-plus birds go in the summer and autumn,
when the Johnston population is at its lowest, is uncertain. Many may
return to the more northerly islands, but many more may remain at sea,
possibly somewhere between Johnston and the northern islands, fishing
in an area too far from land for them to return regularly.

240

Unfortunately, it is not possible to determine accurately the
ratio of fully adult birds in the visiting population. It reaches a
peak in mid-winter, but at this time it is difficult to distinguish
these from the older subadults, which reach their maximum numbers at
the same time. Fully mature birds probably never exceed 50 percent
of the roosting population.

The movement pattern observed probably is correlated with feeding
conditions at sea in the entire region. Food must be abundant within
a day's flight from Johnston during mid-winter, but decreases through
the spring, when heaviest demands by nearly all species are placed
upon it. Minimal numbers of all species occur on Johnston in the
fall, and food must be scareestat this time. The sudden increase in
Red-footed Boobies in late fall occurs at about the same time as
Sooty Terns begin swirling near the islands, which would indicate
that the food resources for these two species are either the same or
at least follow similar cycles of abundance.

The effects of the guywires on Sand Island in drawing unusually
large numbers of Red-footed Boobies was discussed earlier. Whether
‘his causes an unnatural concentration of feeding effort by red-foots
is not known. It may be that the birds fed in the vicinity of
Johnston before the wires were present, but did not bother to come
in to the island for lack of suitable roosting places.

Day-to-day numbers of roosting red-foots vary inversely with wind
velocity and directly with cloud cover. Winds are more effective in
determining numbers than cloud cover, and the effects of cloud cover
are obscured by the more dramatic influences of winds. During periods
of highest wind velocity, daytime numbers may drop to a few adults
sitting on the more protected roosts on nesting sites, and nighttime
roosting birds may number in the low hundreds. By contrast, when
winds are light, daytime numbers of roosting birds may exceed the
nighttime numbers during high winds, and nighttime numbers may range
up to several thousand.

Reasons for the observed effects of clouds and winds on booby
numbers are not well understood, but probably relate to energy re-
quirements for flying and feeding. Energy output necessary for the
birds to feed during calm, cloudy weather may be excessive, if they
depend to any great extent upon upward deflected currents from wave-
tops and thermal convection currents for sustaining flight while
feeding. After several successive days of calm winds numbers begin
to drop, which may indicate that the birds need to feed and leave to
do so in spite of less than optimum flying conditions. It is also
possible that the birds simply have difficulty roosting on the wires
during high winds and find it: easier to stay at sea than to return
to the island. During storms with extremely high winds and heavy
rains, large numbers of red-foots roost on the ground.

During each day, numbers of roosting red-foots on the island
vary from a minimum at mid-day to a maximum well after dark which may

241

be several times the minimum. The increase begins in late afternoon
and reaches its maximum rate at about sunset, when flocks of up to a
few dozen birds stream in from the sea to find roosting places on the
outer guywires. Birds begin leaving before dawn, but fair numbers re-
main on the wires until an hour or more after sunup, especially on
days with moderate or light winds. During this early morning period,
large numbers of red-foots soar over the east hill, riding the upward
deflected trade-winds. These birds often resettle for part of the
day on the ground along the south shore and on the southwest rocks.

Turnover rates of the non-breeding birds probably is fairly slow.
Although the number of birds handled was small in relation to the
total numbers present, streamered birds remained conspicuous in the
population for at least several weeks after periods of major banding
effort. It would appear that during any particular season, birds
return to the island at least every few nights, and many may return
nightly.

Specimens

Appendix Table 7 presents data on the 29 specimens collected from
Johnston Atoll. Of these 29, eight were collected by Wetmore in 1923
and 21 were collected by POBSP personnel; all are located in the USNM.
This is a new specimen record.

Banding and Interisland Movement

In all, 1,047 Red-footed Boobies--180 adults, 844 subadults, and
23 young--were banded at Sand Island through the end of 1973 (Tables
24 and 57).

No interisland recaptures of Johnston-banded birds have been of
known Johnston breeders or young hatched on Johnston. This is not
surprising, for the Johnston area attracts hordes of birds from the
Hawaiian Islands during the non-breeding season, presumably because
food conditions are more favorable around Johnston, and it would be
unusual if the residents regularly went elsewhere for food at that
time. Only two known-age, banded birds hatched on Johnston have been
caught back on the atoll during their second year, and none during
their third year. But if the few young hatched there roosted on the
guywires with the thousands of others the same age, their chances of
being caught would be small. Nearly all the known-age birds caught
in their fourth year, or older, were nesting. Thus, it is doubtful if
any of the breeding birds, or their young, regularly leave the Johnston
area during the non-breeding season.

Conversely, 281 red-foots have made interisland movements between
Johnston Atoll and the Hawaiian Islands (Table 26). A total of 102
banded as non-breeding birds on Johnston have been recaptured to the
north, while 179 banded elsewhere have been recaptured at Johnston.

242

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At-Sea Distribution

Red-footed Boobies were absent from the grid southwest of Johnston
from May through September (Table 21 and Fig. 78). This occurs from
the last of the egg-laying period through the whole of the chick and
fledging period on Johnston Atoll. Thus Johnston breeders are not using
the grid area for food during most of their breeding season. The in-
crease in birds from October through February occurs both in the grid
and on Johnston. On the Atoll, however, numbers remained constant
from February through at least April, whereas in the grid at this time
there was a rapid decrease in numbers.

Three orange-streamered birds from Johnston were observed within
the grid, two in December and one in January. Both of the December
birds were subadults. On 30 January 1966, a banded immature from French
Frigate Shoals was collected a few miles northeast of the grid. Since
84 percent of the grid population consisted of immature or subadult

30

BIRDS PER 100 SQUARE MILES

J F Mea M J J A S 0 N D

Figure 78. Diurnal occurrence of Red-footed Boobies at sea 175 miles southwest
of Johnston Atoll, 1963-1967.

244

individuals it would appear that this area is composed mainly of highly
mobile birds from both Johnston Atoll and the Hawaiian Islands (POBSP,
1967a).

Further analysis of the at-sea distribution of Red-footed Boobies
is being conducted by Brian A. Harrington (in prep.).

GREAT FRIGATEBIRD Fregata mtnor
Status

Common breeding species, as well as a common, transient visitor;
breeders present year-round but most active October through August;
transients present throughout the year but most common in fall. Nests
built of Trtbulus on the ground or in grass clumps at Sand Island;
previously nested at Johnston Island. Transient visitors roost on
antenna guywires, and other raised sites. During POBSP studies several
hundred pairs nested yearly, with up to 113 fledging; transients num-
bered up to 1,000 or more.

Ecological Distribution

Akau Island: POBSP personnel recorded Great Frigatebirds occasionally
perching on towers here during the late 1960's.

Johnston Island: On 15 March 1859 while on Johnston Island, Brooke
(ms.) reported: '"...every now and then frigate birds would sweep with
a roar over head. Perched on some fragments of coral were some frigate
birds, some with flaming red pouches."

Wetmore (ms. a) found 800 here in July 1923 and described the colony
as follows.

Abundant. A few males are pairing and exhibit the inflated
gular sac. Some few have eggs but the majority have young
already on the wing. Birds able to fly are in many cases

still fed by the parent. As the young bird sees the adult
approach it spreads its wings and utters a begging call. The
parent after circling past several times alights on the ground
and opening the mouth regurgitates, while the young bird
plunges its bill down the parent's throat to seize the food.
After feeding, the old bird flies out over the water and swoops
down to dip its bill and wash out its open mouth. A flock of the
birds may be attracted easily by throwing small fish in the air
when the great frigates swoop and circle about, often seizing
the fish in the air. -When one secures such a morsel, others
pursue it and frequently snatch the fish from its bill. It
usually happens that they wrangle until the fish falls into the
water and is lost.

I see them pursuing terns, Wedge-tailed Shearwaters, and boobies,
often driving the unfortunates down into the water.

245

At nightfall they gather in flocks to roost on the eroded
ledges on the peach. They sleep with the head turned on
the back but often spend an hour or more after dark in the
care of their plumage. The adult female has a dull red rim
around the eyelid. In the male this is black.

Many are now molting extensively so that the beach is
scattered with their feathers.

The birds remain active at night and are continually overhead.

Four were seen in May 1945 (Clark, 1945 a and b); 100 were present
in November 1949 but only a few in December 1949 (Jensen, 1949).

Some tried to nest in bushes along the south side of the runway
near the northeast end in 1962, but construction work eliminated the
bushes (Willis G. Morris, pers. comm.). POBSP personnel occasionally
recorded Great Frigatebirds flying over Johnston Island in the 1960's,
but no nesting attempts were observed.

Although the population in 1923 was almost entirely on Johnston
Island, it has been exclusively on Sand since 1962, and it is unlikely
that successful breeding occurred on Johnston for several years before
that--perhaps not since before World War II.

Unfortunately, none of the old photos from Johnston Island shows
frigatebird nests, but several show young birds sitting on the ground
between Lepturus clumps. These all appear to be either along the top
of the bank above the shoreline, or near the top of the 40-foot hill.
These sites probably are those from which the adults can most readily
take flight, since they would have difficulty rising from level ground,
especially from between clumps of Lepturus.

Sand Island: Wetmore (ms. a and b) observed 100 here in July 1923.
Moynihan (1957: 36) in April 1957 recorded the following.

Frigate birds were the second most abundant species on
Sand Island. There were at least 900 adult birds nesting
on the girders of four overturned and partly collapsed
radio towers on the western islet. A few of these were
still incubating but most of them were sitting on chicks,
some of which were half grown. There were also many birds
in juvenal plumage flying overhead or resting on the same
four towers, another collapsed tower a few yards away, and
a few low bushes on the same islet. I would guess that
the total number of Frigatebirds in the area was not less
than 2000. (It might be noted, incidentally, that none of
them ever landed on the bushes, trees, or houses of the
eastern islet.)

POBSP personnel found Great Frigatebirds as common nesters and
transients from 1963 through 1969. They were also present in November
JS /SXe

246

Nesting areas used by Great Frigatebirds on Sand Island during
POBSP studies are shown in Figures 44 and 79. Reasons for changes
from site to site are not clear, but probably the knoll site was
preferred. Intensive disturbance there by POBSP personnel in 1963-
1964 may have caused the move to the south shore. Why more birds did
not nest on the southwest islet, which presents what appears to be the
best raised sites on the island, is not clear. Possibly competition
with Brown Boobies nesting there is responsible.

During POBSP studies, a few Great Frigatebirds attempted to nest
on navigational aids and guywire bases in the lagoon, but the few eggs
laid were lost before hatching.

Because of the almost total lack of bushes, at least in the areas
that are far enough from human disturbance for frigates to be attracted
to them, the birds on Sand Island are forced to nest entirely on the
ground (Figs. 80 and 81). Since the original vegetation of the atoll
contained no shrubs or trees, frigates have never had anything but the
ground or Lepturus clumps on which to nest. Exceptions to this occurred
during the late 1950's after the Navy abandoned Sand Island when frigates
nested on overturned towers and during the early 1960's when they nested
on bushes on Johnston Island. That the Johnston Atoll birds would
readily nest on raised sites is shown by their use of these girders and
towers in 1957 (Moynihan, 1957), and by their attempts to use the
bushes on Johnston in 1962.

Figure 79, Great Frigatebird nesting areas, Sand Island, Johnston Atoll,

1963-1969.

247

A -. 12S eee . Cae
Figure 80. Great Frigatebirds nesting on ground on crest of hill east of
transmitter building, Sand Island, Johnston Atoll, May or June
1964. Sooty Tern adults and chicks surround the colony (POBSP
photo by A. B. Amerson, Jr.).

Ae sis

Figure 81. Great Frigatebird colony on south shore, Sand Island, Johnston
Atoll, 26 February 1969 (POBSP photo by P. C. Shelton).

248

In late 1966 and 1967 nests were built entirely of Trtbulus vines
picked up after Red-footed Boobies had broken up the clumps in their
efforts to gather nesting material. The red-foots landed among the
vines and pulled them apart, while the frigates gathered the material
on the wing, swooping down to pick up loose pieces left by the boobies.
Nest material often was stolen from unattended Red-footed Booby nests,
and was even stolen from red-foots carrying material, much like food
material was stolen. Usually only male frigates gathered nest material,
but at least once (26 February 1957) a female was seen gathering
Trtbulus.

Because of scarcity of material, many nests almost or completely
lacked plant material. The nests of late breeders in the inland colony
of 1967 were nothing more than depressions in the ground and in several
cases, the egg was laid in the middle of Lepturus clumps, which raised
the egg a few inches off the ground. By the time these late birds
nested, Jrtbulus vines were practically non-existent because of removal
by earlier nesting boobies and frigates. Boobies by this time had
switched to Boerhavia for their nests; apparently the frigates made no
attempt to use Boerhavia.

Non-breeding birds roost mainly on the guywires of the LORAN-C
antenna, and in several other locations scattered around the island and
lagoon (Fig. 82) which afford raised sites from which the birds can
become airborne readily. Except for the guywires, most of these sites
are used mainly during the day, and some only at low tide (North
Antenna Rocks). On the guywires, frigates roost on the anchors or just
above the anchors on the wires, or on the steeper parts of the wires,
above the Red-footed Boobies, which apparently are able to drive the
frigates away from the lower perches. In late November 1966, Schreiber
noted that great frigates usually came in to the wires earliest in the
evening and frequently occupied the lower portions of the wires. But
later Red-footed Boobies drove the frigates higher up the wires. In all
cases of conflict observed, the frigate moved. Wires extending from the
tower at right angles to the wind were most heavily used. Thus those
north and south of the tower, including the northeast and southeast
inner wires, were usually most heavily used by both frigates and boobies.

Because they are larger and space themselves on the wires more
widely, and are located higher on the wires, frigates could be counted
in poor light much more accurately than Red-footed Boobies.

Two distinct behavior patterns occurred: sunning, and soaring.
In the sunning behavior, the birds assumed ridiculous-looking positions
with the wings outstretched and turned underside up, or with the body
drooped across the wires as if they were dead. Usually they had their
bills open, probably to help-in cooling. Other aspects of the sunning
postures, especially turning the underwings up, may have been related
to the occurrence of parasites.

Soaring occurred on days with light winds and strong sunshine
causing an updraft to form by air rising from the heated surface of

249

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250

the island. In normal (12-15 mph) or stronger winds, this updraft was
either broken up or blown away from the island at such an angle that

it was not usable by the birds. But with light winds the column
drifted slowly away from the island, and it was invariably marked by
several dozen frigates (and often other species as well, including
Sooty Terns) circling motionless in the column, rising at least as high
as the cloud bases, 1,500 to 2,000 feet up. Frigates could be observed
flying (flapping) slowly from the roosting areas to this column of
rising air, setting their wings and circling upward nearly to the cloud
bases.

During normal trade winds, frigates spent a lot of time soaring
over the east hill, where air forced over the rise afforded ample
upward current to keep the birds aloft indefinitely.

Populations

Like Red-footed Boobies, Great Frigatebirds were difficult to
census because most of them, especially non-breeders, roosted on the
outer guywires and reach maximum numbers only after dark. Because they
were larger, less numerous, and a greater proportion roosted on the
ground, counts of frigates were somewhat more accurate than for red-
foots, perhaps within a 10 percent margin of error.

Figure 83 shows means and extremes of POBSP semimonthly estimates
of Great Frigatebirds on Sand Island, based primarily on roost counts
made at late dusk or at night by moonlight. These figures represent
maximum numbers present at one time during the period, rather than total
birds using the atoll. There was no reliable basis for estimating the
turnover rates in the roosting population, but it would be surprising
if less than two or three times the maximum numbers recorded used the
atoll each year.

In general there was much less seasonal fluctuation in Great
Frigatebird numbers than for most other species on the atoll. The high
of about 1,000 which occurs in early fall was only about twice the low
of 400 to 500 which occurs during the winter. The graph of means of
population estimates (Fig. 83) indicates two lows, one in December and
January and the other in June and July, with broad highs between. The
spring maximum appears to correlate with the breeding season on Johnston,
while the fall maximum may be caused by an influx of transients.

Small period-to-period fluctuations in many cases resulted from
differences in estimating techniques, or of inadequate counting. The _
number of birds present at any given time depends to a large extent upon
the time of day, wind velocity, and cloud cover, and unless several
counts are made during a semimonthly period, the maximum number of birds
present during that period may be poorly estimated. Highest counts
were made at night by moonlight when winds were minimal, but such
conditions often did not occur during a given period. Use of a spot-
light for night counts was not practical because the birds flushed when
the light was turned on them.

7g5)AL

2,000
Adults 2

—— Means e--6
1,500

eeravere Extremes

1,000

NUMBERS

500

— Eggs, 1963-1968
eiatelets Young, 1963-1968

—-— Young, 1969

NUMBERS

J F M A Nitiamere) J A F 0 N D

Figure 83. Means and extremes of semimonthly estimates of Great Frigatebird
numbers, Sand Island, Johnston Atoll, 1963-1969.

252

Birds present in the breeding colonies varied from a low of a few
to a few dozen birds present during late summer, increasing throughout
the fall and winter, and spring as courtship activity increased, to a
maximum of 250 to 350 during the period January through May. Before
egg laying began, the number of birds decreased at night in the nesting
areas, possibly because many of the birds roosted on the guywires, but
later, the number remained constant day and night.

An exception to the above generality occurred in February, 1969,
during the weather disturbance described elsewhere, when numbers in the
colony fell from about 275 birds before the disturbance began to 48 birds
on 18 February. This probably represented only those birds on eggs, but
this could not be determined without disturbing the birds. By the end
of February there were again over 250 birds in the colony each day.

There appears to have been some reduction in numbers between the
times of the pre-POBSP estimates and the mid-1960's. Wetmore's estimate
of nearly 1,000 birds in July 1923 is nearly twice the mean for July
during POBSP studies, and Moynihan's estimate of 2,000 birds in April
1957 is nearly three times the April mean from POBSP figures. Although
large numbers of great frigates roost on the LORAN-C guywires, their
numbers did not appear to increase in response to the availability of
these roosts as did Red-footed Boobies. Reasons for a decline in
frigatebird numbers are not clear. Perhaps recruitment to the adult
population was below normal because of missing age classes from years
of intensive disturbance.

Productivity: Table 58 and Figure 83 list available data on
productivity of Great Frigatebirds on Sand Island during POBSP studies.
The spectacular increase in chicks fledged in 1969 resulted from our
discontinuing all egg and chick counts in that year.

Detailed nesting studies were initiated at the beginning of the 1963-
1964 nesting season, when eggs were counted daily in the nesting colonies.
By mid-April 1964 it had become obvious that the daily disturbance was
causing inordinately high losses of eggs and small chicks (mean fledglings
for 1963 and 1964 = 9). At this time daily counts were discontinued, but
eggs and chicks were counted semimonthly for the remainder of that
season and throughout the nesting seasons of 1965 through 1968. Con-
tinued low productivity continued (mean fledglings for 1965 to 1968 = 26),
however. This prompted Vernon M. Kleen to rope off the south shore
nesting colony in late 1968. Throughout the nesting season of 1968-1969
we stayed far enough away (usually 20 to 30 feet) from the nesting
frigates so that birds on eggs would not flush. Civilian and military
personnel using the island cooperated excellently in this attempt to
minimize disturbance. There were no more birds in the nesting colony
in 1969 than in previous years, and as far as is known, no other factor
changed in 1969 that could account for the increased survival of chicks.
Productivity increased, resulting in 113 chicks fledged. Perhaps the
increase would have been still more spectacular had not the weather
disruption in February reduced the number of birds in the colony to about

725%)

a fifth the usual number for one day. It is not known whether any eggs
were abandoned during this period, but only one could be seen unattended
in a photograph taken of the colony on 18 February, when birds were at
their minimum.

Disturbance to nesting great frigates results in losses of both
eggs and chicks. Adults incubating eggs or brooding small chicks
apparently work the egg or chick up against their abdomen between their
feet in such a way that the egg or chick is carried a few feet before
falling free when the adult flushes. Even if not broken or killed, an
egg or chick so treated is unlikely to be retrieved by the parent.

In addition, small chicks left on the nest when the adult is flushed
may be eaten by other frigates before the parent returns.

Table 58. Productivity of Great Frigatebirds, Sand Island, Johnston
Atoll, 1963-1967*

Colony Area

Knoll 50. shore Islet Inland Total
Year Eggs Young Eggs Young Eggs Young Eggs Young Eggs Young
1963 ? 1k 0 0 0 0 0 0 u iat
1964 400 6 0 0 61 0 0 0 460 6
1965 60 6°? 300 29224 40 1 0 0 (400)** 31
1966 0 OD ZoO. a5 5 0 0 0 (255) 15
1967 0 Oe 250) 6 0 0 s0e 710 (300) 26***

*Numbers of eggs counted daily only in 1964, and then only through
part of season; number of young is number banded and not known to
be lost within 90 days.

**Numbers in parentheses are approximate.

***Observations not made after mid-September; probably several of these
were lost shortly after fledging. Most or all may have been killed
by USCG personnel.

After the chicks become large enough to not be eaten or easily
dislodged from their nests, mortality was relatively low until they
fledged. At the time of fledging, mortality again became high, mostly
because the inexperienced birds hit guywires, breaking their wings.
Almost all the dead birds (destroyed and recovered) listed in Table
59 were birds that hit guywires.

254

If human disturbance continues, the breeding population will no
doubt decline, but if disturbance is minimized the breeding population
will probably increase.

Table 59. Fate of young Great Frigatebirds banded on Sand Island,
Johnston Atoll, 1963-1967

Total

No. Tot. possibly
Year ____ Banded _ Dest. _ Recov. dead Rep._Ret._No Data” alives
1963 15 4 4 8 0 0 7 i
1964 6 0 il ik 2 0 3 5
1965 33 0 3 3) 4 10 16 30
1966 19 4 2 6 0 3 10 13}
1967 Dil 1 0 at mec 0 26 26%
Total 100 9 10 19 6 13 62 81
*Many of these killed by USCG personnel.
*kDest. = Destroyed, Recov. = Recovered, Rep. = Repeat, Ret. = Return.

Food Habits: Little is known of the feeding habits of Johnston
Atoll frigatebirds except that they do commonly pirate food from Red-
footed Boobies and Wedge-tailed Shearwaters, occasionally from Sooty
Terns, as well as from one another, and rarely from Brown Noddies.

No quantitative data as to the amount or proportion of food taken in
this way are available.

An interesting aspect of food habits is the total lack of evidence
that great frigates prey upon Sooty Tern chicks at Johnston Atoll, as
has been noted from French Frigate Shoals (Amerson, 1971). The terns
nest among the nesting frigates, and the young terns hatch during the
time frigates are feeding their own young, which one would expect to be
a time when any available food would be used. In 1967, Shelton even
found several Sooty Tern chicks huddled under incubating and roosting
frigatebirds near the south shore of Sand Island.

Annual Cycle

Breeders: The breeding season of Great Frigatebirds (Table 60
and Figs. 37 and,83) is one of the longest of any species on Johnston
Atoll, both for individual birds and the colony as a whole. Normally
seven to eight months elapse from the time a pair of adults begin
displaying and nest building until their chick fledges, if they are
successful. This includes a month of prelaying activity, about eight

255

weeks of incubation (incubation period for one egg observed in 1964

was 56 days), and four or five months for growth of the chick. Although
no positive evidence is available, birds that lose their first egg may
relay, which would extend the period from displaying to fledging by at
least another month.

Table 60. Extreme dates of significant events in the Great Frigatebird
breeding cycles, Sand Island, Johnston Atoll, 1963-1969

Earliest
displaying

Year males Laying Hatching Fledging
1963- & 27 Nov.-? 1-15 Mar.-? (1-15 Oct.—
1964 1-15 Nov.)*
£964— 1-15 Oct. 1-15 Nov.- 16-31 Mar.- 16-31 Aug.-
1965 1-15 May 1-15 July 1-15 Dec.?

E. hill:(1-15 Apr.- (1-15 June- 16-31 Oct.-

1-15 May) 1-15 Aug.) 1-15 Dec.

1965- ? 1=15 Nov.- 18 Jan- 1-15 Aug.-
1966 early May 16-30 June 16-31 Dec.
1966- 2 4 Dec.—May? 23-28 Feb.- 16-31 Aug.-?
1967 16-31 May

Inland: (8 Apr.-May?) 16-31 May- ?

16-31 July

1967- 2 ?—-May? 9-16 Apr.- 27 Aug.-
1968 1-7 July late Oct.
1968- ? 24 Nov.-? by 32) Mar .= late Aug.-?
1969 June

*Dates in parentheses are approximate.

Neither Wetmore's 1923 observations nor those of Moynihan in 1957
indicate significant variations in the nesting cycle from the pattern
observed in the 1960's. During POBSP studies, the main colonies stayed
on the same schedule year after year, but smaller groups varied from the
larger ones by as much as three months.

Although an occasional male with an inflated gular pouch may be
seen at any time of year, the first marked increase in displaying birds
usually occurred in October. A few eggs were laid in early November of

256

some years but there was not a significant number of eggs present until
January. Displaying birds were common throughout the winter, and in
1965 and 1967 new nesting areas were occupied in early March, five
months after the first nesters began their activities.

In 1963 and 1964 the earliest and largest nesting colony was on
the east hill; in 1965 and subsequent years it was on the south shore.
In mid-March 1965, birds began displaying in the 1963-1964 nesting area
on the east hill, and by early April they had begun egg laying there,
three months after the larger colony on the south shore began laying.

This east hill colony was not occupied in 1966 or 1967, but in
March 1967 a secondary colony was established inland from the main south
shore group, and these birds followed about the same chronology as the
east hill colony in 1965.

Unfortunately, individual birds were not traced from colony to
colony and year to year because banding and recapture operations were
too disruptive to the colonies.

The chronology of nesting on the southwest islet in 1964, 1965,
and 1966 closely followed that of the main colonies in those years.

Most hatching occurred in March and April in the main colonies,
and in May and June in the late colonies of 1965 and 1967. Chicks in
the main colonies were flying by late August, four to five months after
hatching; young in the late colonies remained on the ground until
December, after the first eggs of the next season had been laid in the
larger colonies.

When the young become fully independent is not known, but birds up
to a year old were seen being fed by an adult. If heavy or full depen-
dence of young on their parents extends for more than a few months after
fledging, it is extremely unlikely that these parents could breed again
in the year after successfully raising a chick.

At least some of the young birds continue to use the atoll for
several years. In 1967, several of the 1966 young were seen regularly,
mostly roosting on pilings off the south and east shores. At least one
1965 bird (737-44933) was seen regularly near its hatching site on the
east hill until early 1967. A 1965 bird (737-44932) was photographed
while it roosted on a guywire in April 1967, and a 1963 bird (737-44213)
was found partly decomposed (skeleton saved) on the east beach in March
1967 (see also Banding and Interisland Movement section).

The age at which the young begin breeding is not known, but it
certainly is over five years. Known age five-year-old birds do not appear
fully adult.

Non-breeders: The fall maximum in Great Frigatebird numbers
(see Fig. 83) appears to be caused by an influx of transients, probably

257

birds moving away from the northwestern Hawaiian Islands after the end
of their breeding season. See also the Banding and Interisland Move-
ment section for further details about transients.

Numbers of birds both on the wires and on terrestrial roosts
varied inversely with wind velocity. Also, under any given wind
velocity, there was a two-to-four fold increase in numbers during the
night over those present during the day. Most of the increase occurred
in the evening, and the birds thin out again in the early morning.
During days of high winds there may be no birds on the wires, and only
a few scattered individuals on the more solid roosts, while on days
with little or no wind, up to 500 birds may be found on the wires and
other roosts.

Table 61 shows several samples of ratios of the various recogniz-—
able sex and plumage categories at various locations on the island.
The most serious shortcomings of these data are that they are too few,
do not represent all the roosting locations on the atoll, and do not
cover all possible times. Birds on the outer set of guywires were
too far from shore to properly distinguish the plumage categories and no
counts could be made at night.

There is insufficient evidence to conclude that the sex ratio of
adults departs substantially from a 50-50 ratio.

The outstanding fact emerging from this set of figures is the
preponderance of subadults and immatures in roosting areas (including
all except the one count of 23 March 1967 in the breeding colony on the
south shore). Since so few young are produced each year on Johnston
Atoll, it is obvious that most of those roosting there are from other
islands; presumably all are from the northwestern Hawaiian Islands,
as indicated by the banding records.

What percentage of the adults roosting on the atoll are also from
other islands is unknown, but there are several records of adults
moving between the islands (see Banding and Interisland Movements).

Specimens

In all, 67 specimens of Great Frigatebirds have been collected
at Johnston Atoll. Data for these are presented in Appendix Table 7.
This constitutes a new specimen record.

Banding and Interisland Movement

From 1963 through 1969, POBSP personnel banded 997 Great Frigate-
birds at Sand Island; Amerson banded eight in 1973 (Table 24). Of
these 1,005, 348 have been recaptured back on the atoll (Table 25),
and 14 have been recaptured elsewhere (Table 26). In addition, 29
birds originally banded on other islands were captured at Johnston.

258

Table 61. Plumage ratios of Great Frigatebirds, Sand Island, Johnston

Atoll

Date and Sub-
time Location AM AF AU adult* Imm.** No.
24 Sept. 66 NE & SE
1855-1910 hrs. inner wires 20 23) 36 22 165
29 Jan. 6/7 SE outer
Evening wires Da By) 27 163
Noon SW islet 55 45 I
23 aMareaeoy/ SW islet 32 15 52 40
1435
1530 So. shore

(breeding colony) , 88 3 9 353
1630 SE inner wire 37) 26 37 38
30 Apr. 67 NE and SE
1920-1940 inner wires 33 20*** 9 38 74
24 Feb. 69 SE inner wire
1830 3 Le 43
29 May 69
1430 So. shore 258 il il 260

Inner wires &

ground except

So. shore 95 0 5 100

Outer wires 76 0 8 84
6 Sept.
1945 NE inner wires 2 225) 77

*Head mostly white, mottled with black

*kHead white or white and red.
4

**k*kFive birds had gray breasts.

The interisland birds (Table 26) show that most Great Frigatebird
movements, if not all, involving Johnston are to and from the northwestern

259

Hawaiian Islands. In all, 23 from French Frigate Shoals, the closest
island to Johnston, have been captured at Johnston. Curiously, six of
these 23 were banded as chicks on the same day and same island:

13 June 1963, Whale-Skate Island. It is also noteworthy that two of
these six birds were later retaken on their home island, and that an
adult banded on Pearl and Hermes Reef made a similar round trip to
Johnston and back to its original site. Three others from Pearl and
Hermes Reef were also captured at Johnston. The 15 Johnston birds
captured elsewhere were taken at: French Frigate Shoals (six birds),
Kure Atoll (four), Marshall Islands (four) and Phillipine Islands (one).

At-Sea Distribution

Low grid densities for Great Frigatebirds from May through August
(Fig. 84) correspond to the major breeding season on Johnston Atoll,
while a somewhat higher density is maintained in the grid during the
non-breeding season (November to March). The complete absence of
frigatebirds in June and July may in part be due to the abundance of
other bird species, especially boobies, near Johnston on which frigate-
birds can parasitize (food robbing) without traveling all the way to
the grid (POBSP, 1967a).

BIRDS PER 100 SQUARE MILES

J F M OA M J J A S O N D

Figure 84, Diurnal occurrence of Great Frigatebirds at sea 175 miles southwest
of Johnston Atoll, 1963-1967.

260

LESSER FRIGATEBIRD Fregata artel
Status

Accidental; at least three sight records from Sand Island.
Observations

Harrington of the POBSP identified three adult male Lesser
Frigatebirds over Sand Island on 31 March 1968. Eight others seen at
the same time were thought to be this species, but were not observed
well enough for positive identification. Shelton saw a frigate on
9 August 1969 that probably was an immature Lesser Frigatebird.

Lesser Frigatebirds breed on several islands in the Phoenix,
Line, and Gilbert groups, and normally disperse westward across the
western tropical Pacific (Sibley and Clapp, 1967; Amerson, 1969).
Recent POBSP records from Kure (Woodward, 1972) and Wake Islands have
extended the known non-breeding range in the central Pacific north-
westward.

Specimens

These sightings constitute a new sight record for Johnston
Atoll; no specimen record exists.

CATTLE EGRET Bubuleus tbts
Status

Straggler; six sight records on Sand Island; one sight record
on Johnston Island.

Observations

Four Cattle Egrets were observed by the POBSP at Sand Island; one
from 3 to 12 December 1966, one from 3 to 20 December 1966, one from
2 August to November 1967, one from 11 August 1968 to 29 May 1969.
Kridler (BSFW, 1973) reported two on Sand Island in May 1973.

The first two POBSP birds appeared together following a night of
strong southerly winds. One appeared weakened, with drooping wings,
and disappeared after several days. The other remained until
Schreiber scared it away in attempting to catch it. The other two
birds appeared in August a year apart, and each remained for several
months before disappearing for unknown reasons. No attempt was made
to catch these birds, and unnecessary disturbance was avoided.

All four POBSP birds spent most of their time on Sand; no other
part of the atoll supported sufficient populations of insects and mice

261

to sustain them. The heavily grassed areas on Sand were their primary
feeding places. The 1966 birds fed in the vegetation around the Coast
Guard barracks, but the 1967 and later birds were never seen on the
man-made end of the island. No shallow water feeding was seen. Most
of their food must have been insects, but several times the egrets were
seen eating mice.

During the daytime the egrets frequently roosted on raised perches,
including the old dock, a sign near the dock, the fence around the LORAN-
C transmitter, and an old triangulation stand northeast of the transmitter.
At night, and also for extended periods during daylight, the birds roosted
in sheltered spots on the southwest islet or along the south shore in the
frigate colony. In 1969 the egret often appeared at dusk on the south-
east outer guywire with Red-footed Boobies and may have spent the night
there.

Mobbing by other birds, particularly Brown Noddies, Sooty Terns,
and more rarely boobies, was fairly intense when the two August birds
first arrived. Any time one of the egrets flew over the island, it
was followed by up to a few dozen noddies and terns. No physical con-
tact was made, and the mobbing did not appear greatly to disturb the
egrets. However, the bird that arrived in August 1967 spent most of
its time on Johnston Island for the first week, possibly because of
intense mobbing by terns. After a few days the mobbing action waned,
although it occasionally occurred during the day as long as there were
noddies or sooties on the island. The 1966 birds were not bothered
because virtually no terns were present during the day in December.

In November 1968, when a Peregrine Falcon began roosting on the
LORAN-C tower, the egret avoided Sand Island much of the time. Where
it stayed was not discovered.

All four POBSP egrets were probably adults when they came to the
atoll. All had yellow bills, which, especially in the case of the two
August arrivals, probably means they could not have been hatched the
same year in which they came to the island. The December birds could
have been birds of that year which had just acquired the yellow bills.
The only bird present during the spring acquired full breeding plumage
by late March, with conspicuous buff on the crown and middle of the
back and legs yellow to the feet. The bill and legs did not, however,
turn noticeably reddish or orangish.

The closest breeding population of Cattle Egrets to Johnston Atoll
is that which was introduced to Oahu in 1958. There is no evidence
whether the Johnston birds came from there or from the eastern Pacific.
A Cattle Egret has also been recorded from French Frigate Shoals
(Amerson, 1971).

Specimens

These sightings constitute a new sight record for Johnston Atoll.

262

PINTAIL Anas acuta
Status

Regular fall migrant; four specimen records and many sight
records from Johnston and Sand Islands.

Observations

Pintails were reported at least once each fall during POBSP
studies except 1964. There were two spring records, and one bird
stayed over the winter of 1966-1967 (Tables 20 and 62).

The largest influx of Pintails occurred in late October and early
November 1966 when at least 30 arrived on the atoll. About a third of
these were found dead, probably from exhaustion, and at least one was
struck by an aircraft. The favored habitat on Johnston Island, where
most of the birds were found, was rainpools along the runway. On Sand
they most often stayed on the grassy west side of the original island.
A few birds were seen in the lagoon near the islands.

Healthy birds usually departed a few days after arriving, but one
female stayed on Sand Island from December 1966 until she died on
13 February 1967. She appeared normal and was fully capable of flight
until a few hours before she died, when she was found suffering symptoms
resembling those of botulism--greatly weakened, neck unable to support
her head. She frequently fed on insects around piles of Sooty Terns
killed on the guywires and may have been poisoned by a micro-organism
or the product of a micro-organism picked up from the carcasses.

Specimens

The four Pintail specimens collected on Johnston Atoll by the
POBSP are listed in Appendix Table 7 and constitute a new specimen
record. Three of these had notched, juvenal rectrices, indicating
that they were still in their first year. The fourth, the female
taken in February 1967, would have acquired adult rectrices by the
time she was collected, even if she had hatched the preceding year.
Her oviduct was uncoiled, indicating that she had not bred, and
therefore was probably less than a year old. A bird banded 31 August
1969 also had juvenal rectrices.

One of the males was well advanced in acquiring adult plumage,
with many vermiculate body feathers, and well developed though still
diffuse head and neck pattern. The other male, taken at nearly the
same time of year, had only a few vermiculate feathers and no hint of
the adult head pattern. The most striking characteristic of this
bird was the bright rust color or virtually all the juvenal feathers.
The few new feathers appeared to be normally colored.

263

Banding and Interisland Movement

U.S. Fish and Wildlife Service banding records include a Pintail
banded in California 12 September 1957 which was recovered 10 October
1958 on Johnston Atoll. Three Pintails were banded by the POBSP on
Johnston (Table 24), but none has as yet been recovered.

Pintails breed in the northern parts of North America, Europe, and
Asia, and winter south into the tropics, including Hawaii (AOU, 1957),
where they are the most abundant migrant duck from August to March
(Medeiros, 1958). Pintails are frequent migrants to several of the
northwestern Hawaiian Islands (Amerson, 1971) and the north Pacific
(POBSP, 1964).

Table 62. Observations* of Pintails on Johnston Atoll

Number

Date of Survey of Birds Age/Sex Remarks

ms5o7 £0 Oct. iL Banded 12 September, 1957, California.

1963" LS7Oct: 1 Seen swimming by seaplane ramp, Sand lI.

i965) 34 Mar. if M Described by Johnston I. resident;
dead, not saved.

lon Oct, 5 Flying over east end of Sand I.,
chased by Brown Noddies.

GOuOcE . ip Jy er Collected, Sand I.

i? OG dL F Banded and released, Johnston I.

LOROCE. 9 Flying low over Sand I. Banded bird
not present.

Z90cr. iL Juv. M Collected Johnston I. Wings clipped;
probably held captive before death.

1 Nov. 1 Sight record.

1966 SOGES 1 F S. shore, Sand I. (west end). Sight
record on Johnston same day, possibly
same bird.

5aOct. 2 F Flushed from grass, NW side of Sand I.
>) Reported from Johnston I.

6 Oct. 1 Reported hit by aircraft, Johnston I.

264

Table 62. (continued)

Number
Date of Survey of Birds Age/Sex Remarks
1966 (oreiee 1 M Captured Johnston I.; died, not saved.
14 OChs al Reported on Johnston I.
22 OG. 4 On runway south of snack bar, Johnston
Ibo
29 Oct. 3) On lagoon near Sand I.
2 F Rainpools near runway, Johnston I.
1-13 Nov. 2-9 Seen daily on Sand I.; maximum 9,
7 November. One juvenile male col-
lected 7 Nov.
1-15 Nov. 10+ Reported dead on Johnston I.
7 F Kept in pen, Johnston I. Captured
weakened.
5 Nov. 8 - Rainpools along runway, Johnston I.,
one male in full breeding plumage.
12 Nov. 5 - Along runway, Johnston I.
1-15 Dec. 12 Captured weakened, Johnston 1.3;
kept in pen and all but one probably
eaten (one still in pen through 1967).
i Dec.=
L967» 213" Feb: il AF Present on Sand, probably visited
Johnston also (see text). Died 13
February 1967.
21—29" Mar. il F Present on west side, original por-
tion of Sand I. Probably moved on.
23-26 ‘Oct. 1 - Present on Sand I., died, not saved.
1968 30 Sept.- 1 F? Caught and banded on Sand.
5S 70cGt.
SePOGite..= o+ = Present on Johnston. Possibly cap-
16 Nov. tured, and held until 1969 (see below).

12-13 Nov. 1 - Present on Sand for one day.

265

Table 62. (continued)

Number
Date of Survey of Birds Age/Sex Remarks
£969) “i. Jan.= 3 F In captivity on Johnston I., probably
14 Aug. caught in November 1968. Moved to
Sand I. ca. 16 February. Two died
in June, third was released, and
survived until 14 August.
19 Aug. if = Flew west past Sand I.
29) Aug. = i}: £ Regularly seen around tennis courts,
3..Sept. Sand I. Banded 31 August. Thin,
but healthy.
8 Sept. iL - Reported along runway, Johnston I.

*Al1l observations by POBSP except 10 October 1958 record.

AMERICAN WIGEON Anas [=Mareca] amertcana
Status

Uncommon fall visitor; two specimen records and several possible
sight records from Sand Island.

Observations

Two American Wigeons were collected by POBSP personnel on Sand
Island, the first on 23 October 1964 and the second on 29 October 1966.
Both were first year males, as determined by the presence of juvenal
rectrices. The 1966 bird had several vermiculated feathers on the
sides, and several purplish-pink breast feathers, but the 1964 bird
showed no signs of adult male plumage.

Two birds, probably of this species, were seen 15 October 1964,
and unidentified ducks seen 13 October 1964, 5 September 1964, and 15
November 1965 could have been American Wigeons.

American Wigeons breed inland throughout most of western North
America south to northeastern California, and winter south through Central
America (AOU, 1957). Small numbers regularly visit the main Hawaiian
Islands (Udvardy, 1961) and the POBSP has recorded several from the
northwestern Hawaiians (Clapp and Woodward, 1968).

Specimens

The two collected specimens noted above and in Appendix Table 7 are
the only verified records of this species for Johnston Atoll.

266

NORTHERN SHOVELER Anas [=Spatula] elypeata

Status

Uncommon fall visitor, one specimen record and at least one sight
record from Sand Island.

Observations

One adult female was collected 26 October 1964 on Sand Island by
POBSP personnel. A second record was a sighting by Kleen of a female
on Sand Island 22 September 1968. Another seen by Kleen on 31 October
1968 could have been of the same bird.

Shovelers regularly migrate to the Hawaiian Islands (Medeiros,
1958). They breed across northern North America, Europe, and Asia

and winter south from mid-temperate to tropical latitudes on these
continents and Africa (AOU, 1957).

Specimens

The collected female noted above and in Appendix Table 7 is the
first specimen record of a Shoveler for Johnston Atoll.
DOMESTIC CHICKEN Gallus gallus
Status

Introduced; probably breeds. Occurs in small numbers on Johnston
Island only.

Observations

According to island personnel, a few fighting chickens were
introduced to Johnston Island in early 1973. The cocks were reported
to be caged in November 1973, but at least two hens were said to be
running loose. It is not known if breeding occurs, but most likely

it does. The chickens came from the main Hawaiian Islands.

This is a new sight record.

PEREGRINE FALCON Faleo peregrinus tundrtus
Status

Accidentalg one specimen record from Sand Island, and a sight
record, most likely the same bird, from Johnston Island.

267

Observations

A "hawk" was reported on Johnston Island 8 November 1968 and on
12 November a Peregrine Falcon appeared on Sand Island, no doubt the
same bird. It remained on or near the atoll, returning at least every
few days to roost 50 to 80 feet up on the LORAN-C transmitter tower,
until 2 December when Kleen collected it.

During its stay on the island the Peregrine occasionally chased
Great Frigatebirds, but apparently with no intent to kill. It did
kill and eat young Brown Noddies and Wedge-tailed Shearwaters, and its
presence affected the behavior of several other species. For example,
the Cattle Egret, which had been seen regularly on Sand Island, dis-
appeared to other parts of the atoll for long periods while the
Peregrine was at large. Frigatebirds ceased roosting on the northeast
inner guywire of the LORAN-C antenna, and did not resume normal use of
the roosting area for one or two months after the Peregrine was shot.

Falco p. tundrius breeds on the arctic and subarctic tundra area
of North America and winters from the southern United States to
Argentina. Falco p. pealet breeds along the Pacific coast region to
North America between about 50°N to 58°N and on the Aleutian Islands.
A third race, F. p. japonensts breeds in northeastern Asia southward
to Korea. It winters in Japan and on various western Pacific Islands
(Vaurie, 1961; White, 1968).

A female identified by White as F’. p. pealet, was collected at
Kure Atoll on 7 March 1965 by POBSP personnel (Woodward, 1972). Sight
records are also known from Midway Atoll, Lisianski Island, and the
main Hawaiian Islands (Clapp and Woodward, 1968).

Specimens

The Johnston specimen, a female with heavy fat, was identified by
Clayton M. White (pers. comm.) as F. p. tundrtus on the basis of its
morphology, measurements, and overall appearance. It is a new specimen
record for the atoll. A male Peregrine Falcon collected at sea by
the POBSP on 8 November 1963 at 14°10'N, 171°42'W, 200 miles southwest
of Johnston, was provisionally assigned to this race by White (1968),
but both could just as well have come from Asia as from North America.

AMERICAN GOLDEN PLOVER Pluvitalts domtntea
Status
Common migrant; found singly or in small groups inland and on

beaches on all islands. Most abundant during spring, late summer, and
fall; a few individuals present throughout the year.

268

Ecological Distribution

Akau Island: POBSP personnel found American Golden Plover on
this man-made island from 1964 to 1969.

Johnston Island: Fennell (1948) saw a few "...at the end of the
mat...'' sometime in 1948.

POBSP personnel recorded them in the interior of the island
and on the beaches from 1963 through 1969. Amerson also found it
here in November 1973. American Golden Plovers, as well as other
shorebird species, were especially attracted to Johnston Island during
the 1964 construction period. Throughout the spring of that year
coral was dredged and deposited on the island, resulting in a rich
food source in the form of aquatic organisms (see also Population
section). The decaying remains of these organisms in turn attracted
flies and other insects, providing additional food.

Hikina Island: Small numbers were recorded here from 1964
through 1969 by POBSP personnel.

Sand Island: In 1923 Wetmore (ms. b) noted two American Golden
Plovers and collected one on Sand Island.

POBSP personnel observed this species here during 1963-1969.
Amerson also noted its presence in November 1973. American Golden
Plovers showed less preference for specific habitat than any other
shorebirds. They were almost equally abundant in the interior of the
island and on the beaches. The original portion had the highest con-
centration, probably because of higher insect populations there as a
result of denser vegetation and large numbers of other birds, including
dead ones. They were seldom seen on rocky shorelines or other rough
ground.

Populations and Annual Cycle

Figure 39 shows graphically the monthly mean population fluctuations
recorded during POBSP studies. Figure 85 shows semimonthly population
fluctuations by year, with irregularities smoothed by plotting the
line from moving averages (i.e., each point is plotted at the average
of the actual estimate for that point and the two adjacent points, re-
sulting in smoothed lines which are more easily compared than lines
plotted from the raw data).

Summer and winter numbers were fairly constant because the daily
turnover of birds was small on the atoll as a whole, although there was
movement between the islands. During the periods of migration in fall
and spring, daily fluctuations: were greater. The numbers shown for
these seasons are no more than rough indices to the total number of
birds passing through the atoll because they do not take into account
daily turnover. The reported figures usually were estimates based
on the largest count during the period in question, or in some cases

269

Pi See

NUMBERS
=
oO

50 a
0 5 Be met Bete"
100

PN em P~e 1969

NUMBERS

J F M A Mme J A S 0 N D

Figure 85. Moving averages of semimonthly estimates of Golden Plovers, Sand
and Johnston Islands combined, 1963-1969.

270

only the last count made during the period, which in either case
would result in a figure well below the actual number that had been
on the atoll during the period. During the summer and winter, when
populations were fairly constant, the reported figures probably were
fairly close to the actual number present during the period.

The few former records given provide insufficient data to deter-
mine whether numbers of Golden Plovers visiting the atoll has changed
significantly over the years. Wetmore's count of two birds in July
1923 is within the range of variability in Golden Plover numbers during
POBSP studies, although it is below average.

From a July low which ranged from 50 in 1964 to none in 1967,
numbers began to increase in late August, reaching a peak in late
October or early November. As transients moved on and winter residents
set up territories, numbers declined, then leveled off. During the
spring there was a slight increase some years, but nothing as dramatic
as the fall peak. It is likely, however, that as many birds moved
through the area in spring as in fall, but that individuals spent less
time on the atoll before moving on; thus the numbers recorded were
not as high in spring as in fall.

An exception to this general pattern occurred in the spring of
1964 when numbers were far higher than in any other spring, and higher
than most fall maxima. This unusual spring peak resulted from a
rich food source in the form of aquatic organisms brought up in
dredged coral. During this period 434 acres of land area was added
to the atoll by dredging coral from the lagoon (Bauer, 1965). It was
pumped onto the existing land as a slurry in large pipes, then bull-
dozed into place along the edge of the island, and out into the
lagoon. Pools and slowly flowing sheets of water draining away from
the solid material, and the solid material itself, contained abundant
aquatic organisms; the decaying remains of these plants and animals
attracted flies and other insects, all of which provided abundant food
for shorebirds. It is unlikely that the number of birds visiting the
island was significantly greater then than in other years, but the
birds probably stayed longer. Rather than a constant flow of birds
passing the atoll and staying no more than a day or so, large numbers
Stayed to feed and the daily counts were swelled greatly. This new
land area quickly changed to solid barren coral which held little
attraction for shorebirds.

Two new islands, Akau and Hikina, were built at the same time as
the construction on Johnston Island; the scanty data show that the
numbers of ‘shorebirds found on these islands was greatest during the
construction.

The only other unusual fluctuation occurred in fall 1968 when
counts were as high as in spring 1964; there was no apparent reason
for this unusual abundance. Weather patterns may have affected migra-
tion routes at sea or perhaps there was an unusually successful hatching

ait

year in the Arctic area where birds that migrate via Johnston Atoll
breed.

By early October at least some plovers established territories
on the islands, and fights between adjacent birds were common.
Figure 86 shows the distribution of Golden Plovers on Sand Island
26 December 1965, at which time 35 territories could be distinguished.
The clusters of twos and threes on the southwest and west sides of
the island were probably birds without territories. Territories
usually were maintained no later than January, at least on Sand, but
the breakdown in territoriality probably was forced by the appearance
at that time of hordes of pre-nesting Sooty Terns. In 1969 when
Sooty Tern nesting was delayed until March a few Golden Plovers main-
tained territories until late February.

By late February a few birds had begun to molt into dark breeding
plumage, and by April a third to half were in some stage of molt. As
the breeding plumage developed, so apparently did the migratory urge
and seldom did birds remain on Johnston until they had acquired their
full breeding colors. Plovers remaining on Johnston through June, July,
and August, presumably young of the previous year, sometimes showed at
least a few dark feathers but never developed full breeding attire.
Earliest fall migrants, arriving in late August, often showed remnants
of breeding plumage, and birds with a few black belly feathers were
found until October.

Figure 86. Distribution of Golden Plovers on Sand Island, Johnston Atoll,
26 December 1965.

Golden Plovers showed less preference for specific habitat than
any other shorebirds. They were almost equally abundant in the in-
terior of the islands and on the beaches. The original portion of Sand
Island had the highest concentration of Golden Plovers, probably
because of higher insect populations there as a result of denser
vegetation and large numbers of other birds, including dead ones.
Golden Plovers were seldom seen on rocky shorelines or other rough
ground. Except when on territories these birds appeared to move
freely between the islands of the atoll.

Although Golden Plovers frequently associated with Ruddy Turn-
stones, the two species probably were attracted to common feeding
areas rather than to each other. Plovers were not as gregarious as
turnstones and flocks of more than half a dozen were rare, even during
the height of migration.

Over-winter mortality of Golden Plovers was fairly high, at least
in some years. In 1964, 1966, and 1969 at least ten dead plovers were
found during winter or early spring; lower numbers were reported in
other years. Causes of mortality were not determined, but carcasses
were widely scattered and definitely were not guywire strikes. Some
fall birds appeared to be emaciated and may have been unable to re-
cover from the stress of migration on the food available on Johnston.
In 1964 and 1966, however, the deaths were most frequent in April,
and involved apparently healthy birds. It is quite likely that the
weekly spraying of the islands with insecticides was responsible for
this mortality, but no chemical analysis of dead birds was attempted.
Malathion and Vapono were heavily used on all islands of the atoll in
an attempt to control cockroaches. Changes in metabolism associated
with molt and preparation for migration may have rendered spring birds
highly susceptible to insecticide poisoning.

Specimens

Wetmore collected one American Golden Plover at Johnston Atoll
in 1923; POBSP personnel collected 18 specimens during 1963-1969
(Appendix Table 7). These constitute a new specimen record for the
atoll.

Banding and Interisland Movement

A total of 119 American Golden Plovers were banded by POBSP per-
sonnel at Sand Island (Table 24). Most died on the island: a few
shortly after banding, but two banded in early December 1963 were
recaptured alive in March 1964; two banded in October 1964 were identi-
fied by their streamers in late December 1964 and one was recaptured
alive in January 1965 and another was recovered, recently dead, in
April 1966; one banded in November 1966 died on the island 23 April
1967; and one banded and streamered in 1968 remained on the island
until about 20 March 1969, when it presumably migrated. These data
indicated that individual wintering plovers may remain on the island

273

for several months. They suggest, but do not prove, that plovers
return to the same island in subsequent winters. No plovers banded
elsewhere were recaptured on the atoll, and none banded on Sand Island
has been recovered elsewhere.

At-Sea Distribution

American Golden Plovers were recorded as migrants in the at-sea
grid southwest of Johnston only during October, November, and December
(Zable 21).

BLACK-BELLIED PLOVER Pluvtalts [=Squatarola] squatarola
Status

Accidental; one collected and three reliable sight records from
Johnston and Sand Islands.

Observations

Sundell and Wislocki closely observed, but were unable to collect,
a Black-bellied Plover in winter plumage near the Coast Guard barracks
on 27 November 1963. Two weeks later (11 December) they caught one in
a mist net and collected it. It was also in winter plumage and may
have been the same bird. Wislocki saw another on Johnston Island on
3 January 1964. The final record was Woodward's sighting of one
Black-bellied Plover in partial breeding plumage on the original
portion of Sand Island on 24 July 1965. All sight records, except
that by Wislocki on 3 January 1964, were recorded in detail in field
notes and could hardly be misidentifications.

Black-bellied Plovers are found infrequently in the northwestern
Hawaiian Islands and have been recorded from Kure Atoll, Midway Atoll,
Lisianski, and Laysan (Clapp and Woodward, 1968; Woodward, 1972),
as well as in the main Hawaiians. This species breeds primarily on
tundras of Eurasia and North America and migrates mainly along the
coasts, from mid-temperate latitudes into the tropics (AOU, 1957).

Specimens

The collected specimen noted above and in Appendix Table 7 is a
new specimen record for Johnston Atoll.

SEMIPAIMATED PLOVER Charadrius semtpalmatus
Status

Accidental; two specimen records from Sand Island.

274

Observations

Amerman saw one near the old pier on Sand Island each day from
30 August through 1 September 1963. He saw the same again or another
5 October, and Sundell collected one from the rocks northeast of the
Coast Guard barracks on 15 October. All observations could have been
of the same bird.

Kleen collected the second specimen 30 August 1968 after it had
been run over by the Coast Guard jeep on the causeway.

Specimens

The two collected specimens, both immatures, noted above and in
Appendix Table 7, constitute a new specimen record for Johnston Atoll.

The only other Semipalmated Plover specimen record from the
north-central Pacific was taken at Midway Atoll. Sight records have
been from Midway Atoll, French Frigate Shoals, Oahu, and Maui (Clapp,
1968a; Clapp and Woodward, 1968; Donaghho, 1969; Mull, 1969; and
Amerson, 1971). Two specimens have been taken from Baker Island in
the south-central Pacific (Clapp, 1968a). This species breeds in most
of Alaska and eastward across Canada, and winters from central
California and South Carolina south on both coasts to Patagonia, and
the Galapagos (AOU, 1957).

BRISTLE-THIGHED CURLEW Numentus tahtttensts
Status

Uncommon migrant; found usually singly on Akau, Johnston, and
Sand Islands. Fairly regular fall visitors, having been recorded
every fall from 1963 through 1969, except 1966.

Ecological Distribution

Akau Island: A few were seen here by POBSP personnel.

Johnston Island: There were only three sightings on Johnston
Island from 1963 through 1969.

Sand Island: Bristle-thighed Curlew favored habitat was the
grassy areas within a few yards of shore on the original portion of
Sand Island. Less often they wandered farther inland or used the man-
made portion. The bird present in February and March of 1969 regularly
flew to Akau Island each evening at sundown, returning to Sand each
morning to feed throughout the day. It is likely that most curlews
visited other parts of the atoll occasionally, for they often were
absent from Sand.

275

Populations and Annual Cycle

From one to five Bristle-thighed Curlews were recorded every fall
from 1963 through 1969, except 1966. Birds have been found in all months
except April (Tables 20 and 63).

Table 63. Observations of Bristle-thighed Curlews on Sand Island,
Johnston Atoll, 1963-1969

Number
Date of Survey of Birds Remarks
1963 (Minimum number: 2; maximum: 6; probable:
AOE) is
1 Aug. Seen and heard several times during afternoon.
3) Sept. Reported by several Coast Guardsmen.
1S Sept. Seen at 1500 hours.
16, 17 Sept. In puddle at west end of causeway.
28 Sept. Found dead by dump, skeleton saved.
eos (OCts Reported by Coast Guardsmen.
1964 (Minimum number: 1; maximum: 3; probable:
ily @re\ Pre
i Sept. On causeway, photographed by Lehner.
18 Sept. By barracks, probing among coral rocks in
afternoon.
GuOct. No data.
1965 (Minimum number: 6; maximum: 7-8; probable: 6).
2 Sept. Collected.
14, 15 Sept. No data.
16 Sept. Seen together.
17 Sept. Collected.
19-21 Sept. One or more; no data.
1966 None observed.
1967 (Minimum number: 2; maximum: 3; probable: 2).

28, 31_Aug.

On west end of Sand I.

Table 63. (continued)

Number

Date of Survey of Birds Remarks

MOG 26a. Autor.
mid-Oct.

1968

14 May-
3 June

6, 7 Sept.
20-23 Sept.-
31 Dec.
1969

1 Jan.-

(Joie 7A0) Julebek-
8 Sept.

Oo Sept.

ca. 27 Aug.

il

Seen almost daily on west and southwest sides
of east end of Sand I.; less often on causeway
and west end. One collected in October by
USCG.

(Minimum number: 2; maximum: 4; probable: 4).

One on Johnston I. 20 Sept., on Sand EI. 21
Sept., joined by 2nd bird 23 Sept. Both
present through end of year.

(Minimum number: 3; maximum: 4; probable: 4).
One died about 20 January., the other re-
mained feeding on Sand I., roosting on Akau,

until about 30 Mar.

Seen and heard intermittently on Sand I.

Heard on Johnston I.

John Gossett saw one with bad left leg on
the northeast side of Johnston I.

Most of their food undoubtedly was insects, but three times curlews
were seen attempting to eat mice. Kleen in October 1968 and Shelton in
February 1969 saw a curlew slamming a mouse against rocks, apparently
attempting to break it up enough to swallow it. Kleen was unable to
determine if the mouse was eaten, but Shelton watched for 45 minutes during
which time the mouse was slammed into the rocks several dozen times. The

mouse was finally abandoned.

The axial skeleton of the mouse, an adult

female, was completely crushed and there were small breaks in the skin,
but the Curlew apparently had been unable to swallow it. Brownell re-
ported a curlew eating a mouse in January 1969, but gave no details.

Bristle-thighed Curlews were large and unusual enough to elicit a
weak mobbing response from Brown Noddies--a response observed twice in
1965. In 1969, when the curlew flew from Sand to Akau at sunset, it
passed through or under swirls of prebreeding Sooty Terns which briefly
swarmed after it each evening.

Die

Specimens

Bristle-thighed Curlews were not recorded at Johnston Atoll before
POBSP studies began. The five specimens listed in Appendix Table 7 con-
stitute a new specimen record for the atoll.

Banding and Interisland Movement

No curlews were banded on Johnston Atoll. No interisland movements
involving Johnston are known.

Bristle-thighed Curlews breed in western Alaska and winter from the
Marshall and Hawaiian Islands to Santa Cruz, Tonga, and Tuamotu Islands
CAOU,, 1957).
LESSER YELLOWLEGS Tringa [=Totanus] flavtpes
Status

Accidental; one specimen record from Sand Island.
Observations

A slightly emaciated male Lesser Yellowlegs was collected by
Amerson at Sand Island on 18 August 1963. For two days before being

collected, this bird inhabited the original portion of the island where
it ate insects from a pile of bird carcasses.

Specimens

The specimen noted above and in Appendix Table 7 is a new specimen
rcord for the atoll.

Lesser Yellowlegs have been recorded from Kure Atoll, Midway Atoll,
Laysan, Maui, and Oahu in the Hawaiian Islands (Clapp, 1968a; Clapp and
Woodward, 1968; Woodward, 1972). This species breeds in northern Canada
and Alaska and winters from southern North America to Argentina and
Chile; most migration is inland (AOU, 1957).

SPOTTED SANDPIPER Actitts macularta

Status

Accidental; one specimen record and one sight record from Sand
Island.

Observations
A Spotted Sandpiper in non-breeding plumage was collected by Kepler

and Lehner near the barracks at Sand Island 1 September 1964. A sand-
piper was seen repeatedly by Shelton along the causeway and on the rocky

278

beaches from 28 August through 8 September 1969 probably was this
species, although it could have been the smaller Common Sandpiper
(Tringa hypoleucos) of the Old World.

Specimens

The single collected specimen noted above and in Appendix Table 7
is the only verified record of this species for Johnston Atoll, as well
as for the entire central Pacific Ocean.

Other Pacific records are one collected at Taka Atoll in the Mar-
shall Islands (Amerson, 1969) and a sighting of this or the Common
Sandpiper on Enderbury Island in the Phoenix Islands (Clapp, 1968b).
Spotted Sandpiper breed throughout most of the northern two-thirds of
North America and winter from British Colombia, southern Texas, and
South Carolina, south to Argentina (AOU, 1957).

WILLET Catoptrophorus semtpalmatus
Status

Accidental visitor; one sight record on Johnston Island.
Observations

A large shorebird, 1.5 to 2 times the size of a Golden Plover,
was observed by Capt. Douglas Boucher, USAF, on the northeast shore of
Johnston Island 17 August 1969. It was dark above, with a distinct and
conspicuous white wing bar. The size and wing bar make this bird most
likely a Willet. The call was a distinct whistle, described as some-
thing like a killdeer, which roughly fits the "kay-ee" call of the
Willet as given by Peterson (1961: 110). Captain Boucher was known to
be an astute observer, and was quite familiar with Golden Plovers, with
which he compared this bird's size.

Specimens

This is a new sight record for Johnston Atoll.

Only one other sight record of a Willet, a bird seen on Oahu
(Pyle, 1968), exists for the central Pacific. This species breeds
primarily along the border of Canada and the United States, and, locally,
along the Atlantic and Gulf coasts; it winters from northern California,
the Gulf of Mexico, and south Atlantic coasts of the United States
south to northwestern Peru, Bolivia, and Brazil (AOU, 1957).

279

WANDERING TATTLER Heteroscelus tneanus
Status

Common but scant migrant; found singly or in pairs on rocky shores
of all islands. Maximum numbers occur during spring and fall migrations;
few birds present in summer and winter.

Ecological Distribution

Akau Island: Few were present during POBSP studies.

Hikina Island: POBSP personnel found a few here from 1964 through
1969.

Johnston Island: Wetmore (ms. b) recorded two Wandering Tattlers
in July 1923. POBSP personnel found them inhabiting rocky shores--
never more than a few feet from water.

Sand Island: On Sand Island the favored habitat is the sides of
the causeway which are composed almost entirely of large angular rocks.
They most often feed solitarily, although pairs are common. During
migration the largest aggregations were of only five or six birds.

Populations and Annual Cycle

Figures 39 and 87 show monthly mean population estimates and
semimonthly estimates by year, respectively, based on actual counts of
Wandering Tattlers on Sand and Johnston Islands during POBSP studies.
These must be considered minimum figures because the other two islands,
where one to three tattlers were usually found on the infrequent visits,
were not included. These few birds would not greatly inflate the figures;
it is likely that often birds were using all the islands of the atoll and
counts of the two main islands probably took care of most if not all
birds present. There is even the possibility of counting the same birds
more than once on these two islands.

Major population variations occurred in the spring of 1964. The
unusually large numbers of tattlers probably resulted from the abundance
of food in the freshly dredged coral. Golden Plover and Ruddy Turnstone
populations also increased then, probably for the same reason. The high
number of tattlers present in fall 1968 also corresponded with an un=
usually high number of plovers and turnstones; there is no clue as to
why these birds were more abundant than usual that year. Weather patterns
may influence the number of shorebirds arriving on remote atolls, but no
descriptions of such correlation are available.

Population numbers show slight dips during winter and summer, with
highest numbers during the spring and fall migrations. There appear to
be more tattlers present in the fall than in the spring, but, as in the
case of Golden Plovers and Ruddy Turnstones, this may be only apparent
because of shorter stays by spring migrants. Approximately half the

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281

tattlers passing through in March through May, and a smaller number of
returning fall birds, showed some degree of barring on the breast; the
rest were in typical unbarred non-breeding plumage. Summering birds
probably were pre-breeding young that did not return north until they
were nearly two years old.

Tattlers were more closely tied to shorelines than any other common
shorebird visiting the atoll. They were practically never found more
than a few feet from water, and generally inhabited the most rocky
shores. On Sand Island the favored habitat was the sides of the cause-
way which were composed almost entirely of large angular rocks. They
most often fed solitarily, although pairs were not uncommon. In migra-
tion the largest aggregations noted were of only five or six birds.

Specimens

Appendix Table 7 presents data from the four Wandering Tattlers
collected on Johnston Atoll. These constitute a new specimen record.

Band ing and Interisland Movement

Three Wandering Tattlers were banded and streamered in September
and October 1968 by Kleen (Table 24); one was subsequently sighted in
March 1969. Although it is possible that the bird was on the atoll
throughout the intervening four months, it is more likely that it moved
farther south for the winter and returned north via Johnston in March.

Wandering Tattlers breed above timberline in the mountains of
Alaska and northwestern Canada, and possibly in Siberia. They winter
along the west coast of North America and throughout most of the tropical
Pacific Islands (AOU, 1957).

RUDDY TURNSTONE Arenarta interpres
Status

Common migrant; found in small flocks usually on gravelly areas
along shores of all islands. Most abundant during spring and fall
migrations; some birds present throughout the year.

Ecological Distribution

Akau Island: Ruddy Turnstones were recorded on the gravelly beaches
at Akau from 1964 through 1969 by POBSP personnel. They probably roosted
here at night.

Hikina Island: POBSP personnel reccrded Ruddy Turnstones on the
gravelly beaches at Hikina from 1964 from 1969. Although no night
observations were made, it is likely that turnstones roosted here at
night because of the quiet protected beaches.

282

Johnston Island: POBSP personnel observed this species here
throughout its studies, especially in relatively open areas of coarse,
gravelly coral. They were seen at one time or another on almost any
open, gravelly part of Johnston Island, but largest concentrations were
always on the wide, gravelly beach at the southwest corner, which was
well protected from the surf. This same area was also probably used
as a nightly roosting area.

Sand Island: Wetmore (ms. b) saw two Ruddy Turnstones on Sand
in /July 1923

POBSP personnel found them here during 1963-1969. They were most
frequently found on the newest coral, dredged in 1964, on the man-made
end of the island or on beaches that were similar in texture. At night
roosting flocks of turnstones were frequently flushed from the south-
west islet, but only occasionally did they roost on the northeast
peninsula.

Populations and Annual Cycle

Figure 39 shows monthly mean population estimates of Ruddy Turnstones
recorded during POBSP studies. Figure 88 shows annual fluctuations; lines
are smoothed by the moving averages method, as described in the Golden
Plover account. Comments given in the Golden Plover account on the
relationships between numbers reported in these tables and actual numbers
of birds using the islands apply equally to Ruddy Turnstones.

The general form of the population curves for Ruddy Turnstones is
the same as for Golden Plovers. The highest peaks occur in late fall,
with a decline through winter, and a smaller, less distinct peak in
spring. As with plovers, the actual number moving through in spring may
be as high as in fall; individuals may have spent less time on the atoll
during spring migration; therefore, observed numbers are lower.

Also like Golden Plovers, Ruddy Turnstones showed a higher population
in spring 1964 than in any other spring, because of birds remaining to
feed on freshly dredged coral. Unlike plovers, however, turnstones
showed a similar but smaller increase in spring 1965. Apparently the
new coral still contained abundant food for turnstones, but not for
plovers. _Turnstones reached unusually high numbers in fall 1968, as
did plovers, and the reason is equally unapparent.

Some Ruddy Turnstones began to acquire their breeding plumage by
late January, and by April and May most showed some degree of change
although few, if any, completed the molt before leaving the atoll.

From late June through August the few birds present were in non-breeding
plumage, probably indicating that they were young birds that did not
migrate to the gorthern breeding grounds. Early migrants arriving in
late August and September sometimes showed part of their breeding colors,
but these were soon lost.

Ruddy Turnstones tended to remain near shores more than did Golden
Plovers, but they were often found inland, especially in relatively open

283

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taal M A M J J A S 0 N D

Figure 88. Moving averages of semimonthly estimates of Ruddy Turnstones,
Sand Island, Johnston Atoll, 1963-1969.

areas of coarse gravelly coral. The tendency of turnstones to avoid
vegetated areas may have saved them from the over-winter mortality
described for Golden Plovers. If insecticides were responsible for
Golden Plover mortality, then it would follow that turnstones might
be less severely affected because spraying was primarily directed at
vegetated areas. POBSP field notes and records mention only two dead
turnstones being found: one in November 1965 and one in June 1966.

Feeding areas of plovers and turnstones were by no means mutually
exclusive, and the two species frequently occurred together. The largest
flocks of turnstones usually contained a few plovers, but this associa-
tion appeared to be a mutual attraction to a feeding area rather than an
attraction between the birds. Likewise most Sanderlings observed on
Johnston Island were in the company of turnstones on the southwest
beach, but the attraction for Sanderlings was more likely the beach
than the turnstones.

Individual turnstones were rarely seen, and no territorial behavior
was ever noted. Flock sizes ranged commonly from 3 to 20, but larger
ones up to the 118 birds found 30 September 1968 on the southwest beach
of Johnston Island were not unusual.

At night turnstones gathered to roost in larger groups on Sand
Island. Counts were not possible, but estimates from calls and the
absence of birds from other parts of the island at night indicated that
at least all the birds feeding on Sand, and probably a good many from
other islands in the atoll, were gathered in these roosting flocks. In
November and December 1966 Schreiber noted that turnstones fed during
the day in small flocks, but in late afternoon they gathered into larger
groups on the man-made end of Sand Island where they continued feeding
until dusk, when they all flew off toward Hikina Island. No night
observations were made on Hikina or any other islands on the atoll,
but it is likely that turnstone roosts occurred on islands other than
Sand. Hikina, Akau, and the southwest beach of Johnston all would have
provided quiet protected beaches similar to the southwest islet on Sand.

Specimens

Ten specimens of Ruddy Turnstones have been collected from Johnston
Atoll (Appendix Table 7). These constitute a new specimen record.

Banding and Interisland Movement

The POBSP banded and streamered 17 Ruddy Turnstones on Sand Island
(Table 24). Although marked birds were frequently seen on all the
islands of the atoll, verifying suspected free movement among the islands,
only one was recovered--a bird banded in October 1964 found dead in
November 1965. Birds with streamers were seen in July and November 1965;
the last banding that year had been done in March and April. These data
indicate that turnstones may remain on the atoll for sevéral months.

285

They suggest, but do not prove, that turnstones return to the same island
in subsequent winters. One banded at Sand Island on 19 October 1964 was
recaptured at St. George Island in the Pribilofs on 14 August 1966.

In 1966 Thompson and DeLong (1967) banded and marked (with red
paint and green, plastic, leg streamers) 7,543 Ruddy Turnstones on
St. George from a population of 15,000 to 20,000 birds. Two were later
recaptured at Johnston: one was collected and one was caught alive and
released. Records of marked birds in shorebird counts made from late
September through early November 1966 on both Sand and Johnston Islands
(Table 64) provided data from which crude calculations were made of
the minimum proportion of Johnston Atoll birds that had been on St.
George that year. Since Thompson and DeLong's figures do not account
for turnover, 20,000 would seem to be a reasonable minimum figure for
the population. Thus for each bird banded and marked on St. George,
about two present were not banded. Assuming random mixing of marked and
unmarked birds during migration, the number of turnstones on Johnston
that had been on St. George was about three times the number of marked
birds observed: 3.9 percent x 3 = 11.7 percent. Thus a minimum of about
12 percent of the Ruddy Turnstones on Johnston Atoll that fall had passed
through the St. George Island staging area in the early part of their
migration.

Table 64. Numbers and percentages of Ruddy Turnstones sighted on
Johnston Atoll that were color marked on St. George Island,
Pribilofs, fall 1966*

Period Number sighted Number marked Percent marked
16-30 Sept. 387 17 5) 4%)
J=1'5 Oct. 410 4 0,
16=31 Oct. 401 Zi Dee
1=—15) Nov. 396 al; 4.3
Totals 1,494 59 31658)

*Counts were made every day or every other day on Sand and once weekly
on Johnston. Number of birds involved was far less than totals shown,
because of duplication.

The actual number of marked birds in the population could not be
determined because of duplication of counts and lack of turnover data
for Johnston. The most seen at one time was four, but it is likely that
at least two to four times this number visited Johnston Atoll.

The banding program on St. George extended from 1964 through 1968
and marked birds were observed in years other than 1966, but detailed
ratios were not obtained. Another St. George bird was collected in 1968.

286

Thompson (1973) suggests a circular migration route for Ruddy
Turnstones using islands in the Pacific Ocean. There is a southward
movement through the central Pacific in fall and a northward migration
in the western Pacific in spring through Japan and into Siberia and St.
Lawrence Island, Alaska, to breed.

At-Sea Distribution

Ruddy Turnstones appeared as migrants in the at-sea grid southwest
of Johnston Atoll only during August, September, and October (Table 21).

DOWITCHER species Limnodromus species

Status

Accidental; two fall sight records from Sand Island.
Observations

A dowitcher was observed and photographed on 8 September 1969 on
the beach and a few yards inland in scattered grass at the southwest
corner of the original portion of Sand Island. Mrs. R.C. Laybourne
of the U.S. Fish and Wildlife Service Bird and Mammals Laboratories,
National Museum of Natural History, tentatively identified this bird
from Shelton's photographs (Fig. 89) as a Short-billed Dowitcher on the
basis of spotting on the flank. Another dowitcher, possibly this species,
was seen on Sand on 5 October 1964 by Kepler.

Figure 89. Dowitcher species photographed on Sand Island, Johnston Atoll,
8 September 1969 (POBSP photo by P. C. Shelton).

287

Since 1945, at least 15 sight records of dowitchers or Long-billed
Dowitchers in the Hawaiian Islands have been published, but only two
specimen records exist. A Long-billed Dowitcher was collected at Kure
Atoll and a Short-billed Dowitcher was collected at Midway Atoll, the
latter identified as L. g. courninus (Clapp and Woodward, 1968; Woodward,
1972). Short-billed Dowitchers breed from southern Alaska, southern
Mackenzie, and northern Alberta eastward across Canada, probably to the
Ungava Peninsula. They normally winter from California, the Gulf
Coast and South Carolina to Peru and Brazil. Long-billed Dowitchers
breed from northeastern Siberia and northwestern Alaska to Mackenzie
and normally migrate primarily through the western United States to
Mexico and Guatemala (AOU, 1957).

This is a new sight record for Johnston Atoll.

SANDERLING Croecethta alba
Status

Uncommon migrant; usually found as singles on sandy beaches of
Johnston and Sand Islands. Sporadic but regular visitor usually from
late August through April.

Ecological Distribution

Johnston Island: Most Sanderlings were recorded here by the POBSP
particularly on the sandy beach at the southwest corner. Most were
loosely associated with flocks of Ruddy Turnstones and Golden Plovers.

Sand Island: POBSP personnel observed this species infrequently
on Sand Island.

Populations and Annual Cycle

Sanderlings appeared most regularly from late August through
April, but have been recorded in all months except June (Table 20 and
Fig. 90). Highest numbers occurred in early 1964, when all shorebirds
were unusually numerous because of abundant food from dredging opera-
tions (see Golden Plover and Ruddy Turnstone accounts), and in November
1966, at which time there was nothing unusual on the island to attract
the birds.

Since most Sanderlings recorded were on Johnston Island and not on
Sand Island, POBSP records do not reflect the true regularity of Sander-
ling occurrence on the atoll. In 1964, 1965, and 1966 Johnston was
censused almost weekly, and figures from those years are more reliable
than those from later years when there were long periods in which no
complete shorebird censuses were made on Johnston. None was recorded
from Akau or Hikina, but it is likely they occasionally visited these
islands.

288

NUMBERS NUMBERS NUMBERS NUMBERS NUMBERS NUMBERS NUMBERS
OT

NUMBERS

J
Figure 90.

1969

F M A Mime J Ae S$ 0 N D

Moving averages of semimonthly estimates of Sanderlings, Sand
and Johnston Islands, 1963-1969.

289

Most Sanderlings were loosely associated with flocks of Ruddy
Turnstones and Golden Plovers, which also were most abundant on the
southwest corner of Johnston. Most were in typical light, non-breeding
plumage, but a few fall arrivals still had part of their dark breeding
dress.

Specimens

One specimen was collected by POBSP personnel (Appendix Table 7)
and is a new specimen record for the atoll.

Banding and Interisland Movement

Only one bird was streamered, but was not banded because the bands
were not small enough. No interisland movements are known.

Sanderlings are circumboreal breeders that winter on coasts and
oceanic islands from mid-temperate northern to mid-temperate southern
latitudes (AOU, 1957).
WESTERN SANDPIPER Ereunetes maurt
Status

Accidental; one sight record on Sand Island.
Observations

Kleen identified a bird on the causeway of Sand Island as a
Western Sandpiper on 4 September 1968. It was not collected or seen
again.

One Western Sandpiper has been collected at Kure Atoll and is the
only record for the northwestern Hawaiian Islands (Clapp and Woodward,
1968; Woodward, 1972). This species breeds in Alaska and winters from

California, the Gulf Coast and North Carolina south to Venezuela and
Peru (AOU, 1957).

Specimens

This is a new sight record for Johnston Atoll; no confirmation
specimen record exists, however.
PECTORAL SANDPIPER Calidrts [=Erolia] melanotus
Status

Uncommon but regular fall migrant; eight specimen records and
many sight records from Sand Island.

290

Observations

At least one Pectoral Sandpiper was seen each fall on Sand Island
by POBSP personnel from 1963 through 1969 (Tables 20 and 65). There
were at least 16 fall records, possibly more if birds seen on successive
days were different; eight of these were verified by collecting. Only
two spring records (April and May 1965) exist and these may have been of
the same bird.

Most sightings were of single birds, not associated with other
shorebirds. Habitat preference appeared to be similar to that of Golden
Plovers; they were seen inland more often than along the shores. All
observations were made on Sand Island.

Specimens

The eight Pectoral Sandpiper specimens noted above constitute a
new specimen record for Johnston Atoll (Appendix Table 7).

Pectoral Sandpipers are regular visitors to the central Pacific as
they have been recorded from Kure Atoll, Midway Atoll, Laysan, Oahu,
and Hawaii (Clapp and Woodward, 1968; Woodward, 1972; and Ely and Clapp,
1973). They breed on the Arctic slopes of eastern Siberia and North
America and winter primarily in South America, and casually to southern
and western Pacific islands (AOU, 1957).

Table 65. Observations of Pectoral Sandpipers on Sand Island,
Johnston Atoll, 1963-68

Number
Date of Survey of Birds Remarks
L963, “sr Oct:. IL Northeast peninsula, collected.
1964 25-26 Sept. a Observed first on 25th, collected 26th.
19" Oct. i Observed near several Golden Plovers and
Ruddy Turnstones; collected.
1965 29 Apr. 1 Seen clearly by two observers; not col-
lected.
6-7 May il Seen clearly by two observers; not col-
lected. Possibly samd bird seen 29 April.
14 Sept. il Collected.
27 Sept. 4 2 Collected.
28 Sept. i Sight record; no details.
t—i2 Oct. i Observed daily; probably same as seen

28 September.

Zr

Table 65. (continued)

Number

Date of Survey of Birds Remarks

19657 19-0ct. 1 Sight record; possibly same as earlier in
month.

22-31 Oct. 2 Observed almost daily; one possibly same
as seen earlier.

1966 12-25 Oct. af Seen almost daily, on north shore, east
hill, south of transmitter, and by bar-
racks, all open sandy sites.

1967 (No records; no observations after 12
September).

1968 30 Sept. iL Collected.

24Oet « i Collected.

2-10 Oct. 2 Present on Sand Island.
11-17 Oct. 5 Up to five present daily.
18-31 Oct. 4 Up to four present daily.

4 Nov. 1

6 Nov. iL

12 Nov. 1 Found dead.

SHARP-TAILED SANDPIPER Calidrts [=Erolta] acuminata

Status

Accidental; two specimen records and three sight records from
Johnston and Sand Islands.

Observations

Immature females were collected by Lewis 29 October 1965 and by
Kleen 30 September 1969. Schreiber and Shelton saw, but were unable to
collect, one bird on Sand Island on 23 October 1966, and Kleen saw one
10 October 1968 on Sand and another, or the same bird, on Johnston Island
on 16 October 1968. Pectoral Sandpipers, with which this species is
easily confused, were present on the atoll at the times of all three of
these sightings so comparisons could be made easily; hence these identi-
fications are considered highly reliable.

Specimens

The two specimens noted above and in Appendix Table 7 constitute
a new specimen record for Johnston Atoll.

Sharp-tailed Sandpipers have been infrequently recorded from Kure
Atoll, Midway Atoll, Pearl and Hermes Reef, Laysan (Clapp and Woodward,
1968; Woodward, 1972; Amerson, Clapp, and Wirtz, in press) and the main
Hawaiians. They breed in northern Siberia and winter primarily in the
western Pacific. They occur rarely along the North American Pacific
coast south to California (AOU, 1957).

BUFF-BREASTED SANDPIPER Tryngttes subrtftcollis
Status

Accidental; one specimen record from Sand Island.
Observations

Kleen collected an immature Buff-breasted Sandpiper on the man-
made portion of Sand Island on 14 September 1968.

Specimens

This specimen (see also Appendix Table 7) is a new specimen record
for Johnston Atoll.

The only other known Pacific island records of Buff-breasted
Sandpipers are from Eniwetok, in the Marshalls (Pearson and Knudsen,
1967; Amerson, 1969) and Oahu (Donaghho, 1970). This species breeds
on the Arctic slope of North America and winters in Argentina, migrating
primarily through central North America. They are of casual occurrence
on the Pacific coasts of North America, Japan, and Siberia (AOU, 1957).

RUFF Phtlomachus pugnax
Status

Accidental; one specimen record from Sand Island.
Observations

A Ruff was seen on 10 March 1966 feeding with a small group of
Ruddy Turnstones at the west end of the causeway on Sand Island. At
dusk it roosted with a large flock of turnstones on the northeast

peninsula where Tordoff collected it. The bird, a male, had its crop
full of minute snails, but had no fat deposits.

Speicmens

This is a new specimen record for Johnston Atoll; see also Appendix
Table 7.

293

Previous specimen records from the central Pacific are from Kure
Atoll (Clapp and Woodward, 1968; Woodward, 1972) and Pearl and Hermes
Reef (Amerson, Clapp, and Wirtz, in press). Kaigler (1971) reported a
Sighting from Maui.

Ruffs breed across northern Europe and Asia and winter in the
southern parts of those continents and in Northern Africa. In addition
to the above central Pacific records, there are casual and accidental
records from the western Pacific and from eastern North America (AOU,
LOST).

WILSON'S PHALAROPE Steganopus trtcolor
Status

Accidental; one specimen record from Sand Island.
Observations

Kepler and Lehner collected an emaciated female Wilson's Phalarope
in winter plumage on the original portion of Sand Island 16 August 1964.

Specimens

This collection is a new specimen record for Johnston Atoll
(Appendix Table 7) and the central Pacific Ocean.

Four sight records of Wilson's Phalarope from Maui and Oahu in
the main Hawaiian Islands (Kridler, 1966; Clapp and Pyle, 1968; Kaigler,
1971; and Mull, 1971) are the only other known records from the central
Pacific. This species breeds across the interior of North America, in
southern Canada and northern United States, and winters in southern
South America (AOU, 1957).

GLAUCOUS-WINGED GULL Larus glaucescens
Status

Irregular winter visitor; two specimen records from Sand Island.
Observations

There are two late-winter POBSP records of Glaucous-winged Gulls
from Sand Island. Harrington saw a second-year bird on 1 March 1968 and
collected it 8 March; it proved to be a female with very light fat
deposits and well worn flight feathers. Shelton found a badly decom-
posed immature on the southeast beach on 7 February 1969. A gull,
probably this one, had been reported occasionally during the previous
two weeks.

294

Specimens

The above two specimens constitute a new specimen record for
Johnston Atoll (Appendix Table 7).

Glaucous-winged Gulls have been collected on most of the main
Hawaiian Islands, where they are frequent visitors. There are in-
frequent records in the northwestern Hawaiians from Kure Atoll, Midway
Atoll, Pearl and Hermes Reef, Lisianski, Laysan, French Frigate Shoals,
and Necker (Clapp and Woodward, 1968; Amerson, 1971; Woodward, 1972;
and Amerson, Clapp, and Wirtz, in press), and elsewhere in the central
Pacific (Sibley and McFarlane, 1968).

Glaucous-winged Gulls breed from the Komandorskie, Pribilof, and
Aleutian Islands to northwestern Washington, and winter south-westward
to Japan, and from southeastern Alaska to Baja California. Non-breeders
occur in summer south to California (AOU, 1957).

HERRING GULL Larus argentatus
Status

Rare visitor; various sight records of one bird from Johnston and
Sand Islands.

Observations

A subadult (second-year) Herring Gull was seen by Kleen over Sand
Island in late November and early December 1968 and at Johnston Island
on 31 December 1968 (by Brownell) and 14 March 1969 (by Shelton). There
was no opportunity to collect it.

Specimens

No specimens are known from Johnston Atoll; however, the sight
record is considered new for the atoll.

At least 12 specimens of Herring Gulls have been collected in the
northwestern Hawaiian Islands. All have been identified to the race
Larus argentatus vegae (Clapp and Woodward, 1968) which breeds in Siberia
and winters as far south on the Asiatic coast as central China and
Formosa. It wanders casually to western Alaska, the Aleutians, and
British Columbia (AOU, 1957).

LAUGHING GUL]. Larus atrictlla
Status

Irregular winter visitor; one specimen record and three sightings
from Johnston and Sand Islands.

295

Observations

The first Laughing Gull observed on the atoll landed on the south-
east beach of Johnston Island 7 February 1964. It appeared to have just
made its first landfall, for it flew only reluctantly and persistently
put its head under its wing as if trying to sleep despite the presence
of people. On 13 February it was still quite tame, but by March it had
become more wary. During this time it remained in the same general area,
feeding along the shore where dredging operations were underway. On
7 April it appeared on the northeast peninsula of Sand Island, possibly
attracted by numerous dead fish from dynamiting operations. Amerson
and Amerman collected it there; it proved to be a very fat immature male.
The abundance of food from the dredged coral probably allowed this bird
to maintain itself in good condition.

Second year Laughing Gulls were identified by Bratley and Lehner
on 28 July 1964 and by Heryford and Bratley on 26 February 1965, both
on Sand Island. The first flew over the island during the day, heading
eastward. Sooty Terns harrassed it all the way across the island and it
was not seen again. The second landed about midnight in a small group
of roosting Blue-faced Boobies on the southwest islet. It could not be
caught and was never seen again.

Kleen saw an adult Laughing Gull in non-breeding plumage fly along
the causeway on Sand Island 28 October 1968.

Specimens

The specimen noted above is listed in Appendix Table 7. In
addition to these records from the Atoll, Clapp collected an immature
female 11 miles northwest of Johnston Atoll on 15 January 1965 (Sibley
and McFarlane, 1968).

Most other records are to the south. Specimens are known from at
sea (7 March 1964: 12°41'N, 171°28'W), Christmas Atoll, Palmyra Atoll,
and Baker Island (Sibley and McFarlane, 1968; King, 1967). One specimen
is known from Oahu (Berger, 1972).

Laughing Gulls breed along the Atlantic coast from Nova Scotia
south through the Gulf of Mexico and Caribbean to northern Venezuela,
and on the Pacific coasts of southern California and Mexico. They winter
in the southern half of their breeding range and south to Brazil and
Peru (AOU, 1957).
FRANKLIN'S GULL Larus ptptxean

Status

Straggler; one specimen record from Sand Island.

296

Observations

The only Franklin's Gull recorded from Johnston Atoll was a male
in breeding plumage collected by Shelton on the causeway of Sand Island
on 20 May 1969. When found it was almost dead, probably from starvation
as it was thin and had no fat deposits. It was first sighted on Sand
19 May.

Specimens

This is a new specimen record for Johnston Atoll (Appendix Table 7).

Other records of this species from the central Pacific are from
Maui, Oahu, offshore Kauai (King, 1959), French Frigate Shoals (Clapp
and Woodward, 1968; Amerson, 1969), Palmyra, and Fanning (Sibley and
McFarlane, 1968).

Franklin's Gulls breed from southeastern Alberta and eastern
Oregon to Minnesota and Iowa; they winter along the Pacific coast from
Guatemala to Chile, on the Galapagos Islands, and on the north coast of
the Gulf of Mexico (AOU, 1957).

GULL species Larus spp.
Status

Accidental; one skeleton record (lost) and one sight record from
Johnston Island.

Observations

A large gull, most likely a Western Gull in first winter plumage,
appeared on Johnston Island about 25 January 1964. Despite futile
efforts to collect it, the bird remained there at least through 9 March
1964.

Bratley first saw it lying on the beach on the south side of the
island, during mid-afternoon. When approached to within about 25 feet
it stood up, then flew out and lit on the water 200 yards offshore, only
to return to shore within ten minutes. Attempts were made to collect
the bird the next day, but it was more wary then, probably indicating
some degree of recuperation from its journey to the atoll. On the 28th
of January it was shot and wounded, but escaped by swimming out to sea.
By about mid-February it was able to fly weakly, and it remained too
wary to be collected. It was last seen 9 March. During the entire time
it was on the island it stayed along the south beach.

A skeleton of a large dusky gull, possibly a Western Gull but not
likely the same specimen, was found on Johnston Island 17 December 1965
(USNM field no. 25923). The skeleton, however, was later lost.

297

Specimens

The above records constitute a tentative new sight record for
Johnston Atoll.

Western Gulls breed from northern Washington to Baja California
and winter from southern British Columbia south through the breeding
range (AOU, 1957). There appear to be no authenticated central Pacific
records.

GRAY-BACKED TERN Sterna lunata
Status

Uncommon breeding species; present year-round. About 250 nest
on Sand Island each year from December through July, but only a few
dozen chicks are produced annually. Recent records of nesting on Akau
and Hikina Islands. Previously nested on Johnston Island.

Ecological Distribution

Akau Island: Because high tides destroyed their eggs on Sand
Island in late February 1964, about 200 Gray-backed Terns moved to the
then newly constructed Akau Island in March and laid at least 85 eggs.
Unfortunately, continued construction destroyed this new colony; only
one chick hatched, which did not survive. None has been recorded since.

Hikina Island: Kridler (BSFW, 1973) recorded Gray-backed Terns
nesting here in May 1973. A total of 35 nests was found, 12 of which
contained very small downy chicks. About 80 adults were observed.

Johnston Island: Wetmore (ms. a) recorded 25 well-grown young,
some of which were flying, here in July 1923. Because of human dis-
turbance, none has been recorded since.

Sand Island: Wetmore (ms. a) also recorded 25 with young here in
July 1923. In April 1957, Moynihan (1957: 37) observed "a small diffuse
colony of Gray-backed Terns, including perhaps 200 birds, was stretched
along both sides of the causeway between the islets. Most of the birds
were incubating eggs in nests on the bare ground between boulders or
patches of low vegetation."

From 1963 through 1969 POBSP personnel found Gray-backed Terns
nesting in loose groups of from two up to 200 birds on rough, rocky
ground around the periphery of the original island, including the
southwest islet and (prior to 1968) on the abandoned pier (Figs. 43,
91 and 92). The maximum number of eggs present in 1967 in the various
areas (Fig. 92) indicate approximate relative numbers of birds using
these sites.

Eggs are deposited on the sand, with no prior preparation of site.
Some incubating birds may gather in small stones within reach from the

298

Figure 91, Distribution of Gray-—backed Tern nest sites, Sand Island, Johnston
Atoll, 1966.

DATE: Eggs Present 21 Feb-15 Apr
akSaleal
Maximum Eggs:Chicks Hatched:Fledged
N
|
37:20:10, iF 8 Apr 0 100 200
ya 2+0:0 a |
j Feet
5 Jan-23 Mar
64:20:10

23 Mar-23 Apr
7:4:4

Figure 92. Distribution of Gray-backed Tern nest sites, Sand Island, Johnsto
Atoll, 1967.

299

egg, but usually there is no evidence of any attempt to build even the
most rudimentary nest.

Apparently their choice of peripheral nesting areas and their
beginning to nest before Sooty Terns are both adaptations to avoid
competition for nesting space with that species. Prior to 1967,
gray-backs usually began nesting on the old dock, which was perhaps
the area most secure from competition with Sooty Terns. Unfortunately,
most eggs and chicks were lost from the dock because it provided a poor
base for eggs and practically no hiding places for chicks.

After heavy egg losses occurred on the dock, many birds attempted
to renest on other parts of the island. Some of the second sites
chosen were even poorer than the dock, and many eggs were lost to high
tides (Figs. 93 and 94). In February 1967, two groups began nesting
in areas that appeared to be far better than the dock--in the south-
southwest rocks area, and on the west side of the northeast peninsula.
Unfortunately, these areas were overrun by Sooty Terns in March before
the gray-backs could successfully complete their nesting, and most of
the eggs were lost or abandoned. The gray-backs stayed on their eggs
after sooties moved into the area, but the numbers of eggs declined
steadily with no increase in chicks. Behavioral observations were not
made.

It would seem that if these birds began nesting here instead of
on the old dock, they might be able to have chicks large enough by
the time the Sooty Terns moved in and they would not be lost. It may
be, however, that chicks, even up until just before they fledge, are
highly susceptible to being killed by Sooty Terns, which descend in
hordes and perhaps would kill any small tern within their territories.

In 1957, Moynihan found gray-backs nesting along the causeway
well away from both groups of Sooty Terns that were also nesting then.
Apparently at that time there was enough space so that the gray-backs
could nest unmolested. Unfortunately, there are no records of the
success of the colony then as compared to the period of POBSP studies.

Nesting gray-backs often served as foci for pre-breeding Sooty
Terns when they settled on the island in the evenings. On 4 February
1966, one Sooty Tern incubated all day among Gray-backed Terns on the
west shore; 14 more eggs were laid by early evening and on 5 February
five were incubated a portion of the day; one persisted until 9 February,
by which time no other had begun nesting. In February 1967 as the
swirling sooties flew lower and lower over the southwest grassy portion
of the island, they were attracted to the gray-backs nesting on the
south-southwest shore, and on at least half the evenings, the first
sooties to land did so among the gray-backs. The reactions of the
gray-backs could not be determined. When the sooties began nesting,
they did not do so near the nesting gray-backs; it was a few weeks
before the gray-back nesting areas were taken in by the sooties, at which
time the gray-back eggs began to decline steadily.

300

Figure 93. Gray-—backed Tern on nest (center foreground) among Sooty Terns
in Ipomoea and Lepturus on south shore of Sand Island, Johnston
Atoll, 21 April 1969 (POBSP photo by P. C. Shelton).

Figure 94, Gray-—backed Tern nesting area on southwest islet, Sand Island,
Johnston Atoll, 16 April 1969. Two adults, one chick, and four
eggs are shown (POBSP photo by P. C. Shelton).

301

The peripheral areas used by nesting Gray-backed Terns are usually
shared with Brown Noddies, but the noddies do not nest densely enough
to cause any obvious conflicts with the gray-backs. Close behavi>ral
observations might reveal interactions that have gone unnoticed, .iowever.

Populations

Means and extremes of semimonthly estimates from 1963 through 1969
are shown in Figure 95. Year-to-year variation showed a slight decline
from 1963 to 1969. The trend was evident both in adult and egg numbers,
but was not reflected in number of chicks.

Comparison of both Wetmore's and Moynihan's figures with those
from POBSP observations at the same time of year, however, indicate no
detectable differences in total numbers on the atoll. The population
that originally was divided between Johnston and Sand Island is now
confined to Sand. Since this species occupies habitat marginal to Sooty
Terns, and possibly Brown Noddies, there probably never was much nesting
space available to them, and the population never exceeded a few hundred
birds.

Mortality of both eggs and chicks was extremely high. An average
of 101 eggs (range 63 to 155) was laid each year from 1964 through 1969,
while only an average of 17 fledged (range 11 to 26). These figures do
not show the actual difference, because many eggs were laid and lost
without being reported in the counts. For example, in late February
1964, egg numbers increased to 155, but were reduced by losses to 86
by the end of the month. Also, on the old dock in 1965, about 118 eggs
were laid, but no more than four young fledged. Without daily counts of
new eggs, which were never made over the entire island through a com-
plete nesting cycle, it is not possible to determine the total number
produced each year. Most likely, however, at least 200 eggs were laid
each year.

Numbers of young successfully fledged were not known precisely,
but the figures for numbers banded, which probably were slightly higher
than the number of young surviving to fledging, were: 11 for 1964, 15
for 1965, 13 for 1966, 24 (three of which were known to have died) for
1967, 15 for 1968, and 26 for 1969 (one died; one not banded). Thus,
no more than five to ten percent of eggs laid resulted in fledged young.

Renesting appeared to be common in this species, but the second
nest probably had even less success than the first.

The most dramatic example of renesting occurred in 1964, when about
200 birds--half the population--moved to the newly constructed Akau
Island in March and laid at least 85 eggs, after high tides in late
February destroyed about half the eggs present on Sand. Unfortunately,
continued construction on Akau destroyed the new colony. Only one chick
hatched, which did not survive. An adult found dead in this group had
been banded and streamered earlier in the year on Sand, and several birds

302
600
Adults
500 :
Means D3
eeoeeFX tremes 3
400
Y
[aes
Lu
300
faa)
=
=)
ma
200
100
0
—Y
CE
Lu
faa)
=
=)
ma

J F M A M J J [Nae 0 N D

Figure 95, Means and extremes of semimonthly estimates of Gray-—backed Tern
numbers, Sand Island, Johnston Atoll, 1964-1969.

363

were seen with orange streamers. Streamered birds were also seen in
late established nesting areas on Sand, indicating that these were
birds from the first nesting areas rather than new birds just arriving
on the atoll.

The ultimate cause of nesting failure in this species probably is
competition with Sooty Terns for favorable nesting habitat (see also
Ecological Distribution section).

Annual Cycles

Figures 37 and 95 show the annual cycle for Gray-backed Terns at
Johnston Atoll from 1963 through 1969. Year-to-year variation (Table
66) was similar except for the 1967 and 1969 breeding seasons. For these
two years the first egg laid was about one month early.

After an absence of two or three months, the first birds usually
returned at night only about mid-November; often they were heard several
nights or a few weeks before any were seen. Later they arrived at dusk
and roosted. Arriving birds did not swirl over the island before landing,
but flew directly to the roosting area, settled quickly and remained
relatively quiet through the night. If disturbed, they flew up, calling,
but quickly resettled if left alone.

Numbers present at night continued to build up as the breeding season
approached, and continued to increase until after laying began. In 1966,
the only year in which detailed descriptions of the sequence of events
were made, three or four birds stayed on the dock during the day beginning
ten days before the first egg was laid, and increased to about 40 by the
time the first egg was laid. During this time birds arrived slightly
earlier each evening, and courtship activity increased.

Normally first eggs appeared about mid-January. After laying began,
and throughout the nesting period, no more than a third of the night
population was present during the day; toward the end of the cycle when
numbers began to decline, the day numbers declined more rapidly than
night numbers.

Incubation required about 30 days (1969 data: mean = 30.2 days,
range 29 to 32, n = 25 eggs). Eggs were pipped one day before hatching.
Eggs normally hatched in late February and continued hatching into May
and June. Fledging occurred from mid-March to September. By June,
adults were very inconspicuous on the island, although a few invariably
were present. The unfledged chicks stayed well hidden during the day,
and were visited by the adults only at night. Usually adults tending
chicks were accompanied by a few non-breeding birds, some of which
appeared to be immatures hatched the same year. Others had adult-like
plumage except for speckling on the crown, which may have indicated that
they were subadults in their second year.

After the last chicks fledged, a few birds continued to visit the
island at night for another month or so. These probably were the same
birds that raised the last chicks.

304

Table 66.

Event

Birds first
returned to
Island
Eggs:

First laid
First period

with 20 or
more

Maximum num-—-
ber present

Maximum
number**

Chicks:

First hatched

Last hatched

First fledged

Last fledged

Number
banded

Last birds

seen on Island -

*Except two present 23 Sept.-23 Nov.

1964

1-15
Jan.

16-31
Jan.

1-15
Feb.

16-28
Feb.

145)5)

1=15
Sept.

1965

10 Nov.
(1964)

1-15
Jan.

1-15
Feb.

16-28
Feb.

85

16-28
Feb.

14 June
1-15
Apr.

1-15
July

Le)

p=
Aug.

Nestin

1966

14 Nov.

(1965)

Diller tamns

16-31
Jan.

1-15
Feb.

135

16-20
Feb.

1-15
June

11 Apr.

16=31
July

dls}

16-31*
July

season

1967

25 Nov.
(1966)*

31 Dec;
(1966)

1-15
Jan.

16-31
Jan.

99

ea. 30
Jan.

1-15
May

15 Mar.

10 Sept.

24

After

12 Sept.

1968

(772)

July

Extreme dates of significant events in the Gray-backed Tern
breeding cycles, Sand Island, Johnston Atoll, 1963-1969

1969

22° Oct.
1968

Before 30
Nov. 1968

1-15
Jan.

1-15
Apr. (re-
nesting)

63

7 Feb.

19 May

ca. 1 June

Cae WJuly,

26

After
9 Sept.

**Maximum recorded at any one time; Total number each year estimated 200 or

more.

365

In most years Gray-backed Terns were completely absent from the atoll
or in very low numbers for two or three months--usually from August to
mid-November. But in 1966, two birds arrived back on the atoll in late
September, and continued to roost in the same area near the south-
southeast shore until two days before the first new arrivals appeared
on the old dock on 25 November. The breeding status of these two birds
was unknown, but it seems fairly likely that they joined the new birds
and began roosting with them on the dock. In August and early September
1969, however, several birds appeared on lagoon anchor buoys. An egg
was laid on 9 September; the fate of this out-of-sequence population
was not determined because POBSP personnel left the atoll and ended
their six-year study.

Specimens

Nineteen specimens of Gray-backed Terns have been collected from
Johnston Atoll (Appendix Table 7); these constitute a new published
specimen record for the atoll.

Banding and Interisland Movement

The POBSP banded 271 Gray-backed Terns at Sand Island (Table 24);
23 of these have been recaptured back on the atoll (Table 25). One,
banded as a fledgling on Sand 27 April 1965, was captured some 3,000
nautical miles to the southwest at Wuvulu Island, off the north coast
of New Guinea, in August 1965. No foreign-banded Gray-backed Terns
were recaptured on Johnston Atoll.

At-Sea Distribution

Gray-backed Terns were recorded at sea in the grid southwest of
Johnston only during the months of May, August and October (POBSP,
1967a). These data suggest (1) that during the breeding season this
species usually feeds close to Johnston, and (2) that during the non-
breeding period either is at sea in this area or, more likely, travels
through this area in August going to a distant "wintering ground" and
again in October returning to Johnston.

SOOTY TERN Sterna fuscata
Status

Common breeding species; present from December through August.
Breeds on Sand Island from February through July; previously bred on
Johnston Island. Most abundant spring-summer breeder, numbering about
300,000 breeders producing up to 60,000 fledglings each year; at least
600,000 using the island annually.

\ +. ‘26 ee

306

Ecological Distribution

Johnston Island: Photographs from the 1923 TANAGER expedition
(Wetmore, ms. a) show no distinct Sooty Tern nesting concentrations
and none was mentioned in the reports (Appendix Table 8). There would
be little reason to suspect concentrations, since the habitat was
unusually uniform. Sooty Terns probably nested throughout the Lepturus
on both islands, and non-breeding birds may have concentrated on the
few more open areas such as the small guano digging area on Sand Island.

When the military took over the islands, construction work quickly
destroyed much of the available habitat. Present and past numbers,
however, show no indication that there was a decrease in birds, but
they have had to crowd together much more for nesting, since now they
have available only the original portion of Sand Island--about eight
acres--as compared with the original total of about 50 acres for the
two islands. The density of Lepturus on the original islands un-
doubtedly required wider spacing of nests than is now necessary on the
relatively barren Sand Island, where vegetation is prevented from
becoming too dense by repeated overrunning every few years by men
and equipment and possibly also by the destructive effects of the now
very crowded Sooty Terns.

Unfortunately, data from the war years are almost totally lacking,
but it appears that Sooty Terns must have nested on Johnston Island
through World War II, and afterwards, at least through the 1940's
(Beckham, 1947; Fennell, 1948; Jensen, 1949) and probably until the
mid-1950's (Benson, 1955). Apparently the move to Sand was complete
by 1957, when Moynihan visited the atoll, for he found no Sooty Terns
nesting on Johnston. The reason for the move to Sand Island likely was
the wholesale destruction of eggs on Johnston.

Sand Island: Wetmore (ms. a and b) found Sooty Terns nesting here
in July 1923 (Appendix Table 8). Few probably nested here during
the war years (U.S. Nat. Archives, R.G. 80, U.S. Navy photographs),
because several buildings were on the island and activity was intensive.

In 1957, Moynihan (1957) found Sooty Terns in two groups, one on
each end of the island. The Coast Guard buildings on the western, man-
made end eliminated most of the available habitat there, and repeated
disturbance of the small area probably discouraged all nesting on that
area. POBSP personnel found them nesting only on the original portion
from 1963 through 1969. Harrington (1973 and 1974) also observed nesting
here in spring 1971. Kridler (BSFW, 1973) again recorded them nesting
in May 1973. Amerson saw and heard an occasional Sooty Tern flying at
dusk in November 1973.

Sooty Terns lay a single egg on the grcund without the utilization
of any nesting thaterial. POBSP personnel found Sooty Terns nesting on
almost all the original portion of Sand Island (Fig. 96), with a maximum
area occupied of about 330,000 square feet in 1966 and 1967 (Figs. 35
and 97). The only areas not used were the northwest section, where

*(TTEIIEW *M °Y AQ TeAeT AeM02 JOOF-0Z79 OUI Je Eye
sydeasoj0yd gsqod worz “ar ‘uosazemy *g *y Aq oTesow) 7961 AeW STTOIV UoIsuYyor ‘pueTs]T pues Fo
uofjiod Teurstio |y} UO SeqTs JSeU puNOAS eTQqeTTeAe T[e WSOWTe ezZTTTIN (sq0p AuTq) suzey AJO00S *96 eANn3Tyq

TIOLY NOLSNHOP
GNVISI GNVS

308

possible reasons for their absence were the presence of heavy Lepturus
(which may have been a result, not a cause, of the lack of Sooty Terns),
or the presence of large numbers of ants in this area--possibly more
than any other part of the island--may discourage nesting here. No
objective ant population data were gathered, however.

At least in some years, there were areas too barren of vegetation
to provide Sooty Terns with suitable means for nest orientation.
Ashmole (1963) showed that Sooty Terns will not nest on smooth sand, with
no features by which to orient themselves. The only such bare areas in
1967 were along the periphery of the northeast peninsula, the middle
slope of the east hill, and a spot on the southeast slope under the
southeast inner guywires.

The southwest islet had a few nesting birds in most years, but none
in) 1966 0r 1967 2

Harrington (1973) suggested that the heterogeneity of habitat
features on Sand Island may increase nesting success.

EGGS yf

N
XN cHICKS
°
100 200
———
Reet

Figure 97. Sooty Tern nesting areas, Sand Island, Johnston Atoll, 1967.

309

Nesting density appeared to be highest in areas either bare or
with light vegetation cover, and with enough microrelief in the form
of rocks and other irregularities that the birds could find their
nests, but where there was not enough vegetation to cause increased
spacing of the birds. The only area on Sand where the latter condi-
tions occur was in the dense Lepturus on the southwest portion of the
island (Fig. 98). Otherwise, the spacing of birds appeared to be about
as thick as they will tolerate. Pictures taken from a pole in the
center of the island (Fig. 99) show the distribution of these nesting
birds very well.

Sooty Terns are associated closely with several other nesting
species on the island. The one with which there is a positive inter-
action is the Gray-backed Tern, which begins nesting before Sooty
Terns, and in some cases serves as focal points for arriving sooties
during the swirling, prebreeding phase of their cycle. In no known
case have Sooty Terns actually begun nesting among nesting gray-backs,
but the attraction is fairly well marked in birds settling in the
evenings shortly before they begin nesting. It does not invariably
occur, and the reasons for the variations, and for the gray-backs not
being a nucleus for nesting sooties bear further study. The destructive
effect of nesting sooties on nesting gray-backs was described in that
species’ account.

Sooty Terns and Brown Noddies occupy much of the same habitat,
but little is known of their interrelationships. Noddies generally
nest only around the periphery of the island, in rougher sites than
sooties seem to prefer, but there is considerable overlap, especially
on the east hill (see Figs. 97 and 106). There is some indication that
numbers of Brown Noddy nests on the east hill increase after Sooty
Terns thin out, but little reliable data exist. It may be that counts
of noddy nests on the hill were more complete after the Sooty Terns
left, rather than that they actually increased (see Brown Noddy account).

Sooty Terns also nest among nesting Brown Boobies on the east hill.
Observations indicate that the sooties nested slightly farther from the
boobies than from each other, but measurements were not taken. Several
Brown Boobies stole Sooty Tern eggs and put them in their own nests.
This probably occurred when boobies established nests near already
established Sooty Tern nests, and the tern eggs were within their reach
and were pulled into the nest by the boobies. In 1967 one mateless
male booby used Sooty Tern eggs this way, breaking them all after a few
days of incubation, because the tern eggs were not strong enough to
withstand the trampling of the booby.

Along the south shore, and formerly on the east hill, Sooty Terns
nest close to Great Frigatebirds. Also, formerly on the east hill they
nested close to Red-footed Boobies. In 1967, when the inland great
frigate colony formed, Sooty Terns were nesting densely in the area.
Unfortunately, there are no data to indicate what, if any, reactions
occurred between either the Red-footed Boobies or the Great Frigatebirds
and Sooty Terns. On other islands (Christmas Island, Pacific Ocean,

310

Figure 98. Nesting Sooty Terns cover virtually the entire southwest portion
of Sand Island, Johnston Atoll, 14 April 1969. Most of the
Wedge-tailed Shearwater burrows are under the Lepturus in this
area. The southwest islet and old dock are visible at upper lefq
(POBSP photo by P. C. Shelton).

-
°
ba

s
a

7

Figure 99. Nesting Sooty Terns northeast of the transmitter building and
on the northeast peninsula, Sand Island, Johnston Atolany 30
March 1967 (POBSP photo by P. C. Shelton).

Sime

Schreiber and Ashmole, 1970; and Ascension Island, Atlantic Ocean,
Ashmole, 1963), frigatebirds eat large numbers of Sooty Tern chicks,
but this has never been known to occur on Johnston Atoll. In 1967,
downy Sooty Tern chicks were found huddled under incubating great
frigates in the inland colony. In colonies where frigate feeding on
tern chicks occurs, the behavior is conspicuous, as the frigates hover
over the colony, picking up chicks from on the wing.

Populations

Figure 100 shows semimonthly POBSP population estimates for Sooty
Terns during the years 1963-1969.

In general, maximum numbers present on Sand Island at one time
is about 300,000, which occurs during the breeding peak in late March
and April. Total numbers using the island, however, is at least 600,000
annually. Kridler (BSFW, 1973) estimated 400,000 adults with 192,000
nests in May 1973.

These estimates, however, show gross trends, and cannot be con-
sidered accurate within less than 25 percent; many may be in error by
100 percent. Sooty Tern numbers are practically impossible to determine
accurately, and few of the personnel on the island were sufficiently ex-
perienced to make more than guesses at numbers present. Drastic changes
during a given period are more apt to be caused by changes in personnel
or in techniques used for estimation than to be actual changes in
numbers of the magnitude shown.

The figures presented are supposed to be estimates of the total
numbers of birds using the island during the period in question, based
on numbers seen, day and night (night numbers almost always are greatest),
on egg counts, chick-adult ratios, etc. In general, it appears that
the maximum number ever present is around 300,000, which occurs during
the breeding peak in late March and April.

Another technique used to estimate numbers using the atoll during
the entire breeding season has been the calculation of total popula-
tions based on ratios of banded to unbanded birds in samples. Table
67 shows the results of these attempts.

In summary, it appears that the total number of Sooty Terns using
Johnston Atoll is something over half a million, over half of which may
be present at the peak nesting season in late March through April and
May.

There are no previous estimates or calculations that can be re-
garded as anything more than guesses of the numbers present. Since
all recent attempts to estimate numbers of this species that have been
checked against more objectively calculated figures have been very low,
it is doubted if early estimates for the Johnston population were
anywhere near the actual numbers.

1

?

400

/

300

—— Means

eceoe Extremes

0 @eoeceeee?
eo?
ee
:

=. ceiierereieieiee*

A
Means and extremes of semimonthly estimates of adult Sooty Tern

numbers, Sand Island, Johnston Atoll, 1963-1969.

200
100
Figure 100.

SGNVSNOHL NI SYaaWAN

(o>)
fea
lo

Table 67. Mark-release-capture calculations of Sooty Tern breeding
populations, Sand Island, Johnston Atoll, based on February-
April 1967 recaptures*

No. No. vA Calculated 957%
Manner of Recapture Examined Banded Banded Population C.L.
Netted from air ZS 542 48 342,000 31,000
Guywire strikes 1,193 502 42 388,000 25,000
Ground counts 449 160 36 460,000 +58, 000

*&Calculations based on 163,655 adults banded through the end of 1966.

Wetmore's estimates for 1923, for example, indicate a total
population of 5,500 (3,500 for Johnston and 2,000 for Sand). One
photograph made on Sand Island during that period, however, shows over
1,200 birds. Since this picture probably did not include more than a
fraction of the total, and the total using the island during the day
is never more than a third or half the night population, the actual
number using the island during July 1923 probably was similar to the
numbers using the island during the same periods in the 1960's--15,000
or more.

One would expect that this species might have been reduced by
the disturbance on the atoll, but there is no evidence for this either.
The density of nesting birds is as great as in any colony known in the
central Pacific. There are still small areas not occupied on Sand
Island, however, that cannot be adequately explained in any other way
than that they are not required by the birds. There appears to be an
overfolding of subcolonies, after the initial peak of nesting has
occurred, but this may be entirely renesting birds or late-breeding
young birds that would not require more territory. During the initial
buildup, there probably is more crowding than in most other colonies
where they have room. There is, however, no evidence that the colony
actually suffers from the amount of crowding imposed by the birds having
to nest entirely on the east end of Sand Island. In 1957, Moynihan
found them nesting in two widely separated groups, neither of which
apparently completely covered the respective ends of the island on which
they were found. His description indicates that less area may have
been covered then than during POBSP studies, which may mean the popu-
lation was reduced then and has increased greatly since 1957.
Repeated destruction of eggs, young, and adults, that almost had to
occur throughout the war years, probably reduced the populations con-
siderably and they may still be increasing and readjusting to the
more stable conditions now existing.

Sooty Tern colonies are almost universally unpopular with military
personnel who have to live close to them, so there no doubt was much

314

unrecorded harrassment of thé population on Johnston, which has had to
live closer to humans than any other known. That the colony now is as
large as it is probably indicates a good ability to reproduce rapidly
enough to repopulate the islands soon after afflicting disturbances
were removed.

Nesting success in recent years has been quite high with up to
60,000 fledglings produced each year.

Annual Cycle

Chronology of the major events in the Sooty Tern annual cycle are
shown in Figures 37, 100, 101, and Table 68.

The lowest numbers occur in October and November, when only a few
individuals and small flocks visit the island briefly and irregularly.
These wandering, non-breeding birds almost never land on the island;
there are no data as to the origin of these birds; however, they are
likely Johnston-based birds. Potential breeding birds are first
attracted to the island in late November or December.

110
100
90

——— EGGS
80

eeeee Young
70
60
50

40

NUMBERS IN THOUSANDS

30

20

10

J ie M A M J J A $ 0 N D
Figure 101. Means of semimonthly estimates of Sooty Tern eggs and young,
Sand Island, Johnston Atoll, 1963-1969.

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Sooty Terns at Johnston Atoll breed on an annual cycle. From 1964
through 1971 egg laying usually began in mid-February (+ 15 days) and
reached a peak in March. Laying usually continued into April and vir-
tually ceased by mid-May. Sooty Terns have about a mean incubation
period of 29 days (1969 data: range 27-32 days, 151 eggs) and about a
75-day fledging period. Thus first chicks usually appeared by mid-March
and started fledging by mid-May. Once able to fly, young usually remain
in the vicinity of the colony for about three weeks after which they
departed with at least one parent. Most adults and fledglings leave by
late July but last fledglings usually left by September; a few have
remained into November.

During their nesting season--February through July--Sooty Terns
make up about 97 percent of the bird population on the atoll; they are
almost completely absent from September through November (Harrington,
1973).

Banding data have shown that youngest age of breeding is four years,
but most individuals do not breed until ages six to eight, and a few
probably do not breed until age ten. During their years prior to breed-
ing, adolescents visit Sand Island, but not until they are at least two
years old, and usually not until four or five years old. Most two-year-
old birds return after adults have finished breeding, whereas non-breeding
five- or six-year-old birds return earlier, about one to two months after
most eggs have been laid. Because of very aggressive breeding older
adults, the majority of these visiting adolescents remain flying over
the colony. Some older birds do, however, land in non-breeding areas.
Young adults nest later and in less desirable habitat than experienced
breeders, tend to group with their own peers, and usually have similarly
inexperienced mates (Harrington, 1973 and 1974).

Specimens

In all 236 Sooty Tern specimens have been taken at Johnston Atoll
(Appendix Table 7); these constitute a new published specimen record.

Banding and Interisland Movement

In all, 285,526 Sooty Terns--164,798 adults and 120,728 young--were
banded by the POBSP at Sand Island from 1963 through 1969 (Table 24).
Of these birds, 55,206 were recaptured back on the atoll (Table 25) and
172 were captured elsewhere (Table 26). In addition, 132 banded else-
where were captured at Sand Island.

Movement between Johnston Atoll, and the Line and Phoenix Islands
has been recorded only 17 times. None of these records, however, has
been substantiated by any ‘collections, nor has there been even so much
as a single orange-streamer sighting on land or at sea in that area.
Furthermore, some of the records were obvious errors because of age
discrepancies; others may have been because of band prefix number confusion
by inexperienced banders. Thus few, if any, Sooty Terns from Johnston
move to the Line and Phoenix Islands area.

317

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The remaining Sooty Tern interisland movements have been between
Johnston and (1) the main and northwestern Hawaiian Islands (68 birds
to Johnston, 112 from), (2) the northern Marshall Islands and Wake (58
birds to Johnston, 17 from), and (3) the western Pacific (24 birds from
Johnston).

Gould (1974b) has elaborated on Sooty Tern movement in the north-
central Pacific. As of December 1968, he noted that the POBSP had
processed 1,027 interisland and at-sea movements and 339 at-sea streamer
sightings of Sooty Terns. These represent less than 0.01 percent of the
total number banded. Figure 102 summarizes interisland movements of
Sooty Terns involving Johnston Atoll. Unfortunately, few of these
movements involve known status birds (e.g., breeding, non-breeding), or
elapsed times less than the length of one breeding season. Thus he
suggested that it was impossible to connect the islands of banding and
recapture with meaningful directional arrows.

From recaptures made less than six months after time of banding,
however, Gould found that Johnston-banded adult birds were captured in
the northwestern Hawiian Islands only during June (one bird), July
(three birds), and August (16 birds)-, and that Wake and northwestern
Hawaiian-banded birds were captured at Johnston only during July and
August (three birds each month). Thus during a given breeding season
adult Sooty Terns were moving from island to isiand within the north-
central Pacific only during the summer months, or the latter half of each
breeding season. Furthermore, birds banded at Johnston, the northwestern
Hawaiians, and Wake were captured in the western Pacific primarily during
July (one immature), August (one immature), September (one adult and one
immature), November (one adult), and December (one adult); only one im-
mature was taken in March. This suggests that northcentral Pacific adult
and immature Sooty Terns go to the western Pacific at the end of each
breeding season.

Gould noted that of the 28 POBSP-banded birds (12 of which were
banded at Johnston Atoll) captured in the western Pacific, 20 were probably
driven there by typhoons and tropical storms. By analyzing the origin
and pathways of these storme he was able to determine at least part of
the Sooty Tern non-breeding range. This area was between 5° N and 20°N
and between 170°E and the Philippines.

At-Sea Distribution

The Sooty Tern is the most abundant species in the northern grid,
175 miles southwest of Johnston. There are significant differences
between the annual population curves of the grid and the breeding colony
at Sand Island.

In the grid, (Fig. 103), Sooty Terns arrive in January and build to
high numbers in April and May; density then drops to a lower plateau
until August, but peaks again in September. On Johnston (see Fig. 100),
birds arrive in late December, build to high numbers during the breeding
peak in late March and April, and maintain these high numbers during

Sly)

most of the summer. During the breeding peak most Johnston birds are
laying and incubating eggs. This leaves half the breeding population,

as well as the non-breeding population, free to travel relatively great
distances (probably up to 300 miles) for food. The presence of spring
peak numbers of orange-streamered birds (Table 69) within 50 miles of
Johnston in March and in the grid during April (Fig. 104) support this.
As the young hatch and grow, a greater and greater premium is placed on
feeding nearer to the breeding island thus causing a decrease in grid
numbers to the lower level from June through August. With the fledging
of chicks in August, Sooty Tern numbers near Johnston decreased markedly
especially in October. Immature birds were recorded within the grid from
June through December, with greatest numbers present in September and
October. Orange-streamered immature birds were recorded only during June,
July, and August, with largest numbers in August.

Table 69. Orange-streamered Sooty Tern sightings within a 50-mile-radius
of Johnston Atoll and in the at-sea grid 175 miles southwest
of Johnston Atoll, 1964-1967

Month 50 Mile Radius from Johnston Grid
January 0.18 0.00
February 0.00 0.00
March 0.96 Ons
April 0.26 0.81
May 0.76 O53
June 0.83 0557
July 0.05 0.52
August 5a 0.36
September 0.66 0.00
October 0.00 0.00
November 0.00 0.00
December 0.00 0.00

The September grid peak was probably caused by a transient population
moving through the area. This was supported by the fact that no orange-
streamered birds were observed in September, and that only at this time
were terns most abundant in the southwest half of the grid, which is
farthest from Johnston. In addition, most birds were flying southwest
(48 percent) and south or southeast (26 percent), and not toward Johnston.

The complete lack of diurnal Sooty Tern sightings (see Fig. 103)
but high nocturnal numbers (Fig. 105) in January was a real and consistent
phenomenon. Except for 1967, Sooty Terns were also virtually absent from
the grid during the day in February, but were present in large numbers at
night. At this time large numbers swirled around Sand Island at night,
but disappeared during the day. Many of these were found within a 50-
mile radius of the atoll, but not enough to account for the large
nocturnal numbers over the island.

Food within the grid may not be available during the daytime in
January, and perhaps only very irregularly in February. This was partially

\ ASG B WC

“© f/ iISS/

320

supported by the almost total absence of feeding flocks of any species
at this time. Johnston-based birds may be feeding in other, as yet un- ,
determined, distant areas and returning through the grid at night to i
join the nocturnal pre-breeding swirl over Sand Island. Just why no
Sotty Terns pass through the grid during any part of the day, however,
remains a mystery (POBSP, 1967a; Gould, 1974b).

50

40

3G

20

BIRDS PER 100 SQUARE MILES

10

saber atl ew Winnty bebty: sreAl bog. 69, « Sonmaady

Figure 103. Diurnal occurrence of Sooty Terns at sea 175 miles southwest
of Johnston Atoll, 1963-1967.

321

;
:
2

AUGUST

Figure 104. Orange-streamered Sooty Tern sightings at sea 175 miles southwest
of Johnston Atoll, 1964-1967;“dots: Johnston Atoll, numbers: actual
sightings within each quadrant.

oO (on) ™~S (ee)
(=) i=) oO (=)

WwW
(=)

BIRDS PER 10 HOURS
~~
(=)

ine)
i=)

J F M A M J J A S 0 N D

Figure 105. Nocturnal occurrence of Sooty Terns at sea 175 miles southwest
of Johnston Atoll, 1963-1967.

B22

ELEGANT TERN Thalasseus elegans

Status
Accidental; one specimen record from Sand Island.
Observations

Shelton collected an Elegant Tern at the edge of the Sooty Tern
colony on the west beach of Sand Island 19 April 1969. The bird had
been banded (USF&WS #963-34220) by Monte N. Kirven as a nestling 31 May
1966 at Chula Vista, California, almost 3,000 nautical miles east-northeast
of Johnston. When collected, it was in winter plumage, but had almost
completed a molt of its flight feathers (outer remiges were not quite full
length). The presence of old ruptured follicles in the ovary and a con-
voluted oviduct indicated that the bird had bred previously. Although the
bird was quite thin, its stomach contained a fish 103 mm long.

Specimens

This is a new specimen record- for Johnston Atoll, as well as for
the central Pacific Ocean.

Elegant Terns breed along the coast of Baja California and in the
Gulf of California, wandering in fall north to central California, but
wintering along the Pacific coasts of Peru and Chile (AOU, 1957). There
are no other known records from the central Pacific.

BLUE-GRAY NODDY Procelsterna cerulea
Status

Uncommon visitor; three specimen records and one sight record from
Johnston and Sand Islands.

Observations

Wetmore (ms. a) collected two non-breeding adult female Blue-Gray
Noddies and saw a third 13 July 1923 "among clumps of grass below a low
hill" on Johnston Island. He further noted (ms. b) that "both of those
secured were nonbreeding birds, and I suppose that they had come in from
the sea to rest.'"' A third specimen, a male, was taken at sea.iabout
50 miles west of Johnston (16°45'N, 170°20'W) on 20 July 1923 as the
party was leaving for Wake Island.

There were no further records of this species from Johnston Atoll
until 8 May 1967 when Shelton collected a young female as it roosted on
a broken concrete bunker near the southwest islet.

Another bird appeared at dusk 12 April 1969 and returned to the
island at least ten evenings from that date through 3 May, when it was

523

last seen. For the first few nights it roosted on large rocks along the
south shore, east of the Great Frigatebird colony, with Brown Noddies and

a few Gray-backed Terns. Later it chose a large rock on the south side

of the causeway east of the Signal Building, quite alone. For the last

ten days it was seen only on the southwest islet where it may have roosted
in ammunition slots in the old gun emplacement. Here it was caught, banded
(652-49679), measured (Bill: 23mm; Wing; 171mm; Tail: 90mm; Tarsus: 27mm),
and released on 24 April. It had no brood patch and was just beginning to
replace its primaries and rectrices.

Specimens

The three Johnston Atoll specimens and the one at-sea specimen, noted
above and in Appendix Table 7, are deposited in the USNM. These constitute
a new specimen record for the atoll.

All three specimens, and the banded bird, belong to the light
northern race, P. c. saxattlis,indicating that they came from high, rocky,
northwestern Hawaiian Islands where they breed.

Banding and Interisland Movement

One Blue-Gray Noddy has been banded at Johnston Atoll (Table 24).
No interisland movement of banded birds is known.

At-Sea Distribution

No Blue-gray Noddies were recorded in the grid southwest of Johnston
Atoll. The species is, however, considered to be an uncommon visitor to
the Johnston area. Wetmore's 1923 observations on the atoll and at sea,
as noted above, and recent POBSP observations, suggest that Blue-gray
Noddy visits may be less common now that Johnston is inhabited.

BROWN NODDY Anous’ stoltdus
Status

Common breeding species; present year-round. Breeds on Sand Island
from February through September; bred on Hikina Island for the first time
in 1973; previously bred on Johnston Island. Second most abundant spring-
summer breeder numbering about 3,000 maximum and producing about 1,000
fledglings each year. Nesting is primarily on the rougher areas, peripheral
to the nesting Sooty Terns.

Ecological Distribution

Hikina Island: Brown Noddies nested here in May 1973 (BSFW, 1973);
eight nests containing eggs and 40 adults were observed. Amerson reported
two flocks of about 100 adults each roosting here in November 1973; two
partially constructed nests, containing no eggs, were seen in Sesuviun.

324

Johnston Island: About 1,500 Brown Noddies were observed nesting
here in July 1923 by Wetmore (ms. b). Fennell (1948) saw a few at the
end of the runway in 1948. None has been recorded since, most likely
because of human disturbance.

Sand Island: Wetmore (ms. a) found about 800 here 10-17 July 1923
and described the colony as follows.

About half have eggs; the rest young from newly hatched to
grown. The newly hatched young are about evenly divided
between light and dark.

The nests of these birds are interesting. Some eggs are laid
on bare sand or rock, others are deposited on a platform of
grass with a slight central depression. Frequently the nest
cavity contains ornaments in the form of entire shells, bits
of larger shells, the calcareous opercula of cat's eye
mollusks, bits of bone, a cranium or a mandible, or an odd
shaped piece of wood. It is always of interest to look over
the collection and before now I have been guilty of filching
some pretty shells from the lot.

Mating actions are of interest. The male with neck erect and
bill pointing down patters rapidly off across the sand perhaps
in a semicircle to pause finally and look down as though at a
nest. He then returns to his mate and the two rise and swing
off in erratic circles with a synchronism of movement that
holds them always 15 or 18 inches one above the other.

Moynihan (1957) observed approximately 200, but only a few repro-
ductive activities in April 1957; pairs held territories on either side
of the causeway (alternating with Gray-backed Tern territories) and on
the periphery of the Sooty Tern colony on the original portion. Davis
(1962) saw relatively few on Sand Island in 1959-60, even though it was
not inhabited by man at the time. He noted that they isolated themselves
from the Sooty Terns. POBSP personnel found Brown Noddies to be common
from 1963 through 1969. Amerson also found them here in November 1973.

Brown Noddies on Sand Island nest: almost exclusively on the ground.
The only exceptions were those few that nested on the old dock before it
was burned in late 1967, and the occasional nest found on a concrete
pillar in the intertidal zone of the south shore. Primary nesting areas
were on the periphery of the island, in rougher, more rocky areas than
Sooty Terns, and peripheral to them (Figs. 43 and 106).

PLOT DATE ADULTS EGGS NESTLINGS LOCALS IMMATURES PLOT DATE ADULTS EGGS NESTLINGS LOCALS IMMATURES

A 8 15
15 25
23 25
30 ?
B 8 160
Iey es INe{0)
230 a5 0
30 ?
C 8 26*
15 30
23 30
30 ?
D 8 60
15 75
23 50
30 ?
E 8 40
15 25
23 40
30 2?
F 8 50
15 60
23 75
30 ?
Figure 106.

NUMBER OF NUMBER OF

9 0 0 0 G 8 60 38 0 0 0
10 0 0 0 aye llo}0) 44 1 0 0
14 0 0 0 23), 100 56 1 ] 0

4 0 0 0 30 2 41 6 € 0

104 0 0 0 H 8 6 2 0 0 0
104 1l 0 0 15 1 1 0 0 0
81 5 12 0 23 1 1 0 0 0
50 15 22 0 30 ? 0 1 0 0
18 0 0 0 I 8 1 1 0 0 0
13 0 0 0 15 1 1 0 0 0
17 0 0 0 23 1 1 0 0 0
12 5 0 0 30 ts 1 0 0 0
28 1 1 0 J 8 12 10 2 4 0
31 0 2 0 15 15 11 2 4 0
30 1 2 0 23 10 8 1 3 3
21 3 2 1 30 is 7 0 3 3
14 0 0 0 K 8 100 80 0 3 0
11 0 0 0 15) 50 86 6 3 0
15 0 0 0 Zoe ELSON 4 7 8 1
30 0 0 0 30 2 64 11 23 1
14 0 0 0
18 0 0 0
27 0 0 0
30 0 0 0

Brown Noddy nest distribution and abundance, Sand Island,
Johnston Atoll, April 1967.

325

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326

Nests were described by Wetmore in 1923. Then as now, the nests
varied from nothing at all--egg laid on bare sand--to loose aggregations
of vegetation, shells, and bird bones with a shallow central depression.
Table 70 shows the composition of several 1968 nests. An interesting
addition to the naturally occurring materials were the orange plastic
streamers used by the POBSP for marking birds; these frequently occurred
in the nests. Nests were placed on bare sand, in Tribulus, or Boerhavta
clumps, or more rarely in the top of a Lepturus clump (Figs. 107 and 108).

Roosting Brown Noddies generally preferred exposed peripheral areas,
including the southwest islet, northeast peninsula, and the east slope.
In addition they roosted on the guywires, guywire anchors, and naviga-
tional aids throughout the lagoon. The latter habitat group was heavily
used by non-nesting birds, including recently fledged young. During the
winter, when breeding activity was minimal, roosting was restricted to
the most exposed of these areas, especially the peninsula, islet, east-
southeast slope, and guywires. At the onset of reproductive activity,
more birds occupied the nesting areas themselves.

The relationship between Brown Noddies and Sooty Terns is poorly known.
Noddies begin nesting earlier than sooties, and their first nesting areas
almost invariably are the southwest islet, the old dock, and the northeast
peninsula, indicating that the species has natural affinities for more
isolated areas.

A photograph taken in 1923 shows Brown Noddies roosting thickly on
the bare spot on Sand Island where guano was dug (near center of southwest
area). No Brown Noddies were ever seen nesting this far inland during
POBSP studies, but the area always was covered thickly with nesting Sooty
Terns at that time of year (July). With Sooty Terns able to spread out
more in 1923, Brown Noddies apparently also occupied areas (particularly
in the interior) to which they do not now have access because of the
density of Sooty Terns. Rough peripheral areas were used by the Brown
Noddies in 1923, as indicated by Wetmore's notes. Relative numbers in
the various areas were not precisely indicated.

There was only one instance of Sooty Terns being attracted to Noddies
during the time sooties were swirling and first settling on the ground.
On 31 December 1966 about 50 sooties landed among noddies roosting near
the frigate colony along the south shore. Conflicts for nesting space
did not occur until later in the season, when both species were near
their breeding peak. In September 1967, Shelton observed an adult Brown
Noddy thoroughly thrash an adult Sooty Tern, apparently in defense of a
small noddy chick. Brown Noddies made at least a small part of their
living pirating food from Sooty Terns and other birds. They were observed
flying over the Sooty Tern colony looking for adults feeding young. The
noddy then swooped low and pulled the fish or squid from the Sooty Tern.
They have also been seen snatching food dropped by birds being chased by
Great Frigatebirds.

As Sooty Terns abandoned the northeast peninsula, the east hill, and
southeast slope in early September 1966, there was an apparent move of

ial used by Brown Noddies, Sand Island, Johnston Atoll; observations by

. Harrington, about 1 May 1968

Nesting mater

B.A

Table 70.

Nest sites

aouezind90

snjinqiaf ut ‘Sedots
3seey Nnos—jseqg

snjnqidiy ur Sedots
qyseoyjnos-jseyq

pues o1eq
uo S‘edots 4seq

yoreq
AyD01 uo opts 4see
“eseq ‘e[nsurtueg

untanseg jo yo ued
0} }xoeU YyoRreqg
AYDO1 uo apts 4see
S‘eseq ‘e[nsutueg

yoqed v1OAanag
uo ‘eTnsutuseg

yoqed 07 0AanDag
uo ‘eyTnsutueg

a3pra
Teiod uo eTpnsutued
JO aloys 3SseM

espt2
[Teioo uo epTnsutued
JO ei0Yys 4seM

as3pr1
[e109 uo eTnsutued
JO azoys say

aspra
Te10d uo eTnsutued
JO eioyus 4SseM

dunto

sningdeT yo doq uo
UOTIV9S 4Se9YyIA0U
jo ai0ys yIION
DLADYxeOg UT
uoTjoes 4SPeyzI0U
JO @ez70YS YUIION

Material

397, 10075 8

100%

50%

100%

100%

20%

702%

Tribulus

10%

80%

Lepturus

wet N
x
fon)
+
Dw
oO
i=)
rc

x

j=)

i=)

_

Ss
Saas
Sis
Swoak
See ai
C0 a5 A)
Wi SS)
By = 18s)

10%

30%

90%

100%

25%

Coral & Shell

2%

17%

Bird bones,

Co

1%

crab leg,

=

9
°

eb)

eggshell

Plastic

207

debris

100%

No Material

327

328

(
f ~

Figure 107. Brown Noddies and Sooty Terns on the southwest portion of Sand
Island, Johnston Atoll, July 1923. Johnston Island is visible
in the background (B. P. Bishop Museum photo by Chapman Grant).

\ Sane

Figure 108. Brown Noddies nesting and roosting on the southwest islet, Sand
Island, Johnston Atoll, 9 May 1967 (POBSP photo by P. C Shelton)

329

roosting noddies from the guywires back to the ground in these same
areas. Accurate before and after counts were not made, however, and

in 1967 it did not appear that the change was as significant. Previous
observers had noted an increase in noddy nests on the east hill after
Sooty Terns left, but were unsure the increase was real or resulted from
more accurate counts after the other birds thinned out. Also in early
September 1966 there was an increase in Brown Noddy guywire strikes. In
September, nine were killed while earlier in the summer no more than two
per month were killed.

The relationship between Brown Noddies and Gray-backed Terns is
also poorly known. In 1965 and 1966, Brown Noddies crowded Gray—backed
Terns from the distal portion of the old dock, where the gray-backs had
begun roosting and nesting before the build-up of noddies. In December
1966, gray-backs were roosting on the entire dock early in the evening,
but noddies came in later, crowding the gray-—backs to the proximal one-
third of the dock. On the peninsula in 1965, at least, the two species
apparently nested fairly close together without conflict.

Populations

Using the Brown Noddy population level that existed in 1923 as a
norm, populations appear to have declined markedly during and after the
war years, but seemed to have regained that level during the 1960's.
Wetmore's estimates of 1,500 on Johnston and 800 on Sand Island agrees
closely with POBSP estimates for July, except that all birds were on Sand
Island. Moynihan's observation of 200 birds in 1957 and Davis' observa-
tions of "relatively few" in 1959-1960 indicate that the population must
have been severely depleted, probably as a result of successive nesting
failure during periods of intensive military activity on the islands.
Recovery was rapid after this period, for by 1963-1964 the population had
risen to over 2,000 birds, and to about 3,000 through 1969. This can
only be attributed to the nearly non-disturbed condition that exists on
the original portion of Sand Island.

Figure 109 shows mean semimonthly population estimates by POBSP
from 1964 through 1969. Maximum numbers present at any one time was
about 3,000 adults. Total numbers using the island annually was estimated
to be between 6,000 and 12,000 birds. This species is never entirely
absent from the atoll, and numbers drop significantly only in mid-winter,
with relatively stable numbers throughout the rest of the year.

Variations between day and night numbers are significant, and
differ between seasons. During the mid-winter low, most birds are
present only at night, with only a few (mostly recently fledged young)
on the island during the day. Adults (and presumably several-month-old
young) arrive back on the island mostly within the period one half to one
hour after sunset. These night roosting populations are difficult to
estimate, and they may be larger than the figures indicate. It is pos-
sible, if not probable, that the number of Brown Noddies using the island
does not decrease significantly during the winter, but that the length

5,000
Adults

—— Means
4,000

ec®
esees Extremes ie 5

1,000 +

—— Eggs, 1965-1969

seeee - Young, 1964-1969

NUMBERS
w
=)
=)

J Peal A M Ua) A S 0 N vad

Figure 109. Means and extremes of semimonthly estimates of Brown Noddy
numbers, Sand Island, Johnston Atoll, 1963-1969.

Sie

of time spent away from the island by individual birds increases.

This gives the impression of fewer birds simply because there are fewer
at any given time, although the total number has not decreased. As the
nesting season approaches, the numbers present during the day increased
markedly.

Estimates of the total number of birds using the island were crudely
calculated from banding data.

In March 1967, two samples were counted to determine the proportion
of birds on the population that was banded. On 7 March, 81 birds were
counted on the peninsula and southwest islet, and of these 81, 27 birds
(33 percent) were banded. A second sample on the islet only, made
23 March, showed 17 (34 percent) of 50 birds to be banded. Since the
two samples were not mutually exclusive (parts or all of both were from
the islet), they cannot be added together. Thus, the second, smaller
sample from the islet alone, was deleted. Samples should have been taken
all around the island, on the same day if possible, with the total number
of birds counted at least several hundred.

Banding data show that young were seldom recaptured until they were
three years old. Therefore the number of previously banded birds used
for calculating a population estimate includes all adults banded through
1966 and all young banded through 1964, a total of 2,961 birds. The
calculated population from these figures was 8,880 (42,790) or 6,090 to
11,670.

There are, of course, serious limitations to these figures, chief
among which are the following: (1) the actual rate of recruitment of
young birds into the adult population was not known; (2) the samples
were small (confidence limits wide) and were taken from only two areas
on the island, which may not adequately represent the entire island |
population; (3) the turnover in adults through immigration, emigration,
and deaths was not known; (4) banding was not uniform in the age groups
represented.

The young produced before 1964 were either not banded at all (those
hatched before 1963) or very lightly (1963). These birds would be treated
as adults as they returned to the population, which they should have done.
Even the 1963 birds should have been on the island in fair numbers in
1966, when adult banding was at about the same level it had been in
earlier years. The 1964 cohort was banded more heavily than any other
in this sample, and the presence of large numbers of these in the sample
would inflate the percent banded, which would tend to make the estimate
lower than reality.

In summary, the population of Brown Noddies on Johnston Atoll (Sand
Island) probably possesses the following characteristics. Maximum number
present at any one time: about 3,000 adults. Total number using the
island 1963-1967: 6,000 to 12,000, plus 1,000 young produced each year
in 1965 and 1966.

332

Productivity: Nesting success from 1964 through 1969 averaged
about 1,000 young fledged each year. This is reflected in the number
of young banded each year:

Year Number Young Banded
1964 763
1965 TOV
1966 807
1967 650
1968 1,440
1969 806

The 1967 figure of 650 is minimal since no young were banded after early
September. The 1969 figure of 806 is also minimal for 46 nestlings and

159 eggs remained in early September; the total number of young produced
probably was about 1,000. In all, 2,222 eggs were laid from March 1969
through early September 1969, suggesting a nesting success of approximately
38 percent for that period.

Egg and chick mortality records were kept for a little over a month
during the summer of 1964 on study plots on the south shore and northeast
peninsula. These figures showed losses to be 15 to 20 percent of the 175
eggs present and about 25 percent of the 100 chicks studied being lost
within the first month after hatching.

Other figures for egg and chick loss are not available, but this
species probably has a lower rate of loss than most other birds on the
atoll. Brown Noddies generally appear to be more aggressive in defense —
of their nests than most other species, and losses to abandonment and
natural forces are considered low. Exceptionally high tides do, however,
occasionally inundate nests near the water, especially on the islet and
northeast peninsula. Guywire maintenance work for the LORAN-C tower was
not particularly disruptive of this species in 1966, but they were said
to suffer most in early June 1964. Quantitative data were not recorded
either year. Predation by two cats disrupted two study plots in August
1964; again quantitative data on losses were not recorded.

Color Phases of Young: Brown Noddy young range in color from pure
black to pure white. Whether discreet categories exist is debatable,
and the genetic basis for the variations is totally unknown. Records
kept in 1964, 1965, 1966, and 1967 show that of 573 chicks examined, 34
percent were white, 29 percent were gray, and 3/7 percent were black.
Whether all observers used the same points to distinguish the categories
is not known, but the data are fairly consistent from sample to sample.
There was no evidence of seasonal or year-to-year changes in the ratios,
and no evidence that it varied on different parts of the island. Incuba-
tion periods recorded in 1964 did not vary with the color phase of the
chicks.

3)3)5)

Annual Cycle

Figures 37 and 109, as well as Table 71, show the important aspects
of the Brown Noddy cycle on Sand Island from 1963 through 1969. row
Noddies have perhaps the longest breeding season of any species on the
atoll. A few eggs or chicks may be found at any time, but the largest
numbers occur from February through September. At other times, eggs
occur irregularly and are widely scattered.

In most years courtship and nest site selection became noticeable
by the end of January, and eggs were laid in February, only a few at
first, but increasing to ten to 25 by the end of the month. Egg produc-
tion increases through March and April; numbers hold fairly steady until
July and decline through August and September, the latter being the last
month in which more than a few scattered eggs are found.

The mean incubation period for 93 eggs was 35.6 days (range 30 to
41 days, 1964 data). Fledging required about six weeks. Extensive growth
and development data were taken on this species in 1964, but these data
have not been analyzed.

Daily egg laying, hatching, and losses were recorded for the end of
the 1964 and 1966 breeding seasons on the northeast peninsula (Table 72).
These data show that the number of new eggs, and new chicks per semi-
monthly period, was about the same for these two years, indicating little
annual variation in the breeding season.

Individual cycles have not been followed through the breeding season,
but a few banded adults were recaptured on eggs or chicks at about the
same time in successive years. Also, there were several examples of birds
found on eggs at two different times during the same year, sometimes
separated by more than the normal incubation time. These probably were
birds renesting, but there was no evidence as to what happened to the
first egg or chick in these cases. Two successful nestings in the same
year cannot be ruled out on the basis of POBSP data, although this is con-
sidered unlikely. More careful work with individually marked birds needs
to be done.

Several three- and four-year-old birds (banded as chicks in 1963 and 1964)
nested in 1967. Table 73 shows that all of these were taken in the last
half of the breeding season, although the numbers handled were small in
all cases. Further data are needed.

In summary, present data are insufficient to show why the breeding
season is extended; it could be because of renesting by birds that lost
eggs or chicks, multiple nesting, nesting of younger birds, or even lack
of nest space.

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Table 72. Daily egg laying, hatching, and losses for Brown Noddies,
Sand Island, Johnston Atoll, July-September, 1964 and 966

1964 1966

New New Eggs Eggs New New Eggs Eggs

Period eggs chicks lost remaining eggs chicks lost remaining
July 1-15 27 41 10 72 26 by? 2 Ca.) OO.
16-31 13 7a\s) 2 44 18 t6.eCa.20) - Sea (42
Aug. 1-15 9 16 14 22 8 20 9 21
16-31 4 7 2 17 if 9 2 Le
Sept. 1-15 i 10 Z 6 0 7 2 2
16-30 0 | 3 0 0 2 0 0

Table 73. Known age Brown Noddy recaptures taken from eggs or chicks,
Sand Island, Johnston Atoll, 1967

3-yr .-olds 4-yr.-olds
(1964 chicks: 763 banded) (1963 chicks: 124 banded)

Date Total handled No. Percent No. Percent
4-5 May 10 0) 0 0 0
30 June 9 3 33 0 0
17-24 July iat 6 55 if 9
8 Sept. 7 if 14 2 29

37 10 DLA 3 87%

Banding data suggest that most adults return each year. An insuf-
ficient number of banded one- and two-year-old birds have been recaptured
to determine how many of these birds remain on the island, but at least
some do. They probably spend a greater portion of their time on in-
accessible roosts, making their capture more difficult. Data presented
earlier indicate that many, if not all, three-year-olds return to the
island and nest, probably mostly late in the season.

Table 74 shows available data on primary molt in Brown Noddies on
Sand Island. March and April birds, none of which was known to have eggs
or chicks at the time, were nearly finished replacing their primaries,
indicating that this occurs shortly before egg laying. Birds taken on
eggs during June or later had, however, begun to replace their primaries
before the chicks had hatched. The process apparently continues while
the chick is growing and is completed shortly before egg laying in the
next year. The consistency of occurrence of molt during incubation
argues that this is not a phenomenon confined to renesting birds who

336

lost eggs or chicks earlier in the year, but that it occurs in all
incubating birds. Also, the young adults, hatched in 1964 and nesting
in late 1967, all showed primary molt while they were incubating; it
is virtually certain that these were not renesters.

Table 74. Primary molt scores of Brown Noddies, Sand Island, Johnston

Atoll
Primary Molt score
(Ashmole, 1962)
Date Status or Source No. handled Mean Range
March 1967 Returns 6 82.5 56-100
Guywire kills 5) JOO 100
Apr. 1967 Returns 3 99 96-100
Guywire kills 3) 200 100
June 1967 Returns, on eggs (1964 young) 3 10 6-12
on chicks (banded as
adults) 5) 18 10-26
all other 15) 10.6 0-22
July 1967 Returns, on eggs (1963-64
young) i HORS 0-22
(banded as
adults) 3 24 18-32
on chicks (banded as
an adult) 1 24 9
1-15 Sept.
1966 Guywire strikes 4 2D.) 6-36
16-30 Sept.
1966 Guywire strikes 5 40 26-52
21 Sept.
1966 Returns 2 17 2-32
Sept. 1967 Off eggs ie 24.6 3-41
From air 6 PEST) 1-38
5 irregular or interrupted
Specimens

The POBSP has collected 78 specimens of Brown Noddies from Sand
Island. Wetmore took six skins from the islands in 1923 (Appendix
Table 7). All these are in the National Museum of Natural History
collections. These constitute a new published specimen record.

Banding and Interisland Movement

From 1963 through 1969, the POBSP banded 7,979 Brown Noddies on
Sand Island (Table 24). Of these, 882 recaptures have been made back
on the atoll (Table 25).

In all, there were 31 interisland movements of Brown Noddies
involving Johnston Atoll (Table 26): 24 banded on Johnston have gone
elsewhere and seven banded elsewhere have been captured on Johnston.
Sixteen such movements were between the northwestern Hawaiian Islands
and Johnston, while 15 moved in different directions, including islands
in the western Pacific and islands near or south of the equator.

At=Sea Distribution

Within 100 miles of Johnston Atoll and compared with other Johnston
breeding species, Brown Noddies were fifth in abundance during spring
months. They were absent from at-sea observations from December through
February, and July (POBSP, 1965). In the grid area 175 miles southwest
of the atoll (Table 21), they were absent from January through July,
excepting April (POBSP, 1967a).

BLACK NODDY Anous tenutrostrts
Status

Uncommon breeding species; present year-round. A few pairs nest
on Amaranthus bushes on Sand Island each spring and produce about three
fledglings; up to 100 visiting non-breeding birds roost on antenna guy-
wires. A few nested in Casuarina trees on Johnston Island in 1973.

Ecological Distribution

Johnston Island: Wetmore (ms. a) wrote in 1923: "19 July: One
reported by Grant. I thought I heard one calling on first arrival here
but was not certain.'’ POBSP personnel observed three Black Noddies
roosting here 23 June 1967. Amerson recorded eight adults, one flying
immature, and three inactive nests in the Casuarina trees next to the
tennis courts in November 1973.

Sand Island: Moynihan (1957) recorded 6-10 April 1957: "Some
five or six Black Noddies spent considerable periods of time resting
in two low bushes on the side of the causeway. They showed no reproduc-
tive behavior of any kind."

POBSP personnel found them nesting and roosting only on the original
portion of Sand Island from 1963 through 1969. Amerson observed a few
roosting at night in November 1973. All but one Black Noddy nest were
placed in Amarmthus plants less than two feet high, growing in the areas
southwest and southeast of the transmitter building (Figs. 44, 110, 111,
and 112). One pair in-1967 nested in a high clump of Lepturus southeast
of the transmitter, near other nests in Aimaranthus. Nests seven to ten
inches in diameter, with a shallow center cup about three inches in
diameter, were constructed mainly of Tribulus, with small amounts of
Boerhavia and Lepturus; excreta helped hold each nest together.

Most roosting birds used the northeast outer guywires, but others
used such places as the northeast and southeast inner guywires, rockpiles,

338

the old pier, the fence around the transmitter tower, the rocky west
peninsula shore, and Yournefortta and Amaramthus bushes. No birds were
ever reported from the west portion of Sand Island during POBSP studies.
The presence of good roosts probably attracts more birds to the island
than were present before military occupation.

In 1966 a few Black Noddies roosted on a dead Amaranthus bush near
the old pier throughout July, but abandoned the perch in August when
Sooty Terns stopped using the surrounding area. At all other roosting
places Black Noddies were in company with large numbers of other species,
mostly Brown Noddies.

Populations

Figure 113 shows the means the extremes of semimonthly population
estimates of Black Noddies from 1963 through 1969. The population is
composed of two elements: (1) a small resident breeding population num-
bering no more than 13 pairs, and (2) transient birds numbering about 150
which are probably non-breeders from the northwestern Hawaiian Islands.

(&) Roosting and Nesting

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s (&) 1963-65 ®
we ® 1967
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Feet

Figure 110. Black Noddy nesting and roosting areas, Sand Island, Johnston
Atoll, 1963-1967.

3)3)9)

Figure 111.

Figure 112.

Black Noddy nest in Amaranthus plants south of the transmitter
building, Sand Island, Johnston Atoll, 7 May 1967. Sooty Tern
adults and chicks nearby (POBSP photo by P. C. Shelton).

ri

< > =.

> ake

Black Noody nest with 12-day-old chick in Amaranthus plant south
of the transmitter building, Sand Island, Johnston Atoll, 4 June
1969 (POBSP photo by P. C. Shelton).

ts
ye

340

Although neither Wetmore nor Moynihan found evidence of this species
nesting in 1923 or 1957, it is possible that nesting occurred at least
irregularly in those years. A successful breeding season could have been
completed and most birds dispersed before Wetmore's July 1923 visit, and
a breeding season could have begun after Moynihan's April 1957 visit.

The ability of this species to nest in low vegetation, especially in
Lepturus, here and elsewhere suggests that the lack of bushes or high
vegetation before military occupancy would not necessarily have prevented
breeding. The presence of more suitable bushes since military occupancy,
however, appears to have increased the breeding population.

100

Adults

30 —— Means

eeeee Extremes

26

—— Eggs

eieieiere Young

J F M Ave J J A S 0 N D

Figure 113. Means and extremes of semimonthly estimates of Black Noddy
numbers, Sand Island, Johnston Atoll, 1964-1969.

Ore Mm @® &

NUMBERS

341

From 1964 through 1969, excluding 1966 when none nested, an
average of seven eggs (range three to 13) were laid each year (Table
75). An average of four eggs hatched (range three to seven); hatching
success was 60 percent. An average of only three chicks (range two to
six) fledged during these same years. Thus nesting success during that
period was 46 percent.

Table 75. Nesting success of Black Noddies on Sand Island, Johnston
Atoll, 1963-1969

Eggs Hatched Chicks Fledged
Year Eggs laid No. % No. % of eggs laid % of eggs hatched
1963* -- -- -- 4 -- --
1964 13 7 54 6 46 86
1965 8 3 38 2 25 67
1966 0 0 -- 0 -- =
1967 3 3 100 3) 100 100
1968 7 5 7h 3 60 43
1969 6 4 67 3 50 75
Totals**37 22 SOR a7 46 7

*Data incomplete, observations began 7 July.
*? Excluding 1963.

The maximum number of eggs laid in one year was 13 in 1964; if
all these were first eggs (no renesting), then there would have been a
maximum of 26 breeding birds that year. Whether more of the roosting
birds, particularly those roosting on bushes rather than on guywires,
were potential nesters which failed to nest for lack of suitable
habitat is not known. Separate counts and closer observations should
have been made on these birds.

POBSP population estimates of roosting Black Noddies were based on
counts made in the various roosting areas, or of birds encountered during
routine operations on the island. It is suspected that birds frequently
were missed, especially at times of year ‘when other birds were abundant
and occupied most of the observer's time. Few birds came in to roost
until heavy dusk, and maximum numbers were not present until well after
dark. They could have been easily confused with Brown Noddies that
roosted on many of the same locations.

342

Only in 1967 was there an attempt to determine how many birds may
have used the island during the year. Table 76 shows calculations based
on the number of streamered birds observed on the roosts. The slow
gradual increase in calculated numbers present from 19 March through
16 April (when no additional birds were marked) indicates that the
turnover rate was slow. Also, the figure for 27-29 August was not sig-
nificantly different from those derived in March and April, although no
birds had been handled since May. The presence in August and September
1967 of several birds with streamers put on before 1967 indicates that
many of the same birds may return year after year.

Table 76. Calculations of numbers of Biack Noddies using Sand Island,
Johnston Atoll, 1967

No. No. No. previously Calculated
Date counted marked marked population (95% of C.L.)
19 March 49 6 15 123193
31 March 26 4 aS 98t92
7 April 62 7 15 133t90
16 April 65 5 15 195*168
20 April 83 y 18 1664104
30 April 7a 5 19 2694234
27-29 August 25 4 25 156t143

Although the confidence limits for the estimates in Table 76 are
large, the consistency strengthens their validity. The conclusion drawn
from these data is that the number of Black Noddies using Sand Island
during a year is presently approximately 150 birds, which is about twice
the highest number seen at any one time.

It is fairly likely that roosting Black Noddies have increased
since 1923. Daytime populations are never more than a few birds, and
it is not likely that Wetmore's party overlooked a roosting population
which was present only at night. Moynihan did not visit Sand Island
at night, when maximum numbers would have been present; thus it is pos-
sible that the 1957 population was as large as that found in the 1960's
when daytime numbers also were seldom more than a few birds. It is
possible that few birds used the island until military construction and
subsequent abandonment increased available roosting habitat.

Nearly all birds roosting on guywires during POBSP observations were
non-breeders. In February, March, and April 1967, 18 birds were handled;
17 from the northeast outer guywire and one from the shore of the penin-
sula. All of these birds were in some stage of primary molt, except for

343

one bird which had all new feathers (Table 77). None had bare brood
spots. Presumably most of these were from the northwestern Hawaiian
Islands (one had been banded there).

Annual Cycle

The annual cycle of Black Noddies at Johnston Atoll is shown
in Figures 37 and 113.

Breeders: The nesting cycle shifted slightly from 1964 through
1969 (Table 78). The cycles were similar in 1964 and 1965. In 1966
they did not nest at all. In 1967, first eggs were laid a month and
a half later than in 1964 and 1965, while first eggs for 1968 and 1969
were laid a month later than in 1964 and 1965.

In 1964 and 1965, birds appeared on the nesting areas in February,
and courted and built nests by the end of that month; first eggs, as
well as the egg peak, occurred in the first half of March. First eggs
appeared in late April in 1967 and in early April during 1968 and 1969.
Eggs occurred usually through early July, but did extend through August
in 1964. The few eggs laid later in the year--especially in 1964--
may have been replacements for eggs lost earlier, but no data were ob-
tained. The data from the end of the season in 1963 indicated that the
cycle that year may have been about the same as in 1964 and 1965.

No eggs were known to have been laid in 1966. Only a few birds
were present during the normal time of initiation of nesting, and they
were seen only sporadically on the Amaranthus bushes on which nesting
normally occurred. More intensive activity by POBSP personnel banding
Sooty Terns may have, however, discouraged the Black Noddy nesters.
Disturbance by the antenna maintenance crew in May would have discouraged
only late nesters.

Individual breeding cycles of Black Noddies are among the shortest
of any species on Johnston Atoll. Incubation of three 1965 eggs and
seven 1969 eggs averaged 34.2 days (range 33 to 37 days), and fledging
of two 1965 chicks and three 1969 chicks averaged 38 days (range 34
to 46 days). First chicks appeared in early April in 1964 and 1965,
and late May in 1967, 1968, and 1969. Young were present normally into
August, but have remained until early September. Young usually stayed
around the nest area for at least a few weeks after fledging, but
quickly became difficult to distinguish from adults.

One young bird, banded as a nestling in 1964, returned and nested
in late April 1967. Thus young of this species may nest as early as
three years old. More data would be necessary to determine if this is
the normal age of first breeding.

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Non-breeders: The seasonal fluctuation in numbers of roosting birds
is difficult to discern, but there appears to be a low peak in spring,
probably correlated with the breeding season on Johnston Atoll, and a
slightly higher peak in fall, probably correlated with the post-nesting
dispersal of birds from the northwestern Hawaiian Islands. These fluc-
tuations are not entirely predictable from year to year. Periods of
complete absence occurred only in November and December of 1965, 1966
and 1968. Personnel on the island during these times were familiar with
the Black Noddy roosting areas, and checked them fairly regularly;
therefore, it is not doubted that the birds were absent from the island
during these two periods. The low number present during the first half
of 1966 is not understood. Here also, the observers present were
familiar with roosting and nesting areas, and would not have missed
birds if more were present.

Specimens

POBSP personnel collected six Black Noddy specimens at Johnston
Atoll (Appendix Table 7). This constitutes a new specimen record for
the atoll.

Banding and Interisland Movement

The POBSP banded 152 Black Noddies on Sand Island from 1963 through
1969 (Table 24). Of these 152, 20 have been captured back on the island
(Table 25) and five have been captured elsewhere (Table 26): French
Frigate Shoals (three birds), and Lisianski Island and Kure Atoll (one
each). In addition, five birds banded elsewhere have been taken on Sand
Island; they are from French Frigate Shoals (four birds) and Lisianski
(one). Thus many of the non-breeding birds are presumably from the
northwestern Hawaiian Islands.

At-Sea Distribution

Black Noddies were never observed in the POBSP grid centered
approximately 175 miles southwest of Johnston Atoll (POBSP, 1967a).
They were, however, sighted within 100 miles of Johnston Atoll during
April and May (POBSP, 1965). These data suggest that Black Noddies at
Johnston Atoll do not go far from the Atoll to feed.

WHITE TERN Gygts alba
Status

Uncommon breeding species; present year-round on Johnston Island.
During spring and summer small numbers attempt to nest in trees, on

buildings, or antennas, but few are successful. Previously bred on
Sand Island, but presently only an occasional visitor.

346

Ecological Distribution

Johnston Island: Brooke (ms.) observed White Terns on Johnston
Island on 16 March 1859: "As I returned to the Captains hut I saw a
beautiful white bird perched under the edge of a low cliff. Creeping
cautiously up I took it in my hand. It was very beautiful, a species
of tern, with blue bill black eyes and snow white plumage."

Wetmore (ms. b) in July 1923 observed "About 40 frequented the
larger, higher rock ledges along the beaches of Johnston Island, where
they nested in two small colonies. Nests held eggs or newly hatched
young.'' These are shown in Figure 114.

During or before World War II construction destroyed all of this
rocky habitat. These birds, however, have readily adapted to man-made
structures and tree-nesting. Clark (1945b) recorded a few in May
1945 and Davis (1962) found an egg and a chick on a Casuarina branch
in 1959-1960.

All POBSP observations of nesting were under eaves of buildings,
on water towers, or on antennas until 1969. Birds were most frequently
found roosting in Casuarina trees, especially the ones on the dispen-
sary, northeast of the control tower, and to a lesser extent by the
tennis courts. In February and March 1967, pairs were seen in and
around Coccoloba trees on Johnston Island and hovering over a large
Tournefortta bush on Sand Island, possibly looking for nest sites.

In 1969, Shelton recorded White Tern nest sites located on the
large dish antenna stored on the south side of the taxiway (Fig.
115), in the Casuarina north of the post office, in the Casuarina and
Tournefortta north of the tennis courts, and in the Coccoloba east of
the control tower. Amerson found a few roosting in these same
localities in November 1973.

Remains of White Terns that cats may have killed were found twice
on Johnston Island (March 1965 and February 1967). Cats could prevent
nesting in trees, and certainly would eat any young birds found on the
ground.

Sand Island: Moynihan (1957) found five pairs breeding on the
original portion in trees or on abandoned houses in April 1957. From
1963 through 1969 POBSP observers recorded only occasional birds
visiting Sand Island; none nested here. Few birds actually roosted;
roosting spots included the LORAN-A antenna, the LORAN-C antenna and
fence, and the Casuarina near the barracks. Amerson recorded a pair
roosting in the Casuarina east of the barracks in November 1973.

Populations

There is no evidence for a change in numbers of White Terns using
Johnston Atoll from 1923 to the present. The 40 reported by Wetmore

347

© eee Lie
F “a Hey : te “
wre * eo + ibe hice ee a ee a

Figure 114, White Terns roosting on undercut beach rock on shore of Johnston
Island, Johnston Atoll, July 1923 (B. P. Bishop Museum photo).

| a

io

i i
Figure 115. White Tern chick hatched on framework of old radio telescope,

Johnston Island, Johnston Atoll, 19 June 1969 (POBSP photo by
Pe (Ge 5helton)).

348

for 1923 probably was matched at night during the 1963-1969 POBSP study
period (Fig. 116), although daytime numbers during the 1960's were
usually only half or three-fourths as high as in 1923.

Wetmore did not give counts of eggs or chicks in 1923, but his
"two small colonies' suggest that more birds nested then than the two
or three pairs per year recorded during most POBSP years. In 1957
Moynihan found five pairs incubating. This is also more than ever
found nesting during POBSP studies except for 1969 when 11 pairs nested.
There were no clear indications that populations were higher during
1923, 1957, and 1969; it may be simply that more were able to breed
because of less human disturbance.

Since population estimates for the 1963-1969 period were based on
daytime counts, the estimates presented in Figure 1l6may be lower than
actual numbers using the atoll. Daytime estimates represent no more
than a third to half the numbers roosting on the atoll at night. For
example, in February 1967, when a maximum daytime count of four was
recorded, 59 birds were found at night. Dusk counts in July 1966,
however, showed that the number present was not greatly higher than
those seen by day. Much more needs to be done with this species to
determine the relationship between day and night numbers, and actual
population fluctuations through the year.

Annual Cycle

Figures 37 and 116 show the annual cycle for White Terns at
Johnston Atoll from 1963 through 1969.

The few known eggs during the 1963-1969 period were all found from
February through May, and all chicks fledged by September. In 1969 at
least 11 eggs were laid and six chicks probably fledged. Moynihan's
observations of eggs and almost fully grown young in early April 1957
suggest that egg laying must have occurred by January at the latest.

The presence of eggs in July 1923 extends the nesting season well into
the summer. White Terns at Johnston have a longer breeding period than
their counterparts in the northwestern Hawaiian Islands. There, eggs are
known primarily from April through July, and chicks from May to November.

The mean numbers indicate a broad maximum in mid-summer and a low
in mid-winter; maximum numbers were about four times the lowest numbers
counted. The maximum was roughly correlated with the nesting season, or
followed it by one or two weeks.

Banding was insufficient to yield good information on population
turnover, but suggests that the rate may be rapid. In 1965, when 13
birds were banded and streamered in April, May and June, only one of 20
birds counted in early July had an orange streamer.

Specimens

Fourteen White Tern specimens have been taken from Johnston Atoll
(Appendix Table 7). Six adults and one downy chick were preserved as

349

Adults

= Means

coeee Extremes

NUMBER OF BIRDS

° .*,
2: °@e -@o oOo -@ -@°

— SSS SS ee
J Fae A M J J A S O N D

Figure 116. Means and extremes of semimonthly estimates of diurnal White
Tern numbers, Johnston Atoll, 1963-1969.

BIRDS PER 100 SQUARE MILES

J F M A M J J A S 0 N D

Figure 117. Diurnal occurrence of White Terns at sea 175 miles southwest
of Johnston Atoll, 1963-1967.

350

skins taken by Wetmore in 1923. Six adult skins and one skeleton were
taken by the POBSP. All specimens are in the National Museum of Natural
History. These constitute a new published specimen record for the atoll.

Banding and Interisland Movement

The POBSP banded 35 White Terns on Johnston Island (Table 24).
One of these, banded 29 May 1965, was captured on Tern Island, French
Frigate Shoals--450 nautical miles to the north-northeast--on 6 July
1966. Another banded on French Frigate Shoals 4 August 1965 was found
dead on Johnston Island 14 February 1967. There were no other inter-
island recaptures, and, other than the one streamer observation cited
above, no recaptures back on Johnston Atoll.

At-Sea Distribution

A few White Terns were present in the grid southwest of Johnston
Atoll throughout the year (Fig. 117). Greatest density occurred in
August and September, just after completion of their breeding season
on Johnston Island. A small peak occurred in May at the beginning of
the breeding season. A high March observation was considered to be a
sampling error. Lowest density occurred during the non-breeding season
(December through February), while another low density plateau occurred
during the breeding season (June and July).

It was apparent that more birds utilize the grid than actually nest
at Johnston. Many of these are probably pelagically-oriented non-breeders
from the northwestern Hawaiian Islands, who may also occasionally roost
on Johnston at night (POBSP, 1967a).

ROCK DOVE Columba livta
Status

Introduced breeder. Occurs in relatively small numbers on Johnston
Island only.

Observations

According to island personnel, three pairs of Rock Doves, or
Domestic Pigeons, were introduced from the main Hawaiian Islands to
Johnston Island in 1972. By mid-1973 numbers had increased to about
100 birds. Because they were a nuisance around the military facilities,
the population was reduced’ drastically. None was seen by Kridler (BSFW,
1973) in May and August 1973. As of November 1973 about 30 birds remained.
At that time several flocks of various-colored birds were seen about the
island, with the,largest flock containing 12 birds.

The birds breed on Johnston Island and the young are eaten by island
personnel. None is known from the other three islands.

Specimens

The above observations are considered to be a new sight record for
this species at Johnston Atoll, as well as a new breeding record.

SHORT-EARED OWL Asto flammeus
Status

Occasional visitor; four to six sight records from Akau, Hikina,
Johnston, and Sand Islands.

Observations

Moynihan (1957) recorded a Short-eared Owl at Johnston Island in
April 1957. POBSP records (Table 79) involve four to possibly six dif-
ferent birds unless observations several months apart were of the same
bird.

Intensive observations were made only on the owl present in late
1966 and early 1967 (Schreiber, 1967). From 3 November until the end
of 1966 this bird visited Sand Island fairly regularly. When flushed,
it usually flew toward Johnston or Hikina Island, suggesting that it
roosted on one of these islands when not on Sand. From early January
until 24 February it roosted on Sand each day under a Lepturus clump
about 200 feet southwest of the transmitter building. By mid-January
Sooty Terns were landing at night around the owl's roost. The owl left
its roost each evening just as the sooties settled; encroaching darkness
prevented further observations of its behavior. In late February Sooty
Terns began staying on the island all day. The owl roosted in its
usual place through the 24th, when the nearest terns were 100 feet
away, but abandoned the site on the 25th, when the expanding Sooty Tern
horde engulfed it. Through March and April the owl appeared fairly
regularly at dusk along the road at the northwest corner of the original
portion of Sand Island. During this time it roosted by day on an antenna
on the north side of Akau Island. This owl was last seen on the afternoon
of 27 April being chased by Sooty Terns over the lagoon west of Sand
Island.

From November 1966 through March 1967 owl-eaten carcasses of three
Golden Plovers, four Ruddy Turnstones, 13 Sooty Terns, and 15 Brown
Noddies were found. The noddies were mostly young birds, recently or
not yet fledged. Eight pellets, containing bones of birds and house
mice, were also found during this period. Mice were abundant while the
owl was present and probably made up a large portion of its diet.

This owl was never seen hunting or eating during the day; freshly
eaten carcasses were found only in the mornings, indicating that the
bird fed only at night.

352

Table 79. Observations of Short-eared Owls on Johnston Atoll

Date of Observations Remarks and References

1957. 6-10 Apr. "One Short-eared Owl...flew out of a small
patch of grass one evening" on Johnston I.
(Moynihan, 1957: 35).

1964 24 Nov. One bird flushed from a Wedge-tailed
Shearwater burrow on Sand I., circled island
and disappeared from sight.

10 Dec. One bird, probably the one seen 24 November,
flushed from grass near old pier on Sand I.
and flew to Johnston I.

1966 19 June- One bird roosted regularly during day in
25 July Casuartna tree on Johnston I., and visited
Sand irregularly at night.

3 Nov.- One bird, possibly the one present earlier in
1967 27 Apr. 1966, appeared regularly on Sand I. from Nov.
until its roosting place was overrun by Sooty
Terns in late Feb. Thereafter is apparently
stayed on Akau I. most of time, but visited
Sand I. fairly regularly.

1969 25 May-July Found on Sand I. only on 25 May, eating a
Sooty Tern; reported on Johnston I. until at
least early July.

Reactions of other birds to owls varied greatly. The owl seen
10 December 1964 was chased by frigatebirds and noddies. On Johnston
Island in July 1966 White Terns avoided the tree in which the owl roosted,
although they normally roosted there. In fall and winter 1966-1967,
Red-footed Boobies and Great Frigatebirds roosting on the LORAN-C guy-
wires took flight and circled close together when the owl circled above
the wires. Frigates on nests flushed if the owl flew high over them,
but if it was below about 30 feet the owl passed too quickly for the
frigates to react. Shorebirds flushed, gave alarm calls, and joined in
tight flocks when the owl was near. Brown Noddies and Sooty Terns only
occasionally chased the owl as it flew; they usually ignored it. All
these reactions were observed by day, when the owl was flushed and
would not normally have been flying. No observations were obtained at
night when the owl was hunting.

Specimens

No specimen records are known from Johnston Atoll.

There are resident populations of this circumboreal owl on both
the Caroline Islands (A. f. ponapensts) and the Hawaiian Islands

(A. f. sandwichensts). The Hawaiian population is considerably closer
to Johnston, and is the most likely source of the Johnston visitors.
Owls have also been recorded from the northwestern Hawaiians. The
Short-eared Owl collected on Kure fit measurements of neither the
Caroline nor Hawaiian form and was considered most likely of the
Holarctic race, A. f. flammeus (Clapp and Woodward, 1968; Woodward,
LOT 2) 5

At-Sea Distribution

A Short-eared Owl was recorded in the at-sea grid about 130 miles
west of Johnston Atoll on 16 November 1965 (POBSP, 1965).

SKYLARK Alauda aruensts
Status

Accidental visitor; two sight records on Johnston and Sand Islands.
Observations

On 18 November 1963, Sundell saw a bird on Sand Island which was
most likely a Skylark. A Coast Guardsman first saw it flying eastward
along the causeway in the early evening. Sundell immediately went with
the Coast Guardsman and they flushed it "...from the ground in the bunch
grass [of the original portion] just after leaving the causeway.'' His
complete description follows: "It was the size of a large sparrow and
predominantly brownish streaked on the back and wings and also on at
least part of the breast (upper half). There were rather pronounced white
patches along the outer edges of the tail and it is quite likely that the
bird had a medium-sized crest. The bird flew with an undulating flight
but was extremely wary and impossible to approach. When flushed the bird
gave a single short, musical call note."

Nine days later, 27 November, U.S. Air Force personnel on Johnston
Island reported a small bird of similar description that could have been
the same one Sundell saw on Sand.

Specimens

No specimens are known from Johnston Atoll; however, the above
observations constitute a new sight record.

Skylarks breed across Europe and Asia, and have been introduced
in the main Hawaiian Islands (AOU, 1957). The Johnston Skylark was
most likely from Hawaii. One collected on Kure Atoll, however, was
identified as A a. pekinensts, which breeds in northeastern Siberia,
Kamchatka, and the Kuriles (Clapp and Woodward, 1968; Woodward, 1972).

\ Ase oe

“2 f/ ii iff)

JAPANESE WHITE-EYE Zosterops japonica
Status

Rare visitor; two specimen records and three sightings on Johnston
and Sand Islands.

Observations

Records of Japanese White-eyes for Johnston Atoll include two
specimens and three sightings of single birds. All records but one
were made by POBSP personnel (Table 80).

All birds appeared to be healthy except the one taken 30 November
1965 which was so weak that it was caught by hand; it weighed only 7.9
grams. This bird appeared after four days of unusually strong northeast
winds, which may have accounted for its poor condition. For comparison,
the mean weight of five other POBSP-collected white-eyes--the other
specimen from Johnston Atoll (10 grams) and four taken at sea between
Johnston and Oahu (Ely, 1971)--was 10.4 grams (range 9 to 12.8 grams).

Specimens

The two specimens noted above and in Appendix Table 7 were pre-
viously reported by Ely (1971).

The white-eyes found on Johnston Atoll and the flocks observed at
sea between Hawaii and Johnston presumably came from the population es-
teblished on the main Hawaiian Islands. The occurrences of white-eyes
at sea in October, November, and December, and on Johnston Atoll in
October through March, coincide with the non-nesting season in
Hawaii, where they breed from February through November (Hawaii Audubon
Society, 1967). Although they are migratory only in the northern parts
of their native range in Japan, all populations of Japanese White-eyes
wander extensively after nesting (Austin and Kuroda, 1953).

Table 80. Records of Japanese White-eyes on Johnston Atoll

Date of Observations Remarks and References
1964 11-14 Oct. One bird seen 11, 13, and 14 Oct.; collected
14 Oct., Sand I. (Kepler and Stadel).
1965 29 Mar. One bird flying about in shrubby trees on
Johnston I. (Bratley).
30 Nov. One collected near docks, Johnston I. (Kepler).
15"Dec.. Qne reported by Commanding Officer, Sand I.
1966.31 Jani. One in bushes around barracks, Johnston I.

(Tordoff).

SOCIETY FINCH Lonehura striata
Status
Introduced. Occurs as a caged bird on Sand Island.

Observations

According to Coast Guard personnel, two caged Society (or Bengalese)
Finches were introduced on Sand Island in 1972. By November 1973, one
was kept caged in the Coast Guard barracks building; the other had pre-
viously escaped and no doubt died. No breeding is known.

The Society Finch was probably introduced from the main Hawaiian
Islands where it is commonly found in pet stores. The species is native
to Southeast Asia.

Specimens

This is a new sight record for Johnston Atoll.

Mammals

There are no mammals native to Johnston Atoll, and none was
recorded in the literature before POBSP studies began. A brief summary
of the status of the atoll's mammals through 1964 was published (Kirk-
patrick, 1966) and the following accounts are based on that publication,
with additional notes from 1965 through 1973.

In addition to human occupants, six species of mammals were re-
corded through 1973 (Table 81). The two rodents probably arrived in
ship or plane cargoes, while the dogs, cats, and rabbits were no doubt
purposely introduced by military and civilian personnel; all, except for
the rabbit, probably first appeared during World War II. The seals
arrived from the resident population on the northwestern Hawaiian Islands.

Porpoises probably visit the atoll's lagoon waters, although none
has been recorded. They have been reported by island boat operators as
regularly occurring outside the main channel entrance.

Species Accounts

The introduced species are well known; however, a description and
illustration of the seal can be found in Mammals of Hawatt (Tomich,
1969).

356

Table 81. Distribution and status* of mammals at Johnston Atoll

Sand
Species Akau Hikina Johnston Original Man-made
House Mouse B B B
Roof Rat B
Domestic Dog R R R
Domestic Cat R B R R
Hawaiian Monk Seal R R R B R
European Rabbit R R
*B = Breeding; R = Recorded.
HOUSE MOUSE Mus musculus

Status

Introduced breeder; abundant on Sand Island, smaller numbers on
Johnston Island. Scant data suggest a mid-winter breeding peak;
breeding may, however, be correlated with amount of vegetation.

Observations

No house mice had been recorded from Akau and Hikina Islands as
of the end of 1969.

Johnston Island: House mice are found in small numbers on Johnston
Island. The rodent and insect control program may have indirectly limited
the mouse population size.

Sand Island: House mice were recorded as "very numerous" in 1963,
and the population remained high throughout the POBSP study period. A
live-trapping grid was established in 1964, first with 25 traps placed
at five-foot intervals in a 20 by 20 grid. This was later expanded to
48 traps, then to 72 traps placed at intervals of 20 feet except for two
rows separated by 50 feet through which passed the road to the LORAN-C
transmitter. The final grid covered 27,260 square feet. This grid was
trapped on the first seven nights of each month from May 1964 through
May 1965. After that the grid was trapped intermittently and infrequently
through 1967. Sherman live-traps baited with rolled oats were used.

During 7,329 trap nights from May 1964 through May 1965, trapping
success ranged from 31 to 80 percent, and the population was considered
to remain high throughout the period (Kirkpatrick, 1966). However, it
seems likely that when the trapping success dropped to only 31 percent,
there must have been a decline in the population. These data were ob-
tained by or under the direction of Ralph D. Kirkpatrick, who is
preparing them for publication.

Sioy/

In October 1966 and February 1967, trapping success in this grid
as measured by number of mice caught per trap night was 73 and 74
percent. The calculated population sampled in February 1967 (Hayne,
1949) was about 185 mice, or one mouse per 148 square feet of the grid
area. No attempt was made to correct these figures with buffer zones,
compensations for trap-shyness or addiction, or any other factors.
This area probably was as densely populated as any on the island.

The reproductive condition was analyzed in autopsied mice caught
from October 1964 through March 1965, and external examination of
breeding condition of animals judged to be adult in the October 1966
and February 1967 samples were made. The 1964-1965 data (Table 82)
suggest mid-winter breeding for house mice on Johnston Atoll. However,
the 1966-1967 data (Table 83) indicate that a peak in breeding had taken
place shortly before October 1966, both because of the presence of
lactating females and the high number of young animals that were born
during the summer of early fall of 1966. The lack of obviously pregnant
females in October suggested that breeding had stopped but pregnant
and lactating females in February, as well as more young animals, in-
dicated that breeding probably did not stop.

Table 82. Breeding condition of house mice autopsied, October 1964-

March 1965

Number of Percent Mean No. Number of Percent with
Date Females Pregnant of Embryos Males C.E. Tubules*
October 2 0 - 0 -
November 23 0 - 29 0
December BAL 4.8 4 24 SD
January 19 ZO 5 23 BO
February 27 SIST/ 5 17) 17.6
March 28 0 - 23 ts} 6 7/

*Cauda epididymus tubules macroscopically visible.

Table 83. Evidence of breeding condition in house mice live-trapped
and released, October 1966 and February 1967

% Males
Number Percent Sex ratio % Females % Females with Scrotal
Date Handled Adult M:F Lactating Pregnant* Testes
October 1966 110 58 34:27 ae: 0 Thal
February 1967 72 65 58253 8 min. 10 64

*Only those obviously heavily gravid were detected as being pregnant.

ev was Ee

358

In summary, there are too little data to show if peaks in breeding
occur, whether there are periods of non-breeding, or with what the fluc-
tuations might be correlated. Subjective impressions indicate that mice
are more abundant when the vegetation is more luxurious. For example,
in mid-summer 1966, when the vegetation was at a low point in vigor,
mice were less noticeable than in winter and spring 1967, when the
vegetation was relatively lush.

Although house mice may be found nearly anywhere on Sand Island,
they reach their greatest density in the heavy Lepturus on the eastern
islet, including the area of the grid. Here cover and food are evidently
optimum for these rodents. The density in and around the buildings is
comparatively low, probably because the buildings are concrete and little
if any food is available to them.

Another possible factor influencing mouse abundance may be the
insecticides used against roaches. All parts of Johnston Island, and
all but the eastern portion of Sand Island are sprayed once weekly with
a strong insecticide. It may be more than coincidence that the highest
density of mice is on the eastern, unsprayed portion of Sand Island.
Mice could be secondarily affected by the chemicals through eating
poisoned insects, or simply from contacting it on vegetation and the
ground.

There are no indications that mice have any effect upon bird
populations on the atoll, except in providing occasional food for
three species occurring there--Bristle-thighed Curlews, Short-eared
Owls, and Cattle Egrets. Also, dogs and cats prey on them. The food
of house mice on Johnston Atoll probably consists mainly of seeds and
insects, but no studies have been made.

Specimens

Twelve house mouse skins from Johnston Atoll are preserved in the
National Museum of Natural History (USNM 360999-361010) and a large
collection of skulls, to be deposited in the USNM, are being studied by
Ralph D. Kirkpatrick of Ball State University, Muncie, Indiana. Pre-
liminary study of these specimens by Kirkpatrick indicates that they
are more nearly like Asiatic and Micronesian Mus in body weight than
like North American or European animals. Molt has been studied by Brechner.
and Kirkpatrick, 1970.

ROOF RAT Rattus rattus
Status

Introduced breeder; present, but uncommon, on Johnston Island.
Observations

The roof rat is established on Johnston Island, but not on Akau,
Hikina, and Sand Islands.

359

Johnston Island: Roof rats were reported from Johnston Island
by 1962, but probably were introduced via ships or barges several years
prior to that date.

Rats are most abundant in refuse dumps, along the pier, and in
the warehouses and other buildings near the shipping wharfs. Rat
control measures were initiated by military personnel in late 1964 and
included snap-trapping and poisoning. The success of the program is
not known, but reports of rats around the Navy pier in 1968 and 1969
indicate that they were not exterminated (Kirkpatrick, 1966).

Specimens

Seven specimens were secured from the control operation during
September, October, and November 1964; all are in the National Museum
of Natural History (USNM 360986-360992). One (USNM 360991) is
melanistic. Reproductive data for the females were not recorded, but
testes size and location in the three adult males indicate that they
were in breeding condition.

DOMESTIC DOG Cants famtltaris
Status

Introduced; breeding status unknown; present on Johnston and Sand
Islands.

Observations

Johnston Island: Two dogs, at least one a female, were present
from 1963 through 1965; from 1966 through 1969 numbers were higher.

Sand Island: Two male dogs were on Sand Island in 1963 (Kirk-
patrick, 1966); numbers ranged from two to four through 1969. Two
were present in November 1973. They were kept as pets, but were per-
mitted to run loose.

At times dogs were destructive to nesting and roosting seabirds.
Wedge-tailed Shearwaters were most affected: in November 1964 up to five
nestlings a day were killed, but not eaten, by dogs. At least ten chicks
were killed in October 1966. These chicks were most vulnerable in the
last few weeks before they fledged. During that time adults were rarely
present and no other birds were present in sufficient numbers to dis-
courage the dogs. During the spring and summer, when adult Wedge-
tailed Shearwaters, Sooty Terns, Brown Noddies, and Brown Boobies were
mumerous, the dogs appeared to be afraid to go into the colony. In
early 1967 two Red-tailed Tropicbird chicks were killed on the western
portion of Sand Island, possibly by dogs or cats. During 1968-1969 two
newly introduced dogs never visited the bird colony unaccompanied and
did not molest the birds.

360

DOMESTIC CAT Felts catus
Status

Introduced; breeds on Johnston Island, present on Hikina and
Sand Islands.

Observations
Hikina Island: Two semi-tame cats lived here in February 1969.

Johnston Island: Domestic cats were numerous on Johnston Island,
where a reproducing population existed through 1973. They were sus-
pected of killing and eating White Terns (see White Tern account), but
no evidence was obtained indicating that these animals bothered other
birds. Unattended tropicbird chicks, or any weakened bird, would be
likely victims.

Sand Island: A non-reproducing pair ran wild on the original
portion until early 1964. They preyed on Sooty Terns and Gray-backed
Terns. In 1964 they were lured to the man-made portion and kept as
pets (Kirkpatrick, 1966). Two cats remained on the island through
1973; they did visit the bird colony and occasionally ate mice, but
little evidence of bird predation was found.

HAWAIIAN MONK SEAL Monachus schautnslandt
Status

Visitor; one breeding record.
Observations

The first Hawaiian Monk Seals ever recorded outside the Hawaiian
Islands appeared on Johnston Atoll in 1968 and 1969. The first, a young
animal tagged as a pup in March 1968 on Laysan Island, 547 nautical miles
to the north-northwest by U.S. Fish and Wildlife personnel, appeared on
Johnston Atoll the last week of July 1968 and stayed until late December.
During this time it was reported from nearly every part of the atoll,
but it usually returned to sleep near the Navy docks on Johnston Island
(Schreiber and Kridler, 1969). It reappeared during the last half of
June 1969 and stayed at least to mid-August 1972.

On 29 January 1969 an untagged adult female hauled out on a
protected beach on Sand Island and gave birth to a female pup. The
two animals were tagged.and remained on or near this beach until
3 March, when the adult disappeared. The pup remained and fed on
snails and other marine invertebrates, as well as fish. It died in
1971 from a deep flesh wound, probably from a shark attack.

361

EUROPEAN RABBIT Oryctolagus cunteulus
Status

Introduced; occurs on Johnston and Sand Islands where it is kept
caged.

Observations

Johnston Island: Rabbits were introduced onto Johnston Island
during the early 1970's as a test animal for various chemical agents
stored on the island. The animals are caged.

Sand Island: One rabbit was kept as a pet in a cage near the
Coast Guard barracks in November 1973.

ECOLOGICAL SIGNIFICANCE

The conservation status of Johnston Atoll has been discussed by
King (1973), but no ecological significance was given.

In order to assess the ecological significance of Johnston Atoll,
four well-known ecologists in various fields were asked by Amerson (1973)
to read the manuscript and evaluate the baseline findings. The following
commentaries were obtained.

Ecological Values of Johnston Atoll

(by Ray Dasmann, International Union for Conservation of Nature)

Because of its small size, and extreme isolation Johnston Atoll
was originally of considerable ecological interest as an area in which
it would have been possible to follow the slow process of colonization
and establishment of species on oceanic islands and to study over the
years the processesthat may have led to the development of new races or
species in its limited biota. The opportunity was lost with the ex-
ploitation of the island and later with its development as a naval and
air base. However, human occupancy of the island followed by the
introduction and establishment of many new species of plants and animals
has created an equally interesting ecological situation in which the
interactions among its still limited biota could profitably be studied.
Such situations, however, occur on many other isolated oceanic islands,
and Johnston cannot be considered as particularly unusual or of out-
standing interest from this point of view.

Considering the dry land area of Johnston Atoll, the greatest
concentration of ecological interest is to be found on the remaining
natural island, the ten acres of the eastern portion of Sand Island.
This is the principal breeding area for seabirds and appears to support
the most complex terrestrial biota. Sand Island is of major importance
for its breeding population of Sooty Terns and of significant importance
for breeding populations of Red-footed Boobies, Brown Noddies, Wedge-
tailed Shearwaters and Great Frigatebirds. It is significant also as a

362

wintering area for shorebirds, notably the Golden Plover and Ruddy
Turnstone. Every effort should be made to minimize disturbance of this

area in the future and to maintain it as a refuge for seabirds. With
protection and freedom from disturbance its value as a seabird breeding
center can be expected to increase, and it will achieve greater value

as a site for ecological studies. By contrast, the ecological interest

of the western, man-made portion of Sand Island, of Johnston Island, of

Akau and of Hikina Islands, is slight at the present time, although if
disturbance of these areas were to be greatly reduced in the future,

they would no doubt be colonized in time by breeding populations of seabirds.

The marine area of Johnston Atoll can be considered of equal interest
to the terrestrial area. Although the marine biota has not been thoroughly
studied, the inshore fish population appears to be of considerable bio-
logical interest and it is likely that further studies will reveal a
higher degree of endemism than is now reported, particularly among the
marine invertebrates. Considerable damage to the marine fauna has resulted
from past dredging and filling operations. Future activities of this nature,
when necessary, should be conducted with greater precautions to minimize
damage to reef and lagoon fauna.

Considering that Johnston Atoll may continue to be used for a variety
of purposes in the future, it is recommended that particular attention be
given to protection of the eastern portion of Sand Island for the purpose
of maintaining the reef and lagoon biota in a healthy state, allowing for
its recovery from past disturbance. Avoidance of pollution and siltation
of the reef-lagoon complex should be given priority.

Johnston Atoll: Some Ecological Considerations
(by Robert E. Jenkins, The Nature Conservancy)

The report on the "Ecological Baseline Survey of Johnston Atoll,
Central Pacific Ocean" presents a fascinating picture of what is un-
questionably a unique, interesting, and important scrap of landscape.
Johnston Atoll is unique to begin with because of its small size, extreme
isolation, long (25 to 100,000,000 years) geological and evolutionary
history, tremendous constancy of its oceanically buffered environment
(remarkably small variation of landscape and practically all climatological
variables), recent history of human disturbance and alteration, great
changes in species composition resulting from colonization by introduced
organisms, and, not least, by the amount of scientific research and data
collection which has gone on there. Some of these values and potentials
are considered in more detail below.

The Place of Johnston Atoll in the Natural Biosphere

Johnston Atoll is a focal point of biological activity in the midst
of tremendous expanses of otherwise unbroken ocean. It served as a per-
manent home and habitat for a notably small number of terrestrial organisms
in the past because of its isolation from other sites occupied by potential
colonizers. However, the Atoll always has been and continues to be a vital
link in the life cycles of a variety of pelagic organisms which retire

363

there to breed (especially seabirds) or use it as a haven and way station
along traditional routes of migrations or wandering. In addition to being
the only pieces of dry land over thousands of square miles of ocean, the
submerged platform of the atoll splits and deflects oceanic currents,
causing eddies which produce turnover, upwelling, and nutrient enrichment.
This makes the Atoll system a hot-spot of organic productivity supporting
concentrations of fish and aquatic organisms and the seabirds which depend
upon them for food.

The importance of atolls such as Johnston in determining the faunis-—
tic composition of a very large section of the ocean cannot be overestimated.
Were Johnston physically (or ecologically) removed from its section of the
globe, it would have measurable effects on the ecosystem over a very wide
area. At present, habitats such as Johnston Atoll are particularly
critical because they represent shrinking fragments of relatively predator-
and harassment-free areas of a type becoming increasingly rare as pressure
on the oceanic islands increases. In this capacity, Johnston serves as a
reservoir for a diversity of species which are badly depleted or even en-
dangered over much of the rest of their range.

In addition to acting as a temporary annual stopover for numbers of
migrating species of birds and fish, Johnston Atoll has obviously served
as an important stepping-stone in the longer time sequence of evolutionary
events. Though this whole area of inquiry remains largely unelucidated at
present, Johnston Atoll has clearly been an important focal point in the
dynamic process of faunal exchange.

The Importance of Johnston Atoll as a Scientific Laboratory

Johnston Island has, over the years, attracted the attention of a
large number of competent and even famous scientists who have visited the
island, made their observations, and added to our store of knowledge about
their disciplines and about Johnston Atoll specifically. Most of this
information has been pulled together under the Smithsonian Institution's
Pacific Ocean Biological Survey Program (POBSP). This program has also
added a vast amount of new data of its own for comparison with the more
fragmentary information published by past researchers. This collection of
data will form the foundation of extensive new biological and environmental
baselines which will assume an ever greater importance as we learn more
of the Pacific ecosystems and as the effects of human activity increasingly
influence those systems. The data collected in areas such as Johnston
Atoll, when compared with later measurements in the same area, will be
our only reliable yardsticks whereby we measure changes occurring in inte-
grated ecological and life support systems. Without such comparisons, we
will have no way of distinguishing between trivial environmental fluctua-
tions and evidences of serious perturbation or trends of deterioration.
Without them we would have no way to identify cyclical events and no way
to extend our as yet rudimentary understanding of how natural systems
function.

Of particular importance to Johnston Atoll is the recurring theme
of its seabirds. In spite of all of the "reconstruction" which has

364

severely changed the original natural environments in the area, there is
still a tremendous seabird population using the near-shore feeding grounds
or breeding--primarily on Sand Island. The information collected on these
birds forms a truly impressive body of data. The monumental accomplish-
ment of having banded over 300,000 individual birds in the course of six
years has already added greatly to our knowledge of population dynamics,
distribution, faunal exchange, site constancy, breeding systems, species
composition, etc., and in the years ahead should add even more to our
understanding of some important components of the oceanic system.

There have also been fairly extensive studies made on the effect of
the dramatic and pervasive human alteration of Johnston Atoll. A continu-
ation of these studies will give us new insight into the effect of dredging
on the physical and biological environment of coral reefs, the effects of
greatly enlarging the terrestrial mass in the area through the creation of
entirely new superaquatic platforms from native materials, and the effects
of stocking these (however haphazardly) with a large number of exotic
species of plants and animals. The increase in the vascular plant flora
from three species in 1923 to 127 in 1973 provides us with a very interest-
ing case in point. In the last few years, the relatively young discipline
of island biogeography has been yielding new insights on a number of
ecological and evolutionary phenomena such as colonization, competition,
extinction, community stability, genetic adaptation, ecological exclusion,
niche dimensions, etc., and the Johnston Atoll situation represents a
unique experiment in this field which could richly reward intense scrutiny.
Aside from the population and community phenomena which are favorably
isolated for investigation, the effects of the biota in modifying the raw,
new substrates over time should be carefully observed and documented.
Within the aquatic environment, the same processes of ecological recovery
from the effects of dredging and filling provide us similar opportunities.

Another aspect of the past research which seems very worthy of pro-
tection and continuation is the baseline study on ciguatera-type fish
poisoning. This has become a world-wide problem of growing dimensions,
and the large amount of information, particularly that relating to the
known-time variation in the prevalence of toxic episode and the fish
species affected might eventually help us to unlock the riddle of what
causes ciguatera poisoning and consequently the ways in which we can
avoid or counteract it.

Future Potential

Because of its uniqueness, Johnston Atoll lends itself to so many
different and important uses that a prime difficulty may be simply to
determine--within the constraints imposed by the military, communication,
transportation, physical research, and human habitat factors--which of
the various additional uses should be given priority.

On the one hand, the Atoll area clearly represents a critical species
reservoir. This function could undoubtedly be substantially enhanced
without any significant interference in the "“edificarian" uses of the
island. The report observes, for example, that the Laysan and Black-footed

365

Albatrosses (both depleted species limited by long life cycles and the
limited availability of nesting sites) are unlikely to re-establish
themselves as breeders on Johnston Atoll through natural developments.
However, we have an opportunity to assist by creating favorable c mndi-
tions and otherwise promoting re-establishment. This would require
something on the order of the procedures used to re-establish salmon in
streams from which they have been eradicated. Such a project might in-
clude the hand rearing of nestling albatrosses on the island sites in
question in the hopes of imprinting the young for later return at sexual
maturity. Such activities have been undertaken in behalf of other
endangered species by the Department of Interior which ironically still
retains statutory responsibility for Johnston Atoll as a federal bird
refuge.

On the other hand, parts of the islands might be used in quite
different scientific experiments. The area's isolation offers a
tremendous laboratory for the investigation of species dynamics as
discussed above which could be enhanced through deliberate manipulation.
Parts of the islands might be devoted to the intentional introduction of
a large array of terrestrial biological species. Oceanic barriers would
make control of the experimental design quite extraordinary and would
protect against the sorts of deleterious consequences that frequently
accompany exotic introductions in mainland areas. It would be possible to
eradicate given species or even whole floras or faunas with relative ease
if it became advisable. In this respect, Johnston Atoll may provide an
absolutely unique opportunity to obtain information on maximum community
diversity, competitive interactions in a large species array, etc., which
could be arrived at by no other means. Of course, such uses would have to
be controlled so as not to adversely affect the seabird colonies, but Akau
and Hikina Islands appear to receive little or no use by seabirds and the
same can be said of most of Johnston Island itself.

Ecological Significance of Johnston Atoll
(by Lee M. Talbot, Council on Environmental Quality)

The paper provides exactly what its title calls for, a survey of
baseline ecological data for this important area. It is an extremely
comprehensive job, and I believe an exceedingly valuable one. The detail
provided varies greatly from species to species and from group to group,
reflecting the variations in the data base of the survey. Birds, which
were the focus of the Pacific Ocean Biological Survey Program on which
the author was occupied originally, have been the focus of a greater
amount of research than the other groups of organisms, and this is of
course reflected in the detail presented in the paper.

Johnston Atoll has high ecological significance for two primary
reasons. The first derives from its isolated location in the central
Pacific Ocean. Study of the organisms found there can contribute
significantly to the understanding of migration and distribution mecha-
nisms and evolutionary ecology of a variety of types of organisms. The
inshore fishes are exemplary of this in connection with the distribution,
dispersion, and introduction of warm-water fishes.

366

Another allied source of ecological significance to this isolated
Atoll derives from what studies based there can indicate about the migra-
tory movements of birds, their parasites and pathogens, marine mammals,
reptiles, and fishes. The bird studies have been the most extensive to
date, of course, and the detail in this paper reflects that.

The other, and in my opinion more important, reason for its signifi-
cance, derives from the history and nature of the Atoll. In its present
form it is very largely man-made. Even those parts that have not literally
been constructed by man have been very significantly modified. This history
is well documented with scientific collections, descriptions, and maps and
with extensive photographs. At the same time, it is a relatively simple
ecosystem from the standpoint of topography and other physical aspects. A
high percentage of the terrestrial organisms have been introduced by man
(e.g., 124 out of 127 species of vascular plants, and all of the terres-
trial mammals and reptiles). Since most of these introductions are of
comparatively recent origin and many can be reasonably well dated, the
Atoll provides an almost unique laboratory in which the mechanisms of
dispersion, introduction, adaptation, and development of an ecosystem and
its component species can be studied. The uniformity and simplicity of a
substrate further facilitates study and comprehension of the mechanisms and
isolation and understanding of the dynamic processes involved.

Beyond the basic mechanisms of introduction, dispersion, etc., of
organisms in a new habitat, the Atoll provides opportunities for the study
of the adaptation of species and communities to changed or new habitats.
This derives from the extensive dredging and changes that have been made
and continue to be made in portions of both the terrestrial and aquatic
environment. Again a part of the significance here derives from the ex-
cellent documentation of the changes, including the photographs.

Further, the Atoll provides "control" study opportunities, for
example, as between the dredged and undredged reef areas, and between
the more or less natural and the recently man-made islands. A further
opportunity is provided by the small cluster of different islands, each
with its own well-documented history.

One of the types of study possible at the Atoll because of the condi-
tions noted above is that of succession, or of developmental stages
following the creation (through dredging, filling or clearance) of new
and essentially sterile habitats. The opportunities here involve both
the chance to document the types of succession that occur--and indeed
whether there is a consistent succession or whether there is no consistent
pattern--and the chance to study other processes associated with such a
succession. An example of this is the ciguatera poison problem.

The Atoll offers the opportunity to study the relationship between
ciguatera-type fish poisoning, and disturbance of the environment.
Dredging operations beginning in 1963 provided an opportunity to test
Randall's theory that the toxin responsible for the ciguatera was produced
by algae which occurred early and abundantly during the successional in-
vasion of a newly denuded substrate by algae.

In summary, Johnston Atoll is ecologically significant from the
standpoint of its opportunities for ecological research which prorises
unique contributions to our knowledge of ecological processes. S me very
useful ecological research has been accomplished there.

However, in my opinion, the potential of adding to our ecological
understanding from dynamic study of this Atoll has only just been
scratched. Realization of the research potential of this area requires
the foundation of baseline data which this paper is intended to--and
indeed well does--provide. Because of this, and because of the attention
it focuses on this important area and opportunity, I believe that the
paper is an exceedingly important and valuable contribution.

Johnston Atoll: Ecological Significance
(by George E. Watson, Curator of Birds, National Museum of
Natural History)

Johnston Atoll constitutes the only exposed land in a vast area of
the tropical Pacific Ocean. Although the total land area is about one
square mile, only a small portion of this is available to birds for
undisturbed resting, roosting, and breeding. Thus seabirds that in-
habit many hundred thousands of square miles of open ocean concentrate on
a small island. Likewise, shorebirds breeding on extensive northern con-
tinental areas migrate to the restricted insular ecosystem of Johnston
Atoll.

The birds frequenting the atoll may be classified according to
activity into breeders, offseason or prebreeding migrants, and vagrants.
The ecological significance of the last is nil. The island does not play
any role in the survival of the species and perhaps very little role in
the long-term survival of the errant individual. Far and away the most
important breeding bird is the Sooty Tern which produces about 60,000
chicks a year on Johnston Atoll. Lesser numbers of Brown Noddies and
Wedge-tailed Shearwaters use the island for breeding as do relatively

367

insignificant numbers of other species. None of these species is restricted

to Johnston Atoll nor is the population on Johnston Atoll a significant
fraction of the Pacific Ocean population of the species. There are no
endemic landbirds or seabirds restricted to the islands.

The same is true of the five species of shorebirds that regularly
visit the islands on migration. Most of these are wide-ranging and
scatter from their largely arctic breeding grounds over much of the
tropical oceans of the world. One species, the Bristle-thighed Curlew,
is considered rare and endangered on its breeding grounds in Alaska. It

disperses so extensively to islands in the tropical Pacific Ocean, however,

that even if the Johnston Atoll birds were eliminated, the total species
population would not be jeopardized.

The influence of long distance migrant arctic and subarctic shorebirds
is probably slight at Johnston Atoll. Their numbers are not great and their
dispersal even though over great distance is not concentrated in any given
area. There has been little study of the food supply available to them and

368

although no shorebirds breed on the islands, marking of individuals has
demonstrated that tropical islands, including Johnston Atoll, serve as
year-round tropical nurseries for first-year arctic shorebirds.

In spite of a massive bird-marking program centered on Johnston
Atoll from 1963 to 1969, information on the movements of birds to and
from the atoll is disappointingly meager, except for Sooty Terns. Marked
individuals of that species have been observed at sea in all directions
from the atoll at up to 481 miles distance and over an area of at least
300,000 square miles during the breeding season.

What still remains unknown is the importance of seabirds in the
overall marine environment. Obviously in the waters near the island
concentrations of birds can exert predation pressure on small fish,
crustaceans and squid and thus limit populations in relatively infertile
tropical waters. There is little feeding by seabirds in the lagoon or
other waters near the atoll. Sooty Terns are probably feeding at up to
a full day's flight away from the atoll. Some of the other species may
also have great daily flight ranges. Nor is it known exactly where most
of the individuals that breed on the-island go during the period when
they are not breeding. It is known that the island serves as resting or
roosting ground for numbers of birds that breed elsewhere, particularly
boobies from islands to the north in the tropical Pacific Ocean.

Individuals of most breeding species are present throughout the year
although in nearly all cases the egg and chick stages occupy significantly
less than 12 months. Because of staggered cycles, however, there is no
month when some seabird is not engaged in some stage of breeding. The
three main species, Sooty Tern, Wedge-tailed Shearwater and Brown Noddy,
have eggs and chicks in every month save January and even during that
month the terns and noddies are feeding in order to produce eggs.

The terrestrial insular ecosystem, however, is by no means a closed
one, for on the one hand the seabirds concentrate nutrients on the island
through the guano that they deposit and the rain and, probably to a lesser
extent, wave action washes the nutrients back into the sea for recycling
through the marine environment. There should be a tendency for the inshore
waters of an oceanic island ecosystem to become increasingly richer with
time. This should result in a change in species makeup from wide-ranging
species that gather food far from an island to those whose feeding trips
are short. Likewise, this could also produce a similar long term change
in visitors that use the island for roosting.

Three species of seabirds, the Blue-faced Booby and Laysan and
Black-footed Albatrosses, used to breed on Johnston in small numbers but
no longer do so because of human disturbance. Several other species occur
in smaller numbers than they ever did in 1923 when the island was subject
to periodic visits by feather hunters. The atoll is at the southern limit
of the albatrosses' at-sea feeding range and may be only marginally suit-
able for breeding under the best of circumstances, but several of the other
species could well recover significantly if they were completely undisturbed.

369

SUMMARY

Johnston Atoll, located between the Hawaiian Islands and the Line
and Phoenix Islands, is one of the most isolated coral atolls in the
world. Military activity has greatly altered the atoll: two of the
four islands are man-made and the original two have been greatly changed.
Since World War II, the atoll has been a military base. The wildlife on
the atoll is protected under a little-known 1926 Executive Order.

The flora of Johnston Atoll is well known. There are 67 species
of benthic marine algae known from the lagoon. Increased silt from
dredging activities in 1963 and 1964 decreased the number of algae species
in the dredged areas. Three vascular plants occurred on the original two
islands; man has apparently introduced 124 species since 1923.

The invertebrate fauna is not well known and dredging has further
reduced or eliminated some species. The known groups are: 18 species of
Cnidaria (Coelenterata), 58 species of Mollusca, 12 species of Annelida,
75 species of marine Arthropods, at least 87 species of terrestrial
Arthropoda (including two tick species, five chiggers, two nasal mites,
23 biting lice, and two louse flies), and 37 species of Echinodermata.

The vertebrates are well known. There are at least two species of
pelagic fishes and 194 species of inshore fishes. Dredging operations
have drastically reduced the fish population in certain lagoon areas.
Ciguatera is prevalent among the inshore fishes, with the moray eel,
white-tipped reef shark and grey sand shark being most toxic. Five
species of reptiles are known; all but one were introduced by man.
_Likewise, no mammals are native to the atoll. Of the six species known,
man has introduced five species. Fifty-six species of birds, whose total
population ranges upward to 600,000, are known from the atoll. Of the 22
seabird species, 12 species are breeders, three are former breeders, and
seven are visitors. Of the 34 species of waterfowl, marsh, and land
birds, seven species are regular migrants, six are irregular visitors,
two are stragglers, 16 are accidentals, and three are introductions.
Analysis of 60,932 returns of 303,901 birds comprising 21 species banded
at Johnston Atoll reveals that the atoll is the major focal point for
interisland movements in the north-central Pacific. A total of 733
individual banded birds have moved to or from Johnston Atoll; most inter-
island movement involves the northwestern Hawaiian Islands.

Johnston Atoll is perhaps the most scientifically studied atoll in
the central Pacific. Despite man's disturbance, the atoll is ecologically
Significant because of its isolation and from the standpoint of its oppor-
tunities for island ecological research. Although much ecological research
has been accomplished, the potential of additional ecological understand-
ing of the atoll is great.

ACKNOWLEDGMENTS

The 1963 through 1969 field work was conducted by the Pacific Ocean
Biological Survey Program (POBSP), Department of Vertebrate Zoology,

370

Smithsonian Institution, Washington, D.C. The 1973 field work and
subsequent technical report was conducted by the Ecology Program, Office
of International and Environmental Programs, Smithsonian Instutitior,
Washington, D.C., under U.S. Air Force Contract Number F 44620-67-C-0063.

The number of people who have made contributions to this paper is
enormous. Our apologies first to those whose contributions are unknown
to us, either because we did not personally meet or because we simply
could not keep track of the many persons both on the atoll and elsewhere
who helped with transportation, supply etc.

We would like to especially thank Philip S. Humphrey, University of
Kansas, who had overall responsibility for direction of the POBSP, includ-
ing the work on Johnston Atoll. His continuing interest and stimulation
made possible all the field work and the opportunity for assembling
and publishing the data. His foresight in establishing and planning
the POBSP and in selecting Johnston Atoll as a site of primary interest
has been indispensible in bringing the work to this point. His continuing
interest in seeing the POBSP through six years of field work and in
encouraging and stimulating final writing are without doubt more respon-
sible than the efforts of anyone else for this paper.

The POBSP field directors, Winston Banko, Charles A. Ely, Robert L.
Pyle, and Richard S. Crossin, had close responsibility for logistics and
planning. The latter two in particular made it possible for the junior
author to spend long periods of time on Johnston, with little interference
from outside projects. The work of the field directors was greatly aided
in Honoluly by C. Douglas Hackman and John H. Fitch.

POBSP personnel who were involved in the Johnston studies are listed
in Appendix Table 1, together with dates they worked on the Atoll. Although
the lengths of time spent there varied greatly, it is not possible for us
to single out individuals from this list for special thanks. All made
significant contributions. Most of these people are mentioned in the text
where specific contributions are discussed.

We would like to acknowledge the outstanding contribution of
Alexander Wetmore, Secretary Emeritus, Smithsonian Institution, who
shared with us his field notes, photographs, and brilliant recollections
from his 1923 TANAGER expedition to Johnston Atoll.

For access to various other scientific manuscripts and technical
reports, we thank: Lcdr. L.R. Bauer, U.S. Navy retired; Edwin H. Bryan,
Jr., and Margaret Titcomb, Pacific Scientific Information Center, Bernice
P. Bishop Museum; Bryce Decker, Smithsonian Institution; Brian A. Harrington,§
Manomet Bird Observatory; Philip Helfrich, Hawaii Institute of Marine
Biology; Eugene Kridler, U.S. Fish and Wildlife Service; Ledr.

E.T. Thrall, 14th,Coast Guard District and M.P. Wennekens, State of Alaska,
Department of Fish and Game. We thank Jay Shepard, Bird Banding Laboratory,
U.S. Fish and Wildlife Service, for assistance in obtaining bird band
return data.

S/d

We thank Richard S. Houbrick for researching the invertebrate
literature, and J. Copper Wilson for preparing the invertebrate lists.
For identification of scientific specimens, we thank Fenner A. Chace,
Jr., F. Raymond Fosberg, Richard S. Houbrick, Roxie C. Laybourne, David
L. Pawson, Harald A. Rehder, and George E. Watson, all of the Smithsonian
Institution; Charles H. Lamoureux, University of Hawaii; and Clayton M.
White, Brigham Young University.

We would like to thank the U.S. Air Force and the U.S. Coast Guard,
as well as the U.S. Navy, for full cooperation and assistance in gathering
both field and historical data. Special thanks are extended to the many
past commanding officers and enlistéd men of the Sand Island Coast Guard
LORAN Station who contributed both assistance and information during the
POBSP studies, especially Lt. (jg.) H.A. Allen, Lt. (jg.) H.E.C. Budd,
Peet eoaNielies Haves. Lt..( jms) HA. Hurst, Lt..(je.) Re Moore, Lt. jie.)
mee Moran, Lt. (je.) J.-L. Stapleton, and,Lt., (jg. B.N. .fhurman.,. Thanks
are also extended to Col. Kenneth L. Charpilloz, USAF, Commanding Officer,
Johnston Atoll, and Lt. Don W. Troutt, USCG, Commanding Officer, Sand
Island Coast Guard LORAN Station, for allowing the senior author to visit
the atoll in November 1973.

We are indebted to the following individuals who critically read
various sections of the manuscript: Roger B. Clapp, Brian A. Harrington,
Ralph W. Schreiber, and Paul W. Woodward, former POBSP members; and
F. Raymond Fosberg, Richard S. Houbrick, Dale W. Jenkins, and George E.
Watson of the Smithsonian Institution; and Philip S. Humphrey, University
of Kansas. We thank Harald A. Rehder for writing the Mollusca section;
and Robert E. Jenkins, The Nature Conservancy; Ray Dasmann, International
Union for Conservation of Nature; Lee M. Talbot, Council on Environmental
Quality; and George E. Watson, Smithsonian Institution, for writing the
Ecological Significance section.

We are deeply indebted to the office staff of the POBSP, especially
Administrative Assistants Jane P. Church and Mae H. Esterline, and typists
Anne D. Blanchette, Saverna 0. Gennes, and Helen H. Quinn, who, respectively,
supervised, edited, and typed first draft material. We also thank Pat
Carstater, and J. Copper Wilson, Office of International and Environmental
Programs, Smithsonian Institution, for typing additional first draft
material. Maps and figures were drafted by the senior author and J.

Copper Wilson. The camera copy was typed by Barbara B. Anderson with
funding through a contract with the U.S. Fish and Wildlife Service,
Department of Interior (contract number 14-16-008-596, 3 February 1971).

Sire:
LITERATURE CITED

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Amerson, A.B., Jr. 1967. Incidence and transfer of Rhinonyssidae
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----1968. Tick distribution in the central Pacific as influenced by
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Amerson, A.B., Jr., and K.C. Emerson. 1971. Records of Mallophaga from
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Anderson, W.G. 1954. Notes on food habits of sea birds of the Pacific.
Elepaio 14: 80-84.

Anonymous. 1926. Bureau of Biological Survey creats new bird reserva-
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----1939. Johnston Island to be another American Naval base. Pac.
is eeMon. OCS eeaar

----1945. New Yorker 20: 28-29.

AOQU~~American Ornithologists’ Union. 1957. Checklist of North American
Birds. 5th ed. Lord Baltimore Press, Baltimore.

Ashmole, N.P. 1962. The Black Noddy, Anous tenutrostrts, on Ascension
Island. Part 1. General Biology. Ibis 103b: 235-273.

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383

ADDENDUM

The "Environmental Impact Statement" referred to on page 3 was made
available in November 1974 to the Council on Environmental Quality by the
U.S. Department of the Air Force (1974. Final environmental statement:
Disposition of Orange herbicide by incineration, November 1974. Environ-
mental Health Laboratory, Kelly Air Force Base, Texas. 137 pp + appendices).
This Statement indicated that the Air Force planned to incinerate approxi-
mately 2.3 million gallons of Orange herbicide in a remote area of the
Pacific Ocean downwind of Johnston Atoll. The Air Force proposed to use a
specially designed vessel, the VULCANUS, to convert the Orange herbicide
to its combustion products of carbon dioxide, hydrogen chloride, water,
carbon, and carbon monoxide which would be released into the atmosphere.
Environmentally insignificant amounts on unburned and pyrolyzates of the
herbicide and its impurity 2,3,7, 8-tetrachlorodibenzo-p-dioxin (TCDD)
may also be released into the atmosphere. The Statement concluded that
"there will be no adverse effect on the environment caused by the in-
cineration of Orange herbicide in a remote area of the Pacific."

The Statement further purposed that the principal altertative for
incineration of the Orange herbicide was to be located on Johnston Island
where "the environmental impact of the facility would be minimal." It
further stated, however, that "the potential impact on the delicate
ecosystem of Johnston Atoll and lengthy project duration make this
alternative less desirable than the proposed [primary] action."' Other
alternative methods of disposition considered were: incineration in one
of the 50 states, use of herbicide, return to manufacturer, deep well
disposal, burial in underground nuclear test cavities, sludge burial,
microbial reduction, fractionation, chlorinolysis, soil biodegradation,
and no disposal action.

The Final Environmental Statement was accepted in March 1975 by the
Council on Environmental Quality. The Air Force then sought an Ocean
Dumping Permit from the Environmental Protection Agency (EPA); public
hearings were held in Holululu and San Francisco in late April 1975.
These meetings adjourned with the Air Force further evaluating the pos-
sibility of reformulating the herbicide as an alternative. The meetings
are to reconvene in late summer or fall 1975. If at that time reformula-
tion is considered unfeasible and if the EPA Administrator decides not
to issue a dumping permit, the Air Force will pursue the principal
alternative of incineration in a facility that would be constructed on
Johnston Island. 12 May 1975.

385
Appendix Table 1. POBSP personnel participating in bird studies on
Johnston Atoll, July 1963-September 1969
Inclusive dates present
Name on Johnston Atoll Total Man-days
Amerman, Kenneth E. M9634 (21) Jul 2 Aue.
28 Aug.-22 Oct.
1964 24 Jan.-27 Apr.
1966 17 Jan.-17 May
3 Jun.-21 Jun. 309
Amerson, A. Binion, Jr. 1963 7 Jul.-29 Aug.
1964 12 Feb.-28 Feb.
22 Mar.-12 Jun.
1965 13 Jan.-20 Jan.
L966ye Suda lS Sale. 162
Anderson, Alan H. 1965 3 May.-18 Jun. 46
Bratley, David A. 1963). 22 Dec. —
1964 -24 Mar.
7 Jun.-10 Aug.
1965 17 Feb.-30 Apr. 229
Brownell, Robert L. 1968 16 Dec.-
1969 -11 Jan. 26
Bulmer, Walter 1967 14 Apr.-21 Apr. 7
Burkhalter, David L. 1967 17 Apr.-20 May 33
BLiiott, KR. Patrick 1963 21 Jul<-12 Aue. 22
Ely, Charles A. 1965) ea? Junie — 2) Jum).
1965 13 May-18 May
1966 17 Feb.-22 Feb. IDS;
Harrington, Brian A. 1966 26 May- 7 Jul.
1967 18 May— 2 Jun.
13 Jun.-22 Aug.
1968 1 Mar.-28 May
4 Jun.—19 Jul. 261
Heryford, Norman H. 1965, 13) Jan.— 97 Apr.
17 Jun.-30 Sept. 189
Huber, Lawrence N. 1969 2 Jun.-28 Jun. 26
Kepler, Cameron B. 1964 9 Aug.-19 Oct.

L965) GyApre-ls) Juli.
8 Nov.-17 Dec. 213

ae Ne

386

Appendix Table 1. (Continued)

Inclusive dates present

Name on Johnston Atoll | Total Man-days
Kirkpatrick, Ralph D. 1964 18 Oct.-
1965 -14 Jan. 88
Kleen, Vernon M. 1968 10 May-6 Aug.
13 Aug.-3 Dec. 200
Lehner, Philip N. 1964 26 Jul.-28 Sep.
6 Dec.-
1965 -15 Feb. 135
Lewis, T. James 1965 27 Sep.-5 Nov. 39
Maze, Richard L. 1965 16 Dec.-
1966 22 ebl.
19 Apr.-31 May 110
Merrill, Richard W. 1964 29 Apr.-28 Jul. 90
Pyle, Robert L. 1966 16 Aug.-19 Aug. 3
Schreiber, Ralph W. 1966° 20 Oct .=
1967 -10 Feb.
1968 13 Feb.-16 Feb.
26 Jul.-16 Aug. 17
Shelton. .P hi pC. 1966. 730) Juni 25, Oct.
L967 3 Feb.-9 May
25 Aug.-12 Sep.
1969 24 Jan.= 9 Sep. 458
Smith, frank Hs, Jr. 1966 30 Jun.-30 Aug. 61
Stadel, Dennis L. 1964 9 Oct.-29 Nov. Syak
Sundell, Robert A. 1963 27 Sep.-12 Dec. 76
Tordoff, Jeffery P. 1966 24 Feb.-16 Apr.
1968 13 Feb.- 1 Mar. 67
Wilz, Kenneth J. 1963 ...7.-Jul.-29° Aug. a3
Wislocki, George S. 1963. 20) Now
1964 -22 Jan. 63
Woodward, Paul W. 1965 JoeJule—si Aug. 47

3,216

38i/

Appendix Table 2. Annotated list* of vascular plant species recorded
from Akau Island, Johnston Atoll

Polypodiaceae
Nephrolepts sp.

Zesep. 1967:

Graminea

Cynodon dactylon

ABoeD\. 1967:

6 Sep. 1969:

"Few plants growing from wooden foundation of
building, in fairly well shaded area, close to
plants that are watered regularly." (UH, ?).

Bermuda Grass

One patch about 30 feet in diameter, and a smaller

one, both in a slight depression on the northcentral

portion of the island. Stolons spreading in all
directions. A few fruiting heads.

"...on watered lawn, on imported soil."

Daetyloetentum aegypttum Crowfoot Grass

Zesep. 1967:
Eleustne tndtca

2 Sep. 1967:

Eragrostis tenella

Z Sep. 1967:

Zea mays

6 Sep. 1969:

Fimbristylis cymosa

2 Sep. 1967:

One small patch near the south side of the island.

Goosegrass

Fairly abundant in a few small areas, vigorous,
Bie blalienbo¥een, pom (OSI mEN ye

Lovegrass

Few plants growing in partial shelter of building
near plants watered regularly. (UH, 1).

Corn

One 18 inch pale plant, with tassels, in plot.

Grows over most of the island, in about the same
density as on Hikina. A worker who had been on the
island 15 months said that these plants had become
conspicuous only since his arrival.

*Collections are shown by herbarium designation, followed by the number of

specimens in each collection; UH = University of Hawaii. Data are taken
from POBSP, 1967b and 1969.

—.

— \cwes.

wwe nme USOC X Yh,

388

Appendix Table 2. (Continued)

6 Sep. 1969: Looks much like it did in 1967, possibly thicker.
Fruiting profusely. Thick patch near north shore
in a depression.

Palmae
Cocos nuetfera Coconut Palm
2 Sep. 1967: At least two plants about three feet high, near
the dock; one fairly green, the other mostly dried
up.
6 Sep. 1969: About six trees, up to six feet; not doing too well.
Amaryllidaceae

Crinum or Hymenocallts

2“Sep. 1967: One plant, not blooming.
6 Sep. 1969: One small plant, and one six-foot plant with dried
blooms.
Casuarinaceae
Casuartna equtsettfolta Ironwood
6 Sep. 1969: Two two-foot trees by buildings; heavily watered.

Reportedly have grown five inches per month.

Polygonaceae
Coeceoloba uvtfera Sea-grape
2, Sep. 19677: One small tree; no flowers or fruits.
6 Sep. 1969: One tree, no flowers or fruits.
Amaranthaceae
Amaranthus viridis Pigweed
2° Sep. L967: A few small plants grew in area planted with

several garden species, probably brought in soil
with these plants.

389

Appendix Table 2. (Continued)

Nyctaginaceae
Boerhavia sp.

eo) tts LOGS:

6 Sep. 1969:

Aizoaceae

",.,.there were approximately 20 small Boerhavta
plants establishing themselves, and 4 Boerhavia
plants had runner systems over 4 feet long.

No Boerhavta was seen on 14 April [1965].

All these plants were in the southwest corner
of the island."

Two patches, green and fruiting, in depression,
north side.

Sesuvtum portulacastrum

1-15 Apr. 1965:

ey dul 1965:

2 sep. 11967:

6 Sep. 1969:
Portulacaceae
Portulaca oleracea

2 Sep. 1967:

Caryophyllaceae
Spergularta marina

2 sep. L967:

6 Sep. 1969:

"Several small clumps...are growing in the
southwest corner of the island--not cultivated."

"One clump...was 2 feet in diameter, and many

clumps were several inches across. There may

be as many as 100 separate plants now established,
most of them on the southwest corner of the island."

Scattered small plants, no patch as large as that
found on Hikina Island. No blooms noted, but no

close examination was made.

Scattered, blooming, not abundant.

One small plant seen, near where the Orange was
planted (near office buildings).

Less abundant than on Hikina, but widespread over
the island.

Few scattered pads; blooming.

. we

— \Cwws,

wwICl pe OU SGHOSA YY.

"OC Veer

— en oe

390

Appendix Table 2. (Continued)

Leguminosae
Acacta farnestana Sweet Acacia
2. Sep... 967) One six inch plant found growing from under
wooden foundation of building.
Phaeseolus sp. Bean
2 Sep: 1967: Planted in rich soil and watered; production
unknown.
64Sep. 51969: Growing well in imported, watered soil.
Ptsum sattvum Pea
2 Sep. 1967: Planted in rich soil and watered; production
unknown.
Vigna martina Beach Pea
6 Jul. 1965: Collected by; POBSP../.,(UH,..1).
Ruthaceae
Citrus sinensts Orange
2 Sep. 1967: One plant about three inches high, apparently
healthy, sprouted from seed. Another planted
at the same time died.
6 Sep. 1969: Two or three small trees reported growing.
Euphorbiaceae
Codtaeum vartegatum var. pictumn Croton
2 Spe wI69 One small shrub.
6 Sep. 90969: Several plants, no flowers.
Euphorbta glomertfera Spurge
ZiSCD.> L967: Few plants found in the central portion of the
island a eGUH st),
6 Sep. 1969: Fairly abundant around buildings; blooming.

3911!

Appendix Table 2. (Continued)

Anacardiaceae
Mangtfera indica
6 Sep. 1969:
Malvaceae
Htbtscus sp.

2 Sep. 1967:

Mango

Two planted trees reported.

One plant, few red blooms.

6 Sep. 1969: Three or four plants; red blooms.
Guttiferae
Calophyllum inophyl lum False Kamani
2 Sep. 1967: One small plant, no flowers.
6 Sep. 1969: One ten feet high; no flowers.
Apocynaceae

Plumeria rubra?
2 Sep. 1967:

6 Sep. 1969:

Verbenaceae
Vitex ovata
2 Sep. 1967:
6 Sep. 1969:
Solanaceae
Capsteum frutescens
2 Sep. 1967:
6 Sep. 1969:
Solanum Lycoperstcum

2 Sep. 1967:

Frangipani
One small plant; no flowers.

Two new plants; no flowers. Old one still there;
no flowers.

Few plants; blooming profusely.

One plant seen; blooming profusely.

Pepper

Several’ plants (cultivated).

Two different kinds cultivated in plots.

Few plants.

a a a

B92

Appendix Table 2. (Continued)

Cucurbitaceae

Cttrullus lanatus var. vulgarts Watermelon

2 sep. L967):

Cueumts melo

2 Sep. 1967:

Compositae

Conyza bonartensts

Z Sep. 967:

6 Sep. 1969:

Pluchea tndtca

2 Sep. L9G:

6 Sep. 1969:

Pluchea caroltnensts

2:Sep. 1967:

Tagetes sp.
6 Sep. 1969:
Zimnta elegans
2, Sep. 1967 :

6 Sep. 1969:

Few plants (cultivated).
Muskmelon

According to caretakers, these get about the
size of oranges, ripen and fall off the vines.

One plant, about 15 inches high with flowers and

mature heads, near buildings.

Around dock; fruiting.

One plant about 15 inches high found on the west
central portion of the island.

Few small plants, east southeast side.

A few plants, none over one foot high, were found
near buildings and among the cultivated plants.

Marigold
Blooming (cultivated).
Zinnia
Several plants; blooming profusely.

Blooming.

Appendix Table 3. Annotated list** of vascular plant species recorded
from Hikina Island, Johnston Atoll

Graminea
Eleustne tndica*
21 Feb. 1969:
Lepturus repens*

21 Feb. 1969:

Cyperaceae
Fimbristylis cymosa*

2 Sep. 1967:

21 Feb. 1969:

Palmae
Cocos nuctfera

21 Feb. 1969:

Moraceae

Fteus mtecrocarpa *

2 Sep. 1967:

Goose Grass
Very scarce. Collected from a small clump.
Bunch Grass

Collected from small clump--only one seen--
at top of beach on east shore.

Sedge

The dominant plant on the island. It grows on
almost the entire island, covering an estimated

two to five percent of the surface. Viewed
obliquely the ground almost appeared to be

covered in some areas. Most of the plants were
small tussocks, but there were a few clumps several
inches in diameter, with eight to ten inch fruiting
stalks.

More completely spread over island than in 1967.
Estimated ten percent of ground covered. Nearly all
in small, evenly spaced tussocks. Most with fruiting
stalks. In spots with better moisture, for example
where drainage concentrated water along the edges of
concrete structures, there was fresh green growth.

Coconut Palm
Three trees, about 2.5-3 feet high, fairly recently

planted, and apparently thriving well; two by dock
and one by building.

Banyan

One 15 inch plant, south side of the island;
probably planted.

**Collected specimens (marked *) are in the herbarium of the University

Of Hawaii. = Data are

from POBSP, 1967b and 1969.

394

Appendix Table 3. (Continued)

Aizoaceae

Sesuvtum portulacastrum

eee aloo}:

2 sep. 1967:

21 Feb. 1969:

Several small clumps on the northeast corner of
the island.

This was the second most abundant plant on the
island. The largest clump was six by nine feet,
in which the only blooms were found. Scattered,
smaller non-blooming clumps occurred.

Small, widely scattered patches occurred, mostly
around the perimeter of the island. A few blooms.

Portulaca probably oleracea way Porslane

21 Feb. 1969:

Caryophyllaceae

Spergularta martna*

2 Sep. 1967:
Zi Heb. L969):
Leguminosae

Mucuna sp. *
2 1Sepe, 1967 5
Euphorbiaceae
ATS eo are Lica

2 sep. 1967:

A few plants, with no blooms, by entrance to
office.

Widely scattered over the island, but most plants
appeared to be dead. A flat area of a few square
yards, and a small mound of coral, both. on the
southwest corner near the dock, were almost covered
with living plants, and there were a few blooms on
they plate anea.en (UB asl))).

Scattered along the periphery; scarce. Large patch
found in 1967 gone. Blooming.

One seed found on east beach.

Candlenut, Kukui

Several seeds found on east beach.

Appendix Table 3. (Continued)

Combretaceae
Terminalta catappa *

2 Sep. 1967:

Boraginaceae

Indian Almond, Kamani

Two seeds found on east beach.

Note: Although this species grows on both Sand
and Johnston, these islands are downwind and
west of Hikina, and the seeds were all on the
east beach. Therefore it seems likely that the
seeds came from elsewhere.

Tournefortta argentea Tree Heliotrope

Zirebp. 1969:

Solanaceae
Solanum Lycopersteun

2. 9ep 20967 :

21 Feb. 1969:

One plant, about a foot high, probably planted,
about 100 feet east of entrance to office.

Tomato

The one plant near the north shore had produced
some fruit, according to the caretaker.

Plant(s?) on north shore spread to three or four
feet in diameter; overmature. Several partly
dried fruits hanging to vines.

5395

396

Appendix Table 4. Annotated list* of vascular plant species recorded
from Johnston Island, Johnston Atoll

Polypodiaceae Ferns

Polypodtum scolopendrta a

Apr. 1965: Observed by C.H. Lamoureux, "growing in

bath (?) house."
Araucariaceae
Araucarta heterophylla Norfolk I. Pine

952 "Fosberg saw 3 growing in pots near terminal
in 1952. Now growing in yard of Base Com-
mander's house; one or two planted around
buildings were nearly dead."" (Doty and
Newhouse, ms.).

Apr. 1965: Collected by C.H. Lamoureux, cultivated
(escaped?) northern part of island. (UH, 1).

Ieee yf 2 Two or three relatively small trees.

Pandancaceae
Pandanus tectortus Screw-pine, Hala

Aug. 1944: "Small, but healthy; about the size of a
pineapple plant" (Bryan, 1944).

1950: "In 1950 Fosberg observed one plant cultivated
near a house. Now in the schoolhouse yard."
(Doty and Newhouse, ms.).

Jan. £954; Observed by W.V. Newhouse. Cultivated.

Apr LI6D5: Observed by C.H. Lamoureux. Cultivated.

"Motor pool, and other parts."

*Collections are shown by herbarium designation, followed by the
number of specimens in each collection. BISH = B.P. Bishop Museum;
UH = University of Hawaii; US = U.S. National Herbarium (Smithsonian
Institution). Data for 1967 and 1969 are taken from POBSP, 1967b
and 1969.

Appendix Table 4.

Graminea

Cenchrus echtnatus

1 Nov. 1946:

Nov. 1953:

1954:

ani 958i:

Apr. 1965:

1967:

Chloris barbata
1 Now. 1946:
ILC yAe

Apr. 1965:

Cynodon daetylon
Aug. 1944:

1 Nov. 1946:

1954: "Now growing south of the main runway" (Doty
and Newhouse, ms.).

Apr. 1965: Collected by C.H. Lamoureux. (UH, 1).

O67): Scattered widely, but no well-developed
stands found.

Daetyloetenitum aegypttum Crowfoot grass

Apr. L965! Collected by C.H. Lamoureux, northern part of
island.

LOGT Few plants scattered along runway. Mature

(Continued)

Sandbur
"Abundant" (Fosberg, 1949). (BISH, 1).

Collected by E.H. Walker, by building.
GUS fl).

"Common" (Doty and Newhouse, ms).

Collected by 0. and I. Degener. ''Very
ocaiie 5. CBLSH ad).

Collected by C.H. Lamoureux, northern part
Ohelstand. 7 (He)

Fairly abundant on open disturbed areas.
Mature fruits found all year.

Fingergrass
"Abundant" (Fosberg, 1949). (BISH, 1).
"Common" (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part
Of island. = a(UHL, 2)).

Bermuda Grass
"Small patches" (Bryan, 1944).

"One small patch seen" (Fosberg, 1949).
(BISH.. 1),

heads in September. (UH, 1).

397

Appendix Table 4. (Continued)

Echinochloa crus-galli Barnyard grass

Aug. 1944; Collected. by E.H. Bryan, Jr.,.(BISH,. I). :
Eleusine indica Goose grass

1 Nov. 1946: "Rare" (Fosberg, 1948). (BISH, 1).

1950: "Fosberg noted this to be abundant around

edges of airstrip and along path in 1950."
(Doty and Newhouse, ms.).

Apr. £965: Collected by C.H. Lamoureux, northern part
Of asiand a eaCUH sad).

1967: Scattered, not abundant.

Eragrostis tenella (incl. amabilis)Lovegrass

1954: "Common" (Doty and Newhouse, ms.).
Jan. 1958: Collected (as FE. amabtlts (L) Wight and Arn)
by 0. and I. Degener. (BISH, 1).
Apr. 1965: Collected by C.H. Lamoureux, sandy soil in
north part of island... (UH,)1):.
Lepturus repens Bunch grass
Jul. 1923: "Growing in low, dry, brown bunches, forming

a fairly dense, dominant stand" (Christopher-
sen, 193i)... (BSH +1):

6 Jun. 1942: Collected by Ashley Browne. (BISH, 1).
Aug. 1944; Collected by E.H. Bryan, Jr. © (BISH;, 1).

1 Nov. 1946: "Abundant " (Fosberg, 1949). (BISH, 1).
Nov. 1953: Collected by E.H. Walker. (US, 1).

1954: "Common'' (Doty and Newhouse, ms.).

Apr:. 1965: Collected by C.H. Lamoureux, northern part

Of Visland/) (UH y 3).

E96: A few bunches scattered along runway, south
side.

399

Appendix Table 4. (Continued)

Saeeharum of fictnanum Sugarcane

Apr. 1965:

Setarta vertictllata

Aug. 1944;

Observed by C.H. Lamoureux. "Cultivated, a
few plants only."

Bristlegrass

Collected thy EH aw Bryan, .Jx.. (BISH, 1).

1 Nov. 1946: "Common locally." (Fosberg, 1949). (BISH, 1).
1950-1954: "Occasional around the buildings in 1950
according to Fosberg and common around
dependent's quarters at present" (Doty and
Newhouse, ms.).
Sporobolus virgtinicus Dropseed
1954: "Near quonset hut in officer's dependents

Apr. 1965:

Cyperaceae
Cyperus rotundus
11 Sep. 1967:
Fimbristylus cymosa?

Jan. 1958:

Apr. 1965:

@ecos nuctfera
Aug. 1944:
1 Nov. 1946:

1951-1954:

quarters. This grass is a widespread grass
of brackish beach shore areas and may become
a good cover at Johnston" (Doty and Newhouse,
ms.).

Collected by C.H. Lamoureux, northern part of
island, on sand dune. (UH, 1).

Sedges

Few scattered plants, open coral. (UH, 1).

Collected by 0. and I. Degener, "common about
aieport." » (identified as”. diphylla). (BISH,.2).

Collected by C.H. Lamoureux, northern part
of island. (UHe 2):

Coconut palm
"Small, windblown, no trunk" (Bryan, 1944).
"One seedling seen" (Fosberg, 1949).
"One coconut seedling was seen by Fosberg in
1951. Several are now present, e.g., in the

schoolhouse yard and around the officer's
dependents' quarters'' (Doty and Newhouse, ms.).

mw \C Nw. er S/

— ike <¥21 2 law. Fe] C Bar

OL AMA

400

Appendix Table 4. (Continued)

Apr. 965: Observed by C.H. Lamoureux. Cultivated.
Araceae
Anthurtum andraanum Anthurium
1954: "Potted and growing in the house of Mr. S.A.

Kahoiwai'" (Doty and Newhouse, ms.).

Liliaceae
Allium ftstulosun Welsh onion
1954: "In the yard of Mr. S.A. Kahoiwai' (Doty and
Newhouse, ms.).
Alltum sp. Chives
1954: "In the yard of Mr. E. English" (Doty and
Newhouse, ms.).
Apr. 1965! Observed by C.H. Lamoureux. Cultivated.
Grdyline fruttcosa Cordyline
1954: "In the yard of B.0.Q." (Doty and Newhouse,
ms.).
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Sansevierta trifactata? Bowstring Hemp
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Amaryllidaceae
Crinum astatteum
Aug. 1944; "A few plants" (Bryan, 1944).
Crinum sp.
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Hymenocallts littoralis Spider Lily
Apr. 19651: Observed by C.H. Lamoureux. Cultivated.

401

Appendix Table 4. (Continued)

Zingiberaceae
Alpina sp.

1954:

Musaceae

Heltoconia humtlis
Apr. (L965:

Strelttzta regtnae
Apia 965:

Orchidaceae

Eptdendrum sp,

Ginger

"In the yard of Mr. S.A. Kahoiwai" (Doty
and Newhouse, ms.).

Observed by C.H. Lamoureux. Cultivated.
Bird of Paradise
Observed by C.H. Lamoureux. Cultivated.

Orchids

Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Vanda sp.
1954: "In the yard of Mr. Kahoiwai'' (Doty and
Newhouse, ms.).
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Casuarinaceae
Casuarina equisettfolta Ironwood
Aug. 1944: "Small, brown, windblown" (Bryan, 1944).
1 Nov. 1946: "Commonly planted" (Fosberg, 1949). (BISH, 1).
1951-1954: "Fosberg noted about 25 in the residential
area in 1951. Several over 45 feet high are
now doing well in the B.0.Q. area" (Doty and
Newhouse, ms.).
Apr. 1965: Collected by C.H. Lamoureux, northern part of
islands. (Heo )i.
IG7 i: At least eight to ten trees on the island, up
to 20 feet high, all apparently healthy.
723) Meloy Aeas)e Trees on top of the hospital cut down to about

ten foot stumps. Still alive.

%

Sy oh
IV ty

aw SC/ Ty

\D Ver,

TEV o\ Sea PY een Ss

LS a

402

Appendix Table 4.

Moraceae
Ficus mtcrocarpa

Apri 6 51

Polygonaceae
Coceoloba uvifera
Aug. 1944:
Novier L959):

1950-1954:

Apr. 1965:

1967:

25 Jan. 1969:

Chenopodiaceae

Chenopodium muratle

1954:

Amaranthaceae

Amaranthus dubtus

HOS

Apr. 1965:

Amaranthus sptnosus

Aug. 1944:

(Continued)

Banyan

Collected by C.H. Lamoureux, northern part
of island t.@UH als)

Sea-grape
"Small, but in good leaf" (Bryan, 1944).
Collected by E.H. Walker. (US, 1).
"Seen commonly planted near houses by Fosberg
in 1950. Several trees seen during the present

study" (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part of
istand.. ((UH.o))i.

Several trees, up to 15 feet high, fruiting
profusely in September.

Coeecoloba along bank next to runway are very
dense and much larger than before.

Goosefoot, Pigweed

"Found in the enlisted men's dependents
quonset area'’ (Doty and Newhouse, ms.).

Pigweeds

"Between quonset huts of enlisted men's
dependents quarters" (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part of
island. UH; %2)).

"A few weeds" (Bryan, 1944). (BISH, 1).

Appendix Table 4.

Amaranthus viridis

1 Nov. 1946:

1950-1954:

Nov. 1953:

Anis) 965:

Nyctaginaceae

Boerhavia sp.

O23:

Jun. 1942:
Aug. 1944:

1 Nov. 1946:

Nov. 1953:

1954:

Janes 2958:

Apr. 1965:

1967:

Bougainvillea sp»

1954:

(Continued)

"Common along fences" (Fosberg, 1949).
@irshe 1).

"Fosberg reported this as common around airport
buildings in 1950, between quonset huts at
enlisted men's dependents quarters" (Doty and
Newhouse, ms.).

Collected by E.H. Walker; probably this species.

(usin ADP

Collected by C.H. Lamoureux from northern part
Ofeaisiland= | CUE as ye

1
on the seaward slopes of the highest hill;
being sparingly found elsewhere'' (Christopher-
sens 991)... (BESHo aL).

Collected by Ashley Browne. (BISH, 1).
Collected by Ech. Bryan, Juss (BISH, 1).

"Abundant, especially in places not walked
too much" (Fosberg, 1949). (BISH, 1).

Collected by E.H. Walker. (US, 1).

"Common in yards and along main runway"
(Doty and Newhouse, ms.).

Collected by 0. and I. Degener. ‘''Common along
airport edge" (BISH, 1).

Collected by C.H. Lamoureux, from northern
pake OL) island... “EH. L)..

Scarce, found only in one place.

"In the yard of Mr. S.A. Kahoiwai'" (Doty and
Newhouse, ms.).

',..most abundant at the east end of the island,

403

Sia Oy

L

= NG

\tie,, ves) « GDWHZy

a ~ KF i

tr \O Wey, .

RRS,

Sh F=f PES ON Ba Pf PY ox

qi —™ 7. 46047 cf \<-e,

Lane

404

Appendix Table 4.

Aizoaceae

(Continued)

Sesuvtum portulacastrum

Aug. 1944;

Nov. 1953:

1954:

Jan. 1958:

Apr. 1965:

Portulacaceae
Portulaca oleracea
Aug. 1944;
1 Nov. 1946:

1954:

Apr. -1965:

Caryophyllaceae
Spergularta marina

Apr, 1965:

1967:
Lauraceae

Persea amertcana

1954:

Apr’. 1965:

"One plant seen" (Bryan, 1944). (BISH, 1).

Collected by E.H. Walker, "along shore."
(UH, 1).

"Common south of the main runway" (Doty and
Newhouse, ms.).

Collected by O. and I. Degener
airfield." \=(Bish, 1)

at "edge of

Collected by C.H. Lamoureux from sand-covered
bunker on northern part of island. (UH, 2).

Purslane
"Small patch in good leaf" (Bryan, 1944).
"Common" (Fosberg, 1949). (BISH, 1).

[listed without comment] (Doty and Newhouse,
MS) ee

Collected by C.H. Lamoureux from northern
part of island; —(UH, 1):.

Collected by C.H. Lamoureux from northern part
Of islands =a@UH I).

Scattered, not abundant.

Avocado

"Potted in the yard of Mr. S.A. Kahoiwai"
(Doty and Newhouse, ms.).

Observed by C.H. Lamoureux. Cultivated.

405

Appendix Table 4. (Continued)

Cruciferae
Lobularta marttima

1954:

Leguminosae
Acacta farnestana
Aug. 1944:

1954:

Apr 1965:

1967:

Crotalarta tneana

Apr. 1965:

Leucaena Lattstltqua

Apr. 1965:

Ptthecellobtum dulce
1 Nov. 1946:

1954:

Prosopts pallida
1 Nov. 1946:

ILC IEA

Sweet Alyssum

"In the yard of Mr. E. English" (Doty and
Newhouse, ms.).

Sweet Acacia
Lonalleaverny dry (Bryani 1944) = (BESHS 1): ;
"Only one plant seen. This growing on the
west side of B.0.Q. building" (Doty and

Newhouse, ms.).

Collected by C.H. Lamoureux from northern
part of island. (UH, 2).

One small shrub north of tennis courts, and
a few on bank north of NCO club.

Rattlebox

Collected by C.H. Lamoureux from northern
paGeworets lander. (UHee 1s).

Collected by C.H. Lamoureux from northern
part. ofwasiand.. “(UH 1)r

Manila Tamarind
"Rare, planted" (Fosberg, 1949). (BISH, 1).

"A small tree persisting under drought condi-
tions.'' (Doty and Newhouse, ms.).

Algarobe, Kiawe
"Rare, planted" (Fosberg, 1949). (BISH, 1).

[listed without comment] (Doty and Newhouse,
MS!) ie

ht A

wwIOCl ne SHH:

\S Ve.

SL ay eam eae

SA Vc)\

bo) RN

Bl)

406

Appendix Table 4. (Continued)

Vigna marina Beach Pea

Apr. 1965: Collected by C.H. Lamoureux from sand around
bunker, northern part of island. (UH, 1).

1967: Growing on side of bunker near NCO club,
blooming in May. (UH, 1).

Zygophyllaceae
Trtbulus ctstotdes Puncture Vine
Jul 923): ",.-second in abundance, being scattered
generally among the bunches of Lepturus
repens,"" (Christophersen, 1931). (BISH, 1).
6 Jun. 1942: Collected by Ashley Brown. (BISH, 1).
Aug. 1944; Collected by E.H. Bryan, Jr. (BISH,
1 Nov. 1946: "Abundant" (Fosberg, 1949). (BISH, 1).
1954: "Common" (Doty and Newhouse, ms.).
Apr. 1965: Collected by C.H. Lamoureux, from northern
parteof island...) (UH, =):
Rutaceae
Citrus aurantifolia Lime
1954: "In the yard of Mr. S.A. Kahoiwai" (Doty and
Newhouse, ms.).
Apr. 1965: Citrus sp. observed by C.H. Lamoureux.
Cultivated.
Euphorbiaceae
Codiaeum vartegatum var. ptctum Croton
1954; "Fosberg saw this in a pot at the terminal"
(Doty and Newhouse, ms.).
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Euphorbta atoto? Spurge
12 Nov. 1953% Collected by E.H. Walker, on sand along shore.

(ist -1)2

407

Appendix Table 4. (Continued)

Euphorbia prostrata Spurge
Apr. 1965: Collected by C.H. Lamoureux, from lawn in
northern portion of island. (UH, 1).
Euphorbta prob. heterophylla Spurge
1 Nov. 1946: "Abundant locally on small dune" (Fosberg,
1949), (BISH; 1).
1954: "Common" (Doty and Newhouse, ms.).
Apr. 1965: Collected by C.H. Lamoureux from northern
part of stand... (UH, 2)).
Euphorbia glomertfera Spurge
1 Nov. 1946: "Common along fences and edges of walks"

(Fosberg, 1949). (BISH, 1).

O51: "Fosberg, in 1951, saw many plants of this
species around the buildings" (Doty and
Newhouse, ms.).

1954: "Common near chapel" (Doty and Newhouse, ms.).

30° Jan. 1958: Collected by O. and I. Degener, at edge of
atreiteld. <GhiESH. I),

Apr.) 1965;: Collected by C.H. Lamoureux from northern
part of island.y (HD.

1967: Scattered along runway, in open coral. (UH, 1).
Euphorbta hirta Spurge
1954: "Common at Johnston as in many other places"

(Doty and Newhouse, ms.).

Apr. 1965: Collected by C.H. Lamoureux, northern part
Of a/silands “(UH a).
Euphorbta puleherrima Poinsettia
1954: "Potted plants around dependent's quarters"

(Doty and Newhouse, ms.).

Apr. 1965: Observed by C.H. Lamoureux. Cultivated.

\ Sas £

\
=,
“A

—

/tey

\c er, wr S/ ow

re Oe OT Z/ OG

=a Sa. leu] ae Fie

Atyiylhy

{ (IN Y /S

|

—~we

ln ee

WU

408

Appendix Table 4. (Continued)

Pedilanthus ttthymelotdes Slipper flower

1954: Listed by Lamoureux, not in Doty and
Newhouse (ms.).

Apr. 965% Observed by C.H. Lamoureux. Cultivated.
Rtetnus communts Castor bean
Aug. 1944: "Large and flourishing, as large as some of

the trees" (Bryan, 1944).
1 Nov. 1946: "Common'' (Fosberg, 1949). (BISH, 1).

1954: "Beside the path from PX to chapel" (Doty
and Newhouse, ms.).

Apr’. 1965: Collected by C.H. Lamoureux, northern part of
island 2s 4@UH, 5).

LOG: Many plants formerly (1966) growing near
southwest end of barracks area apparently
were removed during construction of swimming

pool.
Anacardiaceae
Mangtfera indtca Mango
1954: "In the yard of Mr. S.A. Kahoiwai'' (Doty
and Newhouse, ms.).
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Sehtnus terebinthifoltius Christmas Berry Tree
1 Nov. 1946: Observed? [Not in Fosberg, 1949].
1950: "Fosberg in 1950 saw this planted near houses"
(Doty and Newhouse, ms.).
Tiliaceae

Triumfetta procumbens

Aug. 1944: "Small, in good leaf" (Bryan, 1944).

Appendix Table 4. (Continued)

Malvaceae

Htbtscus tiltaceus

mont . 1923):

1g)51e

Ape, L965):
1967:

Htbtseus sp.
Apr. 1965:
Thespesta populnea

Aug. 1944:

1 Nov. 1946:

1954:
Apr. 2965:
1967:

Stda sp.
Aug. 1944:
Sterculiaceae
Waltherta tndica

Apr 2965:

1967:

Hau

"Planted 8 clips of hau (Hibiscus tiltaceus)
on Johnston..." (Wetmore, ms. b).

"Fosberg in 1951 saw large trees of this
around the base buildings which were then

somewhat chlorotic'’ (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part
ofeasilandos “(UH 1).

Tree near NCO club blooming profusely in
early September. (UH, 1).

Observed by C.H. Lamoureux. Cultivated.
Milo tree, Portia-tree

"Leaves were very small" (Bryan, 1944).
(BiESH 1) .

"Commonly planted" (Fosberg, 1949). (BISH, 1).

"Around base buildings and in 1952 growing
at the terminal'' (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part of
fistand. a CUHe)\.

Tree by NCO club with mature fruits in
September. (UH, 1).

"A few plants, growing well" (Bryan, 1944).

Collected by C.H. Lamoureux, northern part
Of mistand = (UH Sl).

Few plants found between taxiway and runway,
northeast end, and on south side of runway
@Hy 1).

409

« YyZ

[=
SY,

FI 4

\e!

a
==)

eo \Ckey er s/

T 3 \ACG 69) CO 0P®RSARL

= <i
FS =

cm A’.

410

Appendix Table 4. (Continued)

Guttiferae
Calophyllum tnophyllum False Kamani
Aug. 1944: "Small, shrubby" (Bryan, 1944).

1951-1954:

Apr. 965%

Combretaceae

Termtnalta catappa
Aug. 1944:
1 Nov. 1946:

1952-1954:

Apr 965)

9678:

Araliaceae

Brassata actinophylla

1 Nov. 1946:

Ug 5i2

1967):
Polysetas guilfoylet

Apr. 1965:

"Fosberg reported this tree in 1951, to be
2-3 meters tall. Now along the north side
of the street between the officer's depen-
dent's quonsets and base commander's house"
(Doty and Newhouse, ms.).

Collected by C.H. Lamoureux. Cultivated on
northern part of island. (UH, 1).

Indian Almond, Kamani

"Small, shrubby" (Bryan, 1944).

"Abundantly planted" (Fosberg, 1959). (BISH, 1).

"Fosberg observed that the large trees of
this species around the base buildings were
somewhat chlorotic. In 1952, he recorded one
as growing in a pot at the terminal" (Doty
and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part of
island. @UHG. 1).

Several trees with mature fruits in September.

Octopus tree
Observed [Not in Fosberg, 1949].

"Fosberg saw one in a pot at the terminal in
1952"" (Doty and Newhouse, ms.).

One plant found north of tennis courts.
Wild coffee

Observed by C.H. Lamoureux. Cultivated.

411

Appendix Table 4. (Continued)

Caricaceae
Cartca papaya ' Papaya
1954: "In the yard of Mr. S.A. Kahoiwai" (Doty
and Newhouse, ms.).
Plumbaginaceae

Plumbago auriculata Plumbago, Leadwort

1954: "In the yard of Mr. E. English" (Doty and
Newhouse, ms.).
Apocynaceae Y
Catharanthus roseus Madagascar Periwinkle 4
1954: "Sprawling shrub in schoolhouse yard" (Doty L
and Newhouse, ms.). Y
Nertum oleander Oleander »
Apr. 1965: Observed by C.H. Lamoureux. Cultivated. ke
Plumerta acuminata Frangipani ”
1954: "In the yard of Mr. S.A. Kahoiwai" (Doty and Bs
Newhouse, ms.). lu’
BIG: One blooming in September near base exchange; tr
a few other small plants seen, not blooming. ‘
Plumeria rubra Frangipani \
17%
APE. 965): Observed by C.H. Lamoureux. Cultivated. ¥
Thevetta peruviana var. aurantica ‘
1 Nov. 1946: "Planted, not common" (Fosberg, 1949). CBESH = s)y. r
1954: Listed, no notes (Doty and Newhouse, ms.). ay
s
Thevetta peruviana (=neretfolia) Yellow Oleander ihe
Ie
Aug. 1944: * "Small, depauperate" (Bryan, 1944). i
1953-1954: "At southwest corner of B.0.Q, building: This and

I, thevettoides should be destroyed as they are
very poisonous plants. One was removed in 1953 at

E.H. Bryan's suggestion, and 11 may be gone now"
(Doty and Newhouse, ms.).

. c Rf

412

Appendix Table 4. (Continued)

Convolvulaceae
Ipomoea pes-caprae
Aug. 1944:

1954:

Apr. 1965)

1967:

Ipomoea maerantha
Aug. 1944:
Merremta tuberosa

1954;

Hydrophyllaceae
Nama sandwicensts

1967:

Boraginaceae
Cordta subcordata
Aug. 1944:
Cordta sebestena
1 Nov. 1946:

1954;

Apr... 1965:

Beach Morning-glory
"A few vines" (Bryan, 1944).

"On trellis work and fence, south side of
commissary" (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux on sand dune,
northern part of island. (UH, 1).

Widely used for binding newly dredged coral;
growing vigorously and blooming profusely in
most locations all year.

Wild Morning Glory
"Probably this species" (Bryan, 1944).

Wood Rose

"In the yard of Mr. S.A. Kahoiwai" (Doty and
Newhouse, ms.).

On open coral between the northeast end of
the taxiway and the main runway. (UH, 1).

Kou

"Small, probably this species" (Bryan, 1944).
Kou, Geiger tree

"Planted, rare’: (Fosberg,, 1949). (BISH Ss

"In the schoolhouse yard'' (Doty and Newhouse,
ms.)

Collected?

413
Appendix Table 4. (Continued)
Heltotropium curassavicum
Ape. 1965: Collected by C.H. Lamoureux, from residential
area... (UH. 1).
1967: Widespread, but not abundant, blooming in
September.
Tournefortta argentea Tree Heliotrope
Aug. 1944; "Stunted, but in good leaf" (Bryan, 1944).
1 Nov. 1946: "Planted, occasional" (Fosberg, 1949). (BISH, 1).
12 Nov. 1953: Collected by E.H. Walker "along shore."' (US, 1).
1954: "Between base commander's house and officer's
club" (Doty and Newhouse, ms.).
Apr. 1965: Collected by C.H. Lamoureux, northern part of
tsiand. (UH; ©).
1967: Several trees, growing vigorously, blooming
all year.
Verbenaceae
Stachytarpheta jamatcensts
1967: Few scattered plants on open coral between the
northeast end of the taxiway and the main run- SS
Welyies) (CEE. 2). “s
\
Vitex ovata 2
Aug. 1944: "small, but hardy" (Bryan, 1944). (BISH, 1). r
1 Nov. 1946: "Planted, occasional" (Fosberg, 1949). (BISH, 1). B
1954: "Across the street north of the infirmary and U
in the B.0.Q. yard" (Doty and Newhouse, ms.). =
7)
Apr. 1965: Collected by C.H. Lamoureux, northern part of .
island. ~(UH, 1):
ESG7': Several large bushes on bank north of NCO club, :
and few on the south side of NCO club, used by
nesting Red-tailed Tropicbirds. With blooms and 4,

fruits in September. (UH, 1).

414

Appendix Table 4. (Continued)

Solanaceae
Capsteum frutescens Pepper
1954: "In the yards of Mr. S.A. Kahoiwai and Mr.

E. English" (Doty and Newhouse, ms.).
Nteottana glauca
2 Nove 71953: Collected by E.H. Walker by building. (US, 1).

1954: ",..near tennis court opposite enlisted men's
dependent's quarters'’ (Doty and Newhouse, ms.).

Apr 2.1965): Collected by C.H. Lamoureux, northern part of
stand... (UH. 1),
Solanum lyeoperstcum Tomato
1954: "Cultivated near enlisted men's dependent's

apartments. Only one plant seen" (Doty and
Newhouse, ms.).

Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Solanum melongena Eggplant
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Bignoniaceae
Tabebuta pentophylla West Indian Boxwood
Apr. 1965: Collected by C.H. Lamoureux. Cultivated,
northern part of island. (UH, 1).
Rubiaceae
Gardenta sp.
1954: "In the yard of Mr. S.A. Kahoiwai" (Doty and

Newhouse, ms.).
Coprosoma sp.

Apr. 1965: Observed by C.H. Lamoureux. Cultivated.

415

Appendix Table 4. (Continued)

Cucurbitaceae

Cttrullus lanatus var. vulgaris

1954:

Goodeniaceae
Seaevola taccada
Aug. 1944:

1 Nov. 1946:

i2eNovi. 2953):
1954:

Apr. 1965:

L967:
Compositae
Btdens pilosa

Ape. 965:

Conyza bonartensis

1954:
Apr. 1965):
1967:

Emtlta sonchtfolia
Apr. 1965: *
Heltanthus annuus

ADE. 2u965

Watermelon

"In the yard of Mr. S.A. Kahoiwai'' (Doty and
Newhouse, ms.).

"Small, but sturdy" (Bryan, 1944).

"Rare. Under Messerschmidia [Tournefortia]
tree" (Fosberg, 1949). (BISH, 1).

Collected by E.H. Walker. (US, 1).
"Near officer's club" (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part of
Teland CUE, al)

Scattered, not abundant, blooming in September.

Bur-marigold

Collected by C.H. Lamoureux, northern part of
is andremmC(UH 2.)

"South of the main runway" (Doty and Newhouse,
mS ic

Collected by C.H. Lamoureux, northern part of
fistands.,(s(UH, os.

Scattered in open area north of NCO club,
mature heads in September.

Observed by C.H. Lamoureux. Weed.
Sunflower

Observed by C.H. Lamoureux. Cultivated.

‘
BR
‘U
i

y
=<

\at. xe Wy.

~
rT

| Cc Sf

416

Appendix Table 4. (Continued)

Pluchea tndtca

Aug. 1944;

1 Nov. 1946:

IZ Novi 9 53):

1950=1951:

Apiee 719.65):

1967:

Pluchea carolinensis
1 Nov. 1946:
12) Nov .* 1953:
1951-1954:

Apr. +1965:

1967:

1969:

Pluchea x Fosbergit

Apr. 1965:

"Dry-looking, growing well" (Bryan, 1944).
(Gaesel > ib).

"Uncommon, five or six plants seen"
(Fosberg, 1949). (BISH, 1).

Collected by E.H. Walker. (US, 1).

"Reported common around airport buildings in
1950 and noted again in 1951 as 'common" by
Fosberg'' (Doty and Newhouse, ms).

Collected by C.H. Lamoureux, northern part of
island. (UH j=).

"Scattered, not abundant, mature heads in
September."

"Occasional (Fosberg, 1949). (BISH, 1).
Collected by E.H. Walker.(US, 1).

"In 1951 Fosberg found this less common than
in 1946. It is still common now in 1954 south
of the main runway" (Doty and Newhouse, ms.).

Collected by C.H. Lamoureux, northern part of
standiouws (UHR w>)r.

Abundant on south side of runway, where it

is attracting nesting Red-tailed Tropicbirds.
Mature heads in September. POBSP recommended
it be removed as a hazard to aircraft.

All bushes along south side of runway are gone.
Most of those remaining in other areas are well
trimmed and present poor habitat for tropicbirds.

Collected by C.H. Lamoureux, northern part of
island (UH 11).

417
Appendix Table 4. (Continued)
Sonechus sp. (oleraceus x asper)? Sow-thistle
1 Nov. 1946: "Occasional" (Fosberg, 1949). (BISH, 1).
1954: "Common around dependent's quarters" (Doty
and Newhouse, ms.).
Apr. 1965: Collected by C.H. Lamoureux, northern part of
Psion CUH.) 1k
"The Sonehus on Johnston Atoll has previously
been called S. oleraceus and S. asper. The
collections I have seen are all t intermediate
between these two species" (Lamoureux, pers.
comm.).
1967: Blooming in open area north of NCO club.
Tagetes sp. Marigold
Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
Vernonta ctinerea Ironweed
1 Nov. 1946: (Not in Fosberg, 1949).
1954: [Listed without comment] (Doty and Newhouse,
ms.).
Apr. 1965: Collected by C.H. Lamoureux, in shade of
buildings near navy decks. (UH, 1). Ne
Xs
1967: Scattered, open coral near barracks between \
NCO club and Base Exchange. (UH, 1). Fe,
Zimnta elegans Zinnia
Apr. 1965: Observed by C.H. Lamoureux. Cultivated. Be
U
1%

418

Appendix Table 5. Annotated list* of vascular plant species recorded
from the man-made portion of Sand Island, Johnston

Atoll
Graminea
Cenchrus echtnatus Sandbur

Jude. 1963 "Found around buildings and on causeway of
man-made portion" (POBSP, 1964).

Apr. 1965: Observed by Lamoureux?

LO G7": In September the only healthy stands were
around the fuel tanks on the west side of the
man-made portion of the island, where con-
tinued disturbance seems to keep competitors
out. Mature- fruits were found on most plants
in September.

Cynodon dactylon Bermuda Grass

Tule 963s "Found only around buildings on man-made
portion" (POBSP, 1964).

Apr. L965: Observed by Lamoureux?

1967: Spread during the winter (1966-1967) both
from previously established plants and from
one inch squares set around buildings by the
Coast Guard in October 1966. Cessation of
close mowing and weekly raking allowed new
stolons to cover much of the bare sand around
the buildings.

1969: Spread greatly around buildings, probably
because of better care by USCG--less frequent
and higher mowing.

Daetyloetentun aegypttum Crowfoot Crass

Apiga e965): Collected by C.H. Lamoureux, weed in
Cynodon lawn. (UH, 1).

May 1967: Collected by P.C. Shelton, open coral dredged

about L963), CUES le).

*Collections are shown by herbarium designation, followed by the
number of specimens in each collection (UH = University of Hawaii).
Data from 1967 and 1969 are taken from POBSP, 1967b and 1969.

419
Appendix Table 5. (Continued)
Digttaria sanguinalts? Crabgrass
Apr: 1965: Collected by C.H. Lamoureux, on causeway.
(UH 92)
Eleustne tndtea Goosegrass
Jul. 1963: "Scattered plants on man-made portion"
(POBSP, 1964).
Apr. 1965: Observed by Lamoureux?
LOG? There was no noticeable change in the scattered
patches on the west portion.
Lepturus repens
made 1963: "Found along causeway and scattered on man-
made portion" (POBSP, 1964).
Apr. 1965: Observed by Lamoureux?
Seterta vertictllata Bristlegrass IS
gy
Jud, 1963's "Scattered plants on original and man-made =
portions, not common" (POBSP, 1964). Py,
Cyperaceae NS
>
Fimbristylts cymosa? Sedge °
Apr. 1965: Collected by C.H. Lamoureux, "rare, few in Ys
lawn and on road to photo shack." (UH, 1). ‘
\
1967: Found only on the newest coral on the man-made
(west) end of the island. All plants had _
mature heads in May and September. (UH, 1). v)
Palmae B
Cocos nuctfera Coconut Palm U
ON
sul. £963: "Planted around buildings only on man-made ¥)
portion" (POBSP, 1964).
v
Apr. L965: Observed by C.H. Lamoureux. Cultivated.
1
J
4
:

Appendix Table 5.

1967:

1969:

Liliaceae
Aloe sp.

Jul. £963):

Apr. 1965:

1967:

1969:

Amaryllidaceae

Crinum sp. or Hymenocallis

1967:

1969:
Bromeliaceae
Ananas comosus

Apr. 1965:

(Continued)

Two mature nuts were produced in early 1967 on
the single 15 foot tree growing at the northwest
corner of the tennis court. By September
several new nuts had grown to two to three
inches, but most of these fell off as they
reached this size. The few seedlings planted

in October 1966, all on the west portion, were
alive through 1967, but none was growing
vigorously.

Several small trees have been set, especially
around the barracks buildings. The larger

tree at the west corner of the tennis court

has several nuts of various sizes, some probably
mature, and others barely past the bloom stage.
Tree has grown noticeably since 1967.

Aloe

"One plant on man-made portion; obvious
introduction" (POBSP, 1964).

Observed by C.H. Lamoureux. Cultivated.
The single plant in front of the Coast Guard
office was still present, but not blooming or

growing vigorously.

The single plant in front of the USCG office
was blooming profusely in February.

Several of these were planted in front of the
Coast Guard office in October 1966; one bloomed
in August 1967.

No trace now.

Pineapple

Observed by C.H. Lamoureux. ''Two tops planted,
unhealthy."

421

Appendix Table 5. (Continued)
Casuarinaceae
Casuarina equtsettfolta Ironwood
Apr. 1965: Observed by C.H. Lamoureux. "Cultivated,

1967:

969:

Moraceae
Ficus mticrocarpa
Apres, 1965):

1967:

1969:

Urticaceae
Pilea mterophylla

Apr. 1965:

Polygonaceae
Coecoloba uvtfera
Apr. 1965:

1967:

doing well."

One tree about ten feet high grows south of

the Coast Guard office. It was transplanted
from near the tennis courts in early 1967, and
has grown little since.

The single tree south of the Coast Guard office
is thriving, and has grown a little since 1967.

Now looks much more healthy than just after
being transplanted.

Banyan
Observed by C.H. Lamoureux. Cultivated.
One tree, about 15 feet high, west side of
the man-made part of the island. In bloom in

September 1967.

The tree by the tennis courts appears to be
healthy.

Artillery Plant
Collected by C.H. Lamoureux, growing as weed

in can with Poinsettia at signal building on
causeway. (UH, 1).

Sea-grape
Observed by C.H. Lamoureux. Cultivated.

One small tree in front of Coast Guard office.

422

Appendix Table 5. (Continued)

Chenopodiaceae

Chenopodtum murale

Apr. 1965)

1967:

Amaranthaceae

Amaranthus dubtus

Apr. 1965:

Amaranthus viridis

REG AEEKoy5}8

Apr. 1965:

Nyctaginaceae
Boerhavta sp.

Sule 903:

Apr. 1965:

Bougatnvtllea sp.

Apr. 1965: Observed by C.H. Lamoureux, "cultivated on
trellis."
Aizoaceae
Tetragonta tetragontotdes New Zealand Spinach
Apr. L965: Observed by C.H. Lamoureux, cultivated near

Goosefoot, Pigweed
Collected by C.H. Lamoureux, sandy soil around
signal building on causeway and as weed in
Cynodon lawn near quarters and boat dock.
(UH, 2).
A small patch grows between the fuel tanks and
dock on the west side of the man-made part of
the island. In bloom in September 1967.

Pigweeds

Collected by Lamoureux, "in coral soil near
signal building on causeway." (UH, 1).

"Few scattered plants around buildings on
man-made portion" (POBSP, 1964).

Collected by Lamoureux in lawn around
quarters... (UH, 1).

"Scattered plants found on original and man-
made portions'' (POBSP, 1964).

Observed by Lamoureux?

dock.

423

Appendix Table 5. (Continued)

Sesuvtum portulacastrum

Jade L963:

Apr. 1965:

1967:

Portulacaceae
Portulaca oleracea

Jia MRE (esis

Apr. 1965:
Caryophyllaceae
Spergularia marina

Jjehs Aussie

Ape. 1965:

1967:

Lauraceae
Persea americana

Apr. 1965:

",...along causeway and around housing
facilities of man-made portion" (POBSP, 1964).

Collected by C.H. Lamoureux near quarters.
CUE)

Along the causeway and on the man-made portion
of the island, seedlings of this species grew
abundantly during September and October,
forming a fine-textured, almost mold-like
covering over the bare coral near clumps of
mature plants.

Purslane

"Occurs as scattered plants on original and
man-made portions" (POBSP, 1964).

Observed by Lamoureux?

"Occurs only on man-made portion around
buildings" (POBSP, 1964).

Collected by C.H. Lamoureux from coral sand
on causeway and near quarters. (UH, 2).

Comparatively rare on the west (man-made) part
of the island, and it seems to be losing in
competition with Sesuvium, Cynodon, and
Euphorbta.

Avocado

Observed by C.H. Lamoureux. "Cultivated,
unhealthy."

”

.
sh
y
—

UU
<=,

\ae, .. wo:

~
ty’

Wo

LRN

| c

424

Appendix Table 5. (Continued)

Cruciferae
Lobularta maritima Sweet Alyssum
Apu. 1965: Collected by C.H. Lamoureux on causeway
leading to old photo shack. (UH, 1).
Rosaceae
Ertobotrya japontca Loquat
Apr. 1965: Collected by C.H. Lamoureux; "cultivated on
man-made part of island, but not surviving
well" (UH. al)
Leguminosae

Leucaena lattsiliqua

Apr .: 1965: Collected by C.H. Lamoureux, south of pump-
house, in Cynodon lawn. (UH, 1).
Zygophyllaceae
Tribulus etstotdes Puncture Vine
Jul. 1963: "Scattered plants found on man-made portion"
(POBSP, 1964).
Apr. 1965: Observed by Lamoureux?
Euphorbiaceae
Codtaeum vartegatum var. pictum Croton
Apr. -1965% Observed by C.H. Lamoureux. Cultivated.
Euphorbta glomertfera Spurge
LO Gwe: Scattered in open new coral on the south side
of the man-made part of the island. Blooming
in May and September. (UH, 2).
Euphorbta hirta Spurge
Apr. 1965: Collected by C.H. Lamoureux, weed in Cynodon

lawn near quarters. (UH, 1).

425

Appendix Table 5. (Continued)

1967:

Euphorbia puleherrima

Apr. 1965:
Malvaceae
Htbtseus sp.

Apr, 1965:

Combretaceae
Terminalta catappa
Apr. 1965:
Myrtaceae
Eucalyptus sp.

Jide 9 O3/s

Apocynaceae
Nertum oleander

jaar. 1963:

Convolvulaceae
Ipomoea pes-caprae

Api.) 1965):

1967:

Dense patches in undisturbed areas around the

buildings and tennis court on the man-made end

of the island. In bloom in September. (UH, 1).
Poinsettia

Observed by C.H. Lamoureux. Cultivated.

Observed by C.H. Lamoureux, "cultivated but
unhealthy."

Indian almond, Kamani

Observed by C.H. Lamoureux, cultivated.

"One plant, found only on man-made portion"
(POBSP, 1964).

Oleander
"Found only around buildings on man-made

portion of island. Definitely introduced"
(POBSP, 1964).

Beach Morning-glory

Observed by C.H. Lamoureux. ‘'Cultivated
around quarters."

A small clump near the south side bloomed
profusely in spring and summer.

426

Appendix Table 5. (Continued)

Boraginaceae
Cordia sebestena Kou, Geiger-tree
Apr’. 1965: Observed by C.H. Lamoureux. Cultivated.

Heltotroptum curassavtecum

Jul, 1963% "Occurs only on man-made portion" (POBSP,
1964).
Apr. 1965: Collected by C.H. Lamoureux, west end of

causeway. (UH, 1).

LOGY. Showed no change from 1966 through 1967.
It was scarce, scattered around the outer
parts of both portions of the island. All
plants were blooming both in spring and late
summer .

Tournefortia argentea Tree Heliotrope

gud. 1963): "One large plant and many very small plants
on man-made portion" (POBSP, 1964).

Apr. 1965: Observed by C.H. Lamoureux. Cultivated.
G7: Planted around buildings on the man-made end

of the island, bloomed and produced fruit
profusely during the spring, summer, and fall.

19169): Present.
Solanaceae
Solanum lycoperstcum Tomato
OG One plant south of the tennis courts flowered

and produced a few three-inch fruits in spring
1967. A few flowers were present in September.

1969): Couldn't find the plant south of the tennis
courts; the area is now covered with large Pluchea
odorata bushes.

427

Appendix Table 5. (Continued)

Goodeniaceae

Seaevola taecada

afd. 2963):

Apr. 1965:

1966:

1567 2

£969:

Compositae
Conyza bonartensts

Apr. 1965:

1967:

Pluchea indica

Apr. 1965:

"Occurs only on man-made portion; appears
to have been planted around buildings; less
than a dozen plants" (POBSP, 1964).

Collected by C.H. Lamoureux, cultivated near
maintenance building. (UH, 1).

Seaevola growing near the Coast Guard buildings
on the west portion of the island bloomed and
bore fruit throughout the summer and early
Pale.

On the man-made end of the island, Seaevola
continued to grow well and provide most of

the nesting habitat for Red-tailed Tropicbirds.
Flowers were produced during the spring, and

in September, both flowers and fruits were
found.

Those around the Coast Guard power house have
grown noticeably since 1967, and had fruits

in February. Those planted along the west
seawall, behind the fuel tanks, in October
1966, had spread laterally to three to five
feet in diameter, but had not attained much
vertical growth. They were green and vigorous.

Collected by C.H. Lamoureux, lawn west of
Eennis court. (UB; 1).

Scattered on the newest coral on the south

side of the man-made end of the island. Plants
that were blooming in the spring were mostly
matured and partly dried up by September.

(UHG 2).

Collected by C.H. Lamoureux, west end of
causeway, in coral sand. (UH, 1).

428

Appendix Table 5. (Continued)

1967:

1969:

Pluchea carolinensts

Apr. 1965%

1967:

1969:

A few plants occur on the southeast side of
the man-made portion of the island; growing
and spreading slowly, but none large enough
to be conspicuous. Blooming in May.

The stand of this species growing on the
eastern part of the area southeast of the
barracks appears to have spread little since
1967, probably because of being crowded by
Pluchea odorata, which is much more aggressive
in this area. The plants, however, appear
vigorous.

Collected by C.H. Lamoureux, coral sand at
west end of causeway. (UH, 1).
",..appeared to have grown over the winter,

but no quantitative data were obtained. During
the summer of 1967 the plants on the south side
of the west portion of the island grew and
thickened rapidly, providing much more
potential tropicbird nesting habitat."

Spectacular increase since 1967.

Sonehus sp. (oleraceus x asper)? Sow-thistle

JuLe963%:

1967:

1969:

Vernonta etnerea

Apr, 965%

1967:

"Occurs only on man-made portion; scattered
plants." (POBSP, 1964).

Occurred scattered around the buildings and
open areas on the man-made end. In early
spring it was in all stages of bloom and fruit,
but by May most plants had mature fruits. By
September many of the plants had partially
dried up.

Scattered plants occur...along the causeway;
also on other areas, especially near pump-
house, etc.

Collected by C.H. Lamoureux, "weed in Cynodon
llawn near quarters... %i7 (UH, 1).

Grew scattered around the tennis court, not
abundant. Blooming in September. (UH, 1).

429

Appendix Table 6. Annotated list* of vascular plant species recorded
from the original portion of Sand Island, Johnston

Atoll
Graminae
Cenchrus echinatus Sandbur
wut. 1963: "...found on original portion along road at
junction of causeway. Plants clumped, sandspur
type” (POBSP, 1964),
Apr. 1965: Collected by C.H. Lamoureux, original part of
island, western part. (UH, 1).
1966: Other grasses, including Cenchrus echtnatus... 4
which grow only along the road on the north- GY
east corner of the island (original portion) w
were scarcely evident in July, but by October
formed a lush cover over the area. UT
D>
T9G7:: Cenehrus echtnatus seems to be losing ground ”
on the island. In the spring of 1967 it was yp
scarce on the original portion of the island, 5S
most of the patches found last year having
been crowded out by other grasses. In
September only a few plants could be found iy
on the northwest corner of the original Z
portion....Mature fruits were found on most py
plants in September. SS
U1
Cynodon dactylon Bermuda grass
SS
Apr. 1965: Collected by C.H. Lamoureux, "mostly beside x
roads, not common." (UH, 1). \
1966: Other grasses, including...Cynodon daetylon ve
...which grow only along the road on the vs
northeast corner of the island (original A
portion) were scarcely evident in July, but 2
by October, they formed a lush cover over the ish
area, with flowering plants.
eS Gre The two patches on the original end of the rR
island, one along the road to the transmitter .
and one on the south side of the island,
showed little change from 1966 through 1967. 1S
All plants had mature heads in September 1967. 14
9, Lae A
*Collections are shown by herbarium designation, followed by the Y,
number of specimens in each collection (BISH= B.P. Bishop Museum; #

UH=University of Hawaii. Data from 1967 and 1969 are taken from
POBSP, 1967b and 1969.

430

Appendix Table 6.

IOS

(Continued)

On original portion, three patches growing
and flowering during February, one on each
side of the road nearly to the transmitter
building, and one next to the northeast side
of the concrete slab near the south shore.
This last seems to be perhaps smaller than it
was in 1967.

Daetyloetentum aegypttum Crowfoot grass

1967:

Eleustne tndtca

Jul. 1963:

Apr. 1965:

1967:

1969:

Other introduced grasses, including...
Daetyloetenium aegyptium, were profusely
flowering in late February and March, but by
May had turned brown and were beaten down by
birds on the east portion. However, they
still afforded many times more ground cover
than in summer 1966.

Dactyloetentum aegypttum formerly had been
found only on the man-made end of Sand Island,
but in September 1967, at least two small
clumps with mature fruits were found on the
southeast side of the original part of the
island.

Goosegrass

"Very few plants on original portion" (POBSP,

1964).

Collected by C.H. Lamoureux, in coral sand
around transmitter building. (UH, 1).

Other introduced grasses, including Eleusine
indica...were profusely flowering in late
February and March, but by May had turned
brown and were beaten down by birds on the
east portion. However, they still afforded
many times more ground cover than in summer
1966.

"Hleustne indica appeared to be losing ground
to the native Lepturus on the original part
of the island."

Spread greatly, especially on areas previously

almost barren along the old roads on the south-
west corner of the island; healthy stands occur
in several other parts of the original portion

of the island, apparently occupying a position

intermediate between Lepturus and Boerhavia in

succession.

431

Appendix Table 6. (Continued)

Lepturus repens

a 923); ",...growing in low, dry, brown bunches,
forming a fairly dense, dominant stand"
(Christophersen, 1931). (BISH, 1).

wal. 1963: "Covers 50 percent vegetated area of original
portion" (POBSP, 1964).

Ape. 1964: Collected by C.H. Lamoureux, west part of
original portion, and southwest islet.
CUE, 2 in

1966: Dry and brown in July with no evidence of

recent new growth. There was no evident
difference between areas used by Sooty Terns
and the small area north of the road to the
transmitter building that was not used by
terns. During August new growth of Lepturus
became evident, and by October, green new
growth overshadowed the dry, decadent old
growth, giving a green appearance to the
west portion of the original island. Most

of the clumps were in flower by then as well.

1967. Spread considerably from October 1966 to
February 1967, but it was relatively dry and
not flowering in February. By the end of
March, it was at least as green as in October
1966 and covered much more area (see especially
photos of Sooty Tern plot #3). It began
blooming in April and May, although by early
May its greenness had begun to decrease. In
September it was green and lush, with blooms
and fruits, even where it had been overrun by
Sooty Terns all summer.

Mare 9 69): Appeared to be less vigorous than other
species, especially Hleustne, which was greener
than Lepturus. In places, new blooming heads
were seen, and in general the species appeared
to have ended a period of growth. Isolated
patches, apparently new, or at least recently
becoming conspicuous, appear on the west side
of the islet, and various places on the east
half of the island. The stand along the east

beach north of the east bunker is all but iu

extinct, for no apparent reason. Christmas

Shearwaters nesting there are now tunneling into 4
U

432

Appendix Table 6. (Continued)

and under dense Boerhavia. Could it be that
Boerhavia actually squeezes out the Lepturus?

Setarta verttctllata Bristlegrass
Jul. 1963; "Scattered plants....not common" (POBSP, 1964).
Apr. -1965: Collected by C.H. Lamoureux, along road,
western portion, original part of island.
QUHG 1D):
1966: Other grasses, including...Setartq...which

grow only along the road on the northeast
corner of the island (original portion) were
scarcely evident in July, but by October,
they formed a lush cover over the area with
flowering plants up to 2.5 feet high.

1967: Profusely flowering in late February and
March, but by May had turned brown and were
beaten down by birds on the east portion.
However, they still afforded many times more
ground cover than in summer 1966.

Also appeared to be losing in competition
with Lepturus on the original part of the

island.
1969: Little found.
Chenopodiaceae
Chenopodium murale Goosefoot, Pigweed
Apr. 1965: Collected by C.H. Lamoureux, beside road,
in’ coral isands: (1 (UHS71).
Amaranthaceae Pigweeds

Amaranthus dubtus
Apr. 1965: Collected by Lamoureux from "SW portion of
original portion of island and west portion
of original partlof: islands) (UH,2)%

Amoranthus virtdts

Jul. 1963: "Found in the central and west area of the
original portion " (POBSP, 1964).

433

Appendix Table 6. (Continued)

Apr. 1965: Collected by C.H. Lamoureux from southwest and
west portions. (UH, 2).

1966: Sparse and low in July, but grew vigorously
in several areas by September and October.

1967: Grew vigorously and bloomed during spring 1967
in central area south-southwest of the trans-
mitter. Patches along the northwest side of
the island, especially along the road, had
been crowded out by grasses. Black Noddies
roosted on the two largest clumps (SSW and SE)
in March, and in April three nests were built
on the southeast clumps, all about a foot off
the ground. By September only the major
clumps were still intact, while seedlings
that were growing around them during the spring
had been destroyed by Sooty Tern trampling.
Smaller clumps north of the transmitter and
near the northwest side of the island were
still intact, but not as vigorous as in 1966
because of more competition from other plants,
especially grasses.

1969: Appeared to have reached maturity and begun

to dry up during February....At the beginning
of the month, most plants were green, but by
the first of March, had turned brown, and were
shedding seeds profusely. Distribution little
changed since 1967, but patch southeast of
transmitter much reduced from 1967 levels.
None there now is capable of supporting a
Black Noddy nest. Also, that north of trans-

Nyctaginaceae mitter has further lost ground to grasses.

Boerhavia sp.

Sree 9235's " ...more abundant than Tribulus ctstotdes,
but neither begins to compare with the bunch
grass in extent" (Christophersen, 1931).
(BISH? a).

Jule 963s "Scattered plants found on original and man-
made portions" (POBSP, 1964).

Apr. 1965: Collected by C,H. Lamoureux from southwest
islet and west part of island. (UH, 2).

434

Appendix Table 6. (Continued)

1966: Green and blooming in July, indicating that
drought was not a great factor in its status
-..econtinued to grow and bloom vigorously
throughout the summer and fall. During and
after August...bore seeds abundantly.

1967: Distribution was not significantly different
from October 1966 to May 1967, but by late
August 1967 it had spread tremendously,
especially on the east half of the original
portion of the island, including the northeast
peninsula, and on the southwest islet. Large
areas of previously bare sand were covered on
the east hill. Blooming and fruiting were
continuous through the spring and summer.
Downy chicks, especially Brown Boobies, fre-
quently had large numbers of seeds stuck to
them. After the supply of Trtbulus on the
island was exhausted by early-nesting Red-
footed Boobies, these birds used Boerhvatia
for nest material. Most Brown Booby nests
contained Boerhavta also.

1969: This species is the most changed from the 1967
level of all plants on the island. It now
carpets perhaps 80 to 90 percent of the east
half of the original portion of the island,
including nearly all areas that were barren in
1967. The diggings by the Coast Guardsmen
for the hospital on the east hill are profusely
covered, as is the ground over most of the rest
of the east half of the island. On the east
slope, Christmas Shearwaters are burrowing
into the thick mat for their nest sites. This
increase had started during the spring of 1967,
and apparently has continued unabated. Thus
this species seems to be an important pioneer,
sand-holding species, at least on this island.
It may be set back periodically by the tower
maintenance and/or drought, but evidently |
there was sufficient rain during the last two
years for it to thrive. During February most
areas were in bloom, and mature fruits were
also abundant, sticking to socks and to birds.
It occurs in scattered clumps throughout the
southwest area also, but does not appear to
harm the Lepturus there.

435

Appendix Table 6. (Continued)

Aizoaceae
Sesuviun portulacas trun

Jul. 1963:

Apr. 1965:

1966:

1967:

1969:

Portulacaceae
Portulaca oleracea

ele 1963's

Apr. 1965:

1966:

"Occurs on northeast peninsula, east shore,
southwest islet and along roadway to trans-
mitter building of the east portion...."
(POBSP, 1964).

Collected by C.H. Lamoureux from southwest
islet, west portion of original island.
(UH, 2).

Sesuvitun portulacastrum was least affected by
the dry weather and disturbance of the early
part of the summer. In July it was in bloom
all along the fringes of the island, especially
on the outer, undistrubed portion of the north-
east peninsula. It continued to bloom and
produce seed throughout the summer and early
autumn.

Sesuvtum portulacastrun underwent no noticeable
change during the period of observation. In
1967 it was in bloom continuously except

where it was badly trampled by Sooty Terns.
Seedlings described in October 1966 did not
survive, if indeed they were even of this
species,

Continues to survive on the edges of the
island, but has been crowded out of the
central portion of the peninsula by Boerhavia.
No really thrifty stands were found. A few
blooms were noted.

Purslane

"Occurs as scattered plants on original and
man-made portions" (POBSP, 1964).

Observed by Lamoureux?

Occurred scattered around the island. These
plants were inconspicuous until they began

to spread and flower in September and October.
By early October, seedlings were abundant

on the nearly bare slope above the bunker on

a APS

Yee
OA thi tt,

Z

wis] ao G

: Vey

\s

\aGe, we] CS

rye

INS
~

~ rer NS] - Bx

me ed]

t= wWs=|) S% WH. ty a |e

Wo); _'

Sy |Cpe

436

Appendix Table 6. (Continued)

1967:

1969:

Caryophyllaceae
Spergularta martna

Apr. »L965%

Leguminosae
Mucuna sp.

Julio 1923

Apr. 1965:
Vigna martna

1967:

the east shore. These probably grew from old
seeds, or possibly from roots of destroyed
plants, for few mature plants grew in this area.

Became fairly conspicuous on the east slope of
the east portion of the island by early 1967,
both on the slope above the east bunker where
it was described in October 1966, and also on
the northeast slope, where none was noticed at
that time. By May 1967 all these plants were
badly beaten down by Sooty Terns, and by
September the areas were largely covered with
Boerhavia. However, the few surviving
Portulaca plants were larger than in May, and
they were blooming more vigorously.

This species apparently lost out to Boerhavia
on the northeast slope and above the east
bunker, where the large patches that appeared
in late 1966 were beaten down by terns and re-
placed by Boerhavia by May 1967. These areas
are now densely covered by Boerhavita, and only
a few scattered small Portulaca plants could

be found. More thrifty patches were found near
the shore northwest of the transmitter, and

on the south side of the transmitter building.

Collected by C.H. Lamoureux beside road on
west part of original portion. (UH, 1).

"A single kukui nut and a Mucuna seed were
picked up on the south beach of Sand Island"
(Christophersen, 1931).

Observed by Lamoureux?
Beach Pea
Plants were found in two places in early

1967: one near the old dock and one at the
edge of the beach on the southeast corner of

437

Appendix Table 6. (Continued)

1969:

Zygophyllaceae
Trtbulus etstotdes

aad x 4923's

Jul. 1963):

Apr. 1965:

1966:

1967:

the original part of the island. Both were
small (one to two feet in diameter) and were
badly trampled by Sooty Terns and Brown
Noddies by May. The southwest plant (near
the dock) could not be found in late summer.
No blooms were found. (UH, 1).

The plant near the southeast inner guywire
appeared to be about the same in size as in
1967, but one near the old dock could not

be found. Leaves looked fresh on the plant
found. On 9 March a very small plant was found
on the east shore near the base of the penin-
sula, and on 24 March a very small plant was
found on the south-southwest shore.

Puncture Vine

"On Sand Island Boerhavia diffusa is more
abundant than Trtbulus cetstotdes, but neither
begins to compare with the bunch grass in
extent" (Christophersen, 1931).

"Comprises 30 percent of the vegetated area
of the original portion" (POBSP, 1964).

Collected by C.H. Lamoureux, southwest islet
and west portion of original part of island.
(UH, 1).

Green and blooming in July, indicating that
drought was not a great factor in its status.
Continued to grow and bloom vigorously through-
out the summer and fall. During and after
August bore seeds abundantly.

One of the few plants showing any vigor during
the dry summer of 1966, was all but wiped out
over most of the island, especially the east
hill, during January and February 1967 by nest-
building Red-footed Boobies and Great Frigate-
birds. Red-foots were most destructive because
of their more vigorous efforts at pulling up
the vines. Frigatebirds usually picked up only
loose vines, often ones torn loose by boobies.
By mid-March new booby nests were being con-
structed almost entirely of Boerhavia, and

S Vue

W)

sjpe Ue Y,
Yity

a

c

Nes,

2
2

(=
=v)

y

\ <a F

c

\

ox

Re A
SSA

NN

438

Appendix Table 6. (Continued)

TIG9:

Euphorbiaceae

Aleurittes molueccana

Jule G23)
Apres, 1965:
Combretaceae

Termtnalta catappa
Apr: 1965:
Convolvulaceae
Ipomoea tndica

1967:

frigatebirds were no longer building nests.

A little Trtbulus was found in Brown Booby
nests, but most of it was gone before the
Brown Boobies started building nests. By May
Trtbulus had made a strong comeback over most
parts of the island, and was blooming, with

a few green fully formed fruits on plants
that boobies had reduced to scattered frayed
stems in February. By September Tribulus was
back at least to the level it attained in
October 1966, and was blooming and fruiting
profusely over the entire island.

This species appeared unusually healthy for
this time of year, probably because Red-footed
Boobies -had not begun nesting and tearing it
apart (this began about 4-5 March). Clumps
were numerous and healthy. Many new blooms
appeared during February. On the northeast
slope, where several large clumps formerly
occurred, there are large barren areas, which
look as if the plants were over-exploited

last year by boobies and frigates. Many

dead or nearly dead roots are visible, but no
growth. Brown Boobies are using this for nests
(7 March) and Red-footed Boobies are
increasing their use.

Candlenut, Kukui
"A single kukui nut...and a Mucuna were
picked up on the south beach" (Christophersen,
i 93d)%

Observed by Lamoureux?

Indian Almond, Kamani

Observed by Lamoureux?

Found in two places on the original portion
of the island in February 1967: one just back

439

Appendix Table 6. (Continued)

1969:

Ipomoea pes-caprae

Jul, Asie

Apr. 1965:

1966:

1967:

1969:

Boraginaceae

of the crest of the east hill, east southeast
of the transmitter, and one a few feet inland
from the west beach about halfway from the
road to the old dock. Both were small plants,
with no blooms. Neither could be found in
September. (UH,1).

Could find neither patch found in February
1967. That on the east hill is overgrown with
Boerhavta, and Lepturus is thick on the south-
west.

"Found only on easter beach of original portion"
(POBSP, 1964).

Collected by C.H. Lamoureux in coral sand at
top of beach on east end of original part of
tsland. (UBS: Lc

Grew in a few spots on the north and east

sides of the island, was alive but decadent

in July. During September and October these
plants grew and spread, but no blooms were seen
by 25 October.

Sent out long runners during the winter, but
few blooms were found on the east portion of
the island during the spring. In September

the plants still looked vigorous, although they
had not spread much during the spring and
summer, and still were not blooming profusely.

Two large patches....which very nearly exclude
all other plants, occur, one on the northwest
corner of the island, and one northeast of the
transmitter. Other plants around the island
appear about the same as formerly.

Heltotroptum curassavicum

APE. LG 5/25)

1967:

Collected by C.H. Lamoureux beside road,
west portion, original part of island. (UH, 1).

"..showed no change from 1966 through 1967.

It was scarce, scattered around the outer parts
of both portions of the island. All plants seen
were blooming both in spring and late summer."

440

Appendix Table 6. (Continued)

Tournefortta argentea

Juli LOG3i:

Apr. 1965:

1966:

1967:

1969:
Goodeniaceae

Seaevola taccada

1966:

1967:

Tree Heliotrope

"Only one small plant on original portion north
of transmitter building" (POBSP, 1964).

Collected by C.H. Lamoureux, north of transmitter
budding. 9 GUE, 9),

The single Messerschmidia argentea growing north
of the transmitter building was almost unrecogni-
zable in Juiy, having been run over by trucks
during tower maintenance, By September it had
produced abundant new leaves and had begun to
bloom.

The single bush growing northeast of the trans-
mitter was 2 1/2, to 3.feet high and 4 to 5Spfect
in diameter by March 196/7--large enough for a
Red-tailed Tropicbird to attempt to nest under
it. By September it had grown slightly larger,
and provided nightly roosts for 20 to 25 noddies
of both species. Wedge-tailed Shearwaters
burrowed at the base of the bush, which may reduce
its vigor. This specimen, as well as all the
larger ones planted around the buildings on the
man-made end of the island bloomed and produced
fruit profusely during the spring, summer, and
False

Present.

Twelve Scaevola sp. plants up to about 6 inches
high, brought from Hawaii by the U.S. Coast
Guard, were planted near the south shore of the
original portion of the island 21 October,
following a heavy rain.

Seaevola taeccada planted in October 1966 on the
south side of the east end of the island were
all alive and growing in February, but by May
had suffered from trampling by Sooty Terns and
Great Frigatebirds. Sooty Tern chicks used them
for shelter from the sun, By September only 8
of the original 12 were still alive, but these
had spread horizontally to a diameter of about

Appendix Table 6.

1969:

Compositae

Pluchea carolinensis

Apr.

1966:

1967:

1969:

1965:

Sonehus sp.

Apr.

1967:

1965:

441

(Continued)

3 feet. Vertical growth was prevented by
roosting terns and frigatebirds. A few flowers
were produced by September.

Only 4 plants, one of which may be dead, remain

of the 12 planted on the south side of the original
portion of the island in October 1966. These are
small, scarcely protruding above the surrounding
grasses, but have spread laterally to perhaps three
feet, and the leaves are green and fresh.

Collected by C.H. Lamoureux, west part of
original portion of island. (UH, 1).

Pluchea growing on the northwest corner of the
original portion of the island (one bush) and
scattered over the man-made portion, showed little
effect of either the dry weather early in the
summer or of the increased moisture in the fall.
These grew comparatively little, and produced no
new flowers.

The single large bush on the original end of the
island and seedlings growing downwind from it are
growing much more slowly than those on the man-
made end, probably because of intensive competition,
especially from Lepturus.

The bushes seeded from the original beside the

road just past the turnaround on the way to the
transmitter building have now grown to 4-6 feet in
height, and as much ‘in diameter, and now attract nest
tropicbirds. These were no more than two feet high
aay AUCXS7/e

Sow-thistle

Collected by C.H. Lamoureux, west part of original
POLE1on of asiland., (UH, 1).

Grew abundantly on the northwest corner of the
original part, where continuous disturbance of
loose sand prevented establishment of competitors,
and occurred scattered around the buildings and
open areas on the man-made end. In early spring

442

Appendix Table 6. (Continued)

it was in all stages of bloom and fruit, but
by May most plants had mature fruits. By
September many of the plants had partially
dried up.

1969: Now relatively scarce on the northwest side
of the island. The area has been taken over
by grasses, both Lepturus and Eleustne.

443

Appendix Table 7. Bird specimens* collected on Johnston Atoll

USNM No. Date Collected Age Sex Remarks

Black-footed Albatross

493512** 12 Feb. 1964 A M 14°46' N, 169944'wW, 120 mi. SSW
of Johnston Atoll. Large brood
patch

494144%% 8 Mar. 1964 A M 16°38'N, 169°38'W, 10 mi. SW
of Johnston Atoll, in sight of
islands

495791%** 15) dan, L965 GES 50UNS 1699°50"W, cas 20 mi.

WNW of Johnston Atoll.
Phoenix Petrel
494139 18 Sept. 1964 A F Kepler and Lehner

Bulwer's Petrel

530877 16> July 1923 A M Wetmore

78 he Pose ye A F Wetmore

719 es, ny A M Wetmore

80 sa len oi = A M Wetmore

81 eee Ae " A F Wetmore

82 ane iu A M Wetmore
300808 1 adky 61923 N F 2/3 grown young; Wetmore

09 a boa N M 1/4 grown young; Wetmore

10 M is x N M Less than 1 week old; Wetmore
493794 1 Aug. 1964 A M

*All specimens are skins collected by POBSP personnel on Sand Island,
unless othemwise noted. All specimens are deposited in the National
Museum of Natural History (USNM).

**Denotes selected specimens collected at sea near Johnston Atoll.

444

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks

Wedge-tailed Shearwater

300736 16 July 1923 A M Gray neck and breast; Wetmore
37 er ages H A M Gray neck and breast; Wetmore
38 LS July 79235 A F Light phase; Wetmore
59 L3*July 1923 A M Dark phase; Wetmore
40 Bron ga i A F Dark phase; Wetmore
Al 18 July 1923 A F Light phase; Wetmore
42 15 July 1923 A F Light phase; Wetmore
484252 20 Aug. 1963 A M Dark phase
492973 21 Aug. 1963 A F
493632 9 July 1964 A F
33 ey : A F
34 mn i A F
35 ae u A F
495059 23 Aug. 1964
60 14 Sept. 1964 A M
61 20 Oct. (1964
62 220 CE L964 A F
63 11 Sept. 1964 A F
64 16 Nov. 1964
65 " i MG A M
66 ae te id
67 eae hed Ne

495831 20 Apr. 1964 A M

445

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
495839 9 Apr. 1965 A F
496233 14 Nov. 1965 A M
496322 20 Dec. 1965 A M
497473 Z5eOCEea 965 A F
74 30 Sept. 1965 A F
75 LOGE. LI65 A F
498323 30 July 1966 A M Skeleton
24 23 Aug. 1966 A F Skeleton
503356 17 Nov. 1965 Alcoholic
508107-
508130 Fall 1963 Alcoholic
508136-
508171 Fall 1963 Alcoholic
544146 1 Apr. 1966 A M
47 4 May 1966 A M
48 9 July 1966 A F
49 12 Sept. 1966 A F
50 2 Dec. 1966 A M
544393 25 Sept. 1968 A M
544596 30 Apr. 1967 A M
544600 13 June 1967 A F

Christmas Shearwater
300701 LG uly LO23 A Wetmore
02 18 July 1923 A F Wetmore

03 LO aly 30923 A Wetmore

446

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
300704 18 July 1923 A M Wetmore
05 16 July 1923 A Wetmore
06 ak ae " A Wetmore
07 14 July 1923 N M Large young, losing down on back
and belly; Wetmore
08 yo et is N F Large young, losing down on back
and belly; Wetmore
09 15 July 1923 A F Wetmore
10 Se ce " A M Wetmore
496323 6 June 1965 N Chick

Newell's Shearwater

492973 21 Aug. 1963 A F Amerson

497980 ee mt A F Trunk skeleton of 492973

544600 13 June 1967 A F Ovary 10 x 8; L. ovum 2.0;
Harrington

Sooty Storm Petrel

544650 21 Dec. 1968 A F

Red-tailed Tropicbird

300992 18 July 1923 M
93 11 July 1923 A F
94 LS Sully 1923 A M
95 16 July 1923 N M Full grown, down-free young
96 iS - Sully 1923 A M
493809 28 Apr. 1964 F
10 21 July 1964 M
495079 23,0ct.. 1963 I M

447

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
495989 1 July 1965 M
496815 26 Feb. 1965 M
16 10 Sept. 1966 A M On egg UT
497213 25 Mar. 1966 Dy
497827 17 Salle, 1966 F Ms
54 22 Dec. 1965 F ry
W " " Ls
56 14 Jan. 1966 M #
i
57 16 Jan. 1966 F =>
=
58 7 Feb. 1966 F
bd
59 16 Feb. 1966 F oe
is
60 8 Apr. 1966 M a
61 13 May 1966 F
ns
62 22 Feb. 1967 F ff
63 31 Mar. 1967 F
64 16 Apr. 1967 F ¥
\
65 18 Mar. 1967 M North I. YW
503030 Mar. 1964 Chick,alcoholic Vv,
31 Apr. 1964 Chick, alcoholic b
544387 5 May 1965 M rL
544398 6 Nov. 1968 M ¥
Blue-faced Booby A
300933 12 July 1923 S oF WOGneds "
34 moo n A F Wetmore $,
es
é
rl
‘s

448

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
300935 Tay foi yy as) 28) A F Wetmore
36 2 Sully, 923 F Wetmore
By sar gts ¢ S F Wetmore
38 13 July 1923 S F Wetmore
39 1 uly 2925 A F Wetmore
495787 29 Mar. 1965 S Banded on Lisianski 21 Aug.
1964
496012 13 -Apx. 965 : F
498122 20 Jan. 1965 Skeleton
Brown Booby
300860 Ze sudiy. 1973 A F
61 Ae a A M
62 i a ye A F
63 bese gat " A 2
64 Loy Julys 923 iE F First year plumage
495078 27 Sept. 1964 M
496626 21 Dec. 1965
75] 15 May 1966 F
28 24 May 1966 N M Chick
29 29 May 1966 N M Chick
497371 20 May 1965 N M Chick
72 22 Mar. 1967 F
D093 73 23 Apr: 1965 N Chick; alcoholic
74 30 Apr. 1965 N Chick; alcoholic

449

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks

Red-footed Booby

300898 Pe eraly 923 A F
99 Ww Ww Ww S F
900 ia; Ww w S ?
01 13 July 1923 Aw OF
02 Ww tw wt S M
03 11 July 1923 SF
04 eh if F Lead colored, young of year
05 W w W S F
493473 13 Apr. 1964 F
495797 19 Feb. 1965 F
98 6 Mar. 1965 M
496249 19 Feb. 1965

50 4 June 1965 F
89 12 Jan. 1966 M
90 25 Jan. 1966 M

91 12 Feb. 1966 F \

Vy

92 17 Feb. 1966 M _

93 20 Feb. 1966 M Ks

Lb

94 22 Feb. 1966 M a

TU

95 25 Feb. 1966 M ‘On

le,

496648 4 Jan. 1966 F :

49 @ fen. 1066 F a

Vu

50 23 Jan. 1966 F b

450

Appendix Table 7.

USNM No.

497367

68

69

498127

544691

544692

ib

27

iS)

6

if

iE

Jan.
Mar.
Apr.
Mar.
Apr.

Apr.

Great Frigatebird

300972
464442
43
44
465207

465213

493908
495068
69
70
71
72
73
74
495809
10

Hab

13 July
11 July

13 July

(Continued)

Date Collected

1966

1967

1967

1965

1969

1969

1923

1923

1923

Li dudiy? £923

spr.-sum. 1923

ital

18

Oct.

Sept. 1964

Oct.
Oct.
Oct.
Oct.
Aug.
Nov.
Feb.
Mar.

Mar.

1964

1964

1964

1964

1964

1964

1964

1965

1965

1965

Sex

M

Remarks

Skeleton
USFW 75/7-37983

USFW 767-53223

Wetmore
Wetmore
Wetmore
Wetmore
Wetmore
Collected on French Frigate

Shoals or Johnston, data
lost; Wetmore.

451

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
496302 19 Aug. 1965 F
03 20 Jan. 1966 M
04 thi de % M
05 nai dae " M
06 bie i, M
07 a oe : F
08 > - 2 M
09 Mi " M
10 ye yy F
498278 6 Aug. 1965 F Skeleton
79 6 Aug. 1965 F Skeleton
498413 24 Mar. 1969 F? Skeleton, USFW 737-44221

banded Sand I., Johnston Atoll
10 July 1963 as N - U.

503043 Apr. 1964 Chick; alcoholic
44 . ms Chick; alcoholic
503322 20 Jan. 1966 F Alcoholic
23 a " x F Alcoholic
24 " " " F Alcoholic
25 " u u M Alcoholic
26 " MS He M Alcoholic
27 Ny m M Alcoholic
28 Ls ut u M Alcoholic
29 " uw "W M Vu
33 He Mi u Alcoholic A
CS,
ru
PS

452

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
503334 24 May 1966 M Alcoholic

35 207 0ct., 1966 M Alcoholic

36 4 Nov. 1964

Si/ 16 Nov. 1965 M Alcoholic

38 19 Nov. 1965 Alcoholic

77 15 July 1965 Alcoholic; chick
508094 21 Nov. 1963 Alcoholic

95 oe mad Alcoholic
543359 20 Dec. 1965 M

60 20 Feb. 1966 M
543361 20 Feb. 1966 F

62 22 Feb. 1966 F

63 5 Mar. 1966 F

64 26 Mar. 1966 M

65 eae M F

66 1 Apr. 1966 F

67 28 May 1966 M

68 18 July 1966 M

69 18 Sept. 1966 M

70 24O0ct. W966 F

ifak 17 Nov. 1966

UP 20 Dec. 1966 F

nS 25 Dec. 1966 M

74 4 Jan. 1967 M

453
Appendix Table 7. (Continued)
USNM No. Date Collected Age Sex Remarks
543375 ji5eJan. «L967, Bisexual.
544396 90ce,. 11965 M
544986 1 Apr. 1968 F
Pintail
495973 HENOCEs L965 J F
496775 29 Oct. 1965 J M Johnston Island, trunk skeleton
saved
497795 1S Reb. 1967, A F
543175 7 Nov. 1966 Jy M Skeleton saved
American Wigeon
495789 Zan Oct. L964 J M
543174 29 Oct. 1966 J) M
Shoveler
495075 26 Oct. 1964 A F
| Peregrine Falcon
493823 ** 8 Nov. 1963 A M 200 mi. SW of Johnston, 14°10'N,
TOA
544948 2 Dec. LIGS | F
Golden Plover
301051 1 ulyeUI2S F Wetmore
495054 I20Ctaw LIGA F
oP) 24 Oct. 1964 M
56 10 Dec. 1964 M od
Sy Nigee te F bi
58 13 Dec. 1964 M A

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
495724 Si Jan. L965 F
25 14 Apr. 1965 M
495892 8 Apr. 1965 F
496085 28 Oct. 1965 F
86 SI Oeb 1965 F
496211 11 Apr. 1964 M
496685 19 Nov. 1965 F
86 22 Apr. 1966 F
87 2 May 1966 F
498219 29 Mar. 1965 Skeleton
20 17 Nov. 1965 M Skeleton
544565 22 Apr... 1969 A M Full breeding plumage
544649 13 Apr. 1966 F

Black-bellied Plover

493340 11 Dec. 1963

Semipalmated Plover

493853 150ct.. 1963 I 2

503671 30 Aug. 1968 if 2 Alcoholic

Bristle-thighed Curlew

496054 2 Sept. 1965 M

2) 17 Sept. 1965 M
498066 28 Sept. 1963 Skeleton
498412 24 Jan. 1969 Skeleton
544942 LSVOct 1967 F

3
455 as
UT
Appendix Table 7. (Continued) YS
LY

USNM No. Date Collected Age Sex Remarks é
Lesser Yellowlegs Tuy
—a ”
493225 18 Aug. 1963 M \
b>
si
Spotted Sandpiper ‘UT
493910 1 Sept. 1964 2
Wandering Tattler .
KS
301026 Lak gi jioulyis ALCS) F At least partial breeding wu

plumage
4
27, 13, July 1923 M Not in breeding condition YY
493223 18 Oct. 1963 M Non-breeding plumage 4
ry
544563 26 Sept. 1968 A M Partial breeding plumage aay
Ruddy Turnstone us
%4 July 1923 9 Wetmore Wy
495909 30 Apr. 1965 M IV)
10 2 May 1965 M us
i: 1 May 1965 F ru
496212 6 May 1964 F \y
SN
497276 10 Mar. 1966 F \\
544397 29 Oct. 1968 F thes
544539 3 May 1965

544557 29 Sept. 1966 F ES
TU
58 16 May 1967 M Johnston Island =
ip
Sanderling :
eo 2 us
496210 25)\ Now... 19165 F Johnston Island 2S
We

/
W,

456

Appendix Table 7. (Continued)

USNM No. Date Collected

Pectoral Sandpiper

493228 3 Oct.) 1963
495076 19 Oct. 1964
77 26 Sept. 1964
496068 14 Sept. 1965
69 27 Sept. 1965
544156 Ree hy Hy
545013 30 Sept. 1968
544395 20ce. 1968

Sharp-tailed Sandpiper

496071 29 Oct. 1965

544394 30° Sept 1969

Buff—breasted Sandpiper

544935 14 Sept. 1968
Ruff
496600 10 Mar. 1966

Wilson's Phalarope
493870 16 Aug. 1964

Glaucous-winged Gull
544989 8 Mar. 1968
? 7 Feb. 1969
Laughing Gull

493503 7 Apr. 1964

Remarks

Ova minute (0.6 x 3 mm)

Skull not ossified

F.N.. 26396

Testis 5 x 23; Amerson and
Amerman

Appendix Table 7.

USNM No.

494981**

Franklin's Gull

544936

Date Collected

(Continued)

13) dieing GOS)

19 May 1969

Large unidentified gull

Z

ALT)

Dec.

Gray—backed Tern

300637
38
39
40
41
495051
52
53
495534
497115
543039
40
41
544388

89

1, daly.

e/ ealy,

14

US)

20

Feb.

Apr.

1965

Sex

F

457

Remarks
Collected at sea 11 mi. NNW of
Johnston, within sight of the

islands; ovary 9 mm granular;
Clapp

Skeleton; F.N. 25923

Wetmore
Wetmore
Wetmore
Wetmore

Wetmore

Chick
Chick

Chick

S
S
oY

S eas

yy

Yu

ie
Y

= iy
3) a YY

ey.

a
=
4

458

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
544566 23 Feb. 1967 M
67 14 May 1967 M
68 28 Feb. 1969 M
69 7 Apr. 1969 F
Sooty Tern
300553 15. July 1923 A M Wetmore
54 ees a w AF Wetmore
5)5) ele i A M Wetmore
56 ak o A F Wetmore
57 sie ie . A U Wetmore
58 ha is A F Wetmore
59 ts ome nt Wetmore; fledged
60 . u WY Wetmore; fledged
61 ut Wy zu Wetmore; fledged
62 ue we Wetmore; fledged
63 43 i m Wetmore; fledged
64 nf " m Wetmore; fledged
493779 1 June 1964
80 healt
81 a ui vs
82 .
83 aR i :

84

459 py

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks

493865 15 Aug. 1964 F
66 23 Aug. 1964 F
67 6 Sept. 1964 M

493871 17 Jul. 1964 F
Te. 23) Jud.) L964 U
13 " eH 4 F
74 ie u M
15 a ah is F
76 10 Aug. 1964 F
ATE 11 Aug. 1964 F
78 15 Aug. 1964 M
79 eda e i M
80 ee es M
81 19 Aug. 1964 M
82 23 Aug. 1964 M
83 24 Aug. 1964 F
84 en ae te M
85 26 Aug. 1964 M
86 2 Sept. 1964 M
87 8 Sept. 1964 F
88 gh : | F
89 oe " M
90 dine ‘i M
91 sie i F

460

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
493892 8 Sept. 1964 F
93 es : M
94 11 Sept. 1964 M
493902 Z23y Tul e964 M
03 vane e M
494983 te eae tt F
84 Santee is F
85 17 July 1964 ae
86 23 July 1964 M
87 se aE F
88 ent ‘\ F
89 25 Aug. 1964 M
90 17 Aug. 1964 M
om 15 Aug. 1964 F
92 10 Aug. 1964 F
93 1 Sept. 1964 M
94 6 Sept. 1964 M
95 7 Sully) W964 M
96 9 Aug. 1964 F
o7 te F
98 17 July 1964 M
Ze) ger ea Mt M
495000 23 July 1964 M

O1 Ww w WwW F

461

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
495002 23 July 1964 F
03 17 July 1964 F
04 23 July 1964 M
05 iy es 4% F
06 17 July 1964 F
07 9 July 1964 F
08 23 July 1964 M
09 Marae dl ‘ M
10 17 July 1964 M
ACE 23 July 1964 F
12 ide iat ri F
13 re i ctl + F
14 1 on ¥ F
15 ae a “ F
16 eo ¥ F
L7 Ne ae F
18 ea M F
19 Or a att rf F
20 No he a + F
2k: 17 July 1964 F
22 9 July 1964 F
23 15 Dec. 1964 M
24 16 Dec. 1964 F
25 20 Dec. 1964 F
26 21 Dec. 1964 F

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks

495027 23 Dec. 1964 F
28 28 Dec. 1964 F
29 ie ‘a A M
30 18 Jan 1964 M
SH re se M
32 NU pe u M
33 eh " F
34 Cae ae B F
35 11 Aug. 1964 M
36 2 Sept. 1964 M
Sy// 2 Sept. 1964 M

495507 12 Feb. 1965 F
08 12) Feb.) 1965
09 et ots iM F
10 ert uy F
Wik ae ll se F
12 a aes M
13 i Mar. £965 F
14 9 Mar. 1965 M
15) 22; Mar.7-1965 F

495737 8 May 1964 M
38 1 June 1964 M
39 1 June 1964
40 1 June 1964

USNM No. Date Collected Age Sex Remarks

Appendix Table 7. (Continued)
495741 3 June 1974 M
42 18 June 1964
43 ey aces Mf
44 23 June 1964
45 2 July 1964
46 8 July 1964 M
47 9 July 1964 F
48 17 July 1964 M
49 27 July 1964
50 9 Aug. 1964 M
a 9 Aug. 1964 M
52 9 Aug. 1964 M
53 9 July 1964 F
495872 2 June 1964 F
13 8 June 1964 F
82 25 Aug. 1964 M
496368 26 Dec. 1965 M
497085 23 Dec. 1965 F
86 24 Dec. 1965 F
87 13 Mar. 1966 M
88 14 Mar. 1966 M
89 alan ts ty F
90 30 Mar. 1966 M
oi ea
92 10 Apr. 1966 M

463

464

Appendix Table 7.

USNM No.

497093
94
95
96
97
98
99
497100
01
02
03
4.98086
87
88
89
503045
46
47
48
49
50
51

503647

(Continued)

Date Collected Age

ital

15

No

29

19

Apr. 1966
" "
Apr. 1966
May 1966
" "
May 1966
" "
" "
June 1966
" "
" "
July 1964
" "
" "
" "
Apr. 1964 N
1964 N
1964 N
Apr. 1964 N
1964 N
Apr. 1964 N
Apr. 1964 N
July 1969

Sex

Skeleton

Skeleton

Skeleton

Skeleton

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Remarks

Appendix Table 7.

USNM No.

503648
49
50
DL
52
33
54
55
56
Sy
58
59
60
61

543001
02
03
04
05
06
07
08
09

10

Date Collected

28

3

17

29

ily)

29

24

14

10

Sil

9

18

30

15

22

July 1969
Sept. 1969
July 1969
July 1969
July 1969
July 1969
July 1969
July 1969
Aug. 1969
July 1969
Aug. 1969
July 1969
July 1969
July 1969

Apr. 1965

(Continued)

Age

Sex

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Alcoholic

Remarks

465

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
543011 22, Apr.) 1965 M
IL ut a i F
13} 27 Dees, 1965
14 fie Marte V9 66 F
15 137 Apr." L966 F
16 22 Apr. 1966 F
IEz/ 24 June 1966 F
18 25 Mar.) 1967 M
544076 127 July, 1966 M
Vd ae a M
78 ae ae us M
79, eee i M
544244 pee J Soyer, | aN)7/ F
544390 10 Aug. 1968 M
ot 10 Sept. 1968 M
92 3 Sept. 1968 M
544454 26 June 1966 F
59 25 May 1967 F
56 9 Apr. 1968 M
57 1 Apr. 1968 M
58 17 July 1968 F
544607 26 Dec. 1966
08 I Jan. 1967 M

09 200 San. 967 F

Appendix Table 7. (Continued)
USNM No. Date Collected Age Sex Remarks
544610 20 Jan. 1967 F
1h 20 Feb. 1967
12 6 Mar. 1967
13 18 Mar. 1967
14 2Z7Man 1967,
15 26 Mar. 1967
16 5 Apr... L967
17 22) Apr. 1967 F
18 8 May 1967
19 TL Ape. 7967
544620 15 Apr. 1967 M
544626 10 Jan. 1967 F
pay 20 Feb. 1967 M
28 27 Aug. 1967 M
544638 2 Ape wel 96s M
39 2 May 1968 F
40 18 June 1968 M
41 23 Mar. 1968 M
42 1 Apr. 1968 M
43 2 Apr. 1968 F
44 26 Mar. 1968 M
45 2 Apr. 1968 M
46 25 Mar. 1968 F
47 20 Mar. 1968 F Five toes on right foot
48 26 June 1965 M

468

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks

Elegant Tern
544937 19 Apr. 1969 F

Blue-gray Noddy

300368 US July o23 A F Wetmore
88 a A 1 Wetmore
8Q*x* 20;-July, 1923 M Taken at sea about 50 miles
west of Johnston (16°45'N,
170° 20'W)
497694 8 May 1967 Ay SF
Brown Noddy
300652 LS Jaly 9923 A M Wetmore
53 rec’ ee . A M Wetmore
54 " m " A M Wetmore
55 a cus A M Wetmore
56 pee A F Wetmore
57 Tee et " A M Wetmore
493756 20 July 1964 F
57 20 June 1964 F
58 ae te 2
Sy) vs i M
60 e u G
61 Me ae ¥ 2
62 20 July 1964 F
63 LE eas i M
64 20 June 1964 ?

65 17 June 1964 ?

469

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
493771 20 June 1974 2
72 ae ?
73 Lay eee 2
74 ia, We MH ?
qe) 6 May 1964 F
493895 18 Aug. 1964 F
96 oe hy " v
97 Wet & q 2
493915 ee * 2
495039 11 Sept. 1964
40 20 Aug. 1964 M
41 18 Aug. 1964 M
42 4 Oct. 1964 M
43 >) OCE L9Gh M
44 4 Sept. 1964 F
45 20 Aug. 1964 F
46 25 Aug. 1964 M
47 20 Aug. 1964 M
48 4 Sept. 1964 M
49 7 Nov. 1964 M
50 20 Aug. 1964 F
495565 12 Feb. 1965
66 ees " M

67 W W WwW F

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
495568 12 Feb. 1965 F
69 bakin iy F
495754 4 June 1964 F
496045 11 June 1965 M
496217 6 May 1964 N U Chick
496394 30 May 1965 N M Chick
497244 3 Jan. 1966 M
45 14 Feb. 1966 ie ae:
46 6 May 1966 M
47 Me sie ue F
497632 9 Feb. 1966 F
33 l Apr. 1966 F
34 25 Feb. 1967 F
35 i Mari.” 1967 M
36 15 Apr. 1967 F
37 21 Apr. 1967 F
497673 18 July 1966 F
74 20 Apr. 1967 M
543151 3° Mar.) £966 M
503054 Apr. 1964 N U Chick; alcoholic
YS) 1964 N U Chick; alcoholic
56 1964 N U Chick; alcoholic
503368 10 Nov. 1965 Alcoholic
69 ZO UNOW: 965 Alcoholic
70 LY Dee. Jb965 Alcoholic

471

Appendix Table 7. (Continued)

USNM No. Date Collected Age Sex Remarks
503662 Sy oul yiel969 Alcoholic
63 16 July 1969 Alcoholic
508172 21 Nov. 1963 Alcoholic
73 eae ei Alcoholic
74 ele es " Alcoholic
75 Mee ta! et Alcoholic
76 he i Alcoholic
Vi eels i Alcoholic
78 Voy Yi UY Alcoholic
79 ere " Alcoholic
80 he ‘ Alcoholic
81 Yaes" 4 Alcoholic
544399 17 Nov. 1968
544588 27 Apr. 1967
89 2 May 1967
544605 10 May 1967
06 3 May 1967
544651 26 June 1968
52 2 May 1968
Black Noddy
495585 26 Jan. 1965
86 ee gt i
87 ea NL a
88 Ne Ne i
89 eae a

472

Appendix Table 7. (Continued)
USNM No. Date Collected Age Sex Remarks
497671 17 Feb. 1967 S) M
White Tern
300409 16: July 1923 A M Wetmore
10 Se July 923 A : Wetmore
uae 6 July 1925 A F Wetmore
eZ it Judy 1923 A M Wetmore
3 12 “July 1923 A F Wetmore
14 i July £923 A F Wetmore
27 ue y N F Wetmore; downy young
493655 24 July 1964 U M Johnston
495900 26 May 1965 A F Johnston
497266 10 Sept. 1966 A M Johnston; Lg. testes, 5 x 3 mm
498128 29 Mar. 1965 U U Johnston; skeleton
503664 1 July 1969 N u Johnston Island; alcoholic;
3-5 day-old chick
65 ue vs A U Johnston Island; alcoholic
? 6 June 1969 Johnston Island; F.N. 26409
Japanese White-eye
495080 14 Oct. 1964 2
496214 30 Nov. 1965 2 Johnston Island

473

Appendix Table 8. Pre-POBSP observations of Sooty Terns on
Johnston Atoll

Year Month Source Remarks

1859 14-15 Mar. John M. Brooke (ms., see also Brook, 1955, and
Findlay 1886: 1034),

"Capt. Piper says that these birds
keep close during the warm part of
the day and go Seaward in the
evening. In fact there are many
flying round us in the moonlight
this evening....The air was filled
with birds screaming very

Loud ycrvac

1859 28 Aug. Pendleton (1859) 7m Ward, 1967: 433).

"The islands are covered with a
wilderness of birds, so that if one
goes ashore he is shaded from the sun
by them. About two months since, the
ground was covered with eggs, so that
it was difficult to walk without
treading on them, but now the young
birds are hatched and are running all
over the island. When they get large
enough to fly, all the birds of one
kind will leave the island together
and go to sea where they will remain
several months, when they return
again and commence laying."

1923 10-20 July A. Wetmore (ms. a and b).

"Abundant. A few have eggs but the
great majority have young on the
wing or nearly ready to fly. Birds
able to fly are still being fed by
the parents and rest with them at
night, the colony at such times
being spaced off as when there are
eggs in it. The call of the young
is high-pitched. Those that I cap-
ture frequently regurgitate squid.

One group that apparently have
finished their breeding activities
gather in a close flock on a stretch
of open beach to sleep. The great

474

Appendix Table 8.

Year Month

1928 April
1945- June
1946

(Continued )

Source

L. Thurston

Lowell Thomas
CEI aALs

Remarks

majority are adults but with them
are a few young.

Partly grown young are pugnacious
and bite each other savagely.

The main impressions left in the
mind by these large colonies are
number of birds and noise. The
clattering calls are tittered in-
cessantly day and night. About
2500 on Johnston Island, about
2000 on Sand Island.

The birds are especially active and
noisy during rains whether by day
or night.

The young when pursued by frigate-
birds give up their fish without
protest.

With the Wedge-tailed Shearwaters
these made the greater part of the
bird population. I estimated those
on Johnston Island at 3500, and on
Sand at 2000. The majority of nests
held fresh eggs. I saw a few young
half grown and a few were able to
fly, but usually did /not?/ attempt
to take wing."

(192804)

"Din audible for half a mile," and
"thousands of sea birds" probably
both refer mainly to Sooty Terns.
No numbers given.

Chapin (1954: 10-11).

"My attention was drawn again to
the Pacific by a film made by
Lowell Thomas in June 1945 on
Johnston Island, near lat. 16°45'
North, in which a colony of Sooty

475

Appendix Table 8. (Continued)

Year Month Source Remarks

Terns was shown close to the air-
port. Correspondence with General
B.E. Nowland and Captain A.S. Hill
of the American Forces revealed
that the terns were back again and
nesting in the same month of 1946.
There was a hint that some had re-
turned to nest on the same island
in November or December. That
seemed curious, but no confirma-
tion could be obtained."

1947 ? G.E. Beckham (1947).

In brochure introducing Johnston
Atoll to incoming and transient
personnel, a map shows "Gooney

Bird Hill" in area on Johnston

where the swimming pool was recently
built. "Johnston Island has about
10,000 birds that live here during
the migratory months. There are
many kinds of birds, but are known
to all the men as gooney birds. If
you should step outside in front of
the terminal and look toward the

end of the island, you would probably
see a small hill covered with them.
This is known as "Gooney Bird Hill."
From the description given, most of
the birds on this hill must have
been Sooty Terns.

1948 1. C. Fennell (CEG YESIR 2 i7/) We

Letter from Seoul, Korea, after
flight which stopped at Johnston
on the way: His glasses were con-
fiscated on landing, and most
information was from asking resi-
dents. "The only important
information I could gather from
several fellows whom I asked about
the birds was that they were a
"damned stinking nuisance," and
that they were destroying all
their eggs in an effort to get

476

Appendix Table 8. (Continued)

Year Month Source
1949 Dec.? S. Jensen
1955 2? J. Benson

1957 6-10 April

M. Moynihan

Remarks

them to move across to Sand Island,
a smaller strip of land about 1/4
mile distant. They all heartily
regretted the fact that Johnston
was designated as a bird sanctuary.
One man named the species found
there as noddy and sooty terns and
moanin' birds."

(1949: 66).

"At this season there are a great
many Sooty Terns flying low over
the south end of the island. They
maintain an incessant and terrific
clamor throughout the night. The
birds fly very low at night, but
during the day they are usually to
be seen at a little distance off
the island at an altitude of
several hundred feet. They number
about 2000, I believe."

G55)

"To make matters worse, it is ad-
visable to wear a hat at all times,
because Johnson [sic] has been a
bird sanctuary since 1926, by
executive order of the late Pres.
Coolidge.

The birds occupy the entire western
end of the island, and when we were
there [date not given], the navy
gathered up the eggs every day and
threw them overboard in an effort
to keep from being crowded off the
island. Took them 2 1/2 hours to
collect all the eggs, so you can
see what they were up against."

(1957)
"This was the overwhelmingly

dominant species on the island, in
point of numbers. There must have

477

Appendix Table 8. (Continued)

Year Month Source Remarks

been at least 10,000 birds present,
almost equally divided between two
colonies, one on the bare ground
and low grass in the center of the
western islet, and the other in

the grass on a low dune in the
north-central part of the eastern
islet (some distance away from a
group of abandoned buildings and
oil tanks). These birds all seemed
to be in the later stages of the
"pairing phase' of the breeding
cycle, Ze. at. the stage, after
Pair formation, when sexual be-
havior is most common and most
vigorous, before the appearance

of the first eggs."

1958 July and Holmes and Drawings and photographs.
August Narver

Engineering drawings of sprinkler
system to protect birds from 1958
Atomic tests show probable approxi-
mate distribution of birds, pre-
sumably mostly Sooty Terns. The
sprinkler was installed on approxi-
mately the west quarter of the
man-made portion of the island
(one picture shows a few birds
scattered over this area, probably
including Noddies, as well as
sooties). The west third of the
causeway was covered with the
sprinkler, but there is no indica-
tion from pictures what birds may
have been there. Approximately
the north half of the original
portion, with extensions to the south
shore in one place, and all along the
east shore was covered with the
sprinkler system. The few pictures
showing the areas around the buildings
indicates that the area of the
sprinkler system roughly corresponded
to the area covered by Sooty Terns.
Two pictures taken 22 July show
several Sooty adults and young.

478

Appendix Table 8. (Continued)

Year Month Source

1959- th C.J. Davis

1960

1959- 2 U.S. Coast Guard
1960

Remarks

The young are all well grown,
but several have short tails.
There is not enough evidence
to indicate any difference
from present timing of the
breeding cycle.

(1962).

"The predominant "nester' on
Sand Island appeared to be the
Sooty Tern (Fig. 5). The
characteristic dull black upper
body and the white forehead and
underparts of wings and body
differentiates it from other
similar terns. The young bird
in the foreground of fig. 6
blends very well with the dirt-
like coral and sand. Unhatched
eggs can be seen. Their density
was about one per square foot.

(Fig. 7) These are the 'wide-
awakes'; observation reveals
that they are active nearly 24
hours each day. It is of inter-
est that their feathers are not
waterproof, so they work hard
for their food, snatching small
fish and squid off the surface
of the water and bringing it to
the waiting young. Their harsh
"quank" and three-syllable 'ker-
wacky-wack' warns the intruder
that they are concerned over the
single egg they have laid on the
ground. Since they range
throughout the tropical Pacific
and are noted for their 'dis-
appearance at sea', it was not
unusual to find them nesting
near remote Johnston Island."

Photographs

Photographs show Sooty Terns,
all adults, some possibly on

479
Appendix Table 8. (Continued)

Year Month Source Remarks

eggs, on various parts of the
original portion of the island.
As best determined from the
photos, the birds covered
almost all the main portion of
the original island, but were
not on the northeast peninsula.

1961 23 May U.S. Coast Guard Photograph

Photograph shows LORAN-C tower
from near east end of causeway.
Sooty Terns are over the entire
island, except possibly the
northeast peninsula, which
cannot be seen well enough to
tell if Sooty Terns are present
or not. Numbers appear about
the same as during the 1966-67
POBSP period.

*x U. S. GOVERNMENT PRINTING OFFICE : 1976 O - 208-933

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