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OS. 193-205

mw ATOLL RESEARCH
BULLETIN-

(194,

195.
196.
197.

198.

A photographic survey down the
seaward reef-front of Aldabra
Atoll

by Edward A. Drew

Topography and coral distribu-
tion of Bushy and Redbill Islands
and surrounding reef, Great Bar-
rier Reef, Queensland

by Carden C. Wallace and E. R. Lovell
Coral Cays of the Capricorn and
Bunker Groups, Great Barrier
Reef Province, Australia

by P. G. Flood

Submarine cementation of grain-
stone fabric, St. Croix, U.S. Vir-
gin Islands

by Lee C. Gerhard

Christmas Island (Pacific Ocean):
Reconnaissance geologic observa-
tions

by Mark J. Valencia

Notes on the vertebrate fauna of
Tongareva Atoll

by Roger B. Clapp

199.

200.

201.

202.

203.

204.

205.

Observations on vegetation of
blue-faced booby colonies on Cos-
moledo Atoll, Western Indian
Ocean

by Mary E. Gillham

Vegetation of sea and shore-bird
colonies on Aldabra Atoll

by Mary E. Gillham

Life history notes on some Aldab-
ran land birds

by C. B. Frith

Climate of Aldabra Atoll

by D. R. Stoddart and L. U. Mole
Annotated check list of corals in
the Mascarene Archipelago, Indi-
an Ocean

by Gérard Faure

Annotated check list of octocoral-
lia in the Mascarene Archipelago,
Indian Ocean

by Gérard Faure

The hosts of the Coral-Associated
Indo-West-Pacific Pontoniine
shrimps

by A. J. Bruce

Issued by

THE SMITHSONIAN INSTITUTION

Washington, D.C., U.S.A.

February 1977

193.

194.

195.

196.

TOT.

198.

ip My god

RESEARCH
BULLETIN

A photographic survey down the
seaward reef-front of Aldabra
Atoll

by Edward A. Drew

Topography and coral distribu-
tion of Bushy and Redbill Islands
and surrounding reef, Great Bar-
rier Reef, Queensland

by Carden C. Wallace and E. R. Lovell
Coral Cays of the Capricorn and
Bunker Groups, Great Barrier
Reef Province, Australia

by P. G. Flood

Submarine cementation of grain-
stone fabric, St. Croix, U.S. Vir-
gin Islands

by Lee C. Gerhard

Christmas Island (Pacific Ocean ):
Reconnaissance geologic observa-
tions

by Mark J. Valencia

Notes on the vertebrate fauna of
Tongareva Atoll

by Roger B. Clapp

Issued by

199.

200.

201.

202.

- 203.

204.

205.

Observations on vegetation of
blue-faced booby colonies on Cos-
moledo Atoll, Western Indian
Ocean

by Mary E. Gillham

Vegetation of sea and shore-bird
colonies on Aldabra Atoll

by Mary E. Gillham

Life history notes on some Aldab-
ran land birds

by C. B. Frith

Climate of Aldabra Atoll

by D. R. Stoddart and L. U. Mole
Annotated check list of corals in
the Mascarene Archipelago, Indi-
an Ocean

by Gérard Faure

Annotated check list of octocoral-
lia in the Mascarene Archipelago,
Indian Ocean

by Gérard Faure

The hosts of the Coral-Associated
Indo-West-Pacific Pontoniine
shrimps

by A. J. Bruce

THE SMITHSONIAN INSTITUTION

Washington, D.C., U.S.A.

February 1977

ACKNOWLEDGMENT

The Atoll Research Bulletin is issued by the Smithsonian Institution
as a part of its Tropical Biology Program. It is co-sponsored by the
Museum of Natural History and the Smithsonian Press, The Press 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 11/7 numbers issued by the
Pacific Science Board, National Academy of Sciences, with financial
support from the Office of Naval Research. Its pages were largely de-
voted 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

ATOLL RESEARCH BULLETIN

NO. 193.

A PHOTOGRAPHIC SURVEY DOWN THE SEAWARD
REEF-FRONT OF ALDABRA ATOLL

by Edward A. Drew

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

February 1977

ae
yg

Mibligen:

Contents

Page

Summary
Introduction
Methods
Establishment of the transect
Photography
Analysis of photographs
Results
Other transects
Discussion
Acknowledgements
References

Figures (following p. 7)

1 Aldabra Atoll, showing location of all transects and
implied submarine contours (modified from Bellamy
Biel: 1970)

Camera orientation on photographic transect

Transect topography and percentage living cover
Depth distribution of various growth types

Horizontal distribution of growth types away from
surf zone

Submarine light intensity and percentage living cover

WO WFwWPd

Plates

1 Phototransect frames illustrating growth types
discussed

a. Millepore

b. Branching coral
ec. Columnar coral
d,s. Braih coral

2 Phototransect frames illustrating growth types
discussed

a. Foliaceous coral

b. Alcyonacean soft coral
ec. Halimeda algae

d. Uncolonised bare areas

3 Representative strips from various important zones

a. Millepore coral zone
b. Branching coral zone
ec. Columnar coral zone
d. Brain coral zone

e. Foliaceous coral zone

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;

A PHOTOGRAPHIC SURVEY DOWN THE SEAWARD
REEF-FRONT OF ALDABRA ATOLL

by Edward A. Drew

SUMMARY

A simple photographic transect method, using aqualung
diving techniques, is described, together with an assessment
of its limitations and possible improvements.

Data obtained from analysis of a 250 metre long transect
(Oo - 40 metres depth) photographed on the seaward reef-front of
Alidabra Atoll are presented. Results are discussed in the con-
text of previous reef zonation data and the possible environ-
Genial factors controlling such zonation.

INTRODUCTION

fhe upper 20 metres of the seaward reef-front of any
barrier, fringing or atoll coral reef has until recently been
impossible to investigate thoroughly due to prevailing sea
conditions so close to the reef ridge.

However, in recent years the introduction of self-contained
diving equipment, and care in selection of the season for field
work to coincide with moderately calm conditions, have allowed
a considerable amount of direct observational data to be acquirec
in this environment.

Nevertheless, such investigations tend to use a variety of
ecological methods, mostly qualitative, and still "much of the
difficulty in comparing reefs stems from the lack of uniformity
in surveying methods" (Stoddart, 1969a).

In this paper a simple method of photographic recording
down a reef transect is described and the results from its use
on the most luxuriant section of the Aldabra reef-front are
presented. The field work can easily be carried out by divers
with no training in coral taxonomy and could be applied to any
Beer System, giving a standard basis for valid comparisons.

a7

(Manuscript received May, 1:9 70-— Bde)

Gatty Marine Laboratory, St. Andrews, Fife, Scotland.

METHODS
Establishment of the transect

This was marked by a string, knotted with a loop at one
metre intervals. Tt was laid down in 50 metre sections from
the reef ridge (mean low water springs - MLWS) to the beginning
of the basal sand plain at 40 metres depth. The strings) tovailed
253 metres in length and was orientated approximately perpendi-
cular to the shore-line although some meandering occurred in the
extensive Shallow parts. Each end was marked with a small buoy
and these were triangulated from the shore; bearings are shown
on the inset of the map in Figure 1.

Depth was recorded at 10 metre intervals using an 'SOS'!
totally enclosed bourdon tube gauge, and corrected to depth
below MLWS according to the state of the tide.

An attempt to assist diyer orientation on the very long
transect by placing numbered polystyrene floats on short strings
at 10 metre intervals proved unsuccessful. The film of algae
(diatoms? ) which rapidly developed on these floats was attractive
to the browsing fish population which rapidly consumed most of
the floats completely.

Photography

Standard 'Calypsophot!' and 'Nikonos' underwater cameras
with 35mm wide angle lenses were used to photograph the transect
every metre from a distance of approximately 2 metres from the
bottom. The open frame viewfinder of the cameras was used to
align the camera as shown in Figure 2. Considerable overlap was
achieved in all frames.

Monochrome 35mm film was used = Kodak Plus X (125 ASA) for
most of the transect photographs, replaced by Ilford HP3 (400
ASA) in the deeper sections. No artificial lighting was used.
The wide aperture setting required in the 30 — 40 metre section
resulted in considerable loss of resolution in many of those
frames.

Films were developed in the field, using a daylight-loading
developing tank and cooling solutions in the refrigerator to
about 20°C before use. In this way it was possible to ensure
complete coverage of the transect with usable negatives.

Although several photographic dives were carried out on the
transect, a large proportion of the usable exposures were made
on a single dive of two hours duration. Three cameras were used,
and each was reloaded twice in the boat, giving a total of over
300 frames.

3

The best negative for each metre section was printed to
i256 20 cm, (5" x 8") in the laboratory in the United Kingdom,
using 'Ilfoprint' materials followed by hypo fixation and high
Sloss finishing.

Analysis of photographs

Sequential prints were oriented and cut to produce complete
but unduplicated coverage down the transect; strips of five
prints were each analysed as follows.

Code numbers were allocated to a total of 26 different
coral growth forms and other recognisable organisms. Some of
these were later lumped together and final analysis involved
the following groups:

Cymodocea (marine angiosperm) Alcyonacean (soft coral)

Branching coral Gorgonian

Columnar coral Halimeda algae (+ traces Caulerpa)
Brain coral Sponges

Foliaceous/encrusting coral Tridacna clams

"Pioneer! coral Anemones

Fungia coral Uncolonised bare areas

Encrusting calcareous red algae could not be distinguished on
the photographs although they represented a significant part

of the living cover in some sections. The organisms and coral
types involved are illustrated in a representative selection of
Prames an Plates 1 and 2.

The outlines of the various colonies in each set of five
contiguous prints were traced onto high quality tracing paper
(uniform density) and the appropriate shapes were cut out and
weighed. In this way the area of each 5 metre transect section
covered by a particular growth type was determined. No attempt
was made to allow for differing colony morphology and therefore
actual surface area of living coral which in some instances
would be several times greater than the area photographed. In
view of the great importance of the symbiotic algae present in
the coral organisms, it seems probable that the area exposed to
direct solar illumination will be of major importance in their
growth.

RESULTS

The topography of the transect studied is shown in Figure 3,
together with an analysis of the percentage cover of living
organisms on both a horizontal and a vertical basis.

this particular transect, the longest of a total of 12
accurately surveyed around the atoll (see Figure 5 provided
two contrasting environments. Firstly there was an extensive
Shallow region where the major variable factor appeared to be

hy

proximity to the reef ridge surf zone, followed by a region of
moderately rapid increase in depth until the growth of herma-—
typic corals ceased at the start of the basal sand plain. This
last appeared to slope away indefinitely at an angle of about
20° from horizontal.

Analysis according to depth has been carried out by
averaging the various series of prints included in each 2 metre
increment of depth. Percentage living cover remains remarkably
constant until below 30 metres, when it drops rapidly to a much
lower value.

The regular alternation between high and low cover values
shown in the horizontal analysis is due mainly to a series of
completely bare areas, on a scale several times greater than the
size of a photographic frame. These were either bare coral rock
or depressions filled with coral debris (mainly branching types)
in shallow water but sandy in deeper parts.

The distribution of the various growth types with depth is
iliustrated ain Piasure,/, again, averaged for: the wepiesmot>
metre strips included in each 2 metre depth increment. Uncolo-
nised bare areas have been disregarded in this analysis which
shows the percentage of the living cover represented by each
type. There is a marked zonation, with the branched and less
massive types in shallow water and the massive brains much
deeper. Alcyonacean soft corals were intermediate in distri-
bution whilst only the foliaceous and encrusting types occurred
in the deepest parts. The significance of this zonation will
be discussed later; representative strips from the major zones
are shown in Plate 3.

Additional information about the response of the shallow
water types to surf action has been obtained by analysis of the
first 60 metres of the transect horizontally, this being restric-—
ted to the upper 2 metres of the depth range. Heavy surf is un-
likely to occur far seaward of low water mark. Figure 5 shows
such a horizontal analysis which indicates restriction of the
millipore corals to the maximum surf zone whereas branching
corals formed a small part of the total cover until some 25
metres from MLWS. .Columnar corals were relatively unimportant
and’ showed little correlation with distance from MLWS, but the
restriction of the Alcyonaceans to the most distant part
suggests that they are relatively sensitive to severe water
movement.

OTHER TRANSECTS

In order to put this transect into perspective compared
with other areas of the Aldabra reef-front, the 14 transects
surveyed around the atoll are shown in Figure 1. Only transect
topography and the approximate coral cover were determined on
all but transect 1, the photographic transect. The submarine

5

contours shown are greatly exaggerated compared with the scale
Ouvthe atoll outline. ~ Also shown is =the ~vertical-extent of
reef—front rocky substrate before the beginning of the basal
sand plain.

The north-west part of the reef-front was the only region
wren luxuriant coral growth, that along the north shore was
generally poor, and that on the south had apparently recently
suffered a major catastrophe with the dead remains of apparently
good growth evident in several places. The eastern end of the
atoll had no coral reef front, but rather a very gently sloping
sand plain from MLWS.

DISCUSSION

Initial structural considerations would suggest that
branching corals of the millepore and acropore types are less
well suited to the rigours of shallow water reef-front environ-
ments than are the more massive brain corals. Indeed, Stoddart
(1969a) cites numerous references to the occurrence of fragile-
branching types on lagoon reefs and in sheltered pools, whilst
stout-branching and more massive types are concentrated on reef
flats and upper seaward slopes.

The data presented in this paper suggest the opposite for
the luxuriant section of Aldabra reef-front, a situation also
noted by Stoddart (1966, 1969b) in the Maldives and the Solomons
where "reefs consist largely of foliaceous and fine branching
forms even in exposed situations". This feature of reef zonation
may be due to the rarity of severe storms in these three regions.
The north-west tip of Aldabra is nevetheless completely exposed*
to oceanic swell and receives a considerable pounding by waves
hor much of the year. Taylor (1968) also reports dominance of
millepore/acropore types in exposed situations on Seychelles
reefs, but with massive species dominant in sheltered areas.

thie wrestrretr1onrot mi’lepore corals “to the heavy surf
region of the reef-front may be associated with the same factor
which allows their predominance in the fast-—-flowing channels
leading into Aldabra's extensive but very shallow lagoon, perhaps
a requirement for constantly moving water or an ability to with-
stand the relatively warm water which flows both off the reef
flat and out of the lagoon. Taylor (1968) suggests that the
vertical inter-connected plates which comprise the surf-zone
millepore colonies are well adapted to absorb the energy of
breaking waves, but it is also necessary to explain their
virtual absence in stiller water.

Foliaceous forms of coral are well suited to intercept a
maximal amount of available light with a minimum volume of
colony, and are thus well suited to the deep dim waters where
they predominated on this transect. It is however surprising
that the total living cover of the reef remained reasonably

6

constant until the very rapid reduction below 30 metres. There
was even a noticeable increase between 15 and 25 metres. Thus,
living cover does not appear fo be correlated directly with
ambient light energy - the two are compared in Figure 6 - although
the interrelations between growth type, efficiency of light
utilisation, rate of primary organic production and rate of cal-
cification are as yet only known in the barest outline. Important
too may be the preferences of coral browsers such as parrot fish,
and their ability to feed in turbulent shallow water.

The simple photographic transect method described here
could provide a basis for rapid accurate assessment of various
types of reef and allow valid quantitative comparisons. The
method has certain limitations in accuracy, such as the edge
distortion produced by uncorrected wide angle lenses, although
most of this is eliminated by cutting off the extremities of each
print for overlap. The expense of using the 28mm corrected under-
water lens available for the Nikonos camera is probably unwarran-
ted, especially as several separate cameras are required for long
transects (see later).

Recognition of various growth forms proved moderately easy
with the monochrome prints used, although artificial illumination
would have allowed increased definition in deep sections whilst
the use of colour print film should in future allow positive
identification of most species, including the encrusting calca-
reous red algae, as well as a further increase in definition.
Unfortunately the prints from colour transparencies usually lack
the sharpness required for this work.

Phototransect methods could also be used in the rapid topo-
graphic survey method used on the majority of transects studied
in Aldabra. This involved two divers, separated by a 10 metre
length of string, leap-frogging each other in a straight line
down the reef and recording depth every 10 metres. A third
diver could photograph the ten metre strip in 1 metre sections
whilst the surveyors recorded their own data. This would require
Six cameras for a long transect -—- say 200 metres —- and may double
the survey time. However, the vast increase in data obtained,
and the saving in time required to establish semi-permanent
transects and relocate them on each dive, would certainly
FUS tahoe hse

Finally, a note on diver safety. Coral reefs are frequently
inhabited by potentially aggressive animals, mainly sharks.
Fortunately these were virtually absent from the Aldabra reef—
front at the time this survey was carried out (December, 1968 to
February, 1969), so that divers could concentrate on their scien-
tific tasks. It would, however, be essential for a survey team
working in more dangerous waters to be accompanied by 'look-outs"',
perhaps involved in more random photography or collection of reef
flora and fauna, but primarily there to allow the team to concen-
trate on their complex tasks knowing they will be warned of
impending danger.

ACKNOWLEDGEMENTS

This work formed a small part of the diving programme of
Phase VI of the Royal Society Expedition to Aldabra, and was
made possible by the organisation and logistic support provided
by the Aldabra Research Committee and the overall expedition
tenger Dra, D. RR. Stoddart;

I would also like to thank all the other members of that
phase of the Expedition, and especially Dr. J. Lythgoe,
Mr. D. Jones and Mr. J. Barnes who accompanied me on most of
the photographic and survey dives, for their help in the field.

The Court of St. Andrews University generously granted me
leave of absence for this expedition.

REFERENCES

Bel vany, D., Drew, E., Jones, D.. and Lythgoe, J. 1970. Aldabra —
a preliminary report of the work of Phase VI of the Royal
Society Expedition. Underwater Ass. Report 1968-69.

Stoddart, D.R. (Editor). 1966. Reef studies at Addu Atoll,
Maladive-Tslands., Atoll Res. Bulli7146: =122.

Stoddart, D.R. 1969a. Ecology and morphology of Recent coral
meets. Biol. Rev., 44: 495-4908:

Stoddart, D.R. 1969b. Geomorphology of the Marovo barrier
reef. Phil. Tran. Roy. See. B. 255: 383-402.

Taylor, J. 1968. Coral reef and associated invertebrate
communities around Mahé, Seychelles. Phil. Trans. Roy.
Soc. B. 254: 129-206.

Figures

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Introduction
Topography
The Islands
The Reef
Coral distribution
Table 1
Table 2
Table 3
Discussion
Acknowledgements
References
Appendix 1
Appendix 2

Contents

TOPOGRAPHY AND CORAL DISTRIBUTION OF
BUSHY AND REDBILL ISLANDS AND SURROUND-
ING REEF, GREAT BARRIER REEF, QUEENSLAND

by Carden C. Wassaceus and E.R. Layela=s

INTRODUCTION

The following description presents information obtained
by the authors over a ten day period, 26 December 1972 to 4
January 1973. The purpose of this paper is to provide an
introduction to an area which was previously unstudied, as an
aid to further investigation.

Locality: The area studied is located on the inner shelf
of the Great Barrier Reef Province, southern region, latitude
20° 57'S, longitude 150° 5'E. This is approximately 92 km
ENE of Mackay, on the Queensland coast (see inset, fies 1).

Orientation and size: The reef is oval, its long axis
running NNW to SSE. Maximum length is approximately 6 km, and
width 4 km. Surrounding water depth ranges from 26 m to 54 m.

Winds and rainfall: The main wind influence is east-—
southeasterly to southeasterly and continues throughout the
year (see inset, fae) Rainfall at nearby Pine Islet

averaged 3478 points per year in the period 1935-1973, the
highest falls being in January to March (information from the
Australian Bureau of Meteorology).

Tidal range: The reef lies within a zone of high tidal
range, this being approximately 5 m (taken by extrapolation
from Maxwell 1968, fig-42).

Otnher descriptions. of the area: The reef and/aslands) are
described briefly in the Australia Pilot Vol.IV (1962). Bushy
Island is mentioned and figured by Steers (1938), who visited

the area during rough weather and made a rapid survey. A
collection of 23 coral specimens, comprising 7 species, in the
ay

Queensland Museum, Gregory Terrace, Fortitude Valley, 4006,
Queensland, Australia.
eS Eye Department, University of Queensland, St. Lucia,
4067, Queensland, Australia.

(Manuscript received September 1973--Eds. )

2

Great Barrier Reef Committee collections at the Queensland
Museum is labelled "Redbill Reef, coll.: Geranium". This was
apparently obtained from the reef flat adjacent to Redbill
Island by Charles Hedley in 1924 (see Hedley, 1925, and also
Appendix 1, this paper).

TOPOGRAPHY
The Islands

Orientation, structure and size: The islands are situated
on the western edge of the reef, Bushy Island 1.2 km NNW of
Redbill. Redbill is a continental island, composed principally
of alkali granite bisected by dykes of microdiorite and trach-
yandesite. It is approximately 21 m high and 1.6 hectares in
area. Bushy Island is a coral cay, of area approximately 7.3
hectares, the beach crest being approximately 4 m above the
inner reef flat. It is composed of coral sand, and a sand pit
is developed at the NNE end of. the island. A ridge of beach
rock extends along the E-SSE edge (see fice ie not on the SW
side as recorded by Steers (1938).

Terrestrial vegetation: Species lists of plants from
both islands are given in Appendix 2. Nineteen species were
recorded from Bushy Island, with three predominating. The
extent of the dominants is shown in fig.2. The most prevalent
species, in terms of area occupied, is Pisonia grandis. The
Pisonia forest excludes other species except on its margin.
Pandanus sp. extends as a continuous margin along the western
perimeter of the Pisonia forest to the north east, and as
clumps around the remainder of the island. Bordering this and
extending to the beach is a band of Tournefortia argentea.
Other conspicuous beach species are Thuarea involuta, Ipomoea
pes-caprae, Salsola kali and Scaevola taccada.

The Reef
Methods: Observations were made along transect lines at
right angles to the reef edge (see alee oye Lines were worked

variously by walking out at low tide, by skin diving and by
diving with compressor unit from a small boat, depending on
accessibility and depth. Contour measurements were made with
a graduated pole. As time and manpower were limited, many
additional observations on the reef were made qualitatively
in areas outside the transect lines.

Transect lines were marked in metres. The method of
observation was to note coral species present within 15 cm on
either side of the transect line in each metre segment. This

was found to be a reasonably satisfactory way of covering a
large cross-section of the reef for a preliminary study,
although it is not proposed as an adequate or standard method
for more detailed survey. Corals were recorded by presence

along the 1m segment. In the early part of the survey,

numbers of colonies were counted, individual colonies being
defined as by Loya (1972); This method proved to be applicable
in inner reef flat areas, but difficult to apply to the arbo-
rescent and layered-plate corals in outer areas. In cases
where the species could not be separated in the field from
related species, corals were identified to genus only. This
was found to be necessary in genera such as Psammocora, Porites,
and Montipora. A representative collection of corals was made,
and these have been tentatively identified (see Appendix 1)

In qualitative work, as much as possible of the non-
transect area was covered by walking out and diving, listing
corals observed, and collecting specimens of each species.
The sectors of the reef least observed were the outer lagoon
and the eastern region (see fig.1).

The authors did not have sufficient opportunity (or
facilities) to study the off-reef floor. Although the floor
immediately adjacent to the reef front (S=A2 m depths ) was
examined on three transect sites and in qualitative work, that
beyond was not studied. It is hoped that this area can be the
focus of further work on a return visit.

Reef structure: The reef can be regarded as consisting
of a central region and three generalized surrounding regions.
These can be seen in the aerial photograph (plate 1) and in
Bee te,

1. The central region is generally lower than the
surrounding regions, and has an inner and an outer lagoon.
These reach low water depths of approximately 3 m, and have a
sandy base, with scattered alcyonarian and coral growth in
small patch reefs. In the non-lagoonal areas microatolls are
developed, and coral growth is generally denser.

2. To the west of the central region is the region of
the reef least exposed to prevailing winds. The largest zone
Pies, resion 1S an "inner flat" zone (in the terminology of
Stoddart 1969) which extends between the islands, gradually
merging with the central region in the east, and delimited by
a series of green algal banks in the west. The algal flora of
these banks has not yet been studied. A single bank approxi-
mately 40 m wide in the vicinity of Bushy Island breaks up into
a series of narrow ledges at the tip of the Bushy Island sand
spit, and into two main banks with ramifying small axillaries
between Bushy and Redbill, these extending to the south west
end of thejreef (see fig.1).: Beyond the algal banks is a "reef
flat proper," which is widest in the NW and much reduced in the
SW end of the region. A small lagoon is developed in this zone
LMeadtavely cow ot, thepysandsspit., The, "outer ireefiiutilat | gan this
western region is characterized by luxuriant alycionarian growth
and sponges, as well as a diverse and luxuriant coral composi-
tion. This zone continues to the reef edge, where very dense

h

coral growth occurs uninterrupted down the face of the reef
slope to the first part of the floor at approximately 10 m.
A very open spur and groove system is developed in the reef
front.

3. In the region most exposed to prevailing winds,
zonation is somewhat different. The widest and innermost zone
is one of dense microatoll development, the microatolls being
very low (approximately 30> cm an height), and elongated ina

direction at right angles to’ the’ reef edge: “in) some aneas
these structures have grown together to form a continuous pave-
ment which crumbles easily underfoot. Beyond’ this is a shallow

"inner flat" zone with oystercovered coral boulders, and some
special characteristics (such as the presence of the commercial
trochus Trochus (Rochia) niloticus). This is followed by a
zone of consolidated coral rubble and sand, with green algal
covering, possibly analogous to the algal banks mentioned
previously, but wider and more gradually sloping. Scattered
coral colonies appear in this zone towards the reef edge and

a narrow reef flat zone at approximately mean low water mark
bordered by the reef edge is apparently analogous to the "algal
ridge" of other reefs - coral cover is low but dense, of
corymbose and encrusting forms, with Acropora cuneata in low
vertical plates perpendicular to the reef edge (see plate Bi)
The reef front in this area drops as a series of terraces,
between which the floor is studded with coral boulders. A

spur and groove system is developed, the grooves often being
secondarily overgrown, forming long slits in the reef.

4. The eastern side of the reef was the least investi-
gated, but in the vicinity of transect 6 (NNE end of reef ) aot
was seen to combine features of the leastexposed and the most-—
exposed regions. A very wide sandy "inner flat" is separated
by broken algal banks from a relatively barren outer flat, then
a zone of flattened coral growth, followed by luxuriant coral
growth on the reef front to the floor at approximately 8 m,
where large coral-covered knolls occur.

CORAL DISTRIBUTION

Distributions are variously described, depending on the
methods and intensity by which zones were studied. The study
concentrated on Scleractinia, but Millepora and Tubipora have >
been included.

"Inner Flat" Zones:

Table 1 gives coral occurrences in sections of transects
running through "inner flat" zones. The zone is characterized
by a small number of species, the commonest being the massive
Porites (mainly P. lutea), Goniastrea australensis and Acropora
palifera. (Note: The classification of Wijsman-Best (1972) is
followed for Faviid species. The authors suspect that a

thorough taxonomic study of A.

alifera and A. cuneata may
prove them to be a single species. )

Algal Banks

The only corals occurring on the green algal banks
occasional very small encrustations of the
Goniastrea and Favites.

in particular Porites,

Lagoons

No lagoonal transects were made,

were

Winner flat" species,

but the following species

were recorded from patch reefs within the lagoons:

Pocillopora damicornis

Seriatopora hystrix

Stylophora mordax

Acropora digitifera

A. palifera

A. hyacinthus

A. cf. syringodes

Montipora (plate and branching species)
Astreopora myriopthalma

Fungia actiniformis

EF. fungites

Porites (massive and branching species)
Goniopora tenuidens

Montastrea curta

Microatoll Zone of the Central Region

No transects were made in the microatoll zone,

following species were recorded:

(a) forming microatolls:

Seriatopora hystrix

Porites andrewsi

Porites sp. 1 and 2 (branching)

Acropora palifera forma a

Pavona frondifera

Favia speciosa
Favites abdita

F. virens

Goniastrea pectinata
Platygyra lamellina
Cyphastrea serailia
Echinopora lamellosa
Leptastrea purpurea
Lobophyllia corymbosa
Acanthastrea echinata
Symphyllia nobilis
Millepora exaesa

M. tenera

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Microatoll Zone of the most exposed Region

This zone was also not studied in detail, but the
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by Acropora palifera forma a.
iteen lat Proper!

The only area corresponding to this zonation studied was
Mimthe vicinity Of Transect 2, and was seen to be of Similar
coral composition to the "inner flat" zone, although depth was
greater.
"Quter Flat" Zone

The whole "outer flat", to the edge of the reef, is here
treated. Table 2 compares coral representation in three tran-

sects. The comparative lengths of the transects should be
noted.

Reef Slope

Four transects passing over the first part of the reef
slope are compared in table 3. Those in the region of transects
2 and 4 were gently sloping Ga m slope to a depth of 10 m) ;
those of transects 5 and 6 dropped sharply to the floor.
Alcyonarians were most prevalent in the region of transect 2
(least exposed side of reef).

Floor adjacent: Mio. tarsi part of Slope

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Aceoporaintermedia, Ay ch. acuminata,” Aji ;pulichra, Hydnophora
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A. murrayensis), and encrusting to plate forms (Montipora,
Echinopora lamellosa, Merulina ampliata). In the region of
transect 2, Pectinia lactuca is common on coral knolls and on
the reef floor; in general, in this area alcyonarian growth
is more luxuriant than scleractinian growth on coral knolls.

The floor is generally sandy, and coral growth includes
seattered "sprawling" Acroporas such as A. cf. syringodes and
various Fungiids.

DISCUSSION

Bushy—-Redbill is a large reef, which presents a variety
of reef associations and zonations. In this brief study some
147 Scleractinian species, from 54 genera, (61 genera and sub-
genera) were recorded, and it could be expected that rarer
species and genera would be found by more intensive survey.

The reef is reasonably accessible from the Queensland mainland,
and Bushy Island is one of the few habitable coral cays in the
central Great Barrier Reef Province. This would appear to be
an excellent reef for further study of coral reef and reef-
associated topics.

ACKNOWLEDGEMENTS

We wish to acknowledge the help of the staff of the
Queensland Herbarium, with identification of plant specimens;
A.B. Cribb, University of Queensland, with botanical advice,
and M. Wilson, University of Queensland, with geological
information. We also thank B. Campbell, Queensland Museum,
who critically read the manuscript, and the following companions
on the trip, who collected specimens and gave criticisms on the
manuscript: R. Elks, W. and G. Gordon, D. Hadely, T. Nelson-
Gracve, Rotand J.~Powell, €.J. Wallace, H. ‘Walsh... R. ‘Pearson
(Fisheries Branch, Queensland Department of Primary Industry )
and P. Beveridge (University of the South Pacific, Fiji)
assisted by discussing the identification of some of the rarer
coral species, and their help is gratefully acknowledged.

REFERENCES

Australia Pilot Vol IV. 1962 (5th ed.). Comprising the eastern
coast of Queensland from Sandy Cape to Cape York, with the
off-—lying islands and reefs, including Great Barrier Reefs.
Hydrographic Department, Admiralty London. 438 pp.

Hedley, C. 1925. Report of the scientific director for 1924.
Trans. Roy. Geog. Soc. Australasia (Qd) Rep. Gt. Barrier
Reef Comm. 1: 157-160.

16

Loya, Y. 1972. Community structure and species diversity of
hermatypic corals at Eilat, Red Sea. Mar. Biol. 13(2):
100-123.

Maxwell, W.G.H. 1968. Atlas of the Great Barrier Reef.
Elsevier. The Netherlands. 258 pp.

Steers, J.A. 1938. Detailed notes on the islands surveyed and
examined by the geographical expedition to the Great
Barrier Reef in 1938. Rep. Gt. Barrier Reef Comm. 4(3):
53-104.

Stephenson, W. and Wells, J.W. 1956. The corals of Low Isles,
Queensland. August 1954. Univ. Qd. Pap. Dept. Zool.
1 (4) al =59R7 plabesr

Stoddart, D.R. 1969. Ecology and morphology of Recent coral
reetis,. Biol. Reve bern 3a=198)

Wells, J.W. 1955. <A survey of.the distribution of reef coral
genera in the Great Barrier Reef region. Rep. Gt. Barrier
Reef Comm. 6(2): 1-9, + chart.

Wijsman-Best, Maya, 1972. Systematics and ecology of New
Caledonian Faviinae (Coelenterata-Scleractinia). BaiqGir:
Dierk.- 42(1): 3-90, 14> plates:

APPENDIX 1

Scleractinian corals and non-scleractinian

hard corals recorded from Bushy—Redbill Reef

The following identifications were made (by C.W.) from
literature descriptions, and wherever possible with the aid
of specimens in the Queensland Museum identified by J.W. Wells
(Stephenson and Wells, 1956; Wells, 1955). Many of the iden-
tifications are thus of a very tentative nature. Where speci-
mens have been entered into the Queensland Museum collections,
the registration numbers are given. Some groups, particularly
the genus Acropora, have not been registered, pending further
study. Species also represented in the Great Barrier Reef
Committee collection are marked with an asterisk.

Other Abbreviations:
(Ss) Sight record only

(Oc) specimen collected, but not deposited in the
Queensland Museum.

CLASS ANTHOZOA
SUBCLASS ZOANTHARTA

ORDER SCLERACTINIA
Suborder Astrocoeniina
Family Thamnasteriidae
Psammocora contigua (Esper) G6931

Pcl. Stephanaria) togianensis Umbgrove G6927
Pe wel . Plesioseris ) haimeana Milne-Edwards and Haime
G69 34

P. sp.1 G6846, G6928, G6935

Pocilloporidae
Stylophora mordax (Dana) G6871

ep pistillata (Bspen) | G6s7Z0

Seriatopora hystrix (Dana) G6874, G7220
Pocillopora damicornis (Linnaeus)* G6867-8, G6872-3

P. verrucosa (Ellis and Solander) (S)

Acroporidae
Acropora abrotanoides (Lamarck).
ein jacuminata Verri Li
aspera (Dana)
ef. brueggemani (Brook)
ef. Clavigera (Brook)
ef. conferta (Quelch)

corymbosa (Lamarck)
cuneata (Dana)

decipiens (Brook)
delicatula (Brook)
digitifera (Dana)

ere Is (Brook)

formosa (Dana)

cf. glochiclados Crossland
haimei Milne-Edwards and Haime
hebes (Dana)

humilis (Dana) *
hyacinthus (Dana)

. intermedia (Brook)

ef tatastel ia (Brook)
cf. murrayensis Vaughan
palifera faassen
palifera forma a (Brook)
ef. polymorpha (Brook)
pulchra (Brook)

cf. quelchi* Crossland
Cf. nosaria (Dana)

ef. rotumana (Gardiner)
Spicifera (Dana)
squamosa (Brook) *

ef. syringodes (Brook)

IFIP IPP lel elle ele lel ele le le lee lle lee le lel lel ll

7

Geo25).

=
ee)

cf. variabilig (Klunzinger)

Spel Gelab map lane)
sp.2 (fructicose)
sp.3 (arborescent)
sp.4 (arborescent)
ep? (=A. rosaria forma 1, Crossland)

sp.6 (ike)
ontipora danae (Milne-Edwards and Haime) G6875, G6877, G6881
divaricata Brueggeman G6917
erythraea von Marenzeller G6913-6
foliosa (Pallas) G6921
foveolata (Dana) G6920

verrucosa (Lamarck) G6879-81
Spice G6923

sp.2 G6922
sp.3 G6884
sp.4 G6885
sp.5 G6924
sp.6 G6883

streopora myriophthalma (Lamarck) G6763, G6951
sp.1 G6946, G6949
sp.2 G6896

Pelee lststet StS shale Sts ts ete le [eee le le le

Suborder Fungiina
Super Family Agariciicae
Agariciidae
"Agariciella" ponderosa (Gardiner) G7219
Pavona cf. clavus (Dana) G6926
P. cf. decussata (Dana) G6843
S frondifera Lamarck G6841, G6844, G6932
(Pseudocolumnastrea) pollicata Wells G6842

oe rugosa (Lamarck) G6838
P. speciosa (Dana) G6839
Leptoseris ?mycetoseroides Wells G6930

Siderastreidae
Anomastraea (Pseudosiderastrea) tayamai (Yabe and Sugiyama)
G7230
Coscinaraea columna (Dana) G6840, G6845

Super Family Fungiicae
Fungiidae

Fungia (Ctenactis) echinata (Pallas) (0C)
F. (Pleuractis) scutaria (Lamarck) 66779
ne (Heliofungia) actiniformis Quoy and Gaimard (0C)
ye (Fungia) fungites (Linnaeus) .G6775, G6777-8
Herpolitha limax (Esper) G6772, G6774
Polyphyllia talpina (Lamarck) G6776
Parahalomitra robusta (Quelch) G6771, G6773
Podobacia crustacea (Pallas) G6933

Wy)

Super Family Poriticae
Poritidae
Goniopora tenuidens (Quelch) G6816, G6939
G. lobata Milne-Edwards and Haime G6817, G6942
Porites andrewsi Vaughan G6832-5
annae Crossland G6823-6
lobata Dana G6827-8
lichen Dana G6831, G6938
lutea Milne-Edwards and Haime* G6829-30, G6936-7
sp.1 (branching) G6818-22
sp.2 (branching) G6837, G6940-1
veopora mortenseni Crossland G6929
BPE Piavil G6836

: ia]

oll

cas

Suborder Faviina
Super Family Faviicae
Faviidae
Subfamily Faviinae

Caulastrea furcata Dana G6897-9, G7O4O
Favia favus (Forskaal ) G7185-6
F. pallida (Dana) G7200
- speciosa (Dana) G7184, G7202
stelligera (Dana) G7183, G7201
eh. mat thax
cf. valenciennesi G7188
Favites abdita (Pllis and Solander) G7205-8
F. flexuosa (Dana) G7197, G7209, G7221
Pemwirens (Dana) G7210=1
Qulophyllia crispa (Lamarck) G6894-5, G7199
Goniastrea pectinata* (Ehrenberg) G6853, G6855, G6904
Goniastrea australensis (Milne-Edwards and Haime) G6903,
G6905, G6907, G6912, G7190-1
G. retiformis (Lamarck) G6851, G6955
Platygyra sinensis (Milne-Edwards and Haime)* G6856-7
P. daedalea Ellis and Solander G6852, G7213
P. lamellina (Ehrenberg) G6854
Leptoria phrygia Wane and Solander)

Hydnophora rigida (Dana) G6762, G6765
H. exesa (Pallas) G6893

Subfamily Montastreinae
Montastrea curta (Dana) G6850, G6901-2, G6911
Plesiastrea versipora (Lamarck) G7187
Leptastrea purpurea (Dana) G6908-10, G7187, G7214-7
Ee eransvensa Khunzangenry G72 Wiz
Cyphastrea ja japonica es and Sugiyama G6886, G6890
C. serailia (Forskaal) G6887-8
Echinopora gemmacea (Lamarck) G6889

E. horrida Dana Dana G6769
E. lamellosa (Esper) G6766-8, G6891-2

Pelee ele

20

Oculinidae

Galaxeinae
Galaxia fascicularis (Linnaeus) G7099
Acrhelia horrescens (Dana) (0C)

Merulinidae
Merulina ampliata (Ellis and Solander) G6866
Clavarina scabricula (Dana) G6761

Mussidae
Acanthastrea echinata (Dana) G6900

Lobophyllia corymbosa (Forskaal) G7101, G7105
L. costata (Dana) G7100, G7103-4, G7107, G7109

L. hemprichii (Ehrenberg) G7102

L. (Palauphyllia) hataii Yabe, Sugiyama and Eguchi 67106
Symphyllia nobilis (Dana) G6764

Parascolymia vitiensis (Brueggeman ) G7039, G7041=2

Cynarina lacrymalis (Milne-Edwards and Haime ) G7043

Pectiniidae
Echinophyllia aspera Ellis and Solander G6859-60, G6944
Mycedium tubifex (Dana) G6862-63, G6865
Pectinia lactuca (Pallas) G6861
Oxypora lacera (Verrill) (S)

Suborder Caryophylliina
Super Family Caryophylliicae
Caryophylliidae
Eusmiliinae
Euphyllia glabrescens (Chamisso and Pas G6876,- G7145
Catalophyllia plicata (Milne-Edwards and Haime G6895
Physogyra lichtensteini (Milne-Edwards and Haime) (S)

Plerogyra sp. OC

Suborder Dendrophylliina
Dendrophylliidae
Dendrophyllia nigrescens (Dana) G6869
Tubastrea sp.
Turbinaria peltata (Esper) G6849, G6954

T. cf. lichenoides Bernard G6848
T. sp.1 GO8h7, G6945

Misp.2) 66952

T. sp.3 G6948

T. sp.4 G6858

Non=-Scleractinian corals
SUBCLASS OCTOCORALLIA

STOLONIFERA
Tubiporidae

Tubipora musica (Linnaeus ) *
CLASS HYDROZOA

MILLEPORINA

Milleporidae
Millepora exaesa Forskaal G7218, G7222
M. tenera Boschma G7223-4

APPENDIX 2

Vascular plant flora of Bushy and Redbill Islands

Bushy Island:

PANDANACEAE
Pandanus sp.

GRAMINEAE
Sporobolus virginicus (L.) Kunth
Thuarea involuta (Forst.) R. &S.

PALMAE
Coces sip).
CHENOPODIACEAE
palsola kali L.
CASUARINACEAE
Casuarina equisetifolia L.
NYCTAGINACEAE
Pisonia ¢randis R. Br.
ATZOACEAE

Sesuvium portulacastrum (Cie) ates

LAURACEAE
Cassytha filiformis L.

TILIACBAE
Triumfetta procumbens Forst.‘f.

LEGUMINOSAE
Sophora tomentosa L.
MALVACEAE
Abutilon albescens Miq.
CONVOLVULACEAE

Ipomoea pes-caprae (L.) R. Br.

2

22 |

BORAGINACEAE
Tournefortia argentea L.f.
Cordia subcordata Lam.

MYOPORACEAE
Myoporum acuminatum R. Br.

VERBENACEAR
ViaGex Ura foliar:

GOODENTACEAE
Scaevola taccada (Gaertn.) Roxb.

COMPOSITAE
Tridax procumbens L.
Redbill Island:

GRAMINEAE

Thuarea involuta (Forst.) R. & S.

MORACEAE
Ficus opposita Miq.

ATZOACEAE
Sesuvium portulacastrum Gi ae 3

Plate 1: Air photograph of Bushy-Redbill Reef. Crown copyright
reserved; reproduced from material supplied by the Department
of Minerals and Energy, Canberra.

Plate 2: Reef flat between Redbill Island (foreground) and Bushy
Island, showing borken algal banks. 4

Plate 3: Reef crest in the vicinity of transect 4 (exposed side of

reef).

General Location
“of Reet.

MPN

Proportionate
Wind Influence
Based on
1964-68 Records.

irmill

ao, ee CM
(_]sand.
Lagoon.
Microatoll Zone.
“inner Flat.

eef Flat Proper.”

ense Microatoll” Zone.
{ll outer Flat to Reef Margin. Ve 0 1Kms.
{J Area Not Investigated.
wa Algal Banks.

SCALE.

Fig. 1: Diagrammatic representation of
Redbill Reef, with position of transect lines indicated.

Insets indicate prevailing winds and relative position of the
reef with reference to the Queensland coast.

general zonation of Bushy-

LEGEND
L] Sand Slope.

[J Beach at level of crest :- Young Pandanus, Tournefortia and |pomoea. SCALE.
Pandanus and Tournefortia. Se eee
Tournefortia. ® 0 on
Pisonia Forest.
FE] Cleared area characterized by Pandanus, grasses, Cocos bounded by
Pisonia and Pandanus.
#4 Cleared area with Pandanus, Pisonia, Tournefortia and grasses.
BB Beach Rock.
Position

Zonation of the dominant vegetation of Bushy Island.
of beach rock also noted.

BiG. ae

‘NOOTd LNON4 W174 4334 Y3LNO : NV «V4 4334 UINNI adO1s tee... ONVIS ieee Ole “NOpoe
4334 Wwao1v How3a HOV3d =—-Y3NNI

*PSpnpToOuT Useq eaey UoT}eIebeA pue souoz Joez HutAkztdAQ sydezbojoyd pejoetes
*@ WOssuezqz go ARTUTOTA 9y3 UT UOCObeT pue j7o0z jUsSe(pe pue puetsz Aysng Fo eTTJOrd OTZeuUSeyos :¢ °Htg

ATOLL RESEARCH BULLETIN
NO. 195.

CORAL CAYS OF THE CAPRICORN AND BUNKER
GROUPS, GREAT BARRIER REEF PROVINCE,
AUSTRALIA

by P. G. Flood

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

February 1977

TOWNSVILLE

MACKAY

QUEENSLAND Tropic of Capricorn |.
GLADSTONE 2
CAPRICORN — BUNKER
GROUP

BRISBANE

CAPRICORN-BUNKER GROUP

Norah KILOMETRES

Tatas ‘

aye Us ~ Broomfield Reef

North West Is. 8." ie Wreck /s.

NS
HERON ISLAND _ ,.o¥Kes\ Reef

Wistari Reef wisi; .
Erskine Is.:2 use Se Fe cee
ca, Masthead Is. \

oe, ie
F200 ev ey,

nies Lamont Reef
Polmaise Reef :

ri Fitzroy Reef
es Llewellyn Reefs

SO Mink ae si Boult Reef

GLADSTONE °& Hoskyn Is. * SK

An syFatrfax Is

2 m

Lady Musgrave Is **

tase

(fH SM

Mast 15 2°E

Fig. 1. Location of the Capricorn and Bunker Groups.

CORAL CAYS OF THE CAPRICORN AND BUNKER
GROUPS, GREAT BARRIER REEF PROVINCE,
AUSTRALIA

by P.G. Flood l

Introduction

The islands and reefs of the Capricorn and Bunker Groups. are
situated astride the Tropic of Capricorn at the southern end of the
Great Barrier Reef Province and approximately 80 kilometres east of
Gladstone which is situated on the central coast of Queensland (Fig. 1).

The Capricorn Group of islands consists of nine coral cays: North
Island, Tryon Island, North West Island, Wilson Island, Wreck Island,
Masthead Island, Heron Island, and One Tree Island. A tourist Resort
and Marine Scientific Research Station have been established on Heron
Island. A manned lighthouse operates at North Island and the
Australian Museum conducts a field research station on One Tree Island.
The Bunker Group consists of five coral cays: Lady Musgrave Island,
Fairfax Islands (West and East), and Hoskyn Islands (West and East).

Morphological changes occurring between 1936 and 1973 are evident
when comparing previous plans of these coral cays (Steers, 1938) with
recent vertical aerial photographs. Changes are catagorised into two
groups; those related to natural phenomena and secondly, those caused
by human interference.

Previous work

The earliest scientific description of the Capricorn and Bunker
Groups is that of Jukes (1847) who visited the area in 1843 on the
voyage of H.M.S. Fly. Saville-Kent (1893) and Agassiz (1898) make
brief references to the Groups. Steers (1937, 1938) provided the
first detailed descriptions and sketches of most islands of the Groups.

l Department of Geology and Mineralogy, University of Queensland,
Brisbane, Australia.

(Manuscript received July 1974 -- Eds.)

The regional location, bathymetry, and physiography of this area
has been discussed by various authors (Maxwell, 1968; Maiklem, 1968,
1970; Maxwell and Maiklem, 1964) and the island vegetation has been
mentioned by several workers (MacGillivray and Rodway, 1931; Fosberg
and Thorne; 1961; Gillham, 1963; Cribb, 1965, 1969; Demm; W9o71)r

Steers (1938) and Domm (1971) provide an introduction to the
Groups and both papers should be read in conjunction with this article.

Reefs with coral cays

Four distinct reef types (Maxwell, 1968) occur within the Capricorn
and Bunker Groups (Figs. 2, 3, 4 and 5):

1. Platform Reefs:- Tyron, Wreck, North, Wilson, and Erskine
Reefs.

2. Lagoonal Platform Reefs:- Heron, and One Tree Reefs.

3. Elongate Platform Reefs:- North West, and Masthead Reefs.

4. Closed Ring Reefs:- Lady Musgrave, Fairfax, and Hoskyn Reefs.

The reader is referred to Maiklem (1968) and Maxwell (1968) for
detailed discussions concerning the zonation of individual reef types.

The coral cays belong to two distinct types (Fairbridge, 1950):

1. Vegetated sand cays:- North West, Heron, Masthead, Erskine,
Wilson, Wreck, Tryon, North, Fairfax (West), and Hoskyn (West).

2. Shingle cays:- One Tree, Lady Musgrave, Fairfax (East), and
Hoskyn (East).

The sand cays are all located to the lee of their reef flat,
whereas the shingle cays, with the exception of Lady Musgrave, are
located on the windward side.

The following brief descriptions are provided to assist
interpretation of the vertical aerial photographs (Plates 1-9) and to
allow comparisons to be drawn between present cay morphology and that
illustrated in sketches made by the Geographical Expedition to the
Great Barrier Reef in 1936 (Figs. 6 and 7).

Capricorn Group

Heron Island (Plate 1)

A sandy beach, 15-30 metres wide at low tide, surrounds the cay.
Exposed beachrock occurs on the southern, northern, and eastern
beaches. Vegetation consists of a central zone of Pisonia grandis
and an outer zone of Casuarina equisetifolia, Scaevola taccada, and
Tournefortia argentea. Pandanus tectorius occurs over the centre of
the cay. A Tourist Resort and Research Station have been developed
on the western end of the cay.

Changes occurring since 1936 are related to the natural elements
and to human interference. Sand erosion on the northern and eastern
beaches has exposed new occurrences of beachrock. The construction
of a retaining wall and a boat harbour have drastically altered the
western end of the cay.

North West Island (Plate 2)

This is the largest sand cay of the Group. A sandy beach
surrounds the cay and beachrock is only exposed at the eastern end.
Vegetation is Similar to that occurring on Heron Island.

The eastern area of exposed beachrock is smaller now than in 1936
and the southwestern outcrop is no longer visible.

Masthead Island (Plate 3)

Beachrock is well developed along the southern beach and an
occurrence of older beachrock is situated some distance from the beach
on the northwestern corner of the cay. Vegetation is similar to that
occurring on Heron Island. Prickly Pear, introduced from the
Mainland, is now well established over the entire cay.

Considerably more beachrock is now exposed on the southern beach
and a spit has formed on the northwestern corner. The shingle and
sand spit which was prominent in 1936 has disappeared.

One Tree Island (Plate 4)

This is a shingle cay. Vegetation consists of scattered
Tournefortia argentea and Scaevola taccada with several small groves of
Pisonia grandis. A small pond of brackish water is situated near the
centre of the cay. A temporary Field Research Station conducted by
the Australian Museum is located towards the northeastern corner of the
cay.

The straight alignment of the western beach in 1936 has been
altered by the addition of a small spit near the northwestern end of
the beach. The northwestern corner of the cay has extended towards
the lagoon.

Tryon Island (Plate 5)

Beachrock occurs along the northwestern and southeastern beaches.
The cay is covered with dense vegetation in the same zonations as on
the larger sand cays.

The areas of outcropping beachrock have altered considerably since
EGaGe No exposures can be seen on the northeastern beach, whereas the
exposures on the northwestern and southwestern beaches have increased
in area.

Wreck Island (Plate 5)

Beachrock is well developed along the southern beach. Vegetation
is similar to that on Tryon Island except that the Pisonia grandis
forest is less well developed.

No apparent changes have occurred since 1936.

North Island (Plate 6)
This is the smallest cay of the Group. A manned lighthouse
operates on this island.

Steers (1938) does not describe this island.

Wilson Island (Plate 6)

This island is not a true sand cay because of the quantity of
coarse coral shingle present in the sediments. Beachrock is well-
developed along the southeastern beach. Pandanus tectorius is the
dominant vegetation with Casuarina equisetifolia restricted to the
western side of the cay.

Significant changes have occurred since 1936. Considerable
quantities of sand have moved, exposing beachrock on the northern
beach and covering formerly exposed beachrock on the southern beach.
The sandy beach on the western side has increased in area.

Erskine Island

Beachrock is well developed on the western and northwestern
beaches. The vegetation differs from that present on the other cays
in that Pandanus tectorius and Casuarina equisetifolia are absent, and
the main vegetation consists of Tournefortia argentea, Scaevola
taccada, and stunted Pisonia grandis.

The cay has increased in size since 1936 and sand now covers
most of the formerly exposed beachrock.

Bunker Group

Lady Musgrave Island (Plate 7)

This island is the only shingle cay situated on the leeward reef
flvayee Beachrock is exposed along the northeastern and eastern
beaches and an outcrop of lithified coral conglomerate, similar to
that forming the core of the cay, occurs near the southeastern corner.
Vegetation consists of Pisonia grandis, Tournefortia argentea,
Casuarina equisetifolia, and Pandanus tectorius. The vegetation is
less dense than that of the larger sand cays of the Capricorn Group.
A small pond of brackish water is located towards the southern end of
the cay.

The shapes of the southeastern and eastern beaches have altered
Since 1936. Erosion has exposed a larger area of beachrock on the
southern beach and the eroded sand has been deposited on the eastern
beach partly obscuring the beachrock exposed in 1936 and forming a
well-defined beach on the northeastern corner of the cay.

Fairfax Islands (Plate 8)

The eastern cay is composed of shingle and the western of sand
and shingle. Interference with the vegetation of the eastern cay
occurred as a result of phosphate mining and secondly during the
period when the Australian Military Forces used the area as a bombing

target. The cay is no longer used for either purposes. Vegetation
is dominated by Pisonia grandis which is restricted to the centre of
the island. Two brackish pools are located towards the eastern end

of the island.

The western cay features an elongated sand spit that supports
vegetation on its western extremity. Vegetation is similar to that
occurring on the larger sand cays of the Capricorn Group. A tin shed,
erected by the Australian Navy when the eastern cay was used for
bombing practice, is located in the centre of the island. This shed
has been occupied for the past three years by Julie Booth who is
studying the behaviour of marine turtles.

Steers (1937) mentions the islands. However, he did not provide
any sketch of the cays.

Hoskyn Islands (Plate 9)

The eastern cay is composed of shingle and supports vegetation
similar to, although less well-developed than, that of Lady Musgrave
Island. The western cay is composed of sand and its vegetation is
Similar to that of the larger sand cays Capricorn Group.

Both cays have increased in size since 1936.

Influences causing morphological changes, 1936-1973

Most cays have changed morphologically as a result of continued
erosion of sand on the eastern, southern, and southeastern beaches and
subsequent deposition of sand on the western and northwestern beaches.
The Southeast Trade Wind which blows for approximately nine months of
the year causes this natural pattern to be common within this region
(Flood, in press) and accounts for the nett westerly migration of sand
on each cay.

Human interference is responsible for significant changes at
Heron Island (Plate 10). A concrete retaining wall constructed in
the early 1960's on the northwestern corner of the island is responsible
for erosion of the western beach (Fig. 8). The wall alignment
reflects and refracts onto the western beach those waves approaching
the island from a northwesterly and northeasterly direction and
considerably increases the erosive energy of waves in that area.

The erosion problem was increased in 1967 by the dredging of a
boat harbour into the reef flat to provide access to the tourist
Hesont “(Plate LO, fig. 92). Even before cyclone "Emily" (April, 1972)

the retaining walls of the harbour were breached in several places,
allowing sand from the western end of the island to move into the
harbour and to the deeper water beyond (Plate 10, fig. 3). The boat
harbour was redredged in late 1972. Approximately 20,000 cubic
metres of sand were placed adjacent to the concrete retaining wall on
the northwestern corner of the island in an attempt to lessen its
erosive influence (Plate 10, fig. 4). The sand is migrating

westward by longshore drift under the influence of the Southeast Trade.

Acknowledgements

The author gratefully acknowledges the use of research facilities
at the Department of Geology and Mineralogy, University of Queensland,
and at the Heron Island Research Station. I am particularly grateful
to Dr. G.R. Orme who organised the fieldtrips and aerial reconnaissance
of most cays of the Capricorn and Bunker Groups in 1972. The
Co-ordinator-General's Department, Queensland is thanked for
reproductions used in Plates 2 and 3. The officers of the No. 2
Sqdn. Royal Australian Air Force, Amberley generously supplied the
aerial photographs of the coral cays illustrated in Plates 1 and 4 to
9 inclusive.

References

Agassiz, A. 1899. A visit to the Great Barrier Reef of Australia in
the steamer "Croyden" during April and May, 1896. Bull. Harvard
Mus. Comp. Zoology 26: 1-203.

Cribb, AJB. =1965e The marine and terrestrial vegetation of Wilson
Island, Great Barrier Reef. PROGieR. SOC. (OG. Wiis 53—05r

@ribb;, AB 969). The vegetation of North West Island. el Neves JL)s
85-93.

DOMMya"SaD ne Odes The Uninhabited Cays of the Capricorn Group, Great
Barrier Reef, Australia. Atoll Res. Bull..142: 1-27

Fairbridge, (RoW... 1950. Recent and Pleistocene coral reefs of
Australia. J. Geol. 58: 330-401.

Flood, P.G. in.press. Sand movement on Heron Island - a vegetated
sand cay, Great Barrier Reef Province, Australia. Proc. 2nd
Intern. Symp. Corals and Coral Reefs, 1973.

Fosberg, F.R. and Thorne, R.F. 1961. Vascular plants of Heron Island,
pp. 5-13 in Fosberg et al., Heron Island, Capricorn Group,
Australia. Atoll Res. Bull. 82: 1-16.

Gillham, M.E. 1963. Coral cay vegetation, Heron Island, Great
Barrier Reef. Proc. R- Soc. Qd 73: 79-92.

Jukes, J.B. 1847. Narrative of the-Surveying Voyage of H.M.S. Fly.
Boone, London. 1: 424 pp.

MacGillivray, W.D.K. and Rodway, F.A. 1931. Plants on islands of
the Bunker and Capricorn Groups. Rep. Gt Barrier Reef Comm.
3(7): 58-63.

Maiklem, W.R. 1968. Thg Capricorn Reef Complex, Great Barrier Reef,
Australia. J. sedim. Petrol. 38(3): 785-798.

Maiklem, W.R. 1970. Carbonate Sediments in the Capricorn Reef
Complex, Great Barrier Reef, Australia. J. sedim. Petrol. 40(1):
55-80.

Maxwell, W.G.H. 1968. Atlas of the Great Barrier Reef. Elsevier,
Amsterdam. 258 pp.

Maxwell, W.G.H. and Maiklem, W.R. 1964. Lithofacies analysis,
southern part of the Great Barrier Reef. Pap. Dep. Geol. Univ.
Domai E21.

Saville-Kent, W. 1893. The Great Barrier Reef of Australia.
‘Allen, London. 387 pp.

Steers, J.A. 1937. The coral islands and associated features of the
Great Barrier Reefs. Geogrl J. 89: 1-28, 119-146.

IMS)e ksi Detailed notes on the islands surveyed and
examined by the Geographical Expedition to the Great Barrier
Reef in 1936. Rep. Gt Barrier Reef Comm. 4(3): 51-96.

LAGOON ,
e
Sy a
®oaTCH REEFS®
Cj o, @

ONE TREE REEF

23° 30'00"
152°05'00"

36) __ wITH NUMEROUS
_- REEF FLAT CORALS

MASTHEAD REEF

23°32'30"
151°44'00" ; METRES

NE,
x

1000

Bu.L UE - LAG. 0 O)N

, SANDY ZONE _—_
HERON REEF —— — rues
23°27'00"
151°57'00" METRE

1000

o silo eee

— SANDY ZONE WITH NUMEROUS REEF FLAT CORALS —

23° 1800"
151°45'00"

NORTH WEST REEF aoecee

Fig. 2. Reef zonation: One Tree, Masthead, Heron, and North West
Reefs.

REEF FLAT-

NCA CORALS

\
CORAL ZONE ‘ é Hair labia

AEH EE

x

x x x x
ALGAL ZONE
x KX x

TRYON REEF ERSKINE REEF

23° 15'00" 4 23°30 00"
151° 46' 30" 151° 46' 20" METRES

NORTH REEF
WILSON REEF 0
23°11' 00" a 23° 18' 10"

pe
ead PET METRES
151°54 30 151° 55 00

WITH NUMEROUS
REE RE ereAT CORALS ums

WRECK REEF

23°20'00" eee ce
ieeaucny METRES

Fig. 3. Reef zonation: Tryon, Erskine, North, Wilson, and Wreck
Reefs.

sal 294 ‘ fo)
Cos hal . \
Leitins LAGOON) » .
WS (Xk 7}
> =< S

Bagi yA

\ &
> AGOON @ LAGOONAL| A

‘

CLOSED RING REEF

Physiographic zonation of the reef types of the Capricorn
and Bunker Groups (modified from Maxwell, 1968, pelO6)

aoa

OF F-REEF

|
|

HOSKYN REEF

23°48'30"
152°18'00"

FAIRFAX REEF

23°51' 30"
152°22'30"

LADY MUSGRAVE REEF

23°54'30"
152°23'30"

Fig. 5. Reef zonation;

BUNKER GROUP

WEST CAY

1e} 500
_——— |
METRES

\
x x “
ALGAL ZONE

——
METRES

ENTRANCE

Py Le

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x
x
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——t
METRES

Hoskyn, Fairfax, and Lady Musgrave Reefs.

HERON ISLAND

ONE TREE ISLAND

SHINGLE

~%. SS ’
| DEPRESSION N2>->. *..
BEACHROCK

0 200

| Ee YARDS

Suinene = ee
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Fig. 6. ‘Shape of coi:al cays in 1936 (\aiver. Steers, 1938). Hernom,

One Tree, Masthead, and North “est Tslands.

WILSON ISLAND

Te

BEACHROCK

HOSKYN ISLAND a
Be pte. BEACHROCK 4%}

he
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ERSKINE ISLAND

BE ACHROCK

: LOW
OSKYN ISLAND €:, SHINGLE

c veky
CEMENTED eee eT
REEF FLAT \ "2

LOW
SHINGLE

Fig. 7. Shape of coral cays in 1936 (after Steers, 1938): Wilson,
Erskine, Hoskyn, Wreck, Tryon, and Lady Musgrave Islands.

R
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Fig. 8. Sequence of events connected with erosion of the western
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25:9°73

PLATE 4

Vertical aerial photograph taken 25.9.73.

One Tree Island.

Plate 4.

~ TRYON ISLAND 0) 300 .eTRES

WRECK ISLAND

RBLATE 5

Plate 5. Tryon Island and Wreck Island. Vertical aerial photograph
taken 2579273:

NORTH ISLAND O 300 WETRES

OE

WILSON ISLAND 25°9'73

PLATE 6

Plate 6. North Island and Wilson Island. Vertical aerial photograph
taken 25.9.73.

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ATOLL RESEARCH BULLETIN
NO. 196.

SUBMARINE CEMENTATION OF GRAINSTONE
FABRIC, ST. CROIX, U.S. VIRGIN ISLANDS

by Lee C. Gerhard

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

February 1977

SUBMARINE CEMENTATION OF GRAINSTONE
FABRIC, ST. CROIX, U.S. VIRGIN ISLANDS

by Lee C. Gerhard

ABSTRACT

Submarine cemented carbonate sand nodules occur off the northeast
coast of St. Croix, U. S. Virgin Islands. Cementation occurs as a 50
micron thick rim of aragonite needles over a thin micrite envelope sur-
rounding the cemented grains. The locality in which the nodules are
found is characterized by large ripples with the sand in nearly constant
motion. The sand is medium-grained with practically no very fine sand
and no micrite; neither are pellets present. Cementation is the result
of physicochemical processes, forming a basic grainstone fabric.

INTRODUCTION

Nodules of cemented carbonate skeletal sand occur several centi-
meters below and at the sediment-water interface in 35 feet of water off
EhesnortEheast coast of St. Croix, U. S. Virgin Islands (Fig. 1), just
west of Cottongarden Point. These nodules lie in loose sand in a chan-
nel between and parallel to the main barrier reef and a second, outer,
reef which does not reach the surface. Actively moving large ripples in
the sand bottom (Fig. 2) characterize the entire bottom of this area, in-
cluding both areas of cemented nodules and areas where no cementation
occurs.

Individual nodules range from a few cemented grains to masses 8
centimeters in maximum diameter (Fig. 3). Grains in cemented nodules,
nodule-bearing sands and non-nodule-bearing sands are medium-grained,
well-sorted, skeletal sands (Fig. 4). Percentage of individual skeletal
constituents are approximately the same in the cemented material, nodule-
bearing sands, and non-nodule-bearing sands (Table 1), and no clastic
rock grains are present in either nodules or sands. Practically no
micrite or pellets are present in any sample.

The cement is exclusively aragonite, determined by SEM and electron
microprobe (Fred T. Mackenzie personal communication). Aragonite
needles rim individual grains or small clusters of several grains. These
rims average about 50 microns in thickness, with individual crystals

a
1 West Indies Laboratory, P. 0. Annex Box 4010
Christiansted, St. Croix, U. S. Virgin Islands 00820

Current address: University Station
North Dakota Geological Survey
Grand Forks, North Dakota 58201

(Manuscript received August 1974--Eds.)

Z

oriented normal or subnormal to grain boundaries (Fig. 5). Nearly every
grain coated with the aragonite rim cement also has a thin micrite en-
velope below the aragonite. Cementation appears to be more complete in
the centers of nodules than towards the exterior. Coarse particles are
not cemented as well as those of near-mode size.

Sand ripples are symmetrical approximately 10 centimeters high from
trough to crest, and the sand is in relatively constant motion during
times of normal ocean swell (Fig. 2). Although the site of cementation
sas not been visited during storm swell times, logically the motion is
even stronger then.

DISCUSSION

Many examples of modern carbcnate cementation are known from a
variety of geographic locations and environments. Nearly all of these
examples are from "special" environments, that is, those with subaerial
exposure, high salinity, or other non-normal marine characteristics.
Most of these examples also involve either pellets in the sediment,
micrite, or both. (A short note is not the place to give an extensive
literature summary, but the papers and references of Macintyre et al,
1968; Bricker, 1971; and Sibley and Murray, 1972 are pertinent). Many
of these previously described types of cementation occur on St. Croix
in addition to the cement described here.

The major unique characteristics of the cementation described here
are the occurrence of the cement in open, normal marine water, lack of
pellets, lack of fine-grained sediment, and lack of a physical stabi-
lizer. Cementation under these circumstances appears to be rare
(Chilingar, et al, 1967, p. 186), although this cement is similar to
that reported by Shinn (1969) in the Persian Gulf.

Granulometric analysis of nodule=-bearing and non-nodule-bearing
sands from the cement area provides little insight into the cause of
cementation (Fig. 4). Both samples are well-sorted. Graphic standard
deviation of the host sand is .40 phi, a well sorted sand (Folk, 1961)
and the same parameter in the non-host sand is .50 phi, still a well-
sorted sand. In both samples less than 5% of the total sample is less
than 1.5 phi size, demonstrating that any fine sand or smaller particles
either have been winowed out or have not been deposited. There is a
sharp inflection in the cumulative curves of these samples between the
most frequently occurring particle size and the next smaller 1/2 phi
interval, although a much less marked inflection occurs to larger phi
sizes. There is nearly no insoluble residue in these samples. The two
samples are virtually identical in size and sorting except for the
broader total size spread of the nodule-bearing area sample compared
with the non-nodule-bearing area sample (-2.0 to 3 phi compared to -1.5
0% 2)5.5; phi.

Constituent particle analysis of the two sand samples and the
cemented sand itself sheds little additional information (Table 1).
Coral content of each sample is about the same, no sample contains any
pellets, and foraminifera, molluscs, and coralline algae form the
remainder of the samples. The category "other'' contains unidentified

grains, Halimeda, echinoderms, and other miscellaneous skeletal
materials. Individual grain skeletal fabrics are unaltered. No dis-
coloration is present nor are any of the cemented grains previously
micritized.

Radiometric dating has not been done, but the cementation is inter-
preted to be modern because the cemented sands are identical to the
sands currently being generated and transported in that environment,
no pre-existing cemented carbonates of this kind have been discovered
in this general area (or anyplace else on the island), no terrigenous
(clastic) grains exist in the cemented or uncemented sands, although
such grains are ubiquitous in nearshore or beach deposits (if the
cemented nodules were remnants of earlier Holocene beachrock, they
would contain large numbers of clastic grains), and grains are not per-
vasively micritized as is common in most of the St. Croix examples of
cemented carbonate sands.

Cementation cause is probably purely physicochemical. Cement mor-
phology is that predicted for normal marine subtidal cement by Badio-
zamani, Mackenzie and Thorstenson (1973) from laboratory studies
(Mackenzie, personal communication). Sands are not stabilized at the
surface, but about 4 centimeters below the troughs of ripples the sand
is not moved except during uncommonly high seas. Cementation apparently
takes place in the temporarily stabilized sand forming nodules, which
are periodically moved by storm waves. No cemented layer has been
found, so the nodules do not lie at rest for really extended periods
of time, but are moved several times each year. Nucleation of cement
appears to be preferential to grains with micrite envelopes. The
resulting fabric (Fig. 5) is the precursor of typical grainstone fabric
such as that of the Lower Ordovician Manitou Limestone (Gerhard, 1972,
Delo).

CONCLUSIONS

Cementation in basically unstabilized carbonate sands which have
no appreciable micrite component and which are deposited well below
low tide occurs. There appears to be no granulometric or constituent
skeletal particle basis for cementation control, as nodule-bearing and
non-nodule-bearing sands are virtually identical in size, sorting, and
constituent particles. Cementation appears to be the result of purely
physicochemical processes in temporarily stabilized carbonate sands.
This example of submarine cementation is significant because it occurs
under normal marine conditions, forms a grainstone fabric, and can be
the source of grainstone fabric intraclasts in other textural type
carbonate rocks.

ACKNOWLEDGEMENTS

I wish to thank Fred T. Mackenzie for analysis of the cement comp-
osition and for his discussions of the problem, and John Milliman who
read the manuscript and made suggestions for its improvement. This is
West Indies Laboratory Contribution No. 20.

REFERENCES CITED

Badiozamani, Khosrow, Mackenzie, F. T., and Thorstenson, D.C., 1973,
Diagnosis of diagenetic environments of carbonate rocks by the
fabric of their cement (Abstract).

Bricker, 0. P., editor, 1971, Carbonate cements: The Johns Hopkins
Univ. Studies in Geology, No. 19, The Johns Hopkins Press, Balti-

more, Md., 376 p.

Chilingar,G.-V., Bissell, ‘H. J., and Wolf, K. H., 1967, Diagenesis ot
carbonate rocks, in Diagenesis in sediments, Chilingar, G. V. and
Larsen, G., eds.; Developments in Sedimentology, v. 8. Elsevier
Publishing Co., New York, p. 179-322.

Folk, R., 1961, Petrology of sedimentary rocks: Wemphill's, The Uni-
versity of Texas, 154 p.

Gerhard, Lee C., 1972, Canadian depositional environments and paleo-
tectonics, central Colorado: Quarterly Colo. School Mines, v. 67,
n. &, pe 1=<30.

Macintyre, I. G., Mountjoy, E. W., and D'Anglejan, B. F., 1968. An
occurrence of submarine cementation of carbonate sediments off the
west coast of Barbados, W.I.: Jour. Sed. Petrology, v. 38,

p. 660-664.

Sibley, D. F., and Murray, R. C., 1972, Marine diagenesis of carbonate
sediment, Bonaire, Netherland Antilles: Jour. Sed. Petrology, v.42,
Pe 168-178.

Shinn, E. A., 1969, Submarine lithification of Holocene carbonate
cements in the Persian Gulf: Sedimentology, v. 12, p. 109-144.

Table 1. Constituent particle analysis of cemented nodules, nodule-
bearing sand and non-nodule bearing sand.

Cemented Nodule Nodule-bearing Non-Nodule-

Sand Bearing Sand
Coral 50.6% SL aT 2 SL SLZ
Foraminifera 20.6 16.5 26.5
Coralline Algae 14.5 18.9 29
Mollusc Les G 8.2 Sel
Other Dial 4.7 Shale)

SUBMERGED
CEMENTED ~1\¢——e™, REEF
NODULES SAND —

BARRIER REEF

64° 45'

TAGUE BAY

£ COTTONGARDEN
aie ee POINT
=—

: lS
Christiansted

eiles. ST. CROIX

F igmal Location map showing St. Croix and enlargement of area
where cementation occurs.

Pies 2 Sand ripples in cemented area, 35 feet deep, showing
crestal sand in motion. Ripples are about 10 cm. high.

Fig «93 Nodule of cemented sand from northwest of Cottongarden
Point, St. Croix.

% %
50 50
40 40
30 30
20 20
10 10
0 ) i
s2.9b 1 Oy bee! SP 2 Or Ir: 2 oS Phi
(A) (B)
Fig. 4 Histograms of sand from non-cemented area (non-nodule

bearing sand) (A) and cemented area (nodule-bearing sand)
(B). Note the abrupt cutoff below 1.5 Phi size. Histo-
gram drawn on 1/2 Phi intervals.

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ATOLL RESEARCH BULLETIN
NO. 197.

CHRISTMAS ISLAND (PACIFIC OCEAN ):
RECONNAISSANCE GEOLOGIC OBSERVATIONS

by Mark J. Valencia

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

February 1977

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ae

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Figure 1. Location of Christmas Island.

CHRISTMAS ISLAND (PACIFIC OCEAN) :
RECONNAISSANCE GEOLOGIC OBSERVATIONS

by Mark J. Valencia“
INTRODUCTION

Christmas Island, the largest atoll in the world in terms of
subaerial surface area, is located 200 km north of the equator in the
equatorial dry zone (Schott, 1933) (Figure 1). The atoll is influenced
by nearly constant 4 m/sec easterly winds and an average annual
precipitation of 873 mm unevenly distributed in time (Jenkin and Foale,
PIGS)": The shape of this southernmost of the northern Line Islands
is like an elongated lobster claw with pinchers open to the northwest
containing a semi-circular lagoon (Figure 2). Two passes, each nearly
2 km wide, connect the main lagoon to the ocean. Numerous sub-
rectangular shaped hypersaline lakes are found to the east of the main
lagoon as well as centrally along the elongated "arm" which extends
18 km to the southeast. Along the southeastern margin of the lagoon,
the lakes are more numerous, and narrow channels connect some of them
to the lagoon.

Wentworth (1931) and Wentworth and Ladd (1931) have reported
reconnaissance observations on the general geology and sediments of
Christmas Island, and detailed analysis of landform, soils, and
hydrology is included in a report on the coconut-growing potential of
the island by Jenkin and Foale (1968). The soils have been further
investigated by Hammond (1969). Geologic data gathered by British
l Hawaii Institute of Geophysics Contribution No. 588.
2 Hawaii Institute of Geophysics University of Hawaii, Honolulu,
Hawaii 96822. Present address: UNDP Regional Offshore Prospecting
in East Asia, c/o ECAFE, Sala Santitham, Bangkok 2, Thailand.

3 First western contact and name courtesy of Captain James Cook,
December 24, 1777.

(Manuscript received June 1974 --Eds.)

2

scientists prior and subsequent to a 1957 nuclear test ("Operation
Grapple") (Jenkin and Foale, 1968) conducted at 9,140 m above the
southeast point, and by private guano and phosphate rock enterprises
at various times, is acknowledged, but such data were for the most
part unavailable to the author.

This paper reports 19 days of field observations and subsequent
laboratory analysis of samples collected on two expeditions during
March and November 1970 which investigated the Artemia (brine shrimp)
aquaculture potential of Christmas Island. The expeditions were
supported by National Sea Grant Program Grant 2-35-243 to P. Helfrich
of the Hawaii Institute of Marine Biology, and by private individuals
Seymour Gaines and Maurice Rakowicz. The aid of Fred Farrel, Glenn
Fredholm, John Hance, and Mayo Ryder during the field investigations
is gratefully acknowledged. Preparation of this paper was done while
the author was with the Hawaii Institute of Geophysics.

General geology

Christmas Atoll surmounts the southeasternmost portion of a

northwest-southeast trending volcanic ridge. The oceanic crust
surrounding the ridge exhibits a broad moat and arch, with the arch
crest some 240 km distant from the ridge axis. An archipelagic apron

forms a smooth curve grading into the insular slope (Menard, 1964).

An echo sounder profile of the southwestern slope shows "cone-like"
features of probable volcanic origin (Ritchie, 1958). From the reef
margin to a depth of 1300 m, the slope is quite steep (Jenkin and Foale,
1968) = Coral thickness is 30 m at Motu Tabu (Figure 2) and elsewhere
exceeds 120 m (Northrup, 1962; Jenkin and Foale, 1968; John Dryden,
oral communication, 1971).

The island can be classified into six subaerial and two submarine
landform units (Jenkin and Foale, 1968) on the basis of morphology and
sediment type. Table 1 and Figure 3 summarize the geomorphologic
characteristics of these landform units. Maximum elevation is 10.7 m
above msl in the coastal dune unit along the Bay of Wrecks (Jenkin and
Foale, 1968), although masses of reef rock distributed throughout the
island's inland area exhibit a rather uniform height not exceeding 4 m
above present sea level (Wentworth, 1931).

On the lagoon flats as much as a meter, but more commonly less
than 10 cm, of beachrock (biosparudite, Folk, 1965) caps extensive areas
of reef rock (coral biolithite and intrasparrudite) and unconsolidated
medium calcirudite. Thicker beachrock sections indicating cycles of
deposition and beachrock formation are exposed along walls of collapsed
spring channels which feed many of the enclosed lakes (e.g., Lakes l6a,
19e, and 27b) and on islet edges within Lake 27a. Incipient beachrock
formation was observed in dry areas of Manulu Lagoon and at 30 cm below
the floor of an inactive spring channel entering Lake 19e.

In situ Acropora and Tridacna reef complexes are exposed along some
lake margins. Repeated attempts at deep probing of the inter-lake
margins betwen Lakes I4 and F7 and on the southeastern shore of 27a met

with animpenetrable horizon at approximately 6.5 m below the surface.

HYPERSALINE LAKES

The approximately 500 isolated and interconnected hypersaline lakes
occupying 25 per cent of the total island surface area are of particular
interest. In area the lakes range from the 16 km? Manulu Lagoon to
puddles a few meters wide and centimeters deep (Figure 2). Frequently,
differences in water level of a meter of more were observed between
adjacent lakes, e.g., the water level of Lake EZ was 1.2 m higher than
that of Lake 16A. These differences may be explained by variations
in the water budgets of the lakes. Successive rings Of shell berms or
a border of evaporite crystals, either gypsum and halite or gypsum only,
around most lakes indicated higher water levels in the past. Large
fluctuations in lake surface area result from water level changes of
less than a meter due to the generally shallow and gently sloping

configuration of the lake basins. The lakes appeared to be ata
relatively low level in their recent history; the expeditions took
place during a prolonged drought. Lake levels were occasionally a

meter or more below inactive spring channels (e.g., Lake 22), and

Manulu Lagoon was observed to be partitioned into smaller interconnected
lakes, in contrast to maps constructed by Jenkin and Foale (1968) from
aerial photographs taken in the 1950's. Asymmetric basin profiles

with recently exposed windward shores indicated active aeolian
deposition in some lakes, e.g., in Lakes 27A and 16A.

A reducing environment, as indicated by the odor of hydrogen
sulphide, was present immediately below the surface sediment in all
lakes sampled. A semidiurnal increase in foam (reported previously
by Wentworth, 1931), was observed along the leeward shores of many of
the enclosed lakes; in the larger interconnected water bodies, linear
patterns of foam extending from leeward to half the lake breadth
indicated active Langmuir cells.

The lakes were of two basic types, subevaporite and evaporite,
depending on the degree and duration of isolation from the main lagoon
and ground water. Lakes actively depositing evaporites could be
further classified according to the nature of the evaporite. Key
indicators of lake types were bottom morphology, sediment type, and the
presence or absence of a red gelatinous alga (S. A. Cattell, personal
communication) apparently associated with medium hypersalinity, i.e.,
in this environment, approximately 200 to 300 o/oo.

Relatively isolated lakes, e.g., 19d, 19e, and 33a, were less than
2m deep and had salinities greater than 300 0o/oo. The lake bottoms
were rough as a result of a network of polygonally arranged ridges
exhibiting nearly 1 m of relief from ridge top to basin floor. The
gypsum and halite crust was only 5 cm thick in Lake 33a and overlay red
algae interspersed with halite crystals. A 1-m-wide band lacking
evaporites or containing only a thin crust of gypsum indicated
relatively low salinity seepage along the lake perimeters. (One could

4

determine thoselakes which were actively receiving shoreline seepage by
the presence or absence, in the evaporite ring, cf halite crystals which
are rapidly dissolved by contact with low salinity water.)

Other evaporitic lakes, e.g., 16a and 27a, had active, relatively
low-salinity springs (36-41 o/oo) flowing from collapse features as
much as 2 m deep eroded in the hardpan-capped reef rock, usually at the

shore nearest the main lagoon. The total input rate of water from the
several springs at the northern end of Lake 16a was estimated as less
than O.O1 m°/sec. Continuous long-term flow from one spring in Lake

16a was indicated by the presence of a distinctive, sharp, inverse
temperature gradient previously noted by Northrup (1962).

These spring-fed evaporitic.lakes, also less than 2 m deep, were
characterized by a smooth bottom consisting of a l-cm thick crust of
halite cubes overlying at least 1 m of red algae mixed with halite
crystals. Core léa-l (Table 2) from the windward shore exhibited a
regressive sequence, whereas a longer core (l16A-2, Table 3) revealed
cyclic variation in sediment type. A core of slightly pebbly, coarse
sandy calcareous mud (Folk, 1965) from the windward shore of:Lake 27a
had a minimum sedimentation rate of 1.8 cm/yr for the past 15 years.
This measurement is based on gieger count detection of the 1957 nuclear
test horizon at 28 cm below the core top (D. Knutsen and R. Buddemeier
personal communication, Figure 4). The sedimentation in this core
appeared normal and continuous for this location (Table 4).

Lakes that lacked gypsum and halite deposits, e.g.,E, F, and I
series, were interconnected by narrow channels to the main lagoon.
Channel morphologies for lakes in the F.series appeared to represent
stages in a developmental cycle. Incipient channel development was
possibly occurring between Lakes Fé and F7 as indicated by a 2-m-wide,
20-cm-deep water connection between the two lakes over the beach-rock-
capped divide. Perhaps during and after infrequent heavy rain, water
of low salinity, undersaturated with respect to calcium carbonate,
initiates dissolution of the hardpan while flowing between lakes.

After an unusually heavy rain, a rise in water levels in the lakes
resulting in submerged interlake boundaries has been observed (George
Krasnick, personal communication). Once the crust is weakened, the

silt and clay would be eroded by the water flow, leaving a sediment
dominated by pelecypod and Acropora fragments. A more advanced stage

in this process was represented by an inactive channel between F2 and

F4. At the F2 end, the beachrock crust was absent along 20 m of channel
length and the channel was filled with unbroken pelecypod shells. Once
the channel bottom is eroded below the water table (0.5 to 1m on the
lagoon flat, Jenkin and Foale, 1968), extensive widening may occur from
groundwater seepage and undercutting, as evidenced by blocks of beachrock
several meters in diameter slumped into one of the channels. An example
of the next stage of this cycle of channel activity was represented by

an inactive channel between F4 and F5. A hypersaline puddle was bounded
by a shell shoal at the F4 end and a shell berm at the F5 end. As the
water evaporates from the puddle, and by capillary evaporation elsewhere,
beachrock will form to stabilize the unconsolidated shells.

Line soundings in Lake F2 recorded depths up to 6 m, and fathometer
records indicated submerged isolated patch reefs in the F series lakes
(Helfrich; vect'al’. ; L973)". A salinity increase from the main lagoon
edge to the interior of interconnected lake series revealed the probable
cause of reef and invertebrate extinction in the lakes farthest from
the lagoon. A general salinity increase in a northeast-southwest
direction for interconnected lake series reflected the influence of
permanent fresh-water lenses in the northeast. For example, springs
entering lake Alb had salinities of 24 and 16 o/oo. (Heikeraatch) ete salle
ibe WS)

Nearshore bottom-sediment type varied considerably in these "open"
lakes but the dominant type for F series lakes was a Sandy, pebble
gravel of pelecypod and Acropora fragments. The berms around the lake
margins consisted of well-sorted pelecypod and gastropod pebble gravel.
In windward nearshore Lake Fla, a core penetrated 16 cm of pink organic
"fluff" down to an extinct pelecypod bed. Windward sedimentation or
reworking is fairly rapid in these open lakes, as evidenced by ruts
partially filled and smoothed by the pink organic "fluff" and aeolian
silts These ruts were made 5 months previously by a landrover used
by the expedition.

Lagoon

The rectilinear pattern of patch reefs and concomitant depositional
upbuilding extended into the lagoon from the southeastern and eastern
Margins (Helfrich, unpublished data). The margin of the lagoon was not
easily defined in these regions due to complexes of broad tidal flats,
peninsulas, reefs, and islands. A maximum lagoon depth of 7 m is
approximately equivalent to that of the F lakes. In addition to aeolian
deposition, large amounts of fine and medium calcareous debris
transported from the seaward leeward shore through the passes results in
a buildup of the lagoon floor (Wentworth, 1931). Continuous rapid
shoaling of the northern pass is evident from its dredging history
(Jenkin and Foale, 1968). In spite of high sedimentation rates, patch
reef development is more frequent in sheltered areas, such as along the
eastern margin, where the substrate is more stable than elsewhere.

Phosphate

Apatite, identified in the field and subsequently confirmed by
X-ray analysis (W. Burnett, personal communication), occurs to a depth
of 0.5 to 1m as peripheral and fracture rinds in a 2-acre outcrop of
beveled and severly eraded beachrock situated in the lagoon flat unit
in the north near Lake 1B (P, Figure 2). This observation is interesting
in relation to Hutchinson's (1950) hypothesis that several thousand
years ago, the equatorial rain belt was centered over Christmas Island,
i.e., farther south than at present. His hypothesis was based on the
supposed present day absence of evidence of phosphatization on
relatively dry Christmas Atoll in contrast to the present day occurrence
of phosphate rock on wetter islands to the north and south.

However, this one occurrence of phosphate rock does not account
for the amount of phosphatization that might be expected from the
combination of low annual precipitation and the present and presumably
past deposition of 200 tons/yr of guano as estimated from bird
populations (Helfrich et al., 1973). Avian Mining Company of Canberra,
Australia drilled a test hole to 40 m on Motu Tabu, apparently without
encountering commercial deposits of apatite (John Dryden, personal
communication).

The curious scarcity of both guano and phosphate rock deposits
could be ascribed to bird behavioral patterns and to leaching by
occassional heavy rains (Helfrich et al., 1973).

Hypothetical geologic development

Variations of rates and directions of spreading within the oceanic
crust emanating from the East Pacific Rise produce zones of crustal
weakness. During the formation of the Hawaiian Ridge, volcanism
progressed from northwest to southeast along such a zone, so the
southernmost volcanically active island of Hawaii is not only the
youngest but also the largest. The northwest-southeast structural
trend of the volcanically extinct Christmas Island ridge and the large
size and position of Christmas Atoll relative to the other Northern
Line Island atolls to the northwest evoke a comparison with the
evolving Hawaiian Ridge. If a similar mechanism of origin applies to
the Christmas Island Ridge, the volcanic complex upon which Christmas
Atoll has formed is\ then, the. youngest of the northern Line Islands,
having developed in early Cenozoic time (Menard, 1964).

The basic shape of an atoll is largely dependant on the morphology
of its volcanic basement (Wiens, 1962) but the surficial detail is
controlled by sea level fluctuations and climatic and tectonic influences.
The irregular shape of Christmas Atoll is probably a result of the
arrangement of volcanic peaks similar to those observed on the southwest
insular slope. The development of the major geomorphologic features is
a result of linear patch reef growth, and a progressive westward and
northward infilling of the main lagoon, perhaps aided by slow northwest-
ward tilting of the entire atoll. Roy and Smith (1970) speculate that
Fanning Atoll, 320 km to the northwest, has tilted up to the west,
whereas the nearest land to the south, Jarvis Atoll, 420 km to the
southwest, has a filled lagoon floor 2.5 m above sea level with evaporite
crystals in the deepest depressions thus indicating recent emergence
(Wiens, 1962).

Wiens (1962) has proposed a hypothetical model of the morphological
cycle of an atoll with changing sea level (Figure 5). His lowest reef
profile represents the appearance of an atoll at the end of the last low
stand of the sea (Wisconsin). At that time the emerged reef, which once
had been 100 m higher than the Wisconsin level, was dissolved and
mechanically eroded so that only portions of the reef periphery (A) and
small lagoon highs (D) remained above sea level. At this stage, the
lagoon bottom (C) was at sea level. Also produced was a fringing reef
flat (B) with an outer algal ridge and a live reef-front with a well-

developed wave-cut bench and a windward spur and groove system.

During the post-Wisconsin eustatic rise of sea level, a rectilinear
pattern of Acropora and Tridacna patch reefs developed on the immersed
lagoon floor of Christmas Island parallel or transverse to northeast
and southeast modes of wind direction. At present, easterly winds
predominate with a minor mode from the southeast, and linear patch reef
development transverse to the easterly mode is occurring in the windward
lagoon. However, the pattern of lake margins, presumably underlain by
reefs, hints at past northeast and southeast wind modes. Linearly
arranged patch reefs oriented downwind or, more rarely, transverse to
the main wind direction are present in several atoll lagoons (Wiens,
1962) including a rectilinear pattern in Fanning Atoll lagoon (Roy and
SuLtth, 197@). Linear patch reef development was due possibly to more
rapid reef growth in divergences of Langmuir cells or to a rectilinear
base of sand dunes developed during the Wisconsin low stand. In the
latter scheme, patch reefs would have been less well-developed in the
leeward than the windward lagoon due to more extensive bar development
nearer the windward sediment source. Differential growth rates of
coral and coralline algae between the seaward and leeward reefs,
possibly aided by northwestward subsidence, resulted in the northwest
passes.

As sea level rose to a maximum 2 m level within the last 5000
years (Fairbridge, 1952, 1961), Christmas Island was almost entirely
submerged, although lagoon patch reef development kept pace with the

rising sea level. During the brief still-stand and the subsequent
eustatic fall of sea level, the reefs were planed off, supplying
sediment to the emerging lakes. Beachrock formation on leveled,

exposed reef tops incorporated bivalves and other biogenic debris.
Aeolian sediment, derived from the emerged windward coastal plain and
the seaward beaches, formed the coastal inland and lagoon dunes and
also accumulated in the lake basins, along with autochthonous biogenic

sediment. Wave transportation and deposition of sediment was
effective in plugging breaches in lake peripheries. The lakes
progressively filled and became isolated from the main lagoon, although
some ephemeral shallow channels were maintained by tidal flow. The

increasing salinity curtailed reef and invertebrate growth, and bivalve
and gastropod shells accumulated in berms on the leeward lake shores.
The windward lagoon is now in the incipient stage of this process of
hypersaline lake formation, whereas the interconnected lakes are
indicative of the next stage. The completely isolated lakes continue
to shoal due to precipitation of evaporites, principally gypsum and
halite.

REFERENCES

Fairbridge, R. W., 1952. Multiple stands of the sea in post-glacial
times. Prac. 7th rPacuLre Sel. Congr. , 3’:

8

Fairbridge, R. W., 1961. Eustatic changes in sea level. Physics and
Chemistry of the Earth, 4: 99-185.

BOL wl cmLieurie 65's Petrology of sedimentary rocks: Hemphill's,
Ausitin,, Texas), 159) pp.

Hammond, Ti. T1969). The characterization and classification of the
soils of Christmas Island: Unpublished, Univ. Hawaii M. S. thesis,
1O9) {pps

Helfrich, Philip, in collaboration with John Ball, Andrew Berger, Paul
Bienfang, S. Allen Cattell, Mari Dell Foster, Glen Fredholm, Brent
Gallagher, Eric Guinther, George Krasnick, Maurice Rakowicz, and
Mark Valencia, 1973. The feasibility of brine shrimp production
on Christmas Island. Univ. Hawaii Sea Grant Tech. Rept.
aM ES OA ALIS) oh o}e

HutchinsonaaG vEi pul 950). The biochemistry of vertebrate excretion.
Bull. Amer. Museum Nat. Hist. 96: 1-554.

Jenkin, R. N. and Foale, M. A., 1968. An investigation of the
coconut-growing potential of Christmas Island. Ministry of
Overseas Development, Resources Division, Directorate of Overseas
Surveys, Tolworth, Surrey, England, Land Resources Study No. 4,
wrewks le

Menard, H. W., 1964. Marine Geology of the Pacific. McGraw-Hill
BooksCo.. , \WONGONn jp, Dis oink:

Northrop, John, 1962. Geophysical observations on Christmas Island.
Atoll Research «Bul lesS9icn - 2.

RittichieyieGe user Sk Sounding profiles between Fiji, Christmas and
Tahiti Islands. Deep-Sea Res., 5: 162-168.

ROY, pen ie ANGE OMitchy aSiy, Vasile Ors Sedimentation and coral reef
development in turbid water: Fanning Lagoon. Hawaii Institute
of Geophysics Report HIG-70-23: 61-76.

Schott inGen OSs. The distribution of rain over the Pacific Ocean.
Proc. SEN vw PaClELe: SCigs CONG TewS 3) L6G kL SIO.

Wentworthia.C.. Koy LOS de: Geology of the Pacific Equatorial Islands.
Bernice P. Bishop Museum Occ. Pap. IX, 15: 3-25.

Wentworth, .GeaksuanGd: addi iytion Sig uo Sdks Pacific Island sediments.
Studs Nat. AUS. Lowa sUNTVicc elo heal

Wiens, Heaiw.,. L962). Atoll environment and ecology: Yale Univ.,
53,2) (PD.

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Table 2. Megascopic description of sediments in Core l6a-l.
Depth (cm) Description
OF a2 Pebble-size halite crystals in groundmass of green

alga-calcium carbonate-halite mud; some pebble-
Size pelecypod fragments.

2 SS e@) Bimodal sediment: whole pebble- and cobble-size
gastropod and pelecypod shells in groundmass of
sand-size shell fragments.

LOU 76 Coarse sand-size gastropods; rounded and subangular
pebble-size shell fragments partially cemented
together with calcium carbonate; occasional entire
leaves.

16 7=: 30 Scattered coarse sand-size gastropods in white
calcium carbonate mud matrix.

30) = 59 Occasional gastropod and pelecypod fragments in pink
calcium carbonate mud matrix.

145)

Table 3. Megascopic description of sediments in Core 1l6a-2.

Depth (cm) Description
Oo- 4 Brownish-green algal mud; occasional pelecypod
fragments.
Ane 9 Brownish-pink algal mud; occasional pelecypod
fragments.
9 = 10 Pink algal mud.
ZO)=: 27 Same; abundant gastropod and pelecypod shell

fragments and intact pelecypod halves.

Bie 29 Gastropod layer.

29 = 30 Brownish-red algal mud.

SO)— 43 Tan-colored algal mud; abundant gastropod and
pelecypod shell fragments and whole large
gastropods.

43 - 49 Yellowish-brown algal mud.

49 - 58 Light-tan-colored algal mud; abundant sand-size

gastropod and pelecypod: shell fragments; green
algal layer at 52 cm.

53) —' 69 Intact pelecypod halves and large gastropods in
tan-colored algal mud matrix.

69 = 79 Tan-colored mud; not algal.

ao "S82 Pelecypod and gastropod shell fragment sand.

14

Table 4. Megascopic description of sediments in Core 27a-l.

Depth (cm) Description
OF =7'6 Sand-size and silt-size gastropod and pelecypod

shell fragments partially cemented together with
calcium carbonate; occasional benthic foraminifera.

6163 Same; abundant rounded beachrock pebbles with
exterior black algal coating.

63 = £20 White calcium carbonate mud with occasional
aeolian pebbles.

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o_O

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REEF AND SEA LEVEL AT PRESENT SEA LEVEL ACCRETION K
CLIMATIC OPTIMUM AND REEF

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PRESENT LAGOON SEDIMENTS
SEA LEVEL AT
GLACIAL LOW REEF AT GLACIAL LOW LEVEL

Figure 5. Hypothetical diagram of atoll evolution. A, undercut

"raised" reef remnant; B, old reef flat; C, old lagoon bottoms;
D, coral patches, knolls, or pinnacles. (From Wiens, 1962, Figure 40.)

ATOLL RESEARCH BULLETIN
NO. 198.

NOTES ON THE VERTEBRATE FAUNA OF
TONGAREVA ATOLL

by Roger B. Clapp

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

February 1977

NOTES ON THE VERTEBRATE FAUNA OF
TONGAREVA ATOLL

by

Roger B. Clapp!

Tongareva (or Penrhyn) Atoll, at 9° S\- 158° W in the south central
Pacific Ocean, is the northernmost of an isolated group of islands
north of the Cook Islands, and like them is administered by New
Zealand. Tongareva is a typical ring atoll about 40 miles in
circumference and contains a lagoon of about 108 square miles (Buck,
1932).

The vertebrate fauna of the atoll has been little studied despite
relatively frequently visits by missionaries, anthropologists, and
zoologists. The atoll was visited by the Kaimaloa Expedition in
December 1924 and in September 1936 and April 1937 by William F.
Coultas and R.W. Smith, respectively. Both Coultas and Smith
collected birds but no formal reports on the collections were ever
written.

On 13 June 1965, Tongareva was visited for 9 hours by a survey
team of the Smithsonian Institution's Pacific Ocean Biological Survey
Program (POBSP). Only the "bird islets" including Vaiari Islet at
the southwestern corner of the lagoon and portions of the northwestern
rim were visited.

Recently Batham and Batham (1973) presented observations on the
birds of the atoll made during a visit from 12 August to 20 September
L968. The present paper presents additional information on the
vertebrates of Tongareva Atoll and comments on Batham and Batham's
useful contribution. Appendix Table 1 lists bird specimens collected
irieLIso,.1937 and: 1965).

1

National Fish and Wildlife Laboratory, U.S. Fish & Wildlife Service,
National Museum of Natural History, Washington, D.C, 20560
(Manuscript received February 1975 -- Eds.)

Mammals

At the time of the POBSP visit, cats (Felis domestica), dogs
(Canis familiaris), and pigs (Sus scrofa) were being raised by the
natives. Comments in Ward (1967) and by Lamont (1867) suggest that
the pigs may have been introduced to Tongareva in 1853 from the ship-
wrecked vessel Chatham. Rats (Rattus sp.) were numerous in 1965, but
as no specimens were collected their identity is unknown. Lamont
(1867) stated that in 1853 "... [the natives] had never seen an animal
larger than a very small rat, that lives principally in the cocoa-nut
EGeCES i751 Lamont's description suggests that the rats on Tongareva
are Rattus elegans, a species widely distributed on the Pacific Islands

Reptiles

Although native informants indicated that turtles (most likely
the Green Turtle [Chelonia mydas] ) were frequently caught at
Tongareva no turtles were observed during the POBSP visit. Lamont
(1867) and Buck (1932) indicate that turtles were caught by the
natives for food.

Burt and Burt (1932) in their report on the herpetological
results of the Whitney Expedition stated that the collections of the
American Museum of Natural History contained one specimen of the
Mourning Gecko (Lepidodactylus lugubris, AMNH 41749) and one of the
Blue-tailed Skink (EHmoia cyanura, AMNH 41748). They also indicated
that E.H. Bryan, Jr., had collected these specimens while a guest of
the Whitney South Sea Expedition in 1925. In fact, these specimens
were collected by the Kaimaloa Expedition in 1924 (E.H. Bryan, Jr.,
in litt.) and mentioned by Gregory (1925). Ball (ms.) added that the
single example of Mourning Gecko seen was collected and that the
skinks were common but difficult to capture.

More recently the POBSP collected 28 specimens of four species of

PEZACdS. The more abundant species, the Mourning Gecko (14 specimens,
USNM 158325-338) and the Blue-tailed Skink: (10 specimens, USNM 158340-
349) were those reported by Burt and Burt. The other two species

collected were the Snake-eyed Skink (Cryptoblepharus boutoni, 1
specimen, USNM 158339) and the Polynesian Gecko (Gehyra oceanica, 3
specimens, USNM 158322-324). Both of the latter species were
widespread on well vegetated central Pacific atolls but neither has
been recorded previously from Tongareva.

Birds

Annotated List

In the following species accounts the presence of brackets about
the name of a bird indicates that the occurrence of the species on

Tongareva is not well documented and that its occurrence there should
be considered hypothetical

[White-tailed Tropicbird Phaethon lepturus |

Native informants indicated that this species is rarely seen at
Tongareva.

Red-tailed Tropicbird Phaethon rubricauda

About half a dozen birds were seen by the Bathams, but no
definite evidence of nesting was obtained by them. One bird seen
under a bush may have been in the pre-laying stage.

Brown Booby Sula leucogaster

The Bathams found a bird incubating two eggs; this is the only
record of nesting for this species on Tongareva. The only other
record of occurrence is that of a Brown Booby banded on Jarvis Island,
30 May 1940 and recovered at Tongareva on 15 February 1941 (Munro
1944).

Red-footed Booby Sula sula

These birds were seen and collected in 1936, 1937, and 1965.
All three of the POBSP specimens were collected from a group of ten
birds seen on the southwestern bird islets. No evidence of nesting
was found.

Two Red-footed Boobies banded elsewhere were recovered at
Tongareva. One was banded 9 August 1938 on Jarvis Island by George
C. Munro and recaptured at Tongareva on 28 February 1941 (Bryan, 1970).
Another banded as a nestling 24 June 1965 on Malden Island by the
POBSP, was recovered at Tongareva on 15 June 1966.

Great Frigatebird Fregata minor

This species, like the Red-footed Booby, was seen and collected
in 1936, 1937, and 1965 but was not recorded by the Bathams. There
is no evidence that the species breeds on Tongareva. Munro (1960)
reported a Great Frigatebird that had been banded on Enderbury Island
of the Phoenix Islands and recovered on Tongareva.

Lesser Frigatebird Fregata ariel

Large immature young were recorded by the Bathams during their
visit, and approximately 150 birds were seen on the southwestern
islets during the visit by the POBSP. The POBSP personnel found about
10 nests with eggs and 2 with nestlings in nests 7-8 feet up in 10 to’
12 foot high trees (Pemphis acidulus). Native informants indicated
that the species formerly nested in much greater numbers.

The Bathams stated that natives told them that band recoveries of
Fregata from Australia and the United States had been obtained at
Tongareva. It seems very unlikely that there have been any band
recoveries from Australia. The "recoveries from the United States"
presumably refers to the Great Frigatebird recovery mentioned above.

Pintail Anas acuta

An adult female banded 16 August 1949 at Tule Lake, California,
was shot 15 November 1949 at Tongareva. This is the only record of
this species from Tongareva.

Domestic Fowl Gallus gallus

Chickens were being raised by the natives when the island was.
visited in June 1965. Probably these birds were first introduced to
Tongareva in September 1853 (Ward, 1967; Lamont, 1867).

Golden Plover Pluvialis dominica

The POBSP saw six Golden Plovers along the west side of the
island. The Bathams reported that this species was widespread on the
atoll. They also stated that natives told them that this species and
the Ruddy Turnstone nest on Tongareva but did not comment on this
obviously erroneous information.

Bristle-thighed Curlew Numenius tahitiensis
The POBSP saw two of these curlews and collected one.
[Asiatic Whimbrel Numenius phaeopus variegatus]

This race of the Whimbrel, listed as Numenius variegatus by the
Bathams, was stated by them to occur in small parties of two to nine
on open areas of old bare coral. Although it is possible that this
western Pacific species was seen, it seems much more likely that this
record (and a similar record by the Bathams for Suvarov Atoll) were of
the Bristle-thighed Curlew, which could be expected to occur on these
atolls in some numbers.

Ruddy Turnstone Arenaria interpres

Turnstones have been recorded only by the Bathams, who found them
on every islet visited. © \
\
\

Wandering Tattler Heteroscelus incanus

Wandering Tattlers were first recorded by the Kaimaloa Expeditio
in 1924 when a few were seen (Gregory, 1925). Subsequently six
specimens were collected in April 1937 by Smith, and three were
recorded along the west side of the atoll during the POBSP visit in
June 1965.

[Black-naped Tern Sterna sumatrana |

Native informants indicated that this species is rarely seen at
Tongareva.

Sooty Tern Sterna fuscata

Three birds were seen offshore in December 1924 and about 40 were
seen flying over the island by the POBSP. The Bathams, received six
eggs which were attributed to this species, but further documentation
of the nesting of this species on Tongareva would be desirable.

Blue-gray Noddy Procelsterna cerulea
A nestling preserved in spirits at the American Museum of Natural
History is the only record of this species from Tongareva. The bird
was collected 26 September 1936 by Coultas.
Brown Noddy _Anous stolidus
Ball (ms.) first recorded these birds on the atoll in December

1924 when they were incubating eggs in nests in the coconut trees.
Brown Noddies were commonly seen when Tongareva was visited by the

Bathams. Nests with young and one with an egg were seen. Only two
large nestlings were found during the visit by the POBSP. Both were
in nests in Pandanus trees. In all, an estimated 300 birds were seen

during the June 1965 visit, about 100 along the southwestern islets
and the rest along the northwestern portion of the atoll.

Black Noddy Anous tenuirostris

These birds were the most abundant seabird on the atoll when it
was visited by the POBSP. An estimated 3,000 birds were seen on the
small islets at the southwestern corner of the lagoon. At this time
between 1,000 and 1,200 nests were present but birds were associated
with only about 150 and all these contained eggs. Nests were
primarily constructed of leaves of Pandanus and grass (Lepturus sp.)
and were located in Pandanus and Tournefortia trees. Later in the
season (of a different vear), the Bathams found both eggs and
nestlings, some in Pandanus and a few others in Pisonia grandis.

White Tern Gygis alba

These terns were fairly abundant when the island was visited by
the POBSP. Sixty-five counted among the islets at the southwestern
corner of the atoll, and another seven were counted at the northwestern
corner. No nests were found during the POBSP visit, but gonad data
from the four specimens collected suggest that the birds were breeding
in June 1965. Both eggs and young were found by the Bathams in
August-September 1968.

Populations evidently decrease considerably during the northern
hemisphere winter as only a few of these terns were seen by the
Kaimaloa Expedition in December 1924 (Gregory, 1925). During this
visit, Ball (ms.) considered the species uncommon.

[New Zealand Cuckoo Eudynamis taitensis |

The Bathams reported a bird that the natives called "Koekoea",
which was purportedly this species, but did not see one. The
occurrence of the New Zealand Cuckoo on this atoll should be watched
EOYs.

Summary

This paper briefly summarizes what is known of the vertebrate
fauna of Tongareva Atoll. Four species of mammals, dog, cat, pig,
and an unidentified rat (Rattus sp.), occur on the atoll as does an
unidentified turtle, probably Chelonia mydas. Four species of
lizards, all of wide distribution in the central Pacific, are present.
Two of these, the Snake-eyed Skink and Polynesian Gecko, are reported
from the atoll for the first time.

Excluding species of hypothetical occurrence 16 species of birds
are known from the atoll. Of these, the Brown Booby, Lesser
Frigatebird, Brown, Black, and Blue-gray Noddies and White Tern are
seabirds that breed or have bred on the atoll. Four other species
of seabirds, Red-tailed Tropicbird, Red-footed Booby, Great Frigate-
bird, and Sooty Tern, visit the island regularly. The Red-tailed
Tropicbird and Sooty Tern may breed on Tongareva.

The remaining six species include four migrant shorebirds, the
Golden Plover, Bristle-thighed Curlew, Ruddy Turnstone, and Wandering
Tattler, a vagrant duck (the Pintail), and the introduced Domestic
Fowl.

Acknowledgements
I am indebted to the members of the POBSP survey team, Fred C.

Sibley, Robert R. Fleet, Lawrence N. Huber, C. Robert Long, Dennis
L. Stadel, and Robert S. Standen, upon whose field work this report

is based. I also thank M. Ralph Browning, John S. Weske, Marshall
A. Howe, and Philip S. Humphrey who read and commented on the
manuscript. This is paper number 110 of the Smithsonian Pacific Ocean

Biological Survey Program.

References
Balla iSec. (ms. 1924) Field notes taken while on the Kaimaloa
Expedition . Original in the B.P. Bishop Museum, Honolulu.
Batham, J. and A. Batham. 1973. Field notes on birds observed on

the 'motus' of two atolls (Penrhyn and Suwarrow) in the northern
Cook Islands, 1968. Notornis 20: 97-101.

Bian ietin, OF. LITO. History of bird banding in) Hawaai 37 32):
Review of articles in the Elepaio. Elepaio 30(11): 100-103

Buck. Pi (1932. Ethnology of Tongareva. BrP YBTShoOp, Mus'ay Su.
O22 25).

Boece, Csr. and MoD Burnt.) 39382: Herpetological results of the
Whitney south sea expedition. W/E. Pacific island amphibians

and reptiles in the collections of the American Museum of
Natural History. Bull. Amer. Mus. Nat. Hist. 63: 461-597.

Gregory, H.E. 1925. Report of the Director for 1924. B.P. Bishop
MahUisNn, 1SYOHATL AES oye

Lamont, E.H. 1867. Wild life among the Pacific Islanders. London,
Hurst and Blackett.

Munro, G.C. 1944. Notes on migration and straggling birds.
Elepaio 4(6): 33-34.

Munro, G.C. 1960. Birds of Hawaii, revised ed. Rutland, Vermont,
Charles E. Tuttle Co.

Ward, R.G. (ed.) 1967. American activities in the central Pacific
1790-1870. vol. 3, Ridgewood, N.J., Gregg Press.

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ATOLL RESEARCH BULLETIN

NO. 199.

OBSERVATIONS ON VEGETATION OF BLUE-FACED
BOOBY COLONIES ON COSMOLEDO ATOLL,
WESTERN INDIAN OCEAN

by Mary E. Gillham

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

February 1977

OBSERVATIONS ON VEGETATION OF BLUE-FACED
BOOBY COLONIES ON COSMOLEDO ATOLL,
WESTERN INDIAN OCEAN

by

Mary E. Gillham!

Localities occupied by boobies on Wizard and Pagoda Islands

_ Cosmoledo Atoll lies approximately 960 km east of the Keny coast
in latitude 9°40' S. It consists of a circle of islands around a 8 km
wide lagoon and spans some 14.5 km by 11 km. Like Aldabra, which is
about 160 km nearer to the African mainland, it consists of an old,
uplifted coral reef, but it is less rugged, much of the rock surface
being hidden under drifts of coral sand. Wizard Island or Grande Ile
is one of the half dozen or so larger islands, second in size only to
Menai Island where the only settlement is situated. eS) Skin Long
and forms part of the eastern rim of the atoll, rising some 3.6-4.6 m
above sea level.

It was possible to visit only the southern part of this island
when the Manihine called there in February 1970. Much of this part is
scrub covered, so that the ground-nesting Blue-faced or Masked Boobies
(Sula dactylatra) are uncommon, but nests were scattered along the
straggling corridors of low heraceous vegetation between the shrubs.

Some of the coastal bushes were occupied by Red-footed Boobies
(Sula sula) and some of the inland ones by Lesser Egrets (Egretta
garzetta) and Cattle Egrets (Bubulcus ibis).

Pagoda Island is representative of the many smaller islets occuring
on the atoll rim. It is perhaps 0.4 km long and less wide, and, being
more exposed to the influence of the sea, supports few shrubs of any
size. Many of these lie below the general ground level around the
rugged margins of a saline pond sunk in the champignon behind the storm
beach on the ocean side, and having only subterranean connections with
the sea.

The falling tide leaves curtains of filamentous algae draped about
the lagoon walls. Vegetation crowning the storm beach is limited by
salty winds to low growths such as the tight wiry rosettes of the sedge,

Extra Mural Department, University College of South Wales, Cardiff
(Revised manuscript received September 1974 --Eds.)

Fimbristylis species, in contrast to the tall Cyperus ligularis and
shrubs of coasts facing in towards the atoll centre.

Bushes on Pagoda Island contained occupied nests of Red-footed
Boobies in mid-February 1970; the low vegetation of the remainder of
the island was sprinkled throughout with those of Blue-faced Boobies.

This islet is too exposed for mangroves and other trees of large
size and the Red-footed Boobies were nesting chiefly on, rather than
in, the fringe of low, windtrimmed Pemphis acidula bushes of the peri-
phery and the even lower growths around the inland lagoon.

On Wizard Island, where the range of nest sites is larger, they
were favouring the white mangrove (Avicennia marina), a species which
is used little if at all on Aldabra, where it gorws in abundance but
where Rhizophora mucronata, Bruguiera gymnorrhiza and Ceriops tagal
seem to be preferred. Other trees such as the scarlet-flowered Cordia
subcordata are also used on Wizard Island.

Other localities occupied in this sector of the Indian Ocean

The status of Blue-faced Boobies on the rest of Cosmoledo is not
completely known. Bourne (1966) records "A few seen on Cosmoledo and
Assumption", giving no details as to which islands. Diamond (in Bayne
et al 1970: 48) says "At least 200 pairs of White Booby S. dactylatra
were occupying clearings in the long grass on the west side of the
island (Wizard) or on the dune ridge to the east" in March 1968. Vesey-
Fitzgerald (1941) records them as breeding in September-November 1940 on
four of the Cosmoledo Islands (West North, East North, Grande Polyte and
South) but does not mention Wizard or Pagoda. Piggott (1961) saw large
numbers of boobies, species unstated, on Grand Polyte and also bird
colonies on Pagoda and South Island.

Aldabra in the same latitude and only approximately 760 km from
the African mainland, is an important sea bird resting site, but Blue-
faced Boobies are not among them. Up to 1966 (Benson, 1966) the only
evidence for their occurrence, even as non-breeders, is from Morris
(1963), who saw both adult and immature birds. Red-footed Boobies,
on the other hand, are abundant in the mangroves, many thousands nesting
among the frigate birds (Fregata minor and F. ariel) on Middle Island.
The absence of the ground-nesting species is surprising in view of the
open platin and grassed areas which would seem eminently suitable.

Assumption, 40 km to the south of Aldabra, supporting a few breed- %
ing Blue-faced Boobies in the late nineteenth century (Ridgeway, 1895;
Nicoll, 1906) but as a result of guano-digging activities it is unlikely
that. any still survive (Stoddart et al 1970).

On a wider scale Sula dactylatra has isolated breeding sites along
the coast of East Africa from the Red Sea to Tanzania, and on Socotra.
Generally these are sited on rocky coats and cliffs 6-9 m above sea
level (Mackworth-Praed and Grant, 1957). Those of Wizard Island and of

Latham Island (65 km off Dar-Es-Salaam and visited by the present author
in January, 1970), however, are on quite smooth terrain, uninterrupted
by cliffy out-crops and not more than about 3 m above sea level.

Early regeneration phase of vegetation: Wizard Island

The areas occupied by ground-nesting boobies on Wizard Island were
limited to patches of open ground seldom more than 20 X 10 m among the
overall cover of shrubs. These sites were scattered along the coasts,
often to the top of the low coral 'cliffs', but usually separated from
the reef by a belt of different vegetation. On and below the jumble
of honeycombed champignon rocks of the coastline were thickets of
brilliant-flowered Cordia subcordata and occasional large white man-
groves (Avicennia marina) with more spherical bushes of Xylocarpus
granatum loaded with big green 'cannon-ball' fruits. Where the shore
was Ssandier these were replaced by lower, denser bushes of white-
flowered Pemphis acidula and yellow-flowered Suriana maritima, both
self-tolerant species, branching copiously from the base and with
characteristically small, sclerophyllous leaves. In areas of mixed
rock and sand were silvery-leaved shrubs of Tournefortia argentia with
speckled orange and white moths massed on their large inflorescences.

Only a small proportion of the guano-impregnated soil of the booby
nesting areas was bare in February 1970. Most, Sometimes up to 90%.
was covered by a flattened, straw-coloured mat of dead grass, probably
Dactyloctenium aegyptium. On this the boobies laid their one or two
eggs (usually white, but occasionally pale blue), more sparsely than on
Latham Island at the same season. No nesting material was used, but
a bare circle of approximately 60 cm diameter was scraped in the dead
grass. This was not one of the peak laying periods and tentative plant
colonisation had begun between nests in areas which the presence of
guano and nest hollows showed to have been recently vacated.

The three pioneer plant species were annuals, growing radially
from their points of origin to form neatly circular clumps up to 1m
across but to occasionally near 3 m by 13 February 1970. One of them,
the succulent Portulaca oleracea, was also the principal species of the
blue-faced colony on Latham Island, where vegetation was generally much
scantier. The other two Wizard Island pioneers were less succulent
and quite prostrate; the minute-leaved, green-flowered creeping spurge
(Euphorbia prostrata) and the yellow-flowered, pinnate-leaved Tribulus
cistoides. Fruits of this usually possess 5 carpels, with 2 large,
upwardly directed prongs. Close after these 3 species came a creeping
ground cover of Boerhavia repens, with umbels of tiny white or pink
flowers protruding from a continuous carpet of dark, red-tinged leaves.
Superimposed in places were the stems of an unidentified cucurbit with
yellow flowers 2.5 cm across. The purple-flowered, green-podded herb,
Cleome strigosa was locally abundant at this stage, principally as
seedlings. Mature plants were more or less prostrate, in contrast to
an average height of 50 cm in parts of Aldabra unaffected by birds.

Other components of this bird flora were the orange flowered, Sida
parvifolia; the yellow-flowered Portulaca species; the yellow-flowered,
trailing Composite, Launaea sarmentosa; the mauve-flowered sea
Convolvulus (Ipomoea pes-caprae) and the grasses, Dactyloctenium
aegyptium, with radiating flower spike and Sporobolus virginicus, ‘with
narrow, erect flower spikes.

Peripheral to these, on less recently disturbed ground, were low
undershrubs of Achyranthes aspera with 30 cm long spikes of mauve |
flowers ripening from the bottom upwards and falling to leave reflexed |
bract scales. Seedlings of another small bush, Acalypha claoxyloides, /
sprouted between them, their soft, balsam-like leaves veined with yellow
among lax catkins of minute yellow-green flowers.

This low and easily negotiable community of undershrubs alternated
with the booby colonies, to form with them a coastal belt 7-10 m wide.
There is every probability that this belt represented areas occupied
at some time or other by boobies and showing an intermediate succes-
sional phase between a fully occupied herbaceous sward and an unoccupied
scrub. Both species, Achyranthes and Acalypha, grew to 2 m in the
main inland scrub, where, along with many other shrubs, such as the
small-flowered, round-leaved Pleurostelma cernuum of the milkweed
family (Asclepiadaceae), they became generously festooned with creepers.
Chief of the lianas was the white-flowered morning glory (Ipomoea
macrantha) and there was also a small, green-flowered passion fruit
(Passiflora suberosa) . The tall brown-headed sedge, (Cyperus
ligularis) occupied sandy hollows.

Later regeneration phase of vegetation: Pagoda Island

Blue-faced Boobies, as indicated, occupied almost the entire area
of Pagoda Island, but spacing of the nests was as wide as on Wizard
(although it was difficult to be certain of this because many of the
well-grown chicks were tucked under bushes and not visible unless
flushed). The ground was of coral gravel further whitened by guano
and with little sand. Not much was bare and there was little grass,
the area being divided between a prostrate herb flora dominated by
Boerhavia repens and a sapling shrub flora dominated by Achyranthes
aspera.

On the open area the Boerhavia carpet formed a ground cover of
about 60%; Euphorbia prostrate covered another 20%, whilst 10% was
bare. The remaining 10% was occupied by two forms of Tribulus, princi-
pally a robust one with larger flowers and fruits than any seen on o
Wizard. With it were Sida parvifolia, Portulaca oleracea, P. australis,
Sesuvium portulacastrum and a little Launaea sarmentosa.

Throughout this sward seedlings of Achyranthes aspera and Acalypha
claoxyloides were growing, especially the former. The Achyranthes
dominating the main part of the island obviously represented the same
community in a more advanced successional phase, the herbaceous sward
which persisted between the 30-100 cm high bushes diminishing until

these finally coalesced.

Apparently not involved in this 'booby succession' were the thick
stands of Plumbago aphylla backing the rocky beaches and land-locked
pond. Most of its grey-green stems were bare, but some patches
retained the ephemeral circular leaves produced during the rains
(December to March), and considerable stands on the ocean side bore
massed white flowers — the plants reminiscent of a less succulent ver-
sion of Sarcostemma viminalis. Cyperus was associated with the
Plumbago in places.

The greater state of advancement of the booby plant succession on
Pagoda is of particular interest when considered in relation to the
more retarded climatic succession of this smaller, more wind-swept and
spray-drenched island. Few bushes exceeded 1 m in height, there were
no mangroves and no Cordia trees around the fringes and little variety
o£ species in the main body of the island. Only one Tournefortia
argentia bush was seen, its main trunk prostrate, living branching to
only 60 cm and dead ones overtopping these by another 60 cm. Pemphis
acidula, the only abundant shrub around the edge of the Boerhavia-
Achyranthes succession, was generally gnarled and mis-shaped by the wind.

The greater growth of the post-booby vegetation could have been
the result of either a sparser total population of boobies on an annual
basis or an earlier start to nesting. Most chicks were 4-5 months
ahead of those on Wizard Island and it is probable that, with this low
bird density, vegetation grew up alongside them, affording useful cover
as their parents left them unattended for longer periods. This cover
may be of greater importance in giving protection from the sun's heat
than from predators, midday temperatures in February reaching to 110 F
on the Aldabra group of islands. (A comparable case is seen with the
lesser black-backed gulls (Larus fuscus) in Britain where eggs and
constantly brooded young chicks occur in open grassland, the bracken
fronds sprouting as the chicks grow, so that they have ample cover
during their vulnerable adolescence. )

The almost universal coverage of dead grass in the Wizard nesting
areas shows that these do not follow quite the same successional

sequence as the Pagoda ones. It seems that the grass invades before
the seedling bushes become established, although the earlier herb phase
of the two successions runs parallel. (Diamond's only reference to

booby vegetation in March is to 'long grass' (Stoddart et al 1970: 49),
but there was little sign of its regeneration in February 1970, so the
sequence may not be the same each year.)

Predation and aggressiveness

Predation of boobies in the air over Wizard was serious, hundreds
of Greater and Lesser Frigate-birds harrying them in flight. It was
not possible to judge whether frigates selected their victims only from
among birds returning with newly caught fish, but many boobies were

seen to regurgitate at the end of a noisy, squawking aerial chase.
The food was invariably caught by one of the pursuing frigates as it

fell. As many as 5 or 6 of these would join forces against a single
booby in a spectacular pursuit from which the quarry's only escape was
to alight, either on land or sea. (Frigates are clumsy on land and

get waterlogged at sea, but are past masters of flight.)

More serious predation on Wizard was by cats. The booby colonies
were thickly littered with cat droppings consisting almost entirely of
booby feathers and down with, sometimes, a little fibrious plant
material. A cat was seen, not resembling the expected feral animal of
domestic origin, but larger, spotted and with a long tail, although how
a wild cat could arrive on an uninhabited coral atoll remains a mystery.

Many adult boobies were on empty nests and, of the remainder, most
had eggs. Only a few chicks were seen, none very large. Chicks would
be very vulnerable to cats, and it is suggested that the regarded state
of breeding as compared with the adjacent Pagoda Island, where no cat
dung was seen, may be due to replacement-laying by robbed parents.

It could be significant that surviving chicks had particularly aggres-
Sive parents, more likely to hold their ground in defence of young.
(There was no indication on Latham Island that birds with chicks were
more aggressive than birds with empty nests or eggs. This seemed to

be an attribute of some individuals only, and on Wizard Island an attri-
bute aiding race survival.) Wizard Island birds with no chicks tended
to take to the air when approached to within 6 m. Those with chicks
stood their ground, lunging at the intruder when handled. While chicks
were very young the parent manages to maintain their positioning — one
on top of each of the grey webbed feet — while swivelling round to face
the disturbance and jab at outstretched hands, quite unintimidated.

Diamond (in Bayne et al., 1970) suggests that cats may be respon-
sible for the partial or complete loss of terns from Wizard Island.
He found only skeletons and feathers in long grass and on dune ridges.
They clearly suffer heavy mortality, whether from predation, starvation
or disease is not known; but the most likely culprits would seem to be
the cats, of which 2 were seen and one shot (A. W. Diamond, personal
communication). Baker (1963) refers to a tern breeding area at the
north of Wizard Island, but none were seen in March 1968 nor February
OVO.

Pagoda Island was occupied by advanced booby chicks progressing
from the downy to the feathered stage and falling about awkwardly as

they exercised their wings. Few were less than half grown and only
3 eggs were seen. These older chicks were left for longer periods
unattended and were exceedingly noisy and aggressive. They started

squawking when the intruder was still a long way off and when he passed
within 2 m he was likely to receive a jab in the leg sufficient to
draw blood.

As far as could be ascertained, there was no predation of boobies
on Pagoda apart from food thieving by frigates, and this was less pre-
valent than on the larger island, only 40-50 frigates being seen.

Landings by man on so small an island surrounded by reef must be
few.

Continuity of bird pressure on vegetation

The most depauperate, bird-suppressed vegetation seen by me in this
part of the Indian Ocean in early 1970 was on Latham Island. Most of
this area was currently occupied by breeding sea-birds in late January
and it is likely that there is a fair continuity of bird pressure
there, because all former records for terns and noddies have been from
June-September, and for boobies from October-November and again in
March (Mackworth-Praed and Grant, 1957). This, in conjunction with
the prevalent salt-laden winds, precludes bush growth and limits the
flowering plants to three low-growing herb species: Portulaca oleracea,
P. australis and Ipomoea pes-caprae.

Local curtailment of bush growth on Wizard Island may be due to a
similar sequence of nesting by different individuals on the same site.
Even if only boobies are involved, it has been established that these
have no regular annual cycle, in either the Indian Ocean or the
Atlantic.

With an incubation period of 42-46 days (6% weeks) and a fledging
period of 120 days (17 weeks), it is likely that the oldest, still down
fledglings on Wizard in February 1970 came from eggs laid in October,
and the predominantly older, half-feathered fledglings on Pagoda
Island from eggs laid in September. Eggs and younger chicks, which
were almost exclusive to Wizard, could have been laid at any time bet-
ween November and February.

The possibility of occupation of the same sites by terns or noddies

at other seasons cannot be excluded. Both have been recorded breeding
on Wizard, and the predominance of dense, creeper-covered bushes limits
the amount of suitably open terrain. All the sites seen without bushes

were occupied to a greater or lesser degree by boobies, but the whiten-
ing by guano on some of these was more than could be attributed to the
present occupants alone. Only future visits to the atoll can estab-
lish such joint use of the open spaces.

Other breeding seabirds observed in the part of Wizard Island
visited in February 1970 were Red-footed Boobies (Sula sula), scattered
in small parties amongst the taller coastal bushes, with eggs and chicks
at all stages, and considerable flocks of free-flying brown immatures.
Several hundred Cattle Egrets (Bubulcus ibis) and Lesser Egrets
(Egretta garzetta) (mostly of the white phase, with no pied intermedia-
tes such as occur on Aldabra) thronged in larger trees further inland,
but were not examined to see if they were nesting.

Other Lesser Egrets fed on the reef with Grey Herons Ardeola
cinerea, Crested Terns Thalasseus bergii, Crab Plovers Dromas ardeola,
Great Sand Plovers Charadrius leschenaultii, Curlew Sandpiper Erolia
testacea, Sanderling Crocethia alba, Whimbrel Numenius phaeopus and

8

Turnstones Arenaria interpres. Of the two species of frigate bird,
the Greater were present in larger numbers.

Very abundant land birds were the Souimanga Sunbirds Nectarinia
sovimanga, taking nectar particularly from the big scarlet flowers of
Cordia subcordata, and the constantly chattering, streaky Madagascar
grass warblers Cisticola cherina, making short, lark-like. flights: over
the bushes and occasionally over the sea. These warblers were present
on Pagoda Island, but not the sunbirds, which would have failed to find
Cordia and the other flowering trees which they favoured in this more
exposed habitat. There were probably 50-100 pairs of Red-footed
Boobies nesting on Pagoda and some 20 Red-tailed Tropic birds (Phaethon
rubicauda) indulging in what appeared to be complicated courtship

flights over~the island. One bird on a nest beneath a dense Pemphis
acidula bush screeched and spread wings and body feathers menacingly
when approached, yielding no ground. Two others had nests under

thickets of Plumbago aphylla.

Several flocks of 20-30 Black-naped Terns Sterna sumatrana had
taken up positions on the rocky shore to which they returned soon after
flushing, as though they might have chicks concealed beneath the rocks,
but no young or eggs were seen. A single flock of about 30 crested
terns fed on one of the sandy beaches. The air was full of terns
above the next island of the atoll, which was almost certainly a breed-
ing area, but was unfortunately not visited. Other shore birds were
Turnstones, some of these leaving the pebbly beaches to forage among
the low Boerhavia and Euphorbia of the booby areas. These and various
other waders also fed around the enclosed tidal pool behind the ocean
storm beach.

The deep hollow in which the pool lay was partly filled with
bushes thronged with Red-footed Boobies. Other slopes carried
Sesuvium portulacastrum near water level (a characteristic halophyte of
mangrove swamps elsewhere), leading up through stands of either Plumbago
or Cyperus to the short Boerhavia turf of the Blue-faced Booby colony.

Little Green Herons Butorides striatus fed from shoreward rocks
and Grey Herons perched in statuesque positions on the tops of bushes,
waiting to move down to the reef to feed. The greater and lesser
frigates never alighted and were presumably from the breeding colonies
in Aldabra, which supply most of the Western Indian Ocean frigates.

The ecological niche normally occupied by scavenging gulls, crows
or kites, was filled by crabs, both land crabs and hermit crabs, hordes
of which were busy devouring regurgitated fish and the occasional dead
bird: The land crabs lived in burrows as much as 20 cm in diamter in
hollows of the champignon.

Postulated vegetation sequence in sea-bird colonies

Even on so small an island as Pagoda, the climatic climax vegeta-
tion consists of bushes sufficiently large to satisfy the breeding needs

of Red-footed Boobies. Close nesting by these causes the dying back of
the branches and eventually the entire shrubs. Heavy pressure by Red-
footed Boobies, therefore, could destroy the scrub and leave the way
open for herb and sapling woody species — i.e. the sort of environment
favoured by Blue-faced Boobies.

The protracted nature of the nesting season on Cosmoledo enables
degenerating and regenerating plant successions to exist side by side,
so that the vegetation fluctuates between a dead or living herb cover

and a sapling cover up to O.5 m high. With the low nesting densities
obtaining in Feburary 1970, this state is likely to be maintained in
the booby colonies. With increasing booby pressure, as on Latham

Island, bareness could previal, to give the sort of habitat favoured
(and induced or maintained) by terns and noddies.

These two smaller species nest sufficiently close together to
inhibit all plant growth in their chosen area, but they do not neces-
sarily return to the same site in successive years. If they move away
the vegetation will return — first the herbs, then the saplings and
finally a full scrub cover, if given long enough without further distur-
bance. This change, however, makes the terrain suitable once again for
the booby species, which may expand into it from nearby colonies.
Colonially nesting sea-birds inevitably modify the vegetation amongst
which they nest, and this may be to their own disadvantage and the
advantage of others.

The vegetative degeneration: regeneration cycle associated with
nesting birds might thus be as follows:

Degeneration:
Mangroves and other trees . .. Red-footed Boobies and egrets
Pemphis and other shrubs .. . Red-footed Boobies and Tropic birds
Boerhavia and other herbs . . . Blue-faced Boobies and Tropic birds
Bare Soil . . » Terns and noddies.

Regeneration:
Herbs and grasses . . . Terns,noddies and Blue-faced Boobies
Shrubs - . « Red-footed Boobies
Trees . . . Red-footed Boobies,egrets,heron,etc.

Comparison between Cosmoledo and Aldabra Atolls

Colonies of ground-nesting sea-birds are more extensive on the
Cosmoledo Atoll than on Aldabra, where the main colonial nesters are
frigate birds and tree-nesting boobies in the mangrove fringes. The
tree nesters defaecate into the intertidal zone, and so have little or
no effect on the land flora.

The only sea-birds on Aldabra which are sufficiently numerous to
affect the terrestrial ecosystem are the noddies of the tiny mushroom

10

islets in the lagoon, and many of the islets are so small and low as to
be devoid of macroscopic vegetation. Crested, Caspian, and Black-
naped and White Terns, Red-tailed and White-tailed Tropic birds, Grey
Herons, Little Egrets and\.Sacred Ibises, if not nesting so diffusely as
to have very localised effects on plants, are tree nesters.

The situation is very different with Cosmoledo's most important
colonial nesters, the Blue-faced Boobies. This species has an immed-
iate and obvious effect on the vegetation in which they nest and have
changed the face of the islands considerably. It does not nest on
Aldabra.

Thus, although Aldabra supports an unparalleled example of rela-
tively undisturbed native scrub, Cosmoledo shows in this respect a much
more interesting and complex interaction between flora and fauna.

Cosmoledo's vegetation is 'spoiled' if looked at in purist terms,
but it is spoiled mainly by natural causes. The fact that the plants
which have taken over in the wake of the birds are also plants which
take over after distrubance by man, detracts nothing from the natural-
ness of their origin. ‘

Modification of natural vegetation by large colonies of nesting
sea-birds cannot fail to be of interest biologically. Intense modi-
fication usually implies the accumulation of guano in sufficient
quantities to be a commercially viable proposition to mining companies,
so that the natural outcome of the association is destroyed. So long
as any of the birds remain, however, they have an inevitable impact on
the existing vegetation, and examples of such bird-dominated vegetation
should be jealously guarded.

Colonial sea-birds in large numbers are all too rare in the
Tropics: In the interests of scientific investigation, it is very
desirable that these semi-exploited islands should be conserved while
birds are still in residence and governing the competitive efficiency
of tolerant plants species over intolerant ones.

The rim of the Aldabra Atoll affords an admirable control area to
show uplifted reef vegetation in the absence of colonial birds, but
its reef islets are scarcely adequate to show reef vegetation in their
presence. Cosmoledo shows this on a big scale, and the double inter-
est, botanical and ornithological, cannot but be a useful adjunct to
our knowledge of the botanical alone.

Tropical sea-birds are so much rarer than temperate and arctic
ones that there would seem to be a very strong case for conferring
reserve status on selected islands of the Cosmoledo Group. These
would be ideal sites for instituting long term investigations on the
delicately fluctuating balance between changing numbers of sea-birds
and vegetation types having different levels of tolerance of trampling
and guano deposition.

Teal

Acknowledgements
I am much indebted to Mr. Basil Bell, formerly Director, of E. A.M.
F. R. O., and Mr. Mike Williams, Captain of the 'Manihine', for sea
passage to Cosmoledo, and to Dr. J. B. Gillett, Dr. Brian Harris and
Mes Jack Frazier for the identification of plants.

Appendix

List of plants in and beside Blue-faced Booby colonies on Cosmoledo

Acalypha claoxyloides
Achyranthes aspera

Avicennia marina
Azima tetracantha
Boerhavia repens (?)
Cassytha filiformis
Cleome strigosa
Cordia subcordata
(Curcurbitaceae)
Euphorbia prostrata
Ipomoea pes-caprae
I. macrantha
Lagrezia sarmentosa
Launaea sarmentosa
Passiflora suberosa
- Pemphis acidula
Pleurostelma cernuum

Plumbago aphylla
Portulaca oleracea

P. australis

Scaevola taccada
Sesuvium portulacastrum
Sida parvifolia

Suriana maritima
Tournefortia argentea
Tribulus cistoides
Xylocarpus granatum
Dactyloctenium aegyptium
Eragrostis eiparia(?)
Fimbristylis cymosa(?)
Cyperus ligularis
Sporobolus virginicus

References

Bovine Cnmeicn mn COGAM, Sc thHiei,. Dlanoncd, As We, EGazten, Wis 7, Grubb), P). 7,

Unies OM weANenrp i LOOGE) le Ma) Hira Dl,

Stoddart, Dawke vance laydora mien Dis

Atoll Res.

US 7O)~ Georgraphy and ecology of Cosmoledo Atoll.
Bull. ISGs SAIS) -

Observations on islands in the Indian Ocean.
44-43.

Bourne We Ro Pe 1966.
Sea Swallow, 18:

DOmRWaTG Dm Ee HOG 2.
brown booby (Sula spp.) at Ascension.

Comparative biology of the white booby and the
Hone, MOI, A422.
Mackwomeh—Pracd, Cs W., and Grant, €. Ho B. L957. Birds of East and
North East Africa. 1: Longmans.

PuiCieionrcie, Co ‘we Meike A report on a visit to the outer islands between
October and November ]960. Directorate of overseas surveys, Land
Resources Div. Typescript 7s Chap. 6) \Cosmoilledel- DYU=IO

S,
\
ff y
‘ Bei

ATOLL RESEARCH BULLETIN
NO. 200.

VEGETATION OF SEA AND SHORE-BIRD COLONIES
ON ALDABRA ATOLL

by Mary E. Gillham

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

February 1977

VEGETATION OF SEA AND SHORE-BIRD COLONIES
ON ALDABRA ATOLL

by

Mary E. Gillham!

Introduction

The present study was carried out during the early months of 1970
on Aldabra Atoll 640 km from the East African coast and 420 km north-
west of Madagascar in lat. 9°24' S, long. 46°20' E.

Ground nesting sea-birds are found almost exclusively on the
smaller islets in the 34 km long lagoon - easily accessible to man (who
seems scarcely to have bothered them) but not to his predatory followers,
the feral cats and rats. On the main island fringing the lagoon the
birds, principally frigate birds and boobies, are tree nesters in the
mangrove fringes, to which overland access is difficult.

Being mostly colonial nesters, sea-birds can have a profound
effect on the vegetation amongst which they live, particularly in hot
dry climates where corrosive guano remains largely undiluted. Compared
with other sites investigated in three continents, both temperate and
tropical and even as close as Cosmoledo Atoll, about 160 km to the east,
these interactions between birds and plants on Aldabra are usually
slight. Noddies (Anous stolidus) are the only birds which can be said
to initiate any profound changes in the vegetation cover. The negli-
gible influence of the rest is thought to be principally due to the
dissipation of the guano, which either falls directly into the lagoon
water or seeps away through the honeycomb structure of the champignon
er uplifted reef rock.

On the neighbouring islands where guano accumulated (and no doubt
had an important bearing on the vegetation type in so doing) this has
been mined, destroying almost all the original plant life so that these
effects cannot be observed. The very fact which saved the flora of
Aldabra from similar destruction at the hands of man thus prevents a
full investigation of plant/sea-bird relationships in this unique type
of environment.

Department of Extra Mural Studies, University College, Cardiff
(Revised manuscript received September 1974 --Eds.)

Much the greatest effect of colonially nesting birds on plants is
chemical, except perhaps in cool moist climates where precipitation
greatly exceeds evaporation. Even there the chemical effect may be
considerable, though beneficial rather than deleterious, the diluted
excrement acting as a fertiliser (phosphatic and nitrogenous) instead
of a too powerful toxin and plasmolysing agent which causes root hairs
to shrivel and leaf tissues to 'scorch'.

Another type of chemical inhibition of plants by birds has been
reported on certain islands further north towards the Seychelles Group
where sooty terns (Sterna fuscata) are said to hover over the massed
Madagascar periwinkles (Catharanthus roseus) and sprinkle them with salt
water carried on their wings, so that they die (Beamish, 1970a). That
this is a motivated activity is highly unlikely, the periwinkle having
a normal growth height of 30 - 60 cm, with large soft leaves and flowers
of a morphological form highly unsuited to survive even light bird
pressure ‘and certainly not the activities of a thriving collony of terns,
whose nests are seldom more than 30 - 40 cm apart. Such bird normally
hover over a nest site for long periods before.finally settling in for
laying.

Tree nésters
White Terns (Gygis alba)

These normally balance their single egg on the branch of a man-
grove — Rhizophora mucronata, Bruguiera gymnorrhiza, Ceriops tagal or
Avicennia marina — among undamaged foliage and are much too sparse on
Aldabra to have any noticeable effect. Adults perch on dead Pemphis
acidula twigs too fragile for egg supports, but this is no indication
that they have killed the twigs. It would be interesting to know
whether the much larger numbers on Cousin Island in the Seychelles,
where fairy terns are said to nest commonly in coconut palms and
Casuarina trees (Penny, 1970), have any local deleterious influence on
these.

Frigate birds (Fregata minor and Fregata ariel)

Although nesting in bushes in some parts of their range, frigate
bird occupy taller trees on Aldabra, Lesser Frigates often in the lower
branches, Great Frigates (with a 7 ft. wingspan) higher up. Usually it
is the various types of mangroves which are used for nesting, but others
are occasionally used for perching, e.g. Casuarina equisetifolia and
Cordia subcordata. Nests are fairly well scattered among the mangrove
foliage, whose glossy leaves take little harm from coatings of white
metabolite. They usually overhang the water of the lagoon. Few
species of ground plant occur beneath nests and these but rarely.

Most, like Plumbago aphylla, have few or no leaves to take harm.

Frigate birds tend often to roost communally and here their
influence seems more potent. Trees used extensively are mostly dead,
with the lesser twigs missing, possibly removed for nesting material.

Because these trees are generally in strategic sites, at channel junc-
tions, on mid creek islands or the ends of promontaries, it is likely
that they are chosen for their position rather than that they have
already been killed by other means, and that their death is secondary
and due to the birds. If so, this bears a close parallel with the many
trees killed by cormorants (Phalacrocoridae), particularly in the
Southern Hemisphere where this group is most numerous. Further evid-
nce of tree death as effect rather than cause of perching is evinced in
local death of utilised branch tips. Upstanding 'elbows' frequented
by birds suffer defoliation while leaves continue to persist on their
pendant tips, proving that the leafless portions are still sufficiently
alive to convey water and solutes.

Sacred Ibis (Threskiornis aethiopicus)

This species was not observed nesting during the February to May
period spent on Aldabra, but Beamish (1970b) refers to "a series of
untidy nests making an almost continuous platform 12 ft. above the
ground" is Takamaka grove, an isolated woodland in the SE.

Tree and bush nesters
Red-footed Boobies (Sula sula)

What has been said of nesting frigate birds applies equally to
these arboreal boobies which build in leafy mangroves. Dead trees for
perching are, however, the exception rather than the rule, possibly
because the boobies usually fly out to sea for their fish eating expedi-
tions and do not need commanding lookout posts from which to spot likely
quarry to be chased and robbed as the frigates do.

Seldom on Aldabra, where big trees are abundant, do Red-footed
Boobies nest in bushes, as they do on Pagoda Island, Cosmoledo Atoll,
in the absence of tall trees. On Wizard Island, Cosmoledo, both trees
and bushes are utilised, the most favoured tree Avicennia marina and the
most favoured bush Pemphis acidula (Avicennia is not a common tree in
the Aldabra colonies and Diamond (1971) states that boobies do not use
it for nesting).

Pemphis is usually set slightly further back from the shore than
are the various mangroves so that excrement falls onto the ground rather
than onto water or intertidal stretches of sand or reef. The density
of the Pemphis bushes, in spite of local killing around the nests, is
often too great, however, for any understorey of vegetation to grow
beneath them. Falling guano can affect only the Pemphis. Tea Cier meat
is probable that little does.

If bushes are sited on champignon, it is likely that roots pene-
trate deep into crevices; if they are rooted in dry sand or chippings
of reef rock, it is likely that the non-liquid guano is held up on the
surface. Rain, when it falls, is usually torrential, so would wash
this off the surface onto the beach or dilute it to a greater extent

than do the gentler rains of less tropical regions. But no doubt some
of the nutriant salts do move downwards past the Pemphis roots during
rain and rise past them again by capillarity during evaporation of
ground moisture in dry spells — particularly in the sandier substrata.

Apart from local killing and twig plucking, it is not easy to
ascribe any deleterious or beneficial changes to booby activity.
Ground-nesting Blue-faced or Masked boobies (Sula dactylatra) which
have such a profound influence on the vegetation of parts of the
Cosmoledo Group, do not nest on Aldabra, in spite of suitable sites
devoid of scrub on platin rock surfaces and tortoise-grazed turf.

Tree, bush and ground nesters
Grey Herons (Ardea cinerea)
Like the British subspecies of grey heron, which nests as readily
on the bare rock of Scottish island cliffs as on the tall trees of

English woodlands, the Aldabran herons-*will accept almost any type of
breeding site.

Some build in trees, some in bushes and some on the ground.
Nests are widely spaced and, although a dozen or more pairs may breed
simultaneously on a small islet of some 20 X 40 m, they seem to be less
colonial on Aldabra than in much of their range — possibly because of
the long drawn out breeding season — all stages of building, eggs,
chicks and fully feathered juveniles being observed during the February
to May period 1970, with no synchronisation of laying as with the
frigates and, to a lesser extent, the boobies.

The fact that most of the treenesting birds seen were in mangroves
of one sort or another, reflects only the fact that mangroves are the
commonest trees in the areas visited. The same can be said of the
Pemphis acidula favoured by the bush breeders. Nests are in, rather
than on these, being normally too bulkly to be supported by the upper-
most twigs. Such sites provide cover for the gawky youngsters as well
as opportunity for acrobatics.

Fairly obviously as a result.of the heron activity many of the
Pemphis bushes are dead, some collapsed into a heap of branches so that
the nests are virtually at ground level and the material of which they
are composed scarcely distinguishable from that of the supporting
plants. Such nests are reminiscent of those of ospreys (Pandion
haliaetus) on Nitraria schoberi bushes in Western Australia and those
of crowned cormorants (Microcarbo coronata) on a variety of bushes on
South African islands where, the host shrubs killed, both ospreys and
cormorants continue to build up the level with annual increments of
twigs to create artificial bushes for themselves (Gillham, 1961, 1963b).
Material added to the Aldabra herons' nests in successive years was not
seen to reach such substantial heights as these.

Around the herons' nests is a fairly thriving growth of Achyranthes
aspera, occupying on average some 45% of the area and reaching to 120 cm
high, evidently stimulated by the diffusing guano. Tufts of the
following herbs occur in clearings: Portulaca oleracea (20%);
Sclerodactylon macrostachyum (15%); Cleome strigosa (4%); Corchorus
aestuans (4%); Boerhavis elegans (1%); Dactyloctenium aegyptium (1%);
with 10% of the rock surface bare.

The Pemphis generally and the occasional Ceriops tagal mangrove
are in poor shape — the whole illustrating a degenerative phase of a
vegetative cover reverting from shrubs through semi-shrubs and herbs to
bare rock.

Nests built deliberately on the ground are seldom in the open but
placed among undershrubs 1/2 to 1 m high — principally Acalypha
claoxyloides but also Achyranthes. These two 'weed' species are
typically associated with biotic disturbance on Aldabra, Acalypha with
disturbance by man and Achyranthes with disturbance by noddies.

Dead stumps of Pemphis suggest that herons in this type of habitat
have continued to return to an acestral site after it has passed from
its optimum of a former bush cover — possibly even a former tree cover,
although no tree remains were found.

The same herbs are associated with these nests except where the
Acalypha-Achyranthes thicket excludes the necessary light. Others
with them are Portulaca australis and Phyllanthus amarus.

Other Ardeiformes

The only other common members of the heron group are Little Egrets
(Egretta garzetta) and these favour the same types of site as grey
herons, with more emphasis on the trees and bushes. Little Green
Herons (Butorides striatus) sometimes occur in surprisingly large
colonies in the mangroves.

Sgquacco Herons (Ardeola ralloides) and Cattle Egrets (Bubulcus
ibis) are still present in numbers too small to have any significant
effect on the vegetation.

Undergrowth nesters
Red-tailed Tropicbirds (Phaethon rubricauda)

Red-tailed Tropic birds or bosun birds nest on the ground under
varying amount of plant cover. Usually the nest is invisible until
the vegetation is parted but they are not hard to find because of the
raucous screeching and lunging with open beak which starts when the
intruder approaches within a few metres. It is unlikely that any land
predator, even feral cats, would stand up to birds so aggressive but,
in fact, both species of tropic bird were found only on the lagoon

islands where such predators are sparse or absent. They are reported,
too, on Polymnie Island, smallest piece of the ellipse which forms the
acolls irsemy:

Pairs may be solitary or in widely spaced colonies of up to about
20 per islet, but seldom closer than 2 m to. neighbouring pairs, so
having only local effects on the plants. Types of site favoured are
as follows:

(a) under bushes

Bushes utilised are usually wind-trimmed to produce a dense
meshwork of branching so that birds can be difficult to reach.
Pemphis acidula, as the commonest bush of the islets, is most used.
Other are Suriana maritima, Desmanthus commersonianus, Sideroxylon
inerme, Ficus reflexa and Lumnitzera racemosa. Birds favour
rocky sites rather than sandy ones and were not recorded under the
three common shrubs of sandy coasts, Tournefourtia argentea, Scaev-
ola taccada and Guettarda species.

(b) among undergrowth under bushes

Birds next under more depauperate specimens of the above
bushes among a thick understorey of semi-shrubs, principally
Acalypha claoxyloides with rather less Achyranthes aspera, the
usually leafless (but flowering) Plumbago aphylla and wire grass
(Sclerodactylon macrostachyum) with the softer grass Dactylocten-
ium aegyptium and a little Eragrostis sp., Portulaca oleracea and
P. australis marginally.

(c) among semi-shrubs

Other birds are in the above understorey species with no shrub
cover, although sometimes with the dead remains of unidentified
shrubs protruding from the low canopy, so this is evidently a
degenerated vegetation phase.

(d) in crevices overarched by plants

Parallel-sided holes in the champignon, overarched by varying
amounts of herbs, grasses and undershrubs are used, the rock forma-
tion supplying the greater cover. Often these holes take the form
of vertical chimneys more than half a metre deep and the diameter
of the oird's head and body length, so that the long tail arches
over the back, its tip beside ‘the beak.

White-tailed Tropicbirds (Phaethon lepturus)
This slightly smaller species of tropic bird lays its purple-

blotched eggs in sites similar to the above and also in more open sites —
although rather less aggressive and therefore apparently more in need of

protection. As both species produce eggs and chicks during the same
period, the smaller birds may be excluded from better sites by the
larger. Nesting occurs throughout the year, with no peak (Diamond,
ROMA Often the two species are mixed in the same colonies, but some

islets seem to be devoted exclusively to one, as the Ile Sylvestre near
Esprit in the Western part of the lagoon where some twenty pairs of

White-tailed Tropic birds were located on 18th April, 1970. Even here
there was no synchronisation of laying eggs, small chicks and fully
grown ones with barred juvenile plumage being present simultaneously.
Two further types of site and types (a) and (d) above are used on this
island.

(e) crevices with no sheltering plants
Chimneys and crevices in the champignon with no overarching
vegetation contain incubating birds fully visible from outside.

(£) open sand

Level sandy sites are usually near the foot of rock walls.
In the rainy season (which is also the observed nesting season)
these support a sparse growth of grass and herbs which has no
concealing function. Commonest is the mauve-flowered member of
the Caper family, Cleome strigosa (which, like Plumbago aphylla
and Acalypha claoxyloides, is a group-endemic found only on the
islands of Aldabra, Assumption, Astove, Cosmoledo, St. Pierre and
Gloriosa in this corner of the Indian Ocean). Two soft-leaved
grasses, particularly Daknopholis boivinii, but also the more
generally distributed Dactyloctenium aegyptium, also grow. A few
Acalypha seedling spring up beside nests in places or, more
correctly nest hollows, no building material being used.

Part of this islet is heavily bushed but there is no evidence
to suggest that the more open sites occupied by birds might once
have been so covered.

Ground nesters
White-capped noddies (Anous stolidus)

Noddies (or noddy terns) are much the commonest species of ground
nesting birds. Chosen sites range from bare rock to a low cover of
semi-shrubs and larger shrubs (up to 3 m high) are used as perches.

In the absence of ground nesting Blue-faced Boobies, they undoubtedly
affect the plant life more than any other birds.

They occur almost exclusively on the smaller undercut 'mushroom
islets' of perforated champignon rock in the lagoon. (Derivation of
the name of 'champignon'for the predominant razor-edged reef rock is
from the eroded mushroom shape of these islets, some of them no bigger
than a large table).

Mangrove and Pemphis-dotted headlands of the larger encircling
islands of the atoll rim are used for perching rather than nesting.
The requisite condition appears to be unrestricted access to open water-
where the noddies fish from the surface, like gulls, instead of diving
for food like the more closely related terns. Even on an island so
small as Latham, forty miles off Dar-Es Salaam on the East African
coast, noddies occupy peripheral areas, leaving more central ones to
sooty terns and blue-faced boobies. Aldabra habitats fall into the

following categories:

(a) on top of low, truncated undershrubs

The fact that noddies, unlike most species of tern, build quite
substantial nests of twigs, mangrove leaves, sea grasses (principally
the common Thalassodendron ciliatum), algae and quill feathers when such
material is available, enables them to nest above ground level. This
is something which is achieved by the tree-nesting fairy terns only at
the expense of high mortality of eggs and chicks, Mary Penny having
recorded only 3 successfully hatched young from 150 pairs studied on
Cousin Island in the Seychelles (Penny, 1970).

It is improbably that mangrove leaves, one of the chief building
materials, are plucked from the trees, as only dry, ginger—-brown ones
are seen in nests and these fall naturally, to float up against the
nesting islets in quantity.

Nests are seldom more than 1/2 m above ground level on Aldabra,
although White-capped Noddies on Heron Island, at the southern end of
Australia's Great Barrier Reef, nest in the lower branches of Pisonia
grandis trees, 3 m or more high, building their nests largely of Pisonia
leaves (Gillham, 1963a).

Achyranthes aspera is the only undershrub seen to support nests on
Aldabra, the plants chosen being dense and flat-topped, due either to
wind-trimming, salt scorch or bird disturbance (including the plucking
of upstanding twigs for nest material). Acalypha claoxyloides, second
commonest plant of this conformation on the islets appears not to he
utilised, due, no doubt, to the laxer mode of branching providing
inadequate support. A parallel can be drawn with the undershrubs used
by nesting crowned cormorants on islands off South Africa where dense
Helichrysum ericaefolium and Lycium ferocissimum bushes are used but
not the post-climax phase of laxly branched Solanum guineense and
Asclepias pubescens which replaces these when the bird pressure has
destroyed them (Gillham, 1963b).

Relatively few nests among the thousands observed on the Aldabra
islets were thus sited, perhaps a dozen in all — the remainder being at
ground level. One island of this type, recently vacated by noddies
shows the following ground cover:

Achyranthes aspera 60% Sesuvium portulacastrum 1%
Dactyloctenium aegyptium 30% Sida fs ates 1%
Boerhavia elegans 2% Corchorus aestuans 1%
Portulaca oleracea 2% Phyllanthus ?amarus 1%

Portulaca australis 1% Bare rock 13

(b) beneath low undershrubs

Although essentially birds of the open, noddies, like the bridled
terns (Sterna anaethaetus) of Western Australian islands (Gillham, 1961),
will occasionally build beneath the dense Achyranthes referred to above.
Seldom are nests found more than 1/2 m in from the edge of the thickets
and the entrance tunnel is always sufficiently wide for the bird to
leave hurriedly with wings spread. Better camouflaged nests would be
more difficult to observe and may occur, but no birds are seen rising
from dense stands of plants.

Type (b) nests are usually on the same islets as type (a), such
sites being less favoured on the whole than sites with sparser vege-
tation. A few nests are under Acalypha plants and all are on fairly
level rock, not in deep pockets of the champignon such as are used by
the closer-sitting tropic birds and form cul-de-sacs from which rapid
escape might be hindered. Noddies are not, however, easily disturbed
by intrusion, often allowing approach to within 1 m of the next before
rising, and 'dive-bombing' the intruder vigorously when young chicks
are present, although less belligerant during the incubation period.
Off-duty birds perch extensively on the truncated surfaces of the
Achyranthes thicket, from where they can communicate danger to those
below.

Twigs at the thicket margins become denuded of leaves, but this
damage appears to be physical rather than physiological and purple
flower spikes are produced prolifically during the rainy season, con-
currently with the observed nesting.

Some of the more robust stands of Achyranthes shelter only moribund
nests, suggesting that this vegetation phase of low semi-shrubs might
be principally a post-nesting phase, stimulated by birds and growing up
on the more open sites which remain the most popular with current
breeders. It is suggested that noddies, which have an obvious pre-
ference here for sites exposed to the sky, may be forced out onto the
top of the scrub (type (a)) as the guano-stimulated 'weed' vegetation
encroaches over their ground sites (type (b)).

A post-nesting phase of vegetation occurs when the Achyranthes,
with increasing proportions of Acalypha, grows to heights of 1m or
more and is used by noddies only for casual perching — preferred
perches being afforded by the more twiggy, smaller-leaved though taller
Pemphis acidula bushes.

White-flowered Ipomoea macrantha and Pleurostelma cernuum twine
among the less disturbed growths.

Some of the islets unoccupied by birds show a drying away of the
Achyranthes-Acalypha undershrubs and colonisation by small Pemphis
saplings with a few young Ceriops tagal mangroves marginally. Other
islets have relict, morbund Achyranthes beneath a thriving Pemphis
scrub — very patently a later phase of the same post-bird succession.
Such islands have no grass or herb flora even in the rainy season,

10

these belonging to an earlier, more open seral phase, which is seen to
advantage only on islets ungrazed by the giant tortoises.

The vegetation of certain islets supporting a combination of
Pemphis with Achyranthes-Acalypha understorey seems to have a different
origin. In these islets the Pemphis is morbund and the undershrubs
thriving. Noddies perching on the upper storey of Pemphis kill back
many of the shoot tips and deposit quantities of excreta which is
apparently appreciated by the robust, white-splashed undergrowth.

These two types of community suggest a divergence of seral direct-
ion during post-nesting phases. If noddies desert the area completely
the Pemphis thrives and shades out the interim stage of Achyranthes-
Acalypha; if they return subsequently to roost in the higher Pemphis
twigs the more guano-tolerant though lower-growing species may return
and gradually come to dominate if pressure is sufficient to cause
extensive die-back of the Pemphis. That this occurs is borne out by
the total death of some of the large central Pemphis bushes on one such
islet used by noddies. That roosting sites change, probably more
frequently than nesting sites, is indicated by the presence of perching
birds in bushes suffering little damage as yet.

Islets undistrubed by birds can, if of sufficient size, support a
mixed scrub which includes Sideroxylon inerme, Ficus nautarum, F.
reflexa, Flacourtia ramontchii, Allophyllus africanus v. aldabricus and
Suriana maritima. Only the larger of these bushes have boughs broad
enough to accommodate fairy terns' eggs, but they are visited by various
of the heron-egret group, Souimanga Sunbirds Nectarinia sovimanga,
Comoro Blue Pigeons Alectroenas sganzini, Madagascar Turtle Doves
Streptopelia picturata, Madagascar Bulbuls Hypsipetes madagascariensis,
and Madagascan White-eyes Zosterops maderaspatana.

(c) on prostrate grass mats

A small proportion of noddies nest on prostrate but still green
mats of the broad-leaved grass, Dactyloctenium aegyptium. Nests in
such sites often have fresh grass shoots incorporated in their struc-
ture. Areas pointed out by the Seychelloise boatman as having been
previously occupied by birds showed this same grass as a dense flowering
stand with shoots erect to heights of 30 - 50 cm and evidently bene-
fitting from the residual nutrients of the droppings.

One islet showed two distinct Dactyloctenium communities in March
1970, part of it being occupied by a densely flowering stand 1/2 m high
and part (three times as much) by a more tufted growth 15 - 30 cm high
and only just coming into flower. This later growth apparently more
recently vacated by birds, had the following floristic composition in
terms of ground cover:

ke
oP

Dactyloctenium aegyptium 80% Sida parvifolia

bh
oe

Boerhavia elegans 10% Phyllanthus amarus?

On)
oe

Portulaca oleracea 3% Bare Ground

iat

Quite commonly noddies in grass communities are among tufts of
fine-leaved Eragrostis sp. or, occasionally, Sporobolus virginicus.
This latter, the pan-tropical grass so common on most sandy or rocky
shores at or just above high tide level, shows no special affinity for
bird colonies, although it is the most generally distributed of the
coastal grasses away from them.

Several mainly bare islands support a little dead and depauperate
Pemphis and a spreading growth of Sporobolus, suggestive of shrub degen-
eration followed by a new secondary succession starting with this halo-
phytic grass. Without some documentation of former bird activity, it
is impossible to say whether this is a biotic or climatic-edaphic
succession, and whether the cyclic changes are due to successive waves
of bird occupation.

(d) on non-succulent herb mats

On slightly larger islets away from the immediate splash of the
sea — which is effectively curtailed throughout at low water because of
the deeply undercut nature of the periphery, ordinary low growing
mesophytes may prevail. These only persist, however, under light
bird pressure. Species concerned are Boerhavia elegans, Cleome
strigosa, Corchorus aestulans, Sida parvifolia and Tephrosia alda-
brensis. The sea convolvulus Ipomoea pescaprae, so much a feature of
tern colonies on the sand of other atolls, was not recorded at all in
the rocky Aldabra colonies, only the less arenicolus I. macrantha.

(e) on succulent herb mats

The three plant species most closely involved with noddies, with
the possible exception of Achyranthes, are the three so-called sea pur-
slanes, Portulaca oleracea, P. australis and Sesuvium portulacastrum
in this order of abundance. The two yellow-flowered Portulaca: species,
together with Ipomoea pes-caprae, are the only plants to withstand
pressure from the hosts of sooty terns, noddies, crested terns and blue-
faced boobies on Latham Island (Gillham, 1972).

The pink-flowered Sesuvium, although very tolerant of guano, is not
necessarily guano-induced, being a typical halophyte of both spray-
splashed champignon and sandy beaches above the mangrove fringes.

Two similar-sized islets of slab-like platin rock, approximately
5 X 8 m, one occupied by noddies and one not, showed the following
flora:

Occupied islet Unoccupied islet
Bare platin 20% 50%
Sesuvium portulacastrum 50% 50%
Portulaca oleracea 20%

Achyranthes aspera seedlings 10%

12

The establishment of young Achyranthes on the occupied islet suggests a
progression to the semi-shrub stage — one which is parallelled by the
semi-shrub stage of Ballota africana on South African bird islands
(Gillham, 1963b).

Another bird-occupied island showed Sesuvium only in pockets
around the seaward edge, the main body of the vegetation being a Portu-
lacetum of the following composition:

Bare champignon rock 2% Dactyloctenium aegyptium 2%
Portulaca oleracea 70% Portulaca australis 1%
Boerhavia elegans 203 Phyllanthus amarus? 0.5%
-Cleome strigosa 4% Sida parvifolia 0.5%

The whole of this community smelt powerfully of guano and was
sprinkled with moulted feathers. All ‘species were ELlowering ue much
of the Dactyloctenium was flattened and dead — as in the booby colonies
of Wizard Island, Cosmoledo (Gillham, 1974).

This island and the next had, my Creole guide informed me, been
quite bare of plants until the onset of the rains 4 - 5 months before.
Evidently heavy bird pressure during droughts is lethal to herbaceous
plants, whereas these will grow under the same bird pressure during the
rainy season and (less well) with no bird pressure during droughts.

The second area showed only a 50% ground cover of the following compo-
Sitiron:

Sesuvium portulacastrum 15% Eragrostis sp. ie
Dactyloctenium aegyptium 12% Sporobolus virginicus 1G
Achyranthes aspera 10% Cleome strigosa 1g
Portulaca oleracea 5% Boerhavia elegans Yr
Portulaca australis 5% Lagrezia oligomeroides 12

(£) on naked champignon rock
Some nesting islets are completely without vegetation or with only

minute patches of Sesuvium or Sporobolus. In the absence of vegetation
the noddies build in shallow depressions of the champignon, sometimes
straddling holes through which the sea is visible below. Eggs in

these more open situations are sometimes destroyed by pied crows (Corvus
albus) the herons also prey on eggs and chicks.

In some instances these islets may be too close to the high tide
mark to support vegetation but, in view of the fact that they are not
too close to support noddy eggs and chicks this seems unlikely (parti-
cularly as Sesuvium and Sporobolus are well able to withstand immersion

3}

in sea water at high springs). Often they are the most crowded, so
could conceivably represent the bare phase of a bird-controlled plant
succession where most plant growth is inhibited. There are no records
available to show whether islets which are bare of both plants and
birds were once occupied by birds and also qualify for inclusion at the
bottom of a biotically controlled sequence.

Few of the noddy islands appear to have any soil or even accumu-
lations of blown sand (hence the absence in this work of what might have
proved useful soil analyses in relation to guano deposition). Obviou-
sly, however, a rudimentary soil must exist, if only invisibly in
crevices and convolutions of the old uplifted reef rock.

Blown sand is at a premium in the lagoon, where only small pocket
beaches are exposed at low water, these often partially sheltered by
mangroves so that the wind cannot pick up sand for transport to the
bird islets. There remain the coral chippings, working gradually
down into a clacareous dust under the influence of weathering, the
considerable amount of organic nest material brought from the driftline
and other vegetated areas by the noddies, and their guano.

From the physico-chemical and textural point of view the birds'
contribution is vital as a prerequisite for plant growth. Noddies,
therefore, which may destroy plants once established, may, nevertheless,
be a necessary step towards their establishment in the first place.
The type of vegetation present is likely to be as dependent upon the
beneficial accumulations of past noddy populations as on the current
increment of plant and animal residues from present populations.
Current increments, undiluted during drought, are likely to prove
detrimental; during light rain they may well prove advantageous to
the growth of species tolerant of mineral salts; during heavy rains
they are likely to be washed off into the sea and have no effect.

Plant communities subjected to varying seasonal pressures from
animals are always delicately balanced — the degeneration - regenera-
tion cycle moving on opposing directions as the pressures fluctuate.

Henns
Black-naped Terns (Sterna sumatrana)

These birds lay their eggs on bare rock in slight hollows scraped
in the limestone shingle. There is no indication that any kind of
plant cover is sought. Often there are no plants in the vicinity,
often only a sparse covering of Sesuvium. One unusually well vegetated
site showed the following seven species: Cleome strigosa, Corchorus
aestuans, Fimbristylis cymosa?, Eragrostis sp., Portulaca australis,
Phyllanthus amarus? and Sclerodactylon macrostachium.

These.small terns are by no means colonial, their nests widely
scattered with usually only one or two pairs per islet, so any effect
which they might have on the vegetation is likely to be negligible.

14

Little Terns (Sterna albifrons)
This species is reported to breed on Aldabra but was not seen.
Sooty Terns (Sterna fuscata) and Bridled Terns (S. anathaetus)

Both these species breed on Cosmoledo Island but no on Aldabra.
In view of the enormous abundance of Sooty Terns on other islands in
this part of the Indian Ocean, 1t is as. surprising -that this) species
has not taken over some of the unused noddy habitats on Aldabra as it
is that the Blue-faced Booby has not come to share the grassed, sandy
and platin areas of South Island with the giant tortoises. Or would
the lumbering passage of tortoises (Geochelone Aldabrachelys gigantea)
weighing 300 - 400 pounds be too great a hazard for eggs and chicks at
ground level and this the reason why ground nesting tropic birds,
noddies and terns prefer the mushroom islets onto which the tortoises
cannot climb? (During the present study tortoises were found only on
Green Island among the smaller islands, a terrain unoccupied by birds
and easily accessible from the lagoon waters through the tangle of
mangrove roots and stems. Tortoises are very bouyant and swim well.)

Crested Terns (Thalasseus bergii)

These birds were not seen to be breeding during the February to
May period, but small flocks were feeding. Several hundred birds
frequent the Avicennia marina mangroves of Ile Esprit, perching only in
completely dead trees and on the dead branches of moribund ones. The
fact that they never seem to perch in living trees indicates that they
cannot themselves be responsible for the death of the chosen perches.
Tree sites are more favoured for resting than ground sites. Perhaps
there is some predation by land crabs (Cardisoma) or coconut crabs
(Birgus), but some terns consort with turnstones (Arenaria interpres)
on the sand below the mangroves.

An ex-nesting site pointed out on Moustique Island was sandy with
marginal Sesuvium and Sporobolus — species needing no encouragement by
birds: ani chas, Some or shabuirteaitt.

Caspian Terns (Hydroprogne caspia)

These are reported to occur but were not observed.

Burrow nesters
Crab Plovers (Dromas ardeola)

These large-billed waders are constantly seen in flocks of many
hundreds among the Palearctic migrant waders from the north, particu-
larly in the western part of the lagoon and on the western sea reefs,
as at AnseVar, but seem to be scarcer in the east end, where most of
the sea-birds congregate. They nest in sandy burrows, but not

15

apparently on Aldabra, although some of the overwintering population
stays on through the summer (Penny, 1971).

Audubon's Shearwater (Puffinus Cherminieri)
This petrel was first discovered breeding on islets in the lagoon
in 1967 and were not seen during the present study.
Exotics in bird-induced vegetation

Renvoize (1971) estimates that the total flora of Aldabra numbers
some 206 species of which only 40 are exotics and as many as 31 are

endemic. Man's influence has been slight and the exotic species have
not spread much beyond the settlement on West Island and a few locali-
sed areas used as fishing camps by the Creoles. The only two having

profound influence on the native flora, the she oak (Casuarina
equisetifolia) and the coconut palm (Cocos nucifera) occur on sandy
flats or more inland areas not used by nesting sea-birds.

The apparent total absence of exotic species from the Aldabra
sea-bird colonies studied in a wide range of habitats, particularly in
the Southern Hemisphere and usually quite remote from direct human
interference, alien plants are conspicuous, often more abundant than
natives and frequently excluding indigenous species altogether
(Gillham, 1961).

The disturbance and trampling, which damages or kills existing
plants, and the addition of phosphatic and nitrogenous nutrients paves
the way for an influx of arable weeds and 'followers of man' — usually
eaSily distributed species which thrive in areas fertilised deliberately
or inadvertently by effluents of one sort of another in man-dominated
areas.

Perhaps more remarkable than the absence of exotic weeds from the
bird colonies it is the abundance of indigenous ones in the settlement.
The native Cleome strigosa and Sida parvifolia compete on equal terms
with the invasive Strachytarpheta jamaicensis and Ricinus communis
seedlings on the open sand: native grasses, particularly Daknopholis
boivinii, and the blue-flowered Evolvulus alsinoides mingle with the
various Euphorbias and Amaranthys

The herbaceous vegetation of certain noddy islands bears some
resemblance to that of the settlement and fishing camps, as does the
local increase of Acalypha claoxyloides in less trampled parts of both.

Both pink and white-flowered Madagascar periwinkle (Catharanthus
roseus) have been planted outside the Creole's shacks and have spread
extensively from the sands of the settlement through the champignon
undergrowth of the neighbouring Casuarina woodland — as at the Anse
Mais and other camps. This species might have been expected in the
sea-birds colonies, in view of its prevalence in nearby sooty tern

16

Table l. Postulated Plant Succession on Lagoon Islets in Relation to
Bird Pressure

This resembles the climax
community utilised by frig-
ates, boobies, fairy terns

and passerines. Only if

defaecation is into the Mixed shrubs

water will this community (or mangroves if sufficiently marine)
remain stable under heavy

bird occupation and not ---No bird disturbance
revert to scrub. In

fact this usually happens

on the lagoon coasts of the Pemphis with little or no understorey

main islands but seldom on
the smaller lagoon islets.

Little bird
~ disturbance
Roosting noddies or
nesting herons Pemphis with understorey of Achyranthes
and- Acalypha
Damage to Pemphis and
return of understorey

of 'weedy' semi-shrubs

Noddies squeezed out
by vegetation.

---Lighter occupation
by tropic birds,

Increasing Heavy occu- herons rets, et
bird pation by Te ees
Pressure oddies .
ressu one ; The semi-shrubs Achyranthes and some
A with a lict herb co i
ieee calypha relic erb community

herons,etc, E :
4 Continued occupation

Return to herb phase pire node at Teenie ses
of tropic birds

Portulaca oleracea, Dactyloctenium

and other herbs and grasses

Drought and

continued Noddies depositing

bird guano and quantities

pressure ---of organic nest
material

Return to lower herb Sesuvium and Sporobolus and pulverised

phase or bare rock reef-limestone, chippings

Terns depositing
---guano but no nesting
material

Bare. rock —‘no, soll) no) plants

iL

colonies, where rank growths spring up each year to be killed back when
the birds return to nest. It was not found, however.

Three explanations are offered as contributing to the 'purity' of
the bird flora:

(a) Lack of bird transport

Most of the bird colonies studied in other parts of the world have
been visited by gulls, if not nested in by them, and gulls, as general
Scavengers and producers of only partly digested crop pellets (Gillham,
1970a) have ample opportunity for transporting viable plant disseminu-
les from man-made habitats to bird-made ones. Not only are gulls
(Lesser Black-backs — Larus fuscus) exceedingly rare on Aldabra, but
mone of the sea and shore-birds were seen to alight in the settlement
apart from cattle egrets, squacco herons and turnstones which do not
nest in the areas studied. Boobies fly over daily to their fishing
grounds at sea — followed by the usual retinue of pirating frigate
birds — and white-tailed tropic birds and noddies pass occasionally, but
none of these alight.

Pied crows are commonest in the settlement and crows are known to
produce crop pellets. Some of these which take eggs and chicks on
the bird islands may hail from there but these Supply the only possible
link for bird transport of weed seeds.

(b) Lack of fertility-demanding ruderals

The usual attraction of bird colonies for weeds of cultivation is
their fertility. Fertility on this notoriously inhospital atoll,
where even coconuts fail to attain a reasonable size, is likely to be
limited to the bird colonies. The weeds of the barren, porous, organic
deficient sands of the settlement cannot, of necessity, be demanding in
their nutrient requirements, so this special attribute of the bird
colonies is no special incentive to establishments should they by
chance arrive.

Sea-birds, no doubt, nest outside the rainy season, but the present
observations (February to May, 1970) were all made during the rains, so
that chick growth and plant growth were proceeding side by side — the
plants benefitting (or otherwise) from fresh, unleached excrement
between showers.

Had the rainy season followed the nesting season instead of coin-
ciding with it — as so often happens in less tropical habitats where
plants invade during the rainy autumn following a dry summer nesting
season, it is likely that the range of plants able to tolerate the
ameliorated chemical conditions would have been wider.

(c) Availability of indigenous ephemerals

The abundance of European annuals in Australasian bird colonies
is facilitated by the paucity of easily-dispersed, fertility-demanding
annuals and ephemerals in the Australasian flora. This does not apply
on Aldabra, where the terrain is so harsh that large areas become bare

18

— |

during the dry season, to become recolonised by short-lived indigenous
plants during the 3 to 4 months of the rains (approximately December to
May) . These pass the adverse drought season as seeds, many of the
shrubs dropping their leaves then, so that shade as well as moisture is
in short supply.

The herbaceous, vetch-like legume, Tephrosia aldabrensis (or T.
pumila v. aldabrensis) and the shrub Allophyllus aldabricus (or A.
africanus v aldabricus) are endemics of the less spray-washed bird
islands. Group endemics found only on Aldabra and the other atolls
within a few hundred miles are Acalypha claoxyloides, Cleome strigosa
and Plumbago aphylla — all frequent members of the Cosmoledo bird
colonies as well as Aldabran ones.

The major natural biotic disturbance on Aldabra is provided by the
giant tortoises and no less than eight of the small herbs of tortoise-
grazed turf feature in the limited flora of the more trampled bird
colonies. These are Boerhavia, Lagrezia, Phyllanthus, Portulaca, Sida
and the grasses Dactyloctenium, Eragrostis and Sporobolus. The wire-
grass, Sclerodactylon, although not grazed sufficiently hard to become
a turf species, is not immune by virtue of its high fibre content bur
is often eaten (Frazier, in litt.). The shrubs and semi-shrubs are
denuded of their leaves up to a browse line set by the height that a
tortoise can reach when propped up on another's carapace. Their
regeneration is hindered by the grazing of seedlings and this may help
to account for the fact that the particularly soft-leaved Acalypha,
although a group endemic, is conspicuously more abundant around the
living quarters of man and on the small sea-bird islets where tortoises
do not normally occur.

Vesey-Fitzgerald reports "a herb mat community" particularly .
beneath dense bird colonies (presumably tree colonies) on other islands
in the west Indian Ocean, but this has no direct parallel on Aldabra
where most of the tree nesters return to the lagoon the quota of
nutrients which they bring from the sea.

SUMMARY

The vegetation of the various types of site occupied by nesting
seabirds on the uplifted atoll of Aldabra in the western Indian Ocean
is discussed. Tree-nesting Frigate birds, Red-footed Boobies, herons,
egrets, and White Terns have little modifying influence on the mangroves
on which they next, their excreta being dispersed in the waters of the
lagoon. Undergrowth nesters, principally Red-tailed and White-tailed *
Tropicbirds, have a slightly more pronounced effect. Terns nest
sparsely and mostly in the open, but the more abundant white-capped
Noddies range from bare champignon rocks to the tops of low Achyranthes
aspera scrub. Their suggested role in the changing cycle of vegetation
from succulent halphytes through mesophytic herbs and guano-tolerant
undershrubs to shrubs is considered and offered in diagrammatic form.
Attention is drawn to the unusual complete absence of exotics! in the

LY)

bird flora and parallels drawn between disturbance by birds, tortoises
and man.

ACKNOWLEDGEMENTS

My grateful thanks are due to the sponsoring and financing bodies
enabling me to visit the atoll — the Royal Society of London and the
Natural Environment Research Council — and to University College Cardiff
for the granting of 5 months' sabbatical leave. Also, to Mr. J. Frazier
of the University of Oxford, Mr. S. Renvoize of the Kew Herbarium,

Mr. S. Kibuwa of the Nairobi Herbarium and Dr. B. Harris of the
University College of Dar Es Salaam for identification of plants, to
Mr. A. W. Diamond for information on where to look for sea-birds, to
the late Cdr. G. Lush, M. B. E. and the others responsible for mainten-
ance of the island outpost prior to the opening of the new research
station and to Harry Charles, my Seychelloise guide.

REFERENCES

Beamish, T. 1970a. Killing of Catharanthus roseus by sooty terns.
BBC Living World television programme.

Beamish, T. 1970b. Aldabra alone. London: Allan and Unwin.

Diamond, A. W. 1971. The ecology of the sea birds of Aldabra. Phil.
Deans. hk. COC. Lond. B 260: 561-571.

Giulihnan, Ms. BE. 1961. Alteration of the breeding habitat by sea-birds

and seals in Western Australia. DeeECOlwI9 2) 239-500)

Gillham, M. E. 1963a. Coral cay vegetation, Heron Island, Great
Barrier Reef. Prec. R. Soc. Queensland 83: 79-92.

Gillham, M. E. 1963b. Some interactions of plants, rabbits and sea-
birds on S. African Islands. Ua lxelolks Iho. 7s eys 2

Gasibam, Me obi. 97/0). Seed distribution by birds. The flora of a
changing Britain (ed. F. Perring) : 90-98.

Gilkinam) Me Ere. 1977.4': Observations on vegetation of Blue-faced booby
colonies on Cosmoledo Atoll, Western Indian Ocean. Atoll Res.

Bull., this issue.

Penny, M. 1970. Breeding success of fairy terns on Cousin Island,
Seychelles. BBC Living World television programme.
Renvoize, S. 1971. The origin and distribution of the flora of

Aldabra. Phi. Trans wR. 1sOCw Lona. 1B) 260s) 22—236..

ete
Tey

ATOLL RESEARCH BULLETIN
NO. 201.

LIFE HISTORY NOTES ON SOME ALDABRAN LAND
BIRDS

by C. B. Frith

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

February 1977

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LIFE HISTORY NOTES ON SOME ALDABRAN LAND
BIRDS

by

C.B. Frith!

Introduction

The following observations were made during one year's residence
(April 1972 - April 1973) on Aldabra Atoll, Indian Ocean (latitude
9° 24' S, longitude 46° 20' E) while I was a member of the Royal
Society Research Station scientific staff. Although collection of
the data presented here was incidental to other studies, daily nest
inspections were made when possible and form the basis of this work.
Notes made on the endemic weaverbird Foudia eminentissima aldabrana
are not included here as they form a separate more comprehensive study
(Frith, in press). Feeding data for most of the species listed also
form another study and only a brief outline is given here. Data on
Centropus toulou insularis are reported elsewhere.

Benson and Penny (1971) completely summarised the existing
knowledge of land birds found on Aldabra and emphasized points
requiring attention. The status of the species was found to be as
given by Benson and Penny unless otherwise indicated. For the
location of places given in this text I refer to a recent paper by
Stoddart (1971).

Malagasy Kestrel, Falco newtoni

Sparse throughout the atoll, one or two birds being seen each day
at various locations about the islands. Although being recorded as
not found on Ile Michel by Benson and Penny three birds were seen in
June and a pair with young were observed on this small island in
October.

Coconut palm, Cocos nucifera, and stands of Casuarina equisetifolia
trees appear to be favoured breeding sites, no nests being found in any
other situation during this, or previous, studies. Benson and Penny
expressed the view that the kestrel possibly colonized Aldabra
subsequent to the introduction of palm trees by man. They concluded

1 Formerly Royal Society Aldabra Research Station
(Revised manuscript received January 1975 -- Eds.)

Oo

this as the only known nest at that time was found in the crown cf a
palm tree. Casuarina trees are, however, equally used and there is
some doubt as to whether this tree was introduced or is naturally
present on Aldabra.

Feeding

The two endemic geckos, mostly the diurnal species Phelsuma
abbotti, and the'endemic skink, Ablepharus boutonii, plus various
large insects form the basis of the diet. Twice I saw a kestrel
eating a small rat (Rattus rattus). Benson and Penny suggest that
small birds are taken but I did not see any evidence of this, nor
did I see a small bird showing any fearful reaction to kestrels.

' Behaviour

Unlike the observation of Benson and Penny more than two
individuals in close association were seen. Three adult birds were
observed flying and gliding together in a playful fashion in strong
winds during June. During an inspection of an active nest the
resident pair was assisted by a neighbouring pair (which had recently
lost a clutch to predators) in mobbing me, all four birds flying at,
and sometimes striking me. Pied Crows, Corvus albus, and vagrant
Eleonora's Falcons, F. eleonorae, and Broad-billed Rollers, Eurystomus
glaucurus, were mobbed by kestrels.

Breeding

On 18 August a pair was seen to copulate in the upper dead
branches of a tall Casuarina tree in palm grove. Both birds held
wings extended and dropping and called continuously.

A pair was found nesting in the crown of a palm tree at Anse Mais
on 28 October but the tree could not be climbed. Another nest .
containing one egg was found thirty feet high in the crown of a palm
near Old Settlement on 28 August: on being examined twenty days later
it was found to be empty. A nest found on Ile Michel on 21 October .
was ina hole in a Casuarina tree approximately forty feet high. It
contained three nestlings in off-white down, the youngest being three
or four days old.

A fourth nest was located on 10 September when the remains of one
egg were seen to be carried from it by a Pied Crow. On the ll
September this nest contained three eggs, the original clutch with
little doubt, therefore, consisted of four. The eggs were laid in a
hole in a large Casuarina, formed by the loss of a large bough,
approximately forty feet above the ground. The tree was situated in
the centre of human dwellings, directly between two occupied houses.
The history of this nest follows:

10 October 1972 3 eggs

Wye Ye 1 egg, two nestlings

13 u 3 nestlings

21 MY Empty, almost certainly predated by
Pied Crows.

9 November 1 egg

12 December 1 egg

22 December Empty

The incubation period for the original clutch was at least 31 days.
Throughout this pair's breeding attempts they were continuously
disturbed by Pied Crows, causing them to spend much time in nest
defence. The population of crows about Old Settlement is relatively
very dense (ca.40) and is supported by food made available by man.
One parent kestrel was so continuously chased and attacked by crows
at the nest that it fell into a fresh-water tank and nearly drowned.
It was helped out of the water by a woman then spent several days
perched at the nest entrance recovering.

White-throated Rail, Dryolimnas cuvieri aldabranus
Feeding

I have little to add to what Penny and Diamond (1971) published
Save one or two observations confirming more unusual food items.

An immature rail was seen to kill and eat a small ghost crab,
Ocypode cordimana, measuring approximately one inch across the carapace.
Odd legs detached during killing were swallowed first, the remainder
being swallowed whole.

Five rails were watched attacking and eating Green Turtle
hatchlings, Chelonia mydas, which had erupted from a sandy beach in
bright sunshine (Frith, in preparation). Unhatched turtle eggs,
exposed by me, were also immediately taken by rails and eaten. The
turtle hatchlings were not eaten whole but were pecked about the
carapace and eaten in small pieces. The hatchling turtles broke
surface at the same spot and their appearance was anticipated by the
rails, the birds standing over the hatching point and probing the sand
to the full length of the bill. Such probing was performed with an
open bill, with the nictitating membrane covering the eye during
deeper thrusts. Upon contact with a submerged turtle the bill was
tightly closed about the prey, the head shaken sideways and the turtle
extracted vertically. Details of this activity were obtained by
simulating erupting turtles. This I did by burying an extended leg

in the beach sand with my toes pointing upward several inches beneath
the surface. By slowly wagging my toes a very slight movement in the
surface sand was caused and attracted four rails from up to 3 metres
away. They rushed to the disturbed spot and probed in the above
described way. Two individuals returned to the spot after an absence
of several minutes and probed again, without any movement being
produced to attract them.

I once saw a rail disturbing the nest of a red wood ant,
Camponotus maculatus, by pecking at it and feeding upon eggs, larvae
and some adults.

Behaviour

The White-throated Rail apparently thrives in areas supporting
considerable numbers of introduced rats, Rattus rattus (Penny and
Diamond). I observed one interaction between these two animals. An
adult and a large immature rail were observed feeding upon camp scraps
when a large male rat appeared and slowly approached the food. As
soon as it was observed by the rails the adult bird walked directly
at it and pecked it severely on the nose. This action was attempted
at each advance of the rat but after the initial peck the rat avoided
further physical contact and slowly retreated, finally leaving
altogether. Throughout this eviction the immature rail continued to
feed, showing no fear of the rat a couple of feet away.

Short copulation sequences were witnessed twice in May (out of
breeding season) performed by the same two birds, the ceremony being
the same in both instances. The female bird had the brighter pink bill
base which agrees with the findings of Penny and Diamond. The apparent
soliciting posture of the female consisted of spreading and lowering
the white outer tail feathers and lowering the bill until the tip
touched the ground. The male mounted and dropped a wing either side
of the female, gave two or three thrusts of the body and hopped down.
Upon dismounting the male walked about the female with head held
stretched forward and downward and wings spread and slightly raised.

As active nesting is recorded for November and December (Penny and
Diamond; C. Huxley, pers.comm.) it is considered unlikely that the
above copulation postures had anything to do with reproduction. Of a
number of copulations observed by Penny and Diamond only two were
followed by a display, apparently identical to the above described.
These were performed by a male after mounting a female to which he was
not paired. It would seem possible, therefore, that some ritualised
mounting takes place between unpaired birds which is directly followed
with a postcoital display from the male. This is supported by the fact
that other copulations between known paired birds were followed by no
displays, the birds simply separating to feed (Penny and Diamond).
Penny and Diamond suggests the postcoital display is connected with the
status of a male as an intruder in an occupied territory.

Penny and Diamond recorded that the White-throated Rail was never
seen to use the feet as weapons but referred to an account by Abbott,
in Ridgway (1895) of birds "flying at each other like game cocks". An
instance of the latter was observed once. The two birds concerned were
of the same brood, being ringed as large young in the company of an
adult on 17 April 1972. The fight took place on 30 May 1972 and was
apparently a dispute for presence in the immediate area of some
resting people and their food scraps (i.e. feeding territory). The
birds faced each other, a foot apart, with bill lowered and giving an
occasional low grunt accompanied by a short upward flick of the bill.
Suddenly, both would leap at the other feet first, beating the wings
and vigorously kicking. Should one bird get a good grip both tumble
the ground severely pecking at each other. Fights such as this could
result in permanent damage. Whilst marking one bird my hand was quite
deeply cut as a result of the kicking action. It is possible the
bastard wing claw of this species is utilized during fights, particu-
larly as wing beating was conspicuous.

Notes

During the period October 1967 - February 1968 Penny and
Diamond ringed 23 birds at Middle Island Camp and 13 at Anse Coco.
During the term of the present study the author and other workers
visited the two localities fairly frequently, usually for at least
several days at a time. Only one marked bird was noted at each
location. The individual seen at Anse Coco in May 1972 was marked
as an adult during June 1968. The marked bird seen at Middle Island
in October 1972 was originally ringed in October 1967 as an adult.
Thus these two birds had lived at least five and six years
respectively. Lack of sightings of any other marked birds is not
necessarily indicative of mortality. Penny and Diamond found that
the rail population was to some degree transient. It is also possible
the camps of these workers attracted large numbers of rails which,
after the departure of the men, dispersed from the area and would not
have again been attracted to the smaller, more temporary, camps since.

Comoro Blue Pigeon, Alectroenas sganzini minor

The Blue Pigeon is noticeably more common at localities as close
to West Island as Anse Mais and Anse Polymnie than at the former
locality. It is possible this is due to a lack of suitable foods on
West Island but it is equally possibly due to predation by man in the
recent past. Small flocks were commonly seen on the lagoon islands,
Ile Michel and Ile Esprit, and were to be frequently seen flying
strongly about the atoll. Of particular note (at least during August)
were very large numbers of birds flying from Middle Island to South
Island, across East Channel, in the morning and returning in the
evening. Birds flew across the channel singly or in small flocks of

up to fifteen, at a height of approximately 25 to 40 feet. Many
hundreds of birds performed these flights daily.

Feeding

This species was never seen on the ground. Lone birds and flocks
of up to twenty were observed feeding on various fruits and berries
which were picked from the plant. The arboreal feeding of this species
considerably limits any competition between it and the only other
pigeon on Aldabra, Streptopelia picturata coppingeri, which is a
ground frequenting seed eater.

Behaviour

When not feeding birds were typically seen in bare, higher,
branches. A type of displaywas observed, given by one bird to another,
on two occasions (once in upper bare branches of Casuarina in April
and once in mangrove in May). One bird flew to perch a foot or two
from the other, lowered the head gently until horizontal with its feet,
raised the rather elongate feathers of the neck, and shook the head.
The head was shaken perhaps one inch either side of the normal
position, and was reminiscent of a pigeon regurgitating food. No
distinct bowing of the whole body took place, but a low grunt-like
"weow-weow" was uttered. This was undoubtedly a directed display
which is possibly a ritualized regurgitation, or feeding, action or
ritualized nest building ("nodding" Goodwin, 1967). Benson and Penny
record nesting during January and February (1968) and it is therefore
probable that the above display was given toward the end, or after,
breeding.

Notes

Two individuals observed feeding closely were seen to have
slightly aberrant plumage. Both had a noticeable amount of white in
the upper wing when extended, though not extensive, and one had the
central tail feathers pure white and so gave the tail the appearance
of being white bordered with blue instead of uniform blue.

Aldabra Drongo, Dicrurus aldabranus
Feeding

Benson and Penny considered this species to show a marked
preference for mangrove but my observations indicate that stands of
Casuarina are equally favoured. Some authorities consider the
Casuarina recently introduced onto Aldabra, in which case it might
be used as a suitable substitute for mangroves, particularly as
reptiles are available in the former but not the latter habitat.

Observed feeding activity suggests that, in habitats other than
Mangroves, small reptiles are eaten as much as are insects. Geckoes,
skinks and larger insects are pounced upon, taken to a perch upon which
they are pounded and are then ripped apart whilst held to the perch by
the feet. Insects are commonly hunted in flight, often high above the
tallest trees. No records of feeding in mangroves were made, possibly
due to limited visibility and time spent in the habitat. Benson and
Penny's suggestion that this species eats other birds is considered
most unlikely. No fear of drongoes was noted.

Behaviour

Commonly seen in family parties (immatures having a grey and
whitish plumage) of about four birds, feeding together by perching in
exposed branches and pouncing on prey within a small area of
relatively open ground. Lone birds or pairs were seen to severely mob
and strike fruit bats Pteropus seychellensis, Green-backed Herons
Butorides striatus, Pied Crows Corvus albus, kestrels Falco newtoni
and the migrant Eleonora's Falcon F. eleonorae and Grey Cuckoo Cuculus
canorus (presumably reacted to as if a falcon as the Drongo has evolved
without the presence of a parasitic cuckoo) at various times of the
year. Human intruders near an active nest were also mobbed and
sometimes struck.

One odd observation of pair activity was made in the middle of the
non-breeding season (June). Two adult birds were seen perched by a
fresh water pool. They then flew to a nearby tree where one bird
picked off a dead leaf and offered it to the other. The second bird
took the leaf and then dropped it.

A simple display was observed to be performed by an established
pair of birds involved in nest building. The smaller bird added
material to the nest and then flew to perch nearby where it was joined
by its mate. The birds faced each other holding the bills slightly
above horizontal and shook the wings vigorously and wagged tails from
side to side. During this display both birds produced soft high
pitched squeaky twitterings. This was not unlike young birds begging
and possibly functions in maintaining the pair bond for part, or all,
of the breeding season.

Breeding

Breeding activity was first noticed in mid September and
continued until at least early January after which the species was not
studied due to other work. Three pairs were observed at the nest, two
only temporarily but one over a period of three months. All six nests
involved were built in the fork of twigs of Casuarina trees. Nest
examinations were made by use of a 'mirror-stick'.

One nest found on 5 November 1972 was twenty-five feet above
ground in palm grove, with some Casuarina trees, and contained a
single egg. An examination on 18 November found the nest empty.
Another nest was found on 4 December twelve feet high, in palm grove
with Casuarinas, containing one egg. Subsequent examination on
7 December found only one egg, which was gone on 12 December.

The breeding of a pair found close to the research station is ,
worth recording as a number of interesting points were established.
The very beginning of a nest was found on 25 September 1972 in
Casuarina woodland. It was placed twenty-five feet above the ground
and consisted of spider webs attached to the supporting fork and a
very sparse bowl of fine twiglets. At this early stage birds were
seen to sit in the very flimsy bowl and shape it by vigorously pushing
down and outward with the breast.

The (continued) persistence of this pair of drongoes was
remarkable. At the start of breeding a nest was built in little more
than twenty days and a clutch of three laid which was predated. A
second nest was built, utilizing the last, and was completed fifteen
days after predation upon the first clutch. This second nest was
found under 'deconstruction' four days after previous examination and
it 1s possible a second clutch was laid\and. predated) in the sinterime
The third nest took certainly less than fifteen days, to complete in
which a clutch of two was laid. This nest was predated and a fourth
built in a period of ten days or less in which at least one egg was
laid and predated. A fifth structure was not attempted, but the fourth
was added to and a clutch of two eggs produced no more than fourteen
days after predation upon the previous. This final clutch was
predated and no further breeding attempt by this pair was noted as
the birds apparently left the area. At. least four, possibly five,
clutches were laid by the female of this pair. It is interesting to
note that as*the Season progressed: nests were constructed in® faster
time, material from the previous structure being utilized despite an
apparent abundance of suitable materials in the immediate area. Also
of interest is the original clutch of three followed by clutches of
two.

Eggs examined agree in colour with those described by Benson and
Penny. A clutch of two measured 26.2 x 19.4 and 26.3 x 19.0 mm. Nest
building and incubation of eggs was observed to be carried out by both
sexes, no calls or displays being given during ‘take-over of the Vavcer
duty.

Pied Crow, Corvus albus
Feeding

Other than from human settlement and temporary camps food is almost
exclusively obtained on the beaches, open beach crest and coastal
grassland areas, where various Crustacea appear to form the basic diet
(in particular ghost crabs Ocypode spp., and hermit crabs Coenobita
spp.). The fact that approximately fifty per cent of the entire crow
population relies upon man's discarded food suggests that natural food
sources may be limited for this species. Pied Crows are extremely
predatory upon eggs and nestlings of other land bird species. Diamond
(in Benson and Penny) found them taking eggs of tern species. Crows
were often seen perched on top of the nests of Foudia eminentissima,
ripping through the roof of the chamber or pulling the whole structure
to the ground. As described above crows severely predated upon
kestrels in the vicinity of Old Settlement. They were also seen to
extensively mob migrating falcons, Falco eleonorae chasing them to
great heights and over considerable distances.

Dying and dead giant tortoises, Geochelone gigantea, would
obviously provide temporary large food sources for crows. Pied crows
were twice observed to quickly appear at the location of a daylight
eruption of Green Turtle hatchlings, Chelonia mydas, and to eat them
(Hisashi 975)”.

Breeding

Benson and Penny recorded a nest containing three eggs ina
coconut palm during December and one inaccessible nest was seen,
apparently with attendant birds, during each of November, December and
January. Two nests were found on West Island during the present study,
both approximately forty feet high in the tops of Casuarina trees on
beach crests. One, which could not be reached, was seen to have a
pair in attendance on 1 July but was considered unlikely to contain
eggs or young at that time. The second nest was discovered on
9 November containing three eggs and two nestlings. On 19 November
only three eggs remained one of which was smashed, these were obviously
addled. The nestlings would not have been old enough to have fledged.
This same nest was examined on 17 December and contained four fresh
looking eggs but no further examinations were made. During twelve
months casual observation of the Pied Crow population on West Island
only three young birds were noted (other than nestlings), on 12
November, in close association within a large flock. Possibly limited
food and longevity restricts the reproductive rate.

LO

Souimanga Sunbird, Nectarinia sovimanga aldabrensis
Feeding

Feeding takes place in all habitats but to a far lesser degree in
mangroves than elsewhere, food consisting primarily of flower nectar
and insects in that order.

Behaviour

Out of the breeding season flocks of from three or four to
approximately thirty birds, in both male and female plumage, were
commonly seen engaged in some form of chasing activity. This took the
form of flying in very close association swiftly through vegetation
giving excited twittering calls. Such behaviour was most commonly
observed just after breeding and may therefore be the result, or cause,
of territorial breakdown.

Males in full breeding plumage typically perch on conspicuous
twigs and sing loudly. A 'bill up' display was commonly given by males
to both females and other males, facing the recipient and raising the
bill to, almost vertical and ‘thrusting -the, chest out. ) This, daspiltay
probably functions in both territorial and initial courtship
confrontations.

A simple nest-site selection ceremony was observed several times.
A male and female would fly to a potential nest site and perching on,
or clinging to, available material situate themselves very close to
each other. The material to which the intended nest was to be
attached was pecked and billed, mostly by the female. During this
mutual inspection, lasting approximately two minutes, a soft
twittering was produced by both birds. Suddenly both birds would
leave the nest site, the male to perch nearby and the female to
collect spider web for the nest foundation.

Breeding

Active breeding was observed from August 1972 through to April
1973. This very early start to breeding (compared with other
passerine species) preceded the start of any increase in insect
abundance (fig. 1), unlike other species more dependant upon insect
food. As this species feeds predominately upon nectar its breeding
possibly correlates more closely with available nectar but
unfortunately the latter was not assessed in any way. It was noted
however that earlier breeding attempts were less successful until,
that is, insects became more numerous (for feeding young).
Nevertheless several clutches were laid in early September and one
brood had fledged before the end of that month.

alias

Breeding occurs in all habitats with vegetation higher than
grasses. Five successful nests were located in and upon buildings
constantly in use by people, one being in an occupied bedroom and
another in a noisy and well used workshop. Benson and Penny give
detailed descriptions of nests and sites.

Only the female nest builds, the male often following her as she
searches for material and adds it to the structure. One female
observed during her first day of nest construction flew between the
nest and spiders webs forty feet away thirteen times in seven minutes.
Only four successful nests were timed in their construction from start
to finish (egg laying), but it is interesting to note, however, that
structures built earlier in the season took longer than those built
during the height of it:

Nest started No. of days to egg laying
21 August Sil
11 October 21
16 November 8
9 December 9

Nests were observed to be constructed much faster than eight days,
but were not timed. Nests predated early in the season were abandoned
and another built closeby. Nests predated in the height, or toward the
end, of the season were usually added to for a day or two and re-used.

Of twenty-four clutches examined twenty-three consisted of two
eggs and the other of one egg. As predation of part clutches was
found to be common it is probable that the one egg clutch was the
result of predation between nest inspections. Eggs are laid at
approximately twenty-four hour intervals, mostly during the early
morning hours.

Males take no part in incubation. Length of incubation, from the
date of second egg laying, was found in five clutches to be thirteen
days, in three clutches fourteen days and in two clutches fourteen to
fifteen days. These periods fall within the variation given by Skead
(1967) for a single sunbird species. The usual pattern is for one egg
to hatch one day and the second the next, (in one clutch only did both
eggs hatch on the same day), incubation starting with the first egg.
Twenty-two eggs averaged 15.7 x 11.1 mm. the variation being
Ae elite NO) a) = Gmc

Nestling sunbirds react to the arrival of a parent immediately
after hatching, a nestling being seen to beg vigorously as the nest
was touched although still freeing itself from the egg. Nestlings are
fed by both parents although several hourly observations indicated that -

We

the male contributes! only a ithird or less to feeding, wisitss, Nestling
period varied from fourteen to seventeen days, averaging fifteen days.
Young of the only two successful broods of two fledged on the same day.
In four clutches one egg did not hatch and disappeared a few days after
the other egg hatched. In one brood of two, produced early in the
season when insects were relatively sparse (fig. 1), one nestling died
of starvation.

Notes

Only a single family of sunbirds was marked with colour rings but
subsequent observations of them proved interesting and indicated that
established pairs return to the same breeding territory season after
season. The family concerned consisted of a pair and their single
nestling. All three were individually marked at the nest site on
12 April 1972 (when the young bird fledged). The immature bird was
still begging on 4 May, twenty-two days later, but on 10 May appeared
quite independent and was feeding itself extensively. All three birds
were regularly seen, together or separately, within a hundred foot
radius of the original nest location until 26 June after which the
immature was absent (next being seen on 13 December 250 yards from
parents' territory). The parent pair was continuously observed
within their original territory, and nested again in almost exactly the
same spot as the original nest, until I left the atoll.

Malagasy White-eye, Zosterops maderaspatana aldabrensis
Status

This bird does not benefit from the groves of coconut palms as do
several other passerine species and was rarely seen in that habitat
unless Casuarina trees or bushes and shrubs (i.e. introduced cotton)
were plentiful. It was abundant in mangroves, where many other land
bird species were not but, like Benson and Penny, I found no nest in
mangroves.

Feeding

Diet consists predominantly of insects save for the taking of some
flower nectar, fruits and a little seed. Insects are mostly taken from
foliage in typical Zosterops fashion, being picked from the surfaces of
leaves. Birds feed in loosely associated flocks of from three or four
to about thirty birds and move through vegetation quickly. Benson and
Penny considered the White-eye to have no particular habitat
preference but data obtained by myself indicate that Casuarina woodland
firstly and mangrove secondly are preferred feeding grounds (Frith -
in preparation).

3

Breeding

Reproductive activity was not noted until early December (fig. 1),
considerably later than Nectarinia, and eggs were still being laid
during late March 1973. Nests have been described in detail by Benson
and Penny. I found them in all habitats except mangroves, and only
rarely in palm groves.

Five complete clutches consisted of three eggs. A previous record
of a two egg clutch (Abbot - in Ridgway, 1895) was possibly an
incomplete or part predated one. Six clutches were observed and all
were predated, almost certainly all by rats (particularly abundant
about the research station where the nests concerned were located).

One pair of birds was observed to produce three successive clutches, a
new nest being built, quite close to the last, for each clutch.

The pale green eggs are laid on successive days. Average size of
nine eggs (three complete clutches) were 15.5 x 12.0 mm., the extreme
length and width being 17.1 and 11.5 mm. respectively. Both sexes
incubate. Average weight of twenty-two live birds was 8.3 g. varying
between 7.2 and 10.2 g.

Summary

ie Various life history notes on some Aldabra land birds were
collected, incidentally to other work, during a twelve month residence
on Aldabra Atoll. The majority of data presented concerns breeding
biology and is new or supplementary to previous knowledge.

Bis Data are given for Falco newtoni, Dryolimnas cuvieri, Alectroenas
Sganzini, (Centropus toulou), Dicrurus aldabranus, Corvus albus,
Nectarinia sovimanga and Zosterops maderaspatana.

8}, Breeding seasons of the more insectivorous bird species were found
to coincide with an increase in insect abundance except in Nectarinia
sovimanga. It is suggested that the latter species may commence
breeding as a result of an increase in nectar abundance, prior to an
increase in insects.

14

Acknowledgements

I thank all those present with me on Aldabra for bringing many and
various points to my attention. In particular I thank Mr. L. Mole who
utilised much unintended time on Aldabra searching for bird nests,
including a total of fourteen hours spent watching activity at a coucal
nest. Thanks also to Antonio Constance and Harry Charles whose
observations and skill with ropes enabled me to frequently examine
kestrel nests.

I am very grateful to staff of The Royal Society of London and to
the staff of the British Museum (Natural History) concerned with
Aldabra, and the ornithological department, for assistance in numerous
respects. Thanks to Dawn W. Frith for unpublished figures of insect
abundance for Aldabra.

tks)

References

Benson, C.W., 1967. The birds of Aldabra and their Status. Atoll
RES sees Gian oles (GS — Mela

Benson, ©.W.; and Penny, M.J: 1971. “The ‘land birds of Alidabra. Phil.
Buanse ROU, SOC. Lond. Bi: \260: 417-527.

Beith, (C.B- 1974. New and interesting observations of migrant and
vagrant birds for Aldabra, Farquhar and Astove atolls, Indian
Ocean awe. Beit. ‘Ornith. Club.’ 12-19.

Frith, C.B. In press. A twelve month study of the Aldabra Fody,
Foudia eminentissima aldabrana (Ploceidae).

Frith, €.B. 1975... Predation upon hatchlings and eggs of the Green
Turtle, Chelonia mydas, on Aldabra Atoll, Indian Ocean. Atoll
RES maeBUdelk | M6 Sian LD"

Frith, C.B. In preparation. Feeding ecology of some land birds on
West Island, Aldabra Atoll, Indian Ocean: a preliminary survey.

Goodwin, D. 1967. Pigeons and doves of the world. London. Trustees
of the British Museum (Natural History).

Penny, M.J. and Diamond, A.W. 1971. The White-throated Rail
Dryolmnasicuvrler!t (on Alidabra. Phil. Trans. Roy. Soc. fond. Bi,
260: , 529-548.

Ridgway, R. 1895. On birds collected by Dr. W.L. Abbott in the
Seychelles, Amirantes, Gloriosa, Assumption, Aldabra, and
adjacent islands, with notes on habits, etc.,by the collector.
PEOG. UsSi. Maen Mus.) 18s) 509-546).

Skead, C.J. 1967. The sunbirds of southern Africa. Cape Town.
Trustees of the South African Bird Book Fund.

Stoddanst, DoR. 1971. ‘Place names’ of Aldabra. |Phil. Trans. Roy. Sec.
HOnd=a Be 260%) 63-632):

WarSOny (Gabe, AUS, Rd. and Storer, Rob. 1963. Preliminary field
guide to the birds of the Indian Ocean. Washington. Smithsonian
Institution.

Malora ake Monthly insect abundance on West Island, Aldabra Atoll.
Totals are numbers caught during daylight hours by a
Johnson-Taylor Suction Trap during one week of each month
(by kind permission of Dr. Dawn W. Frith). The breeding
periods for Nectarinia, Zosterops and Foudia are illustrated.
Note that Foudia and Zosterops commence breeding with an
increase of insects but Nectarinia starts remarkably
earlier (see text).

Nectarinia

Foudia

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ATOLL RESEARCH BULLETIN
NO. 202.

CLIMATE OF ALDABRA ATOLL
by D. R. Stoddart and L. U. Mole

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

February 1977

td WOOT LID

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14.

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List of Tables

Mean monthly atmospheric pressure, mb
Mean monthly wind speed, kts

Number of occurrences of calms and winds in excess of 10 and
20 kts, 1967-1974

Maximum wind speeds recorded in each month, 1967-1974 (kts)
Highest maximum temperatures in each month, °C.

Mean maximum monthly temperatures, °C.

Mean minimum monthly temperatures, °C.

Lowest minimum temperatures in each month, °C.

Aldabra rainfall records before 1967: monthly totals in mm.
Midabrarcainnal records ) L96/—197/ 4: monthly totaliseeim mms
Frequency of daily rainfalls at Aldabra Atoll, 1968-1972

Rainfalls occurring in intensities exceeding 10, 25, 50 and
100 mm/day, 1968-1973

Duration of wet (rain recorded) and dry (no rain) spells at
Aldabra, 1968-1972

Duration of wet (more than 5mm/day) and dry (less than 5 mm/day)

spells at Aldabra, 1968-1972

Local variability of Aldabra rainfall, 1969-70 (West Island and

Dune Jean-Louis)

Mean monthly rainfall at Aldabra, Assumption and Iles Glorieuses,

mm

Records at Aldabra of (a) absolute maximum monthly temperature,

(b) mean maximum monthly temperature, (c) mean minimum monthly
temperature, (d) absolute minimum monthly temperature, (e) monthly
rainfaill, (f) mean monthly wind speed, and (g) mean monthly
atmospheric pressure, 1967-1973.

Annual records of (a) mean monthly wind speed and (b) mean monthly
atmospheric pressure at Aldabra, 1968-1973.

Mean monthly rainfall at Aldabra based on pre-1959 records,
1967-1973 records, and all available data.

Monthly rainfall records and mean monthly rainfall for Aldabra,
all available records.

Relationship between annual rainfall and number of days in the
year with raintalls of LO-25 mn, 1968-197 3.

Frequency and duration of wet and dry periods at Aldabra,

IGS VOW

ee een

CLIMATE OF ALDABRA ATOLL

by

D. R. Stoddart! and L. U. Mole?

Introduction

Weather records have been maintained by the Royal Society at
Aldabra Atoll (9°24'S., 46°20'E.), southwest Indian Ocean, continuously
Since mid-October 1967. The recording station is located on West
Island, on the northwest side of the atoll, first at the Settlement,
but after 1970 at the Royal Society Research Station 1 km to the south.
These are both leeward and rather protected situations. A single
synoptic observation is taken at O600 GMT (0900 local time).

Records for the period November 1967 to Octobe: 1968 were very
fully analysed by Farrow (1971), in the light of yeneral information
available for the southwest Indian Ocean, and the rainfall records up
to 1970 were placed in a regional context by Stoddart (1971). Records
now available since 1967 in some respects modify these preliminary
reports and also permit a more detailed analysis of ecologically signi-
ficant parameters such as rainfall frequency and duration of drought.
This paper presents an abstract of the climatic data to the end of
1974 (seven complete years), with fulleranalysis of the daily rainfall
records for the five years 1968-1972. Daily observations for the
period of record are available on request from the Royal Society.

In addition to these records J. Frazier maintained rainfall records
at Dune Jean-Louis on the exposed south coast for most of the period
July 1969-June 1970, and G. E. Farrow made records at East Channel on
the northeast side of the atoll during September-October 1968. A
more detailed investigation of local and microclimate is now being made
by Red). Hnatiuk

l Department of Geography, Cambridge University, England.
2 The Royal Society, London, England.
(Manuscript received May 1975 --Eds.)

Pressure and Winds

Farrow (1971, 69) points out that during winter pressure is high
and the Southeast Trades blow strongly, while in summer pressure is

low and winds are lighter and northwesterly. Records for 1968-74
(1969 is absent because of equipment failure) confirm this pattern
(Table. T-and 2; Hagures) Ih, lgvand 2): These show substantial agree-

ment from year to year in pressure distribution, with a maximum in

July, but considerable variability from year to year in mean monthly
wind speed. 1970, 1973 and 1974 were years with generally low wind
velocities, and 1972 a year of high velocities. 1971 was characterised
by abnormally high wind speeds in February and March. Although both
1970 and 1973 had similar wind patterns, the former was an exceptionally
dry year and the latter exceptionally wet.

Table 3 gives the number of occurrences in each month of calms,
of winds in excess of 10 kts, and of winds in excess of 20 kts at the
daily recording time. Calms are rare (less than 10 days per year
except in 1971). The frequency of winds higher than 10 kts varies
from year to year, being low in 1969, 1970, 1973 and 1974 and high in
197 1 anew G72" As with mean monthly wind speed there is no obvious
correlation with rainfall, though moderately high winds (10-20 kts) are
most characteristic of the Trade Wind months August-October. Table 4
lists the maximum wind speeds recorded in each month. Occasional
summer squalls bring speeds greater than 30 kts. The maximum wind
speed recorded is 44 kts on 21 September 1968; the maximum in most
months is less than 20 kts.

Temperature

Figure 1 a-d gives curves of absolute monthly maximum (a), mean
monthly maximum (b), mean monthly minimum (c), and absolute monthly
minimum (d) temperatures over the period of record; maximum
temperatures are missing for the period November 1969 to March 1970
because of instrument failure. Corresponding data are tabulated in
Tables 5-8. The annual range in mean monthly temperature is about
Ao. The monthly range is least in winter (July-August) and greatest
in summer (January). The 1967-74 averages of mean maximum and minimum
monthly temperatures do not rise about 31.24°C (December) or fall below
22.15°C (August), respectively. The highest temperature recorded is
3673°C.and: the; Lowest oi725°C. Temperatures fell below 20°C on only
eight occasions during the seven years 1968-74; four of these occasions
were during the winter of 1969.

Rainfall

Three complete and four incomplete years of rainfall record were
available for Aldabra before the Royal Society began recording in 1967
(Table 9). The complete years gave a mean annual rainfall of 640 mm,
with both 1958 and 1959 having less than 400 mm. Records since 1967
show a variable but substantially greater annual rainfall, with several
years in excess of 1000 mm (Table 10). The mean annual rainfall for

the Royal Society period of record is almost exactly 1070 mm, and all
records to the end of 1973 give a mean of 940.6 mm. Figures 3 gives
histograms for the pre-1967 and post-1967 records and for all data
combined. The earlier records showed considerably driers conditions
during the Trade Wind months May-August, and also during November-
December: during the last few years only the months of September and
October have been consistently dry, whereas according to earlier
records a comparably dry season had extended for nearly six months of
the year.

Figure le gives the sequence of monthly rainfalls since 1967 and
Figure 4 the actual monthly totals over the period of record. The
former shows great variability in incidence and amount from year to
year; the latter demonstrates that whereas the driest months of the
Trades are consistently dry, the wet months can be highly variable.
The main control of total annual rainfall appears to be the extent to
which rainfall during January and February is high (1969, 1973) or is
suppressed (1968, 1970), though in 1974 a substantial part of the total
fell during March and April. There is no clear correlation between
monthly rainfall and other monthly mean characteristics such as
temperature, pressure and wind. The highest monthly total so far
recorded is 423 mm (March 1950); several months have recorded zero
rainfall, but all these records are before 1967. Annual totals vary
from 547 mm (1968) to 1467 mm (1974).

Daily rainfall records permit the analysis of daily intensities

and frequencies. Table 1l gives the frequency of occurrence of

daily rainfalls in 5 mm/day class intervals for 1968-1972. 7O per
cent of all rain-days have less than 5 mm rainfall, and 90 per cent
less than 15 mm. Three days in five years have recorded more than

100 mm of rain, and the highest daily fall recorded is 165.5 mm on

7 April 1969, almost the same as the 6 inches recorded on 12 January
HSI (Spurs 8927548) During 1968-72 rain fell on 38 per cent of
the days.

Data on the frequency of daily falls in excess of 10, 25, 50 and
100 mm are given in Table 12. Normally three-quarters of all annual
rainfall comes in falls greater than 10 mm/day. There is a close
positive correlation between the annual rainfall in any year and the
number of days in the year with 10-25 mm/day rainfall (Figure 5); the
correlation with higher intensity falls is, however, erratic, presumably
because of the brevity of the record.

An important ecological correlate of these rainfall patterns is
the duration of wet and dry periods. If wetness is simply defined as
a day with recorded rain and dryness a day with no recorded rain, then
most wet and dry periods (i.e. sequences of such days) are of short
duration (Table 13, Figure 6): only one wet spell and seven dry spells
lasted more than 14 days during 1968-72 (though according to earlier
records three consecutive months passed without recorded rainfall in
1949). If, however, 5 mm/day is taken as the threshold dividing 'wet'
and 'dry' days (Table 14, Figure 6), most wet spells lasted only one

day and virtually all three days or less, whereas there was no less
than 30 dry spells lasting more than 14 days, 24 lasting more than 3
weeks, 16 more than 4 weeks, 8 more than 5 weeks, 5 more than 6 weeks,
4 more than 7 weeks, and 2 more than 8 weeks. The longest dry spell
so defined lasted 88 days. By contrast the longest wet spell lasted
7 days. These long periods of dry weather, especially during dry
years, must have profound ecological consequences.

All these rainfall records come from the station on the leeward
Side of the Atoll. Farrow (1971) drew attention to the need for
comparative data from different parts of Aldabra, to establish local
variability, and presented one month's comparison in 1968 between West
Island and East Channel. In 1969-70 J. Frazier maintained records
for most of the period July 1969 to June 1970 at Dune Jean-Louis on
the south coast. It is possible to compare the days for which he has
records (221) with Station records (Table 15). The total rainfall at
the Station over this period was 70.5 per cent of that at Dune Jean-
Louis, and the total number of rain-days at the Station was 69.5 per
cent of that at Dune Jean-Louis. The correlation between daily rain-
fall at the two sites is only 0.35. The mean rainfall per rain-day
is, however, very Similar (6.33 mm at the Station and 6.24 mm at Dune
Jean-Louis), and the higher totals on the south coast clearly arise
from rain falling there more frequently than on the leeward coast.
These data are, of course, preliminary, and more detailed work on local
climatic variability is now being carried out by R. J. Hnatiuk.

These new rainfall data modify the pattern of Indian Ocean rainfall
north of Madagascar presented by Stoddart (1971, Figure 2), making the

arid zone in the Aldabra area less evident. The Aldabra records are
strikingly parallel to those for the Iles Glorieuses to the south,
which were also not included in the earlier analysis. Battistini and

Cremers (1972, 1) show that the mean annual rainfall for the Iles
Glorieuses is almost identical to that for Aldabra over the Royal
Society period of record, though the summer period is drier on Aldabra
and the onset of the Trades is wetter (Table 16).

Othe~ Data

Daily records of maximum, minimum and dry bulb temperatures, dew
point, relative humidity, pressure, wind speed, wind direction, cloud
cover and rainfall are available from the Royal Society for the period
Since October 1967. In addition, daily records are also maintained
of visibility, vapour pressure, form and height of cloud, and ground
temperature at 1 ft depth, and could be provided on request. IBS atts}
planned to increase the scope of the recording station to World Weather
Watch standards during 1975. The standard Station records will also
be extended by additional screen and microclimate records being

maintained on a temporary basis in different parts of the atoll during
NOT 374s

Acknowledgements

The data recorded here were obtained by volunteers from the Royal
Society Expedition to Aldabra 1967-69, and subsequently by the staff
of the Royal Society Aldabra Research Station. We thank Dr. J.
Frazier for making available his daily rainfall records for Dune
Jean-Louis for 1969-70.

References

Battistini, R. and Cremers, G. 1972. Geomorphology and vegetation of
Iles Glorieuses. Atoll Res. Ball eso ollO; Eig | 9) pili:

Haceow, (Gs Ei. 1971. The climate of Aldabra Atoll. Phil. Frans:
Roy.- SOCGe EOnd 4 B E260.) 467-9 6 Las.

SPUES eh. Jia L892: Report on Aldabra to T. Risely Griffiths, Esq.,
Administrator of Seychelles Islands. Port Victoria,
Seychelles, July 19, 1891. Colonial Reports - Annual, No. 40:

Mauritius (Seychelles and Rodriguez), Annual Reports for 1889 and
1890, with a report on the island of Aldabra (London: H.M.S.O.),
46-50.

Stoddart, D.oOR.. LO7L. Rainfall on Indian Ocean coral islands.
AtLOl’ (Res. Bude. 4:7 b=215, lL- fags.

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16

Table ll. Frequency of daily rainfalls at Aldabra Atoll, 1968-1972

Percent total

Daily rainfall, mm Number of days number of
raindays
OS 487 70.38
61— ho 86 V2543
HES aL) 43 6321
167 20 14 Za@2
2 S25 15 Zio
26— 130 8 LG
Sil a 1335) 6 0.87
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18

Table 13. Duration of wet (rain recorded)

and dry

(no rain). spells at. Aldabra,..1968-1972

Number of occurrences

Duration of spell

days Wet
1 164
2 69
3} 32
4 19
5) 22
6 3
7 3
8 5
9 3
10 1
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Table 14. Duration of wet (more than 5mm/day) and dry
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Number of occurrences

Duration of spell
days Wet Dry

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ul
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i

ATOLL RESEARCH BULLETIN

NO. 203.

ANNOTATED CHECK LIST OF CORALS IN THE
MASCARENE ARCHIPELAGO, INDIAN OCEAN

by Gérard Faure

Issued by
THE SMITHSONUAN INSTUPU TION
Washington. D.C... U.S.A.

February 1977

=)

Nee

vial HALE nt

ANNOTATED CHECK LIST OF CORALS IN THE
MASCARENE ARCHIPELAGO, INDIAN OCEAN

by

Gérard Faure!

Summary’

1500 samples of corals have been collected by the author since
1969, in the different reef communities of the Mascarene archipelago,
from O to 50 meters by scuba diving. A total of 135 species (includ-
ing 6 species of non-scleractinian corals, belonging to 58 genera
(including the genera, Millepora, Heliopora, Distichopora) known to
date are listed in this paper.

Variations in the distribution of genera and species between the
three islands can be attributed primarily to a more or less great
diversity in the biota of the islands, and secondarily to the disparity
in the present status of our knowledge of the different islands.

Introduction

The Mascarene archipelago is situated in the western Indian ocean
about 700 km, to the east of Madagascar, in the longitudes 55°23' -
Gsic20) and Watttnudess 2 1o7"—" 19°40": It is composed of three volcanic
islands: Réunion, Mauritius, Rodriguez which form geologically
topographical units.

General Mascarene reef studies have been carried out by : Gardiner
(1936), Baissac and al. (1962), Pichon (1967, 1971), Faure and
Monitzacig avon COMO remeOi/aar cay ON ire m Si 4) at auae "(ClO Shae OMe)
Monitaggiuonie (lOO. O73):

There are many papers dealing with the systematics of corals in
the Mascarene archipelago (see references), specially in Mauritius,
but most of them are old and incomplete, except those of Michel (1974),
Stoddart (970), sony Maunrittuus-) = Beuggemanm (S79) tor Rodraguez, and

1 Station Marine da'Endoume, Marseille; Centre Universitaire,

St. Denis-., Reuniom:
(Manuscript wecerved June 1975 -—-Eds-)

Rosen (1971) for the three islands. The present is intended to serve
as a first annotated list of the corals of the Mascarene archipelago.

Class ANTHOZOA Ehrenberg, 1834
Subclass HEXACORALLIA Haeckel, 1896
Order SCLERACTINIA Bourne, 1900
Suborder ASTROCOENIINA Vaughan and Wells, 1943 |
Family THAMNASTERIDAE Vaughan and Wells, 1943

Genus Psammocora Dana, 1846
Subgenus Psammocora Dana, 1846

Psammocora (Psammocora) contigua Esper : Reunion, Mauritius, Rodriguez

common: Inner reef, lagoon coral formations, reek pool simmer
reef, passe (O - 5 m).
Recorded from Mauritius : Crossland (1952)
Psammocora (Psammocora) nierstrasZi Van Der Horst : Reunion (uncommon) ,
Mauritius (rare) base of spurs and grooves zone, volcanic flagstone
(5S 30) a): |
Psammocora (Psammocora) explanulata Van der Horst : Reunion
sporadic : Outer slope (base of grooves and overhanging sidewall)
(10. =,.25. mm)

Subgenus Plesioseris Duncan, 1884

Psammocora (Plesioseris) haimiana-Milne-Edwards and Haime : Reunion,

Mauritius

sporadic : Outer slope (spurs and grooves zone O- 5 m), reef \
front, outer reef flat, furrow and outfall.

Recorded from Mauritius : Ortmann (1888), Stoddart (1970),
Michel (1974).

Other species recorded from Mauritius : P. folium Umbgrove,
P. planipora Milne-Edwards and Haime, (Michel, 1974);
P. profondicella Gardiner, Crossland (1948)
P. Vaughani Yabe and Eguchi, (Michel, 1974;* + Steddanit,
INS)TAO)))-2

Family POCILLOPORIDAE Gray, 1842

Genus Seriatopora Lamarck, 1816
Seriatopora hystrix Dana : Mauritius

common : reef pool, inner slope, innér reef slope, 0 - 10 m,
outer Slope (rare,. —.30 m).
Seriatopora angulata Klunzinger : Mauritius
sporadic. \innergsillope, seety pool). inner exeek, ) O.— a Om :

Recorded from Mauritius : Ortmann -(1888), Stoddart (1970),
Machel i(ho74 ie
Other species recorded from Mauritius : S. stellata Quelch
(Steddact,. 1970; —Michel.,« 1974) 3 S. calrendnum
Ehrenberg (Ortmann, 1888).

Genus Stylophora Schweigger, 1819
Stylophora pistillata (Esper) : Mauritius, Reunion, Rodriguez :
abundant : Outer slope (O - 20 m), reef front, outer and inner
Beery tlaes ; lagoon Conall fonmatronse
Recorded from Mauritius :. Stock (1966), Michel (1974).

Stylophora mordax (Dana) : Reunion, Mauritius, Rodriguez
common : Outer slope ( O - 20 m), reef front, outer reef flat.
Recorded from Rodriguez : Brtiggemann (1879), as S. palmata
Milne-Edwards and Haime.
Stylophora subseriata Ehrenberg : Mauritius
sporadic : Inner reef, inner reef slope ( O - 20m).
Recorded from Mauritius : Stock (1966), Michel 1974).
Stylophora sp. Reunion
rare : Outer slope (volcanic flagstone, 25 - 40 m).

Genus Pocillopora Lamarck, 1818
Pocillopora brevicornis setchelli Hoffmeister : Reunion, Mauritius,

Rodriguez
sporadic Outer slope @O)— 5m), “eet! front:
Pocillopora damicornis (Linnaeus) : Reunion, Mauritius, Rodriguez

sporadic : Outer slope 20 - 40 m.
Recorded from Mauritius : Crossland (1948), Crossland (1952),
Steddane (1970) Michel (1974):
Recorded from Rodriguez : Britiggemann (1879), as P. favosa Dana
Pocillopora danae Verrill : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, O - 5 m), reef
front, outer and inner flats, reef pool.
Recorded from Mauritius : Crossland (1952), Stoddart (1970),
Michel (1974).
Pocillopora cespitosa Dana : Reunion, Mauritius, Rodriguez
common : Outer and inner reef flats, lagoon coral formations,
inner slope ( O - 10 m), outer slope (sheltered reefs,

© - 5m).
Pocillopora eydouxi Milne-Edwards and Haime : Rodriguez
rare : Reef front.
Pocillopora verrucosa (Ellis and Solander) : Reunion, Mauritius,
Rodriguez

common : Reef front, outer slope ( O - 30 m).

Recorded from Mauritius : Crossland (1952), Stoddart (1970),
Michel (1974).

Recorded from Reunion : Ortmann (1888).

Other species recorded : from Mauritius, P. damicornis favosa ?,

Stock (1966), Michel (1974); BP. grandis Dana,
Michel (1974); P. mauritiana Brltiggemann, Briiggemann
(1877), Michel (1974); P. elegans (Dana), Ortmann (1888).

from Rodriguez, P. favosa Ehrenberg (non
P. favosa Dana), Briiggemann (1879); P. grandis Dana,
Britiggemann (1879).

from Reunion, P. grandis Dana, Ortmann
(1888).

Other Genus recorded from Mauritius : Genus Madracis (Milne-

Edwards and Haime, 1849), M. kauaiensis Vaughan,
Michel, 1974).

Family ASTROCOENIDAE Koby, 1890

Genus Stylocoeniella Yabe and Sugiyama, 1935
Stylocoeniella armata Ehrenberg : Reunion, Maritius
sporadic : Outer slope ( 10:- 30m).
Recorded from Mauritius : Wells (1954), as Astocoenia stylifera
Pourtales.

Family ACROPORIDAE Verrill, 1902

Genus Acropora Okey, 1815

Acropora arbuscula (Dana) : Mauritius, Rodriguez
Sporadic ; reef pool ,, Lagoon:coral formations: ‘()O°>=254m)*<
Acropora corymbosa (Lamarck) : Reunion, Mauritius, Rodriguez
common : top of spurs (spurs and grooves zone, O - 2m), reef
FrOmite

Recorded from Mauritius : Lamarck (1816), Freycinet (1828),
Stoddart (1970), Michel (1974).
from Rodriguez : Brook (1893), Brtiggemann (1879), Vaughan (1918),
Gravier) (1911)\,/ Hoffmeister! (doZ5)
Acropora danae (Milne-Edwards and Haime) : Reunion, Mauritius,
Rodriguez:
common : Outer slope (Spurs and grooves zone, O - 10 m).
Acropora humilis (Dana) : Reunion, Mauritius, Rodriquez
common : Reef front (spur upper platform), outer reef flat.
Acropora hyacinthus (Dana) : Reunion, Mauritius, Rodriguez
common : Outer slope (O - 10 m), inner slope (O - 10 m), reef pool.
Recorded from Mauritius : as A. cytherea Dana ( = A. hyacinthus
(Dana), Wells (1954)), by Mobius and
al. (1880), Ortmann (1888), Michael.
(STA)
as A. arcuata (Brook) (= A. hyacinthus
(Dana), Hoffmeister (1925)), by
Stoddart. (1970):,..Michel (1974) %
Acropora reticulata Brook. : Reunion, Mauritius, Rodriguez:
common : Outer slope (sheltered reefs, O - 10 m), lagoon coral
formations “(© (= (27m).
Acropora pharaonis Milne-Edwards and Haime : Reunion, Mauritius,
Rodriguez
dominant : Outer slope (sheltered.reefs,O - 10 m), inner reef flat,
lagoon coral formations, inner slope, reef pool.
Recorded from Mauritius : Stoddart (1970), Michel (1974).
from Rodriguez : Brtiggemann (1879).
Acropora hemprichii (Ehrenberg) : Mauritius, Rodriguez
sporadic : Outer slope (sheltered reefs, O - 10 m), channel of the
reef flats, reef pool.
Acropora syringodes (Brook) : Reunion, Mauritius, Rodriguez:
Common : Outer slope (base of spurs and grooves zone, volcanic
Flagstone 18 .-- 50 m).
Other species recorded from Mauritius : A. appressa (Ehrenberg),
Michel (1974);

baeodactyla (Briiggemann), Stoddart (1970), Michel
(1974).
cerealis (Dana), Brook (1893), Vaughan (1918),
Michel (1974).
clathrata (Brook), Brook (1891, 1892), Michel (1974).
conccina (Brook), Brook (1891, 1892), Michel (1974).
conferta (Brook), Brook (1891, 1892), Michel (1974).
erythraea (Klunzinger), Mobius and al. (1880).
gonagra (Milne-Edwards and Haime), Briiggemann (1877),
Michel (1974).
grandis (Brook), Stoddart (1970), Michel (1974).
gravida Dana, Michel (1974).
haimei (Milne-Edwards and Haime), Mobius and al.
(1880), Michel (1974).
irregularis (Brtiggemann), Stoddart (1970), Michel
(L974).
macrostoma (Brook), Brook (1891-1892), Michel (1974).
multicaulis (Brook), Stoddart (1970), Michel (1974).
kenti (Brook), Stoddart (1970).
muricata cervicornis (Lamarck), Mobius and al. (1880),
Michel (1974).
muricata prolifera (Lamarck), Freycinet (1828),
Michel (1974).
oligocyathus (Brook), Brook (1892, 1893), Michel
(1974).
polystoma (Brook), Brook (1891, 1893), Michel (1974).
Pyramidalis (Klunzinger), Mobius and al. (1880),
Michel (1974).
rousseaui (Milne-Edwards and Haime), Michel (1974).
seriata (Ehrenberg), Brook (1893), Michel (1974).
stigmataria Milne-Edwards and Haime, Michel (1974).
symmetrica Brook, Brook (1891), Stoddart (1970),
Michel (1974).
tuberculosa Milne-Edwards and Haime, Michel (1974).
tylostoma Ehrenberg, Brook (1893), Michel (1974).
Sp, steddart (1970) .<
scherzeriana Briiggemann, Klunzinger (1879).

from Rodriguez

alces (Dana), Briiggemann (1879).

- baeodactyla (Brtiggemann), Vaughan (1918).

brevicollis Brook, Vaughan (1918).

conferta (Quelch), Vaughan (1918), Wells (1954) ?

corymbosa (Lamarck), Briiggemann (1879).

flabelliformisMilne-Edwards and Haime, Brtiggemann
(S79)

gonagra (Milne-Edwards and Haime), Brtiggemann (1879).

haimei (Milne-Edwards and Haime), Brtiggemann (1879).

microclados (Ehrenberg), Brtiggemann (1879).

plantaginea (Lamarck), Brtiggemann (1879).

pustulosa (Milne-Edwards and Haime), Brtiggemann (1879).

seriata (Ehrenberg), Brtiggemann (1879).

from Reunion:

A. granulosa Milne-Edwards and Haime, Milne-Edwards and
Haime (1860).

Genus Astreopora de Blainville, 1830.

Astreopora myriophtalma (Lamarck) : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, volcanic flagstone,
OF == 40" mir

Recorded from Mauritius : Bernard (1896), Michel (1974).

Genus Montipora Quoy and Gaimard, 1830.
Montipora edwardsi Bernard : Rodriguez
uncommon : Inner reef flat, lagoon coral formations.
Montipora foliosa (Pallas) : Reunion, Mauritius, Rodriguez.
sporadic : Lagoon coral formations, reef pool, passe, inner slope,
outer slope (sheltered reef).
Recorded from Mauritius : Ortmann (1888), Vaughan (1918), Thiel
(1932), Crossland ((1952)-, Bernard (1897)x%
Montipora informis Bernard : Reunion
rare : Inner reef flat, lagoon coral formations.
Montipora verrucosa (Lamarck) : Reunion, Mauritius.
sporadic : Outer slope (spurs and grooves zone, volcanic flagstone,
12-- 50 :m)-.
Recorded from Mauritius : Ortmann (1888, 1892).
Montipora cf tuberculosa (Lamarck) : Reunion, Mauritius, Rodriguez
Recorded from Mauritius : Ortmann (1888-1889-1892).
sporadic : Outer slope(spurs and grooves zone), outer and inner

reef flats.
Montipora cf millepora Crossland : Mauritius
rare : outer slope (volcanic flagstone - 48 m).

Other species recorded from Mauritius

M. explanata Brliggemann, Bernard (1897), Brtiggemann (1879),
Michel (1974).

M. expansa Dana, Ortmann (1888).

M. guppyi Bernard, Stoddart (1970), Michel (1974).

M. lanuginosa Bernard, Bernard (1897), Michel (1974).

M. lobulata Bernard, Bernard (1897), Michel (1974).

M. mammifera Bernard, Michel (1974).

M. rus (Forskal), Ortmann (1888).

M. solanderi Bernard, Bernard (1897), Michel (1974).

M. stylosa (Ehrenberg), Ortmann (1888-1889).

M. undata Bernard, Stoddart (1970), Michel (1974).

From Rodriguez

M. explanata Brtiggemann, Brtiggemann (1879), Bernard
GLS97.)s

M. divaricata Britiggemann, -Briiggemann (1879), Bernard
(ESO7)::

M. incrustans Briiggemann, Briiggemann (1879), Bernard

(ESISIT NN
M. lichen Dana, Britiggemann (1879), Bernard (1897).
M. perforata Bernard, Bernard (1897) = M. explanata
Brtiggemann.

M. solanderi Bernard, Bernard (1897) non M. foliosa
(Pallas) Brtiggemann (1879).

from Mascarene

M. spongodes Bernard, Bernard (1897).
M. subtilis Bernard, Bernard (1897), Wells (1954).

Suborder FUNGIINA Verrill, 1865.
Superfamily AGARICIIDAE Gray, 1847.
Family AGARICIIDAE Gray, 1847.

Genus Pavona Lamarck, 1801.
Subgenus Pavona Lamarck, 1801.
Pavona (Pavona) cactus (Forskal) : Reunion, Mauritius, Rodriguez
Sporadic : Inner reef flat, reef pool, O - 5 m.
Recorded from Mauritius : Michel (1974).
Pavona danai Milne-Edwards and Haime : Reunion, Rodriguez
rare 7 Inner nreet flat, reef pool, O —- 5 m.
Pavona (Pavona) decussata Dana : Reunion, Mauritius
sporadic : Inner reef flat, reef pool, O - 5 m.
Pavona (Pavona) clavus Dana : Reunion, Mauritius, Rodriguez
sporadic : Outer slope, O - 30 m (spurs and grooves zone,
volcanic flagstone), lagoon coral formations (O - 5 m).
Pavona (Pavona) divaricata (Lamarck) : Reunion, Mauritius Rodriguez
common : Inner reef flat, reef pool, lagoon reef.
Pavona (Pavona) explanulata (Lamarck) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs with overhanging sidewall, reef
gallery and reef tunnel) - 5 - 30 m.
Pavona (Pavona) variens (Verrill) : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, O - 15 m).
Outer and inner reef flat.
Recorded from Mauritius : Ortmann (1889), as P. repens Brtiggemann.
Other species recorded from Mauritius

P. cristata Ellis and Solander, Michel (1974) = P. cactus
(Forskal).

P. cristata Milne-Edwards and Haime, Michel (1974)
= P. cactus (Forskdl).

P. frondifera Lamarck, Stoddart (1970), Michel (1974).

P. praetorta Dana, Stoddart (1970), Michel (1974).

P. seriata Briiggemann, Stoddart (1970), Michel (1974).

from Rodriguez
P. lata Dana, Matthai, 1974.

P. cristata Ellis and Solander, Brtiggemann (1879)
= P. cactus (Forskal).

Subgenus Pseudocolumnastraea Yabe and Sugiyama, 1933.
Pavona (Pseudocolumnastraea) pollicata Wells : Reunion:
rare : Outer slope of sheltered reefs, 5 - 15 m.

Genus Leptoseris Milne-Edwards and Haime, 1849.
Leptoseris columna Yabe and Sugiyama : Reunion
rare : Outer slope (volcanic flagstone, stone and nodule
pavement, 35 - 60m).
Leptoseris hawaiensis Vaughan : Reunion, Mauritius

rare : Outer slope (volcanic flagstone, stone and nodule
pavement, 35 - 60 m).
Leptoseris incrustans (Quelch) : Reunion, Mauritius, Rodriguez

sporadic : Outer slope (overhaning sidewall, reef tunnel, reef
gallery, volcanic flagstone, 15 - 45 m).
Leptoseris scabra Vaughan : Reunion

rare : Outer slope (stone and nodule pavement, - 60 Mm).
Leptoseris sp : Mauritius :
rare : Outer slope (volcanic flagstone, - 45 m).

Genus Leptoseris ? Milne-Edwards and Haime
Leptoseris ? mycetoseroide Wells : Reunion, Mauritius
sporadic : Outer slope (reef tunnel, reef gallery, overhanging
sidewall, 15 - 40 m).
Other species recorded from Mauritius : Leptoseris digitata
Vaughan, Stoddart (1970),
Michel (1974).
from Reunion : Leptoseris fragilis Milne-
Edwards and Haime, Milne-
Edwards and Haime (1860).,
Matthai (1924).

Genus Agariciella Ma,
Agariciella ponderosa Gardiner : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone, 10 - 30 m), passe
(2 Ee Ol mii

Genus Pachyseris Milne-Edwards and Haime, 1849.
Pachyseris speciosa (Dana) : Reunion, Mauritius, Rodriguez.
common : Outer slope, 10 - 40 m (Spurs and grooves zone, volcanic
flagstone), passe (2 - 15 m).
Recorded from Mauritius : Stoddart (1970), Michel (1974).
Pachyseris rugosa (Milne-Edwards and Haime) : Reunion
rare : Outer slope (15 - 40 m).
Other species recorded from Mauritius : P. laevicollis (Dana),
Michele (L974) %

Family SIDERASTREIDAE Vaughan and Wells, 1943.

Genus Coscinarea Milne-Edwards and Haime, 1848.
Coscinarea cf. columna Dana : Reunion, Rodriguez
rare : Outer slope (spurs and grooves zone of sheltered reefs),
reef pool, passe.

Coscinarea monile (Forskal) : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, volcanic flagstone,
10 = 30 m), passe.
Recorded from Mauritius : Ortmann (1892).
Coscinarea sp. : Mauritius
rare SS elnner Geek) — os) m.

Genus Horastrea Pichon, 1971.
Horastrea indica Pichon : Reunion, Mauritius
sporadic : Outer slope (volcanic flagstone, 15 - 40 m).
Recorded from Reunion : Pichon, 1971.

Superfamily FUNGIICAE Dana, 1846
Family FUNGIIDAE Dana, 1846.

Genus Cycloseris Milne-Edwards and Haime, 1849.
Cycloseris cyclolites (Lamarck) : Reunion, Mauritius
sporadic : Outer slope (base of spurs and grooves zone, and
specially volcanic flagstone, 25 - 40 m).
Recorded from Mauritius and Rodriguez : Matthai, (1924).

Genus Fungia Lamarck, 1801.

Subgenus Fungia Lamarck, 1801.
Fungia (Fungia) fungites (Linnaeus) : Reunion, Mauritius, Rodriguez:
common : (Mauritius and Rodriguez) : Branched coral reef flat,
reef pool, inner reef flat, lagoon coral formations,
inner slope, O - 5 m.
rare : Reunion.
Recorded from Mauritius : Stoddart (1970), Michel (1974).

Subgenus Danafungia Wells, 1956.

Fungia (Danafungia) danai Milne-Edwards and Haime : Reunion, Mauritius,
Rodriguez
sporadic : Outer slope (O - 10 m), branched coral reef flat, passe.

Subgenus Pleuractis Verrill, 1864.

Fungia (Pleuractis) scutaria Lamarck : Reunion, Mauritius
common : Outer slope (20 - 40 m).
Recorded from Mauritius : Michel (1974).
Fungia ? somervillei Gardiner : Mauritius
rare >, Outer sillope, —.50.m.
Other species recorded from Mauritius : F. agariciformis Lamarck,
Michel (1974).
F. madagascarensis Vaughan,
Stoddart (1970), Michel
(1974).
F. patella (Ellis and
Solander), Matthai (1924);

10

F. samboangensis Vaughan,
Stoddart, (970),
Michel (1974).

from Rodriguez:

F. dentigera Leuckart,
Brluggemann (1879);
F. haimei Verrill,
Briiggemann (1879).

Genus Herpolitha Eschscholtz, 1826.

Herpolitha limax (Esper) : Reunion, Mauritius, Rodriguez
common : (Mauritius : Outer slope (sheltered reef, 5 - 15 m), reef
pool, lagoon, "inners “slope (O'=. 15am)
rare (Reunion, Rodriguez) : Outer slope (-20 - 40 m).

Recorded from Mauritius : Matthai (1924), Yabe and Sugiyama (1941),
Stoddart (1970), Michel (1974).

Genus Podabacia Milne-Edwards and Haime, 1849.
Podabacia crustacea Pallas : Reunion, Mauritius
rare : Outer and inner reef slopes, passe, 15 - 20 m.

Superfamily PORITICAE Gray, 1842.
Family PORITIDAE Gray, 1842.

Genus Porites Link, 1807.
Porites (Porites) somaliensis Gravier : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, volcanic flagstone,
O - 50m), reef flat (outer and inner reef flats, micro-
atoll zone, lagoon coral formations, reef pool, passe,

O - 20 n.
Recorded from Mauritius : Michel (1974).
Porites (Porites) solida (Forskdl) : Reunion, Mauritius, Rodriguez:

common : Outer slope (spurs and grooves zone, volcanic flagstone,
O - 40 m., reef flat, lagoon coral formations, “eek (pool,
On—s ESmmt.
Recorded from Mauritius : Ortmann (1892, 1888), Stoddart (1970).,
Crossland (1948), Michel (1974).
Porites (Porites) pukoensis Vaughan : Reunion, Rodriguez
rare : Reef flat with scattered coral growth.
Porites (Porites) nigrescens Dana : Reunion, Mauritius
sporadic : Lagoon coral formations, reef pool, passe, outer slope
(sheltered reefs, O - 10 m).
Porites (Porites) andrewsi Vaughan : Mauritius
rare : Lagoon coral formations, outer slope (- 45 m).
Other species recorded from Mauritius
P. columnaris Klunzinger, Ortmann (1888).
P. lichen Dana, Stoddart (1970), Michel (1974).
P. lobata Dana, Stoddart (1970), Michel (1974).
P. lutea Milne-Edwards and Haime, Stoddart (1970),
Michel (1974).

dal

P. mauritiensis prima Bernard, Bernard (1905),
Michel (1974).
P. mauritiensis secunda Bernard, Bernard (1905),
Michel (1974).
P. mauritiensis quarta Bernard, Bernard (1905),
Michel (1974).
P. mauritiensis guinta Bernard, Bernard (1905),
Michel (1974).
Recorded from Rodriguez:
P. arenosa (Esper), Briiggemann (1879) = P. rodericensis
secunda Bernard,
Bernard, (1905),
Crossland (1948).
P. lutea Milne-Edwards and Haime, Britiggemann 1879 =
P. rodericensis
prima Bernard,
Bernard, (1905).
from Reunion :
P. arenosa (Esper), Milne-Edwards and Haime (1860).

Subgenus Synarea Verrill, 1864.
Porites (Synarea) iwayamaensis Eguchi : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone of sheltered reefs,
O - 25 m), passe, inner reef flat, microatoll zone,
reef pool.
Porites «<Synarea) cf. danae Milne-Edwards and Haime : Reunion, Mauritus,
Rodriguez
sporadic : Lagoon coral formations, microatoll zone.
Recorded from Mauritius : Ortmann (1888), Michel (1974).

Genus Goniopora Blainville, 1830.
Goniopora lobata Milne-Edwards and Haime : Reunion, Mauritius,
Rodriguez
sporadic : Outer slope (Spurs and grooves zone of sheltered reefs
(C= 20 m)F,, passe.
Goniopora savigny Milne-Edwards and Haime : Reunion
uncommon : Outer slope (Spurs and grooves zone of sheltered reefs,
el Siem) lagoon (Or — 2 1m)
Recorded from Mauritius : Michel (1974).
Goniopora cf somaliensis Vaughan : Reunion

rare 7 Outre slope, — 15 m:
Goniopora tenuidens Quelch : Reunion, Mauritius
rare : Outer slope (spurs and grooves zone, 5 - 15 m).

Other species recorded from Mauritius : G. mauritiensis prima
Bernard, Bernard (1903),
Michel (1974).

Genus Alveopora Blainville, 1830.
Alveopora cf mortenseni Crossland : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone of sheltered reefs,
O - 20 m), passe.
Recorded from Mauritius : Crossland (1952).

2)

Alveopora sp : Reunion, Mauritius

rare : Outer slope (volcanic flagstone, 25 - 40.m).

Other species recorded from Mauritius
A. daedala Blainville, Ortmann (1888), Michel (1974).
A. retepora Blainville, Ortmann (1888), Michel (1974).
A. verilliana Dana = A. daedala Blainville, Stoddart

(1970), Michel (1974).

A. octoformis Blainville, Ortmann (1888).

Suborder FAVIINA Vaughan and Wells, 1943.
Superfamily FAVIICAE Gregory, 1900.
Family FAVIIDAE Gregory, 1900.
Subfamily FAVIINAE Gregory, 1900.

Genus Plesiastrea Milne-Edwards and Haime, 1848.
Plesiastrea versipora Lamarck : Reunion, Mauritius
rare : Outer slope (spurs and grooves zone, 10 - 20 m.).
Recorded from Rodriguez : Brtiggemann (1879), as P. quatrefagesana
Milne-Edwards and Haime, Chevalier
CAO Aes
Other species recorded from Mauritius : P. peroni Milne-Edwards
and Haime (near
P. versipora Lamarck),
Ortmann (1888).

Genus Favia Oken, 1815.
Favia favus (Forskdal) : Reunion, Mauritius, Rodriguez
sporadic : Inner reef flat (reef flat with scattered coral growth),
lagoon coral formations, reef pool, inner slope,
O2—-5..mé
Recorded from Mauritius : Ma (1959), Chevalier (1971), Wijsman-
Best (1972), Michel (1974).
from Rodriguez : Brtiggemann (1879), as F. denticulata
Ellis and Solander and F. affinis
Milne-Edwards and Haime.
Favia pallida (Dana) : Reunion, Mauritius, Rodriguez
common : Outer slope (Spurs and grooves zone, volcanic flagstone,
O - 40 m), Outer and inner reef flats.
Favia speciosa (Dana) : Reunion, Mauritius, Rodriguez
sporadic : Same repartition as F. pallida.
Recorded from Mauritius : Stoddart (1970), Michel (1974).

Favia stelligera (Dana) : Reunion, Mauritius, Rodriguez
sporadic : Reef front, outer slope (spurs and grooves zone, X
©; = WoOgm)r.

Recorded from Mauritius : Ortmann (1888), as F. lobata Milne-
Edwards and Haime, Chevalier (1971).
from Rodriguez : Briiggemann (1879), as F. lobata Milne-
Edwards and Haime.

Favia laxa (Klunzinger) : Reunion:
rare : Outer slope (O - 10 m).
Favia cf hombroni (Rousseau) : Reunion, Mauritius, Rodriguez

sporadic : Outer slope (spurs and grooves zone, 5 - 20 m).

13

Genus Favites Link, 1807.

Favites abdita (Ellis and Solander) : Reunion, Mauritius, Rodriguez
Sporadic : Outer slope (O - 40 m), outer and inner reef flats,
passe.

Recorded from Mauritius : Ortmann (1888), as Prionastraea gibbosa
Klunzinger, Ortmann (1892), as
Goniastrea seychellenis! Milne-Edwards
and Haime, Crossland (1952), Chevalier
(eID e
Favites pentagona (Esper) : Reunion, Mauritius, Rodriguez
common : Outer slope (O - 40 m).
Favites vivens (Dana) : Reunion, Mauritius, Rodriguez :
sporadic : Outer slope (O - 40 m).
Other species recorded from Rodriguez : Prionastraea rodericana
Briiggemann, Briiggemann
(1879); BP. scabra
Britiggemann, Briiggemann
(2879)-.

Genus Goniastrea Milne-Edwards and Haime, 1848.
Goniastrea retiformis (Lamarck) : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, O-15 m), reef front,
outer and inner reef flat.
Recorded from Mauritius : Ma (1959), Chevalier (1971), Michel
(1974).
Goniastrea pectinata (Ehrenberg) : Reunion, Mauritius, Rodriguez
abundant : Outer slope (spurs and grooves zone, O - 20 m), reef
front, outer and inner reef flats, reef pool, passe.
Recorded from Mauritius : Stoddart (1970), Michel (1974).
Goniastrea of incrustans Duncan : Reunion, Mauritius, Rodriguez :
sporadic : Outer slope (5 - 30m).

Genus Caulastrea Dana, 1846.
Caulastrea tumida Matthai : Reunion, Mauritius

sporadic : Outer slope (volcanic flagstone, 25 - 40 m).

Genus Oulophyllia Milne-Edwards and Haime, 1848.

Oulophyllia crispa (Lamarck) : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, volcanic flagstone,
1G) SO lam)
Oulophyllia cf aspera Quelch : Mauritius
rare : Outer slope (volcanic flagstone, - 35m).

Genus Platygyra Ehrenberg, 1834.
Platygyra daedala (Ellis and Solander) : Reunion, Mauritius, Rodriguez
abundant : Outer slope (O - 40 m), reef front, inner and outer reef
flats, lagoon coral formations, reef pool, passe.
Recorded from Mauritius : Stoddart (1970), Wijsman-Best (1972),
Michel (1974).

laccording to Chevalier (1971), Wijsman-Best (1972).

14

Recorded from Mauritius and Mascarene islands : Yabe and al.
(1936).
Recorded from Rodriguez : Matthai 1928, Brltiggemann (1879), as
P. esperi Milne-Edwards and Haime.

Platygyra lamellina (Ehrenberg) : Reunion, Mauritius, Rodriguez
sporadic : Inner reef flat, lagoon coral formations, passe, reef
pool.

Recorded from Mauritius : Michel (1974).

Genus Leptoria Milne-Edwards and Haime, 1848.
Leptoria phrygia (Ellis and Solander) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone, O - 15 m), reef
front, outer and inner reef flats.
Recorded from Mauritius : Matthai (1928), Crossland (1952),
Michel (1974).
Other species recorded from Mauritius : Leptoria gracilis (Dana),
Ortmann (1888), Matthai
(1928), Yabe and al.
G986)
From Rodriguez : Leptoria gracilis (Dana),
Briiggemann (1879), as
L. tenuis Milne-Edwards
and Haime, Matthai (1928).

Genus Hydnophora Fischer de Waldheim, 1807.
Hydnophora microconos (Lamarck) : Reunion, Mauritius, Rodriguez
common : Outer slope (spurs and grooves zone, volcanic flagstone,
© '="35 .m)",> reef Eront,. Outer and/ inner reek flatsr
Recorded from Rodriguez : Brliggemann (1879).
Hydnophora exesa (Pallas) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone, O - 15 m), lagoon
coral formations, reef pool, passe.
Recorded from Mauritius and Rodriguez : Matthai (1928).
Recorded from Mauritius and Mascarene islands : Yabe and al. (1936).

Subfamily MONTASTREINAE Vaughan and Wells, 1943.

Genus Montastrea

Montastrea curta (Dana) : Reunion, Mauritius
rare : Outer slope (spurs and grooves zone, O - 15 m), reef
front.

Genus Leptastrea Milne-Edwards and Haime, 1848.
Leptastrea purpurea (Dana) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone, O - 15 m), reef
front, outer and inner reef flats.
Leptastrea transversa (Klunzinger) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves, O - 20 m), reef front.
Leptastrea bottae (Milne-Edwards and Haime) : Reunion, Mauritius
rare : Reef flat with scattered coral growth.
Other species recorded from Mauritius : L. ehrenbergiana Milne-
Edwards and Haime, Ortmann
(1888).

15,

Genus Cyphastrea Milne-Edwards and Haime, 1848.
Cyphastrea microphtalma : Reunion, Mauritius, Rodriguez
common : Inner reef flat, lagoon coral formations, O - 5 m.
Cyphastrea sairailia (Forskdl) : Reunion, Mauritius, Rodriguez
Sporadi¢ : Outer slope (spurs and grooves zone, O - 15 m)-.
Recorded from Mauritius : Ortmann (1888).
Other species recorded from Mauritius : C. chalcidicum (Forsk4l),
Michel (1974).

Genus Echinopora Lamarck, 1816.
Echinopora gemmacea (Lamarck) : Reunion, Mauritius, Rodriguez
Abundant : Outer slope (O - 30 m), outer and inner reef flats,
lagoon coral formations, reef pool, passe, O - 15 m.
Recorded from Mauritius : Milne-Edwards and Haime (1857), as
E. ehrenbergii Milne-Edwards and Haime,
Ortmann (1888); as E. ehrenbergii
Milne-Edwards and Haime, Thiel (1932),
Michel (1974).
Other species recorded from Mauritius : FE. lamellosa (Esper),
Stock (1966), Michel
(1974).
from Rodriguez : E. spinulosa Briiggemann,
Britiggemann (1879), near
E. gemmacea Lamarck.

Family ASTRANGIIDAE Verrill, 1869.

Genus Culicia Dana, 1846.
Culicia cuticula Klunzinger : Reunion, Mauritius, Rodriguez
uncommon : On the under face of massive and encrusting forms,
outer slope (O - 30 m), reef front, passe.

Family OCULINIDAE Gray, 1847.
Subfamily Galaxeinae Vaughan and Wells, 1943.

Genus Galaxea Oken, 1815.
Galaxea fascicularis (Lamarck) : Reunion, Mauritius, Rodriguez
common : Outer slope (O - 30 m), reef front, outer and inner reef
flats, reef pool, passe.
Recorded from Mauritius : Ortmann (1888, 1892), Thiel (1932),
Stoddart (1970), Chevalier (1971),
Michel (1974).

from Mauritius and Mascarene islands : Yabe and al. (1936).

from Rodriguez : Brtiggemann (1879), Thiel (1932),
Chevalier (1971).
Other species recorded from Mauritius : G. musicalis (Linné) =
G. clavus (Dana), Stoddart
(1970), “Michel (1974) ;
G. hexagonalis Milne-

Edwards and Haime, Stoddart

(1970), Michel (1974).

]
|
|

|

|
|
a
|

16
Family MUSSIDAE Ortmann, 1890.

Genus Acanthastrea (Milne-Edwards and Haime, 1848).
Acanthastrea echinata (Dana) : Reunion, Mauritius, Rodriguez
sporadic ; Outer slope (O = 30m), reef: pool, passe innerusilope;
O) Sy ILS). tan
Other species from Rodriguez : A. angulosa Brtiggemann, Britiggemann
(auish7/S)))

Genus Lobophyllia de Blainville, 1830.

Lobophyllia corymbosa (Forskdal) : Reunion, Mauritius, Rodriguez :
sporadic : Outer ‘slope (sheltered reef, O -— 20 m), pasSe;.neet
pool.

Recorded from Mauritius : Milne-Edwards and Haime (1849), as
L. rudis Milne-Edwards and Haime,
Ortmann (1888), Matthai (1928),
Crossland (1952).
from Mauritius and Mascarene islands : Yabe and al. (1936).
from Rodriguez : Brtiggemann (1879), as L. umbellata
Brtiggemann, Matthai (1928), Crossland

G1iSI512))) =
Lobophyllia costata (Dana) : Mauritius
rare 7eOuceq ESOpes(S =" 20) m)
Lobophyllia hemprichii (Ehrenberg) : Reunion, Mauritius, Rodriguez

common : Outer slope (10 - 40 m), passe (O - 10 m).
Recorded from Mauritius : Stoddart (1970), Michel (1974).

Genus Symphyllia Milne-Edwards and Haime, 1848.
Symphyllia recta (Dana) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves, volcanic flagstone,
i571 40m); passe’, (@) >25 m)r:

Genus Parascolymia Wells
Parascolymia vitiensis (Brtiggemann) : Reunion, Mauritius
common : Outer slope (volcanic flagstone, 20 - 40 m).

Family PECTINIIDAE Vaughan and Wells, 1943.

Genus Echinophyllia Klunzinger, 1879.

Echinophyllia aspera (Ellis and Solander) : Reunion, Mauritius,
Rodriguez
sporadic : Outer slope (spurs and grooves zone, volcanic flagstone,
LOV=74Oxm) ke.

Genus Oxypora Saville Kent, 1871.
Oxypora lacera (Verrill) : Reunion, Mauritius, Rodriguez
sporadic : Outer slope (spurs and grooves zone, volcanic flagstone,
LOe=4Onm)y, passe™ (5e— 205 em)

Genus Mycedium Oken, 1815.
Mycedium tenuicostatum (Verrill) : Rodriguez
rare : Outer slope (sheltered reef, 15 - 20 m).

17

Recorded from Mauritius : Stoddart (1970), Michel (1974).
Mycedium elephantotum (Pallas) : Mauritius
rare : Passe and reef pool, 5 - 15 m.

Genus Pectinia Oken, 1815.
Pectinia cf. lactuca (Pallas) : Reunion
rare : Outer slope (volcanic flagstone, -35 m).

Suborder CARYOPHYLLIINA Vaughan and Wells, 1943.
Superfamily CARYOPHYLLIICAE Gray, 1847.
Family CAROPHYLLIIDAE Gray, 1847.

Genus Euphyllia Dana, 1846.
Euphyllia glabrescens (Chamisso and Eysen Hardt) : Mauritius
rare ? Passe, -ree& pools, 2) — 5m.

Genus Plerogyra Milne-Edwards and Haime, 1848.
Plerogyra sinuosa Dana : Mauritius
rare : Reef pool, passe, 2 - 15 m.

Genus Gyrosmilia Milne-Edwards and Haime, 1851.
Gyrosmilia interrupta (Ehrenberg) : Reunion, Mauritius
uncommon : Outer slope (sheltered reef, 5 - 40 m), passe, reef
pool, creek

Suborder DENDROPHYLLIINA Vaughan and Wells, 1943.
Family DENDROPHYLLIIDAE Gray, 1847.

Genus Dendrophyllia de Blainville, 1830.
Dendrophyllia nigrescens Dana : Mauritius
rare : Passe, 15 - 20 m.
Recorded from Mauritius : Stoddart (1970), Michel (1974).
Dendrophyllia sp. 1 : Reunion

rare : Outer slope (sheltered reef) - 15 m.
Dendrophyllia sp. 2 : Mauritius
rare : Passe, - 20 m.

Other species recorded from Mauritius : D. ehrenbergiana Milne-
Edwards and Haime, Ortmann
(1889); D. manni (Verrill),
Stoddart (1970), Michel
(1974), D. profunda
(Pourtales), Stoddart
(O70) ye Michelan(LS 74) x

from Rodriguez : D. ehrenbergiana Milne-
Edwards and Haime,
Brtiggemann (1879).

: Genus Tubastrea Lesson, 1834.
Tubastrea aurea (Quoy and Gaimard) : Mauritius, Rodriguez
sporadic : Reef front, outer slope (O - 5 m).
Recorded from Mauritius as D. coccinea (Michelin) by : Michelin
(L845), Stoddart’ (970) Michele (1974):

18

Tubastrea micrantha (Ehrenberg) : Mauritius, Rodriguez
uncommon : Passe (10 —- 20 m).
Recorded from Mauritius : Ortmann (1888), Stoddart (1970),
Michel (1974).

Genus Turbinaria Oken, 1815.

Turbinaria mesenterina (Lamarck) : Reunion, Mauritius, Rodriguez

sporadic : Outer slope (sheltered reef, O - 5 m), reef front,

outer reef flat.
Recorded from Mauritius : Ortmann (1888).
from Rodriguez : Brtiggemann (1879), Wells (1954),
Bernard (1896), Vaughan (1918).

Turbinaria crater (Pallas) : Mauritius

rare : Outer slope (volcanic flagstone, 40 m).
Turbinaria peltata Esper : Reunion, Mauritius

common : Outer slope (volcanic flagstone, 20 - 50m).

Recorded from Mauritius : Bernard (1896), Vaughan (1918).
Turbinaria cf. irregularis : Reunion, Mauritius, Rodriguez

sporadic : Outer slope (volcanic flagstone, 20 - 35 m).

Recorded from Mauritius : Wells (1954), Bernard (1896).
Turbinaria cf. porcellanea Bernard : Rodriguez

rare : Outer slope (sheltered reef, - 20 m), passe, - 20 m.
Turbinaria sp. : Mauritius
Gare : Outer slope (spurs and grooves zone, - 20 m).

Genus Heteropsammia Milne-Edwards and Haime, 1848.
Heteropsammia michelini Milne-Edwards and Haime.
Recorded from Mauritius : Stoddart (1970), Michel (1974).

Subclass OCTOCORALLIA Haeckel
Order COENOTHECALIA Bourne, 1895.
Family HELIOPORIDAE Moseley, 1876.

Genus Heliopora Moseley, 1876.

Heliopora coerulea (Pallas) : Mauritius, Rodriguez
uncommon : Outer slope (8 - 12 m), outer reef flat, lagoon coral
formations.

Recorded from Mauritius : Michelin (1845), Michel (1974).
from Rodriguez : Tixier-Durivault (1972).

Class HYDROZOA Owen
Order MILLEPORINA Hickson
Family MILLEPORIDAE Flemming

Genus Millepora Linnaeus
Millepora platyphylla Ehrenberg and Hemprich : Reunion, Mauritius,
Rodriguez
common : Outer slope (spurs and grooves zone, O - 15 m), reef
front, outer reef flat.

Recorded from Mauritius : Ortmann (1892), as M. verrucosa Milne-
Edwards and Haime, Boschma (1957),
Michel (1974).

iL!)

from Rodriguez : Briiggemann (1879), as M. verrucosa |
Milne-Edwards and Haime.
Millepora exaesa Forskdl : Rodriguez
rare : Inner reef flat
Recorded from Rodriguez : Britiggemann (1879), as M. gonagra Milne-
Edwards and Hajime.
Millepora intricata Milne-Edwards and Haime : Mauritius, Rodriguez
sporadic : Inner slope, reef pool, 2 - 8 m.
Millepora tenera Boschma : Mauritius, Rodriguez
sporadic : Inner slope, reef pool, lagoon coral formations, 2 - 5m.
Recorded from Mauritius : Boschma (1957), Michel (1974).
Other species recorded from Mauritius : M. dichotoma Forskdl,
Michel (1974); M. truncata
Dana, Michel (1974);
M. tuberosa Boschma,
Boschma (1966), Michel
(1974).

Order STYLASTERINA Hickson and England
Family STYLASTERIDAE Gray
Subfamily DISTICHOPORINAE Stechow

Genus Distichopora Lamarck |
Distichopora violacea (Pallas) : Reunion ?, Mauritius, Rodriguez
uncommon : Outer slope (spurs and grooves zone, reef tunnel
overhanging sidewall, 2 - 10 m).
Recorded from Mauritius : Michelin (1845), Michel (1974).

Conclusions

The list given here cannot be exhaustive for various reasons:
first some of the specimens collected are now being identified and do
not appear here. Second some genera and species still await discovery
in this part of the Indian Ocean. Thus only in the most recent studies
of the outer slope of the reefs of the archipelago has the presence of
the following genera been detected : Stylocoeniella, Culicia,
Caulastrea, Gyrosmilia, Pectinia, Pseudocolumnastrea.

In order not to increase the risk of errors we have mentioned
separately the species appearing in the lists of Stoddard (1970) and
Michel (1974), with the exception of those species actually seen during
the present survey. In fact in Michel's list only 50 species, belong-
ing to 22 genera, exist in the collections of the Mauritius Institute,
Port-Louis (Mauritius) where they are represented by one or only a few
specimens of which the names originally provided by P. Boshoff. Further
48 other species (27 of which occur in the genera Acropora and Porites)
are listed or described in old papers and are in need of revision.

Though it might be premature to draw final conclusions from the
present survey (see table) some findings are evident:

20

The greater range of genera seen in Mauritius is due to a greater
variety of biotopes in the reef than exists in Réunion.

The relatively high number of genera and species found in Réunion
however is due to a better knowledge of the outer slope of the reefs of
that island.

The relatively low figures given for Rodriguez are due on the one
hand to the fact that through lack of facilities studies have been less
intensive than on the other two islands; on the other hand the presence
of an important "soil pollution" which extends from the lagoons to the
outer Slope constitutes an obstacle to the development of the coral
population both qualitatively and quantitatively. The cause of the
"pollution" is the extensive clearing of forests on the island which
moreover lies right in the path of the main tropical cyclones.

With the exception of Mauritius the lack of data concerning.the
abundance of genera and species has long led to the belief that the
Mascarene archipelago was relatively poor in scleratinian corals.

The present check list, however, shows a wealth comparable to that of
the neighbouring islands of the S.W. Indian Ocean. Rosen (1971) gives
(including inferred and doubtful genera records) the following figures
Maldives, South (60); Seychelles (57); Aldabra (55); Chagos (56);
Cargados (47); Farquhar (52); Comoros (48); Amirantes (54). Further
the results obtained to date in the Mascarenes confirm the generic
diversity model for reefs in the Indian Ocean of the same author.

The specimens on which the present provisional list have been
based are currently being compared with material previously collected
in the same part of the Indian Ocean and located in the main European
museums.

Acknowledgements

Thanks are due to my friend and diving partner Dr. L. Montaggioni

(Dept of Geology, Centre Universitaire, La Réunion). I should like to
thank Dr. M. Pichon (James Cook University, Townsville, Australia) for
identifying some scleractinian corals in the present list. I am

grateful to C. *Michel,: Director, Mauritius institute; Dp. Ardniieor
the Mauritius Fisheries Office; and J. Forget, formerly Magistrate of
Rodriguez for their assistance in many ways while I was in Mauritius
and Rodriguez.

71k

References
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Baissac, J. de B, Lubet, P. and Michel, C. (1962). Les biocoenoses
benthiques littorales de 1'Ile Maurice. Rec. Trav. Stn. Mar.
Endoume 25, 39 : 253-291.

Faure, G. (1973). Morphology and bionomy of the Coral reef dis-
continuities in Rodriguez island (Mascarene Archipelago, Indian
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(in press).

Faure, G. (1974). Etude comparative des récifs coralliens de 1'Archipel
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Faure, G. and Montaggioni, L. (1970). Le récif corallien de St-Pierre
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Oy BAe

Faure, G. and Montaggioni, L. (197la). Les récifs coralliens de 1'Ile

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Faure, G. ‘and Montaggioni, L. (1971 b). Les récifs coralliens sous le
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Faure, G. and Montaggioni, L. (1974) : Les récifs coralliens au vent de
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Gardiner; wisse @L9S6)F: The reefs of the western Indian Ocean. IAL
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Monitagg@onisy, Tees (97/5). Coral reefs and Quaternary shore-lines in the
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Pichon), Mien (s9 6H.) Caractéres généraux des peuplements benthiques des
récifs et lagons de 1'Ile Maurice. Cahiers) O-R.S.fJ0.M. ,
Océanographie, V, 4: 31-45.

22

Pachon yess ieee) Comparative study of the main features of some
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coral reefs. Symp)... ,;Z0o0l. Soc, London, \280 185-2615

ROSen.BaRewa (97 )c. The distribution of reef coral genera in the
Indian Ocean. SYMP.) ZOOL. SOC «5 LONdON 728) }\ 263-299.

Bibliography of works recording reef corals from Mascarene Archipelago.
Bernard, H.M. (1896). Catalogue of the madreporarian corals in the

British Museum (Natural History), 2 : The genus Turbinaria, the
genus Astreopora London : British Museum (Natural History).

Bernard, H.M. (1897). Catalogue of the madreporarian corals in the
British Museum (Natural History), 3 : The genus Montipora, The
genus Anacropora. London : British Museum (Natural History).

Bernard, H.M.;. (L903), . Catalogue of the madreporarian corals in the
British Museum (Natural History), 4 : The family PORITIDAE. I.
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Bernard), H.M.: <(1905)). Catalogue of the madreporarian corals in the

British Museum (Natural History), 5 : The family PORITIDAE. II.
The genus Porites. I. Porites of the Indopacific region. London
British Museum (Natural History).

Boschma, H. (1948). The species problem in Millepora. Zool. Verhand.
Leiden. ees <a lalb oye

Boschma, H. (1966). On a new species of Millepora from Mauritius with
notes on the specific characters of Millepora exaesa. PLOCe We
ned= Akad. Wet. (€)7 69): 409-4197

Brook;|).Ga: (189L).. Description of new species of Madrepora in the
collection of British Museum. Ann..~Mag:., Nat. Hist:: (6)i.8
458-471.

Brook: Gz (lSo2r. Preliminary descriptions of 40 new species of
Madrepora in the collection of British Museum. Panta bk eAnns

Mag: “Nat. Hist. 16), LlOn ss) 45i=465~

BLEOOk Gee CUS93) Catalogue of the madreporarian corals in the
British Museum (Natural History), 1 : The genus Madrepora. London
British Musuem Natural History.

Bruggemann, F. (1878). Neue Korallen-Arten aus dem Rothen Meer und
von Mauritius. Abh. naturw. Ver. Bremen 5 : 395-400.
Brtiggemann, F. (1879). Corals (in the collections from Rodriguez

ZOOLOGY), Phil. “Trans Rs, SOC. LOSms. 569 = 59).

23

Ghevalser* ¢ Ji Be o2(197 1.) Les Scleractiniares de la Mélanaisie
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Grossiland,, €-.. (1948). Reef corals of the South African coast. Ann.
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Crossland, C. (1952). Madreporaria, Hydrocorallinae, Heliopora and
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250-252.
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oe (3) = La 1e8

Hoffmeister, J.E. (1925). Some corals ‘from American Samoa and the
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Klunzinger, C.B. (1879). Die Korallenthiere des rothen Meeres.
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2

Maeeee ee Ha (LOS). Effect of water temperatures on growth of reef
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24

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Yabe, H., Sugiyama, T. and Eguchi, M. (1936). Recent reef building
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25

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26

ATOLL RESEARCH BULLETIN
NO. 204.

ANNOTATED CHECK LIST OF OCTOCORALLIA IN
THE MASCARENE ARCHIPELAGO, INDIAN OCEAN

by Gerard Faure

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

February 1977

ANNOTATED CHECK LIST OF OCTOCORALLIA IN
THE MASCARENE ARCHIPELAGO, INDIAN OCEAN

by

Gérard Faure!

Summary

A total of 112 species belonging to 37 genera of the Mascarene

Octocorallia known to date are listed in this paper. Some of them
were collected by the author, by means of scuba, and determined by
Tixier-Durivault (Museum National d'Histoire Naturelle, Paris). The

others were recorded in former papers, specially by Tixier-Durivault
(1966, 1972) and Michel (1974).

Introduction

The Mascarene Archipelago is situated in the western Indian Ocean
about 700 km, to the east of Madagascar, in the longitudes 55°32' -
ESe2o rand latitudes 21°7" — a9e4o" : It is composed of three volcanic
islands : Réunion, Mauritius, Rodriguez, forming separate geological
topographical units. The total surface of areas of the coral reefs
are very different according to the islands : negligible (12 km?) ,
compared with that of the island (2500 km*) in Reunion, more important
in Mauritius (300 km? for 1800 km?j and very important in Rodriguez
where the reef is overlapping 200 km? as compared to 110 km? in ter-
restrial surface.

General Mascarene reef studies were done by : Gardiner (1936), Baissac
andeals (!962) ," Pichon’ (967, 1971), Faure and Montaggioni (11970). °197La,
OVA pL OV A) yp baure 97s) VO)», Montaggzons. (COTO 9i3))i. There are
some records of Octocorallia from the Mascarene Archipelago in the
works of : Ellis and Solander (1786), Lamarck (1816), Templeton (1835),
Michelin (1845), Milne-Edwards and Haime (1857), Studer (1878),

Brtiggemann (1879), Ridley (1882a, 1882b, 1882c), Thomson and MacKinnon
(1910), Simpson (1910), Stiasny (1940), but there are very few works
concerning the systematics of Octocorallia from the Archipelago,
except those of Tixier-Durivault (1966, 1972), Michel (1974).

Istation Marine d'Endoume, Marseille; Centre Universitaire,
St. Clothilde, Réunion.
(Manuscript received June 1975-- Eds.)

This paper intends to Serve as’ a first provisional dist (of (ene
Octocorallia in the Mascarene islands.

Order ALCYONACEA Lamouroux, 1816
Family ALCYONIIDAE Lamouroux, 1812

Genus Alcyonium Linné, 1758
Alcyonium aspiculatum Tixier-Durivault, 1965

RO 65 A! : Passe of Port Sud-Est, - 8m : Rodriguez

RO 7 : Branched coral reef flat (Acropora pharaonis), Port
Mathurin : Rodriguez

RO) 22 : Reef flat with scattered coral growth, (Inner reef flat),
Pointe au Sable : Rodriguez

MAU 34 : Lagoon. of Trou aux Biches, -— 2m.:-Mauritius

Alcyonium legitimum Tixier-Durivault, 1970

RO. 51 : Outer slope, reef of Anse aux Anglais, - 20m : Rodriguez

ROiS2 : Passe Grenade, - 18m : Rodriguez

Genus Cladiella Gray, 1869
Cladiella australis (MacFadyen, 1936)

MAU 67 : Reef front, La Preneuse : Mauritius
Cladiella ceylonica (Pratt, 1905)

MAU 29 : Reef front, Trou aux Biches : Mauritius
Cladiella hicksoni (Tixier-Durivault, 1944)

MAU 3 A : Outer reef flat, Pointe d'Esny : Mauritius
Cladiella krempfi (Hickson, 1919)

RO 10 : Reef front, Pointe aux Cornes : Rodriguez

Mauritius : in Tixier-Durivault (1972), Michel (1974)
Cladiella laciniosa (Tixier-Durivault, 1944)

MAU 3 B : Outer reef flat, Pointe d'Esny : Mauritius

Cladiella latissima (Tixier-Durivault, 1944)

MAU 26 : Outer reef flat, Trou aux Biches : Mauritius

Cladiella pachyclados (Klunzinger, 1877)

RO 80 B : Outer slope reef of Quatre Vingts Brisants, - 20 m
Rodriguez

Cladiella ramosa Tixier-Durivault, 1970

MAU 46 > Bay .of Arsenal, = 5m : Mauritius

RO 7 : Reef front Anse Tamarins : Rodriguez

Cladiella sphaerophora (Ehrenberg, 1834)
Rodriguez : in Brtiggemann (1879)

Genus Lobophytum Marenzeller, 1886
Lobophytum altum Tixier-—Durivault, 1956

ROSS : Compact reef flat (inner reef flat) Anse aux Anglais
Rodriguez

Lobophytum batarum Moser, 1919

MAU 41 : Outer slope’, Trou aux Biches, ‘—"20-m'/y “Mauritius

1 Author's collection.

Lobophytum
MAU 68
Lobophytum
REU 172
Mauritius
Lobophytum
RO14 A

RO: 50! B
REU 173
REU 189 B
Lobophytum
RO 39
Lobophytum
MAU 1
Lobophytum
REU 171A
Lobophytum
RO 66 B
Lobophytum
MAU 42

REU 180

Lobophytum
MAU 12
Lobophytum
REU 151
REU 169
Lobophytum
MAU 58
Lobophytum
REW 55)
REU 156)

compactum Tixier-Durivault, 1956
Outer slope, reef of Flic en Flac, - 15 m : Mauritius
crassum Marenzeller, 1886

: Outer slope, reef of cap Lahoussaye - 15 m : Réunion

in Tixier=—Durivault (1956), Michel (1974)
crebriplicatum Marenzeller, 1886

; Outer slope, reek of Port-Mathurin, »=—~lO%m* 2 Rodriguez
Quter slope, reef~ of Anse’ aux Anglais; - 10. m»: Rodriguez
: Outer slope, reef of cap Lahoussaye, - 15 m : Réunion

Outer slope, reef Trois Bassins, - 30 m : Réunion
hedleyi Whitelegge, 1897

: Reef flat with microatolls, Port Sud-Est : Rodriguez

lamarcki Tixier-Durivault, 1956
Outer reef flat, Pointe d'Esny : Mauritius
mirabile Tixier-Durivault, 1956

Outer slope, reef cap Lahoussaye, - 15 m : Réunion
patulum Tixier-Durivault, 1956

Passe of Port Sud-Est, - 20 m : Rodriguez
pauciflorum (Ehrenberg, 1834)

Outer slope, Troux aux Biches, - 20 m : Mauritius

Volcanic flagstone (outer slope), reef of 1'Hermitage,
- 35 m : Réunion
robustum Tixier-Durivault, 1957

Backreef channel, Ile aux Aigrettes, - 5 m : Mauritius
sarcophytoides Moser, 1919

Reef front of St-Pierre : Réunion

Outer slope, cap Lahoussaye, - 15 m : Réunion

schoedei Moser, 1919

Outer slope, reef of Pointe aux Sables, - 10 m : Mauritius

variatum Tixier-Durivault, 1957

Reef front of St-Pierre : Réunion

Genus Parerythropodium Ktikenthal, 1916

Parerythropodium fulvum (Forskal, 1775)

ROO 59) A

: Outer slope, reef of Grande Pointe, - 15 m : Rodriguez

Genus Sarcophyton Lesson, 1834

Sarcophyton crassocaule Tixier-Durivault, 1946

RO 16 B Outer slope, reef of Port-Mathurin, - 15 m : Rodriguez
RO 40 B Outer reef flat, Port Sud-Est : Rodriguez

RO 48 A Outer slope, Middle ground reef, - 8 m : Rodriguez
Sarcophyton glaucum (Quoy et Gaimard, 1833)

REU 189 A Outer slope, reef of Trois Bassins, - 30 m : Réunion
RO 40 A : Reef flat with microatolls, Port Sud-Est : Rodriguez
RO 46 = Outer Slope, Post—Mathunin, — 15 m 1s Redriiguez

RO 58 Outer slope, reef of Grande Pointe, - 15 m : Rodriguez

Sarcophyton latum (Dana 1846)

Rodriguez

in Brtiggemann (1879)

Sarcophyton molle Tixier-Durivault, 1946

REU 160

: Volcanic flagstone (outer slope), reef of 1'Hermitage,

- 25 m : Réunion

4

REU 161 Volcanic flagstone (outer slope), reef of 1'Hermitage,
- 30 m : Réunion
REU 184 : Volcanic flagstone (outer slope), reef of La Saline,
- 35 m : Réunion
Sarcophyton regulare Tixier-Durivault, 1958
REU »170 Outer slope, reef of cap Lahoussaye, - 15 m : Réunion
ROM S/7/ : Outer slope, reef of Grande Pointe, - 15 m Rodriguez
Sarcophyton solidum Tixier-Durivault, 1958
REU 182 B Volcanic flagstone (outer slope), reef of 1'Hermitage,
- 35m Réunion
Sarcophyton spongiosum Thomson and Dean, 1931
REU 153 Lagoon of St-Pierre Réunion
Sarcophyton stolidum Verseveldt, 1971
MAU 40 : Outer slope, Trou aux Biches, - 20 m Mauritius
Sarcophyton trocheliphorum Marenzeller, 1886
Mauritius in Tixier-Durivault (1972), Michel (1974)
Genus Sinularia May 1898
Sinularia arborea Verseveldt, 1971
MAU 40 Outer slope, reef of Trou aux Biches, - 25 m : Mauritius
Sinularia crassa Tixier-Durivault, 1945
MAU 2 Outer reef flat, Pointe d'Esny Mauritius
Sinularia densa (Whitelegge, 1897)
MAU 66 Reef front, La Preneuse Mauritius
REU 179 Outer slope, reef of St-Gilles, - 25 m Réunion
RO 14 B Outer slope, reef of Port-Mathurin, - 10 m : Rodriguez
Sinularia dura; (Pratt ,. 1.903)
MAU 17 C : Outer slope, Ile de La Passe, - 10m Mauritius
Sinularia erecta Tixier-Durivault, 1945
MAU 17 A Outer slope, Ile de La Passe, - 10m Mauritius
Sinularia grayi Tixier-Durivault, 1945
REU 159 Outer slope, reef of 1'Hermitage, - 20 m : Réunion
RO 56 Outer slope, reef of Grand Baie, - 18m Rodriguez
Sinularia leptoclados (Ehrenberg, 1834)
ROP6Se8 Passe of Port. Sud-Esit,))=) 8m Rodriguez
Sinularia macropodia (Hickson and Hiles, 1900)
REU 181 Volcanic flagstone (outer slope), reef of l'Hermitage, -
35 m : Réunion
RO 49 Outer slope, reef of Middle Ground, - 10m Rodriguez
Mauritius in Tixier-Durivault (1972), Michel (1974)
Sinularia marenzelleri (Wright and Studer, 1889)
MAU 16 Channel of Mahebourg, - 20m Mauritius
RO 48 B Outer slope, reef of Middle Ground, - 10m Rodriguez
Sinularia pedunculata Tixier-Durivault, 1945
MAU 37 Outer slope, reef of Trou aux Biches, - 20 m Mauritius
RO 16 A Outer slope, reef of Port-Mathurin, - 10m Rodriguez
Sinularia polydactyla Tixier-Durivault, 1945
MAU 22 A Outer slope, Horseshoe reef, - 8m Mauritius
MAU 45 Bay of Arsenal, - 3 m: Mauritius
MAU 49 Outer slope, reef of Trou aux Biches, - 28 m : Mauritius
MAU 54 Reef front, Pointe au Sable Mauritius
REU 150 Reef front of St-Pierre Réunion
REU 178 Outer slope, reef of St-Gilles, - 22m : Réunion

REU 154
RO 60 B
Sinularia
RO 66 A
Sinularia
Réunion
Sinularia
MAU 30
Sinularia
REU 152
RO 50 A
ROE 53" B
Sinularia
MAU 39
Réunion
Sinularia
REU 158
Sinularia
RO 13

RO 53 A
MAU 4
MAU 5

MAU 10
MAU 17 B

Lagoon of St-Pierre : Réunion

Passe of Grand-Bassin, - 12 m Rodriguez
querciformis (Pratt, 1903)
Passe of Port Sud-Est, - 8 m Rodriguez

rigida var. amboinensis (Burchardt, 1902)
in Tixier-Durivault (1972)

robusta MacFadyen, 1936
Reef front, reef of Trou aux Biches

simpsoni Tixier-Durivault, 1945
Lagoon, reef of St-Pierre Réunion

Mauritius

Outer Slope, reef of Anse aux Anglais, - 6 m : Rodriguez
Outer slope, reef of Port-Mathurin, - 12 m Rodriguez
triaena Kolonko, 1926
Outer slope, reef of Trou aux Biches, - 20 m Mauritius
in Tixier-Durivault (1972)
venusta Tixier-Durivault, 1970
Outer slope, reef of Boucan Canot, - 15m Réunion
whiteleggei Ltittschwagger, 1914
Reef front, reef of Pointe aux Cornes Rodriguez
Outer slope, reef of Port-Mathurin, - 12 m Rodriguez
Outer reef flat, reef of Pointe d'Esny Mauritius
Outer reef flat, reef of Pointe d'Esny Mauritius
Backreef channel of Ile aux Aigrettes, - 5m Mauritius

Reef front, outer slope, reef of Ile de la Passe, O - 6m
Mauritius

Family NEPTHEIDAE Gray, 1862

Genus Nephthea Savigny, 1817

Nephthea cupressiformis Ktikenthal, 1904

ROMA

Passe of Port Sud-Est, - 20 m Rodriguez

Genus Roxasia Tixier-Durivault and Prevorsek, 1957

Roxasia gravieri (Ktikenthal, 1910)

Mauritius

in Tixier-Durivault (1972), Michel (1974), Tixier-Durivault

and d'Hondt 1973 (1974)

Roxasia hirsuta Tixier-Durivault and Prevorsek, 1960

REU 188 Volcanic flagstone (outer slope), reef of Pointe des
Aigrettes, - 40m Réunion

Roxasia mirabilis (Henderson, 1909)

Mauritius in Tixier-Durivault (1972), Michel (1974)

Genus Stereonephthya Ktikenthal, 1905

Stereonephthya unicolor (Gray, 1862)
REU 157 Outer slope, reef of Boucan-Canot, - 15 m Réunion,
Stereonephthya sp.
RO 80 A Outer slope, reef of Quatre-Vingts Brisants, - 20 m
Rodriguez
Genus Dendronephthya
Dendronephthya divaricata (Gray)
Mauritius in Thomson and MacKinnon (1910), Michel (1974)

Family XENIIDAE Ehrenberg, 1828

Genus Anthelia Lamarck, 1816
Anthelia desjardiana Templeton, 1841
Mauritius : in Milne-Edwards and Haime (1857), Michel (1974)
Anthelia flava (May, 1899)
Mauritius : in Tixier-Durivault (1972), Michel (1974)

Genus Cespitularia Milne-Edwards and Haime, 1850
Cespitularia densa Tixier-Durivault, 1966
Mauritius -2 in; Taxier-Durnivaul ty, (1972). Mechel (1974)
Cespitularia mantoni Hickson, 1931
Mauritius : in Tixier-Durivault (1972), Michel (1974)

Genus Sympodium Ehrenberg, 1834
Sympodium coeruleum Ehrenberg, 1834
MAU 6 : Outer reef flat, reef of Pointe d'Esny : Mauritius

Genus Xenia Lamarck, 1816
Xenia blumi Schenk, 1896
Mauritius : in Tixier-Durivault (1972), Michel (1974)
Xenia distorta Tixier-Durivault, 1966
Mauritius : in Tixier-Durivault (1972), Michel (1974)
Xenia elongata Dana, 1846
REU 149 : Reef front, reef of St-Pierre : Réunion
Xenia florida (Lesson, 1826)
Mauritius : in Tixier-Durivault (1972), Michel (1974)
Xenia garciae Bourne, 1894
Mauritius : in Michel (1974)
Xenia plicata Schenk, 1896
Mauritius : in Tixier-Durivault (1972), Michel (1974)
Xenia ternatana Schenk, 1896
Mauritius : in Tixier-Durivault (1972)

Order COENOTHECALIA Bourne, 1895
Family HELIOPORIDAE Moseley, 1876

Genus Heliopora Blainville, 1830
Heliopora coerulea (Pallas, 1766)

RO 110 : Outer reef flat, reef of Grande Pointe : Rodriguez

RO 302 : Outer Slope, reef of Quatre-Vingts Brisants, - 12 m :
Rodriguez

MAU 155 : Outer slope, reef of Ile Plate, - 8 m: Mauritius

MAU 404 : Lagoon, reef Pointe Bambou : Mauritius

Order GORGONACEA Lamouroux, 1816
Suborder SCLERAXONIA Studer, 1887
Family SUBERGORGIIDAE Gray, 1859

Genus Subergorgia Gray, 1857
Subergorgia mollis (Nutting, 1910)
Mauritius : in Tixier-Durivault (1972), Michel (1974)

Subergorgia reticulata (Ellis and Solander, 1786)
Mauritius : in Tixier-Durivault (1972), Michel (1974)
Subergorgia suberosa (Pallas, 1766)

Mauritius : in Tixier-Durivault (1972), Michel (1974)

Family CORALLIIDAE Lamouroux, 1812

Genus Corallium Cuvier, 1798
Corallium stylasteroides Ridley, 1882
Mauritius : in Ridley (1882a), Tixier-Durivault (1972), Michel (1974)

Family MELITHAEIDAE Gray, 1870

Genus Melithaea Milne-Edwards and Haime, 1857
Melithaea ochracea (Lamarck, 1816)
Mauritius : in Michelin (1845), Michel (1974)

Genus Wrightella Gray, 1870
Wrightella coccinea (Ellis and Solander, 1786)
REU 185 A : Volcanic flagstone (outer slope), reef of Pointe aux

Aigrettes, - 40 m : Réunion
Mauritius : in Ellis and Solander (1786), Milne-Edwards and Haime
(1857)

Family PARASIDIDAE Aurivillius, 1931

Genus Parisis Verrill, 1864
Parisis fruticosa Verrill, 1864
Mauritius : in Wright and Studer (1889), Tixier-Durivault (1972),
Michel (1974)

Suborder HOLAXONIA Studer, 1887
Family ACANTHOGORGIIDAE Gray, 1859

Genus Acanthogorgia Verrill, 1868
Acanthogorgia spinosa Hiles
Mauritius : in Thomson and Russel (1909), Michel (1974)

Genus Anthogorgia Verrill, 1868
Anthogorgia divaricata Verrill, 1868
Mauritius : in Tixier-Durivault (1972), Michel (1974)

Family PARAMURICEIDAE Bayer, 1956

Genus Muricella Verrill, 1869
Muricella grandis Nutting, 1910
Mauritius : in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)
Muricella perramosa Ridley, 1882
Mauritius : in Ridley (1182c), Tixier-Durivault (1972), Michel (1974)
Muricella rubra Thomson, 1905
Mauritius : in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)

8

Muricella
Mauritius

Paracis orientalis

Mauritius

tenera Ridley, 1884
in. Stiasny ((l940), Tixier-Durivault (972), Michel“ (1974)
Genus Paracis Ktikenthal, 1919
(Ridley, 1882)
in*Wright ‘and ;studer -(1889)i,; Tixter—Durcivauli) (i972)i,
Michel (1974)

Genus Villogorgia Duchassaing and Michelotti, 1860

Villogorgia alternans (Wright and Studer, 1889)

Mauritius

in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)

Villogorgia ceylonensis (Thomson and Henderson, 1905)

REU 185 B

: Volcanic flagstone
Aigrettes, - 40m:

(outer slope), reef of Pointe aux
Réunion

Villogorgia mauritienis Ridley, 1882

Mauritius

in Ridley (1882b), Stiasny
Michel (1974)

(1940), Tixier-Durivault (1972),

Family PLEXAURIDAE Gray, 1859

Genus Echinogorgia Kélliker, 1865

Echinogorgia lami Stiasny, 1940

Mauritius

in Stiasny' (1940), Tixier—Durivault: (1972), Michell (1974)

Echinogorgia pinnata Studer, 1878

Mauritius

Leptogorgia boryana Milne-Edwards and Haime,

Réunion

in Studer (1878), Tixier-Durivault (1972), Michel (1974)

Genus Leptogorgia Milne-Edwards and Haime, 1857
1857
in Milne-Edwards and Haime (1957)

Family ELLISELLIDAE Gray, 1859

Genus Ellisella Gray, 1868

Elisella candida (Ridley, 1882)

Réunion
Mauritius

Ellisella
Réunion
Ellisella
Réunion
Ellisella
Mauritius

Junceella
REU" 185; €

Mauritius
Junceella
Réunion
Junceella
Réunion

in Tixier-Durivault (1972)

in Ridley (1882a), Michel (1974) as Verrucella candida,
Tixier-Durivault (1972)
laevis (Verrill, 1865)
in Tixier-Durivault (1972)
ramosa Simpson, 1910
in Tixier-Durivault (1972)
vaughani Stiasny, 1940

in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)
Genus Junceella Valenciennes, 1855
juncea (Pallas, 1766)
: Volcanic flagstone (outer slope), reef of Pointe aux
Aigrettes, - 40 m Réunion
im?’ Simpson «(.190)), “Tixier—-Durni vault
vVinem Valenciennes, 1855
in Tixier-Durivault (1972), Milne-Edwards and Haime (1857)
caliculata Valenciennes, 1855
in Milne-Edwards and Haime

(1972), Michel (1974)

(1857)

Genus Nicella Gray, 1870
Nicella carinata Nutting, 1970
Mauritius : in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)
Nicella dichotoma (Gray, 1859)
Mauritius : in Gray (1859), (1870), Simpson (1910), Stiasny (1940),
Tixier-Durivault (1972), Michel (1974)
Nicella granifera (Kélliker, 1865)
Mauritius : in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)

Genus Verrucella Milne-Edwards and Haime, 1857

Verrucella flexuosa Klunzinger, 1877

Mauritius : in Klunzinger (1877), Thomson and MacKinnon (1910),
Michel (1974)

Verrucella verriculata (Milne-Edwards and Haime, 1857)

Mauritius : in Milne-Edwards and Haime (1857), Lamarck (1816), Michel
(1974), Tixier-Durivault (1972)

Family ISIDIDAE Lamouroux, 1812

Genus Isis Linnaeus, 1766
Isis hippuris (Linnaeus, 1766)
Mauritius : in Michelin (1845), Dana (1846), Milne-Durivault and
Haime (1857), Michel (1974)

Genus Keratoisis Wright, 1869
Keratoisis grayi Wright, 1869
Mauritius : in Michel (1974)

Family PRIMNOIDAE Gray, 1857
Subfamily PRIMNOINAE Gray, 1857

Genus Callogorgia Gray, 1858
Callogorgia flabellum (Ehrenberg, 1834)
Mauritius : in Stiasny (1940), Tixier-Durivault (1972), Michel (1974)

Genus Pseudoplumarella Ktikenthal, 1915
Pseudoplumarella plumatilis (Milne-Edwards and Haime, 1857)
Mauritius : in Tixier-Durivault (1972)
Réunion : in Milne-Edwards and Haime (1857), Tixier-Durivault (1972)

Subfamily CALYPTROPHORINAE Gray, 1870

Genus Calyptrophora
Calyptrophora japonica Gray, 1886
Réunion : in Tixier-Durivault (1972)

Order PENNATULACEA Verrill, 1865
Suborder SUBSELLIFLORAE Ktikenthal, 1915
Family VIRGULARIIDAE Verrill, 1868

Genus Virgularia Lamarck, 1816
Virgularia densa Tixier-Durivault, 1966
Mauritius : in Tixier-Durivault (1966), (1972), Michel (1974)

10
Conclusions

From the 112 species listed in this paper, 76 are known from
Mauritius, 32 from Réunion, and 27 from Rodriguez. Rather than to a
greater diversity of Octocorallia biota, the greater abundance of
species in Mauritius must be attributed to a better knowledge from the
numerous former records. The lack of affinity of Octocorallia species
between the three islands (only 8 species in common for Mauritius and
Réunion, 7 Mauritius-Rodriguez, 6 Réunion-Rodriguez, and 2 for the
three islands), must be chiefly ascribed to the lack of data in this
part of western Indian Ocean. In the meantime it should be observed
that more than 70% of scleractinian corals (more than 120 species
belonging 55 genera) are common to Mauritius, Rodriguez and Réunion.

If it is too early to compare the Octocorallia distribution with
the scleractinian corals, it is a good argument for carrying on eco-
logical and systematical investigations of both in the Archipelago.

References

General studies on the Coral reefs from Mascarene Archipelago

Baissac, J. de B., Lubet, P. and Michel, C. (1962). Les biocoenoses

benthiques littorales de 1'Ile Maurice. Rec). Trav. "Stnw Mare,
Endoume 25, 39": 253-291.

Faure, G.2 (973); Morphology and bionomy of the coral reef discon-
tinuities in Rodriguez island (Mascarene Archipelago, Indian
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(in press).

Faure, G. (1974). Etude comparative des récifs coralliens de
l'Archipel des Mascareignes (Océan Indien). Colloque Commerson,
St-Denis (Réunion), Oct. 1973, Bull. Mauritius Inst. (in press).

Faure, G. and Montaggioni, L. (1970). Le récif corallien de St-Pierre-
de-La-Réunion (Océan Indien) : Géomorphologie et répartition des
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VO eZee 2 OA

Faure, G. and Montaggioni, L. (197la). Les récifs coralliens de
1'Ile Rodrigue (Archipel des Mascareignes, Océan Indien)
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rauney, G.- and Montaggdionasys, bet (el 7Aao) a Les récifs coralliens sous le
vent de 1l'Ile Maurice (Archipel des Mascareignes, Océan Indien)
Morphologie et bionomie de la pente externe. Ge ReEWACad sr iSGae
Parisl, (2730 i) OMA —ealLONGs

ie

Hause, Gand, Montaggvon.,, i. (1974)). Les récifs coralliens au vent
de 1l'Ile Maurice (Archipel des Mascareignes, Océan Indien)
Morphologie et bionomie de la pente externe. Cah. wACaAd = SCL.

Paris (in press).

Gardiner! o2S." (1936)! The reefs of the Western Indian Ocean. II,
the Mascarene region. Transe Linn SOC -mLOndOny, 2nd Sere ,. 19
426-436.

Montaggioni, L. (1970). Répartition et zonation géomorphologique des
structures récifales de 1'Ile de La Réunion (Océan Indien). GaRe
Acad.iSer. Larvs, 270 = 663-665.

Montaggioni, L. (1973). Coral reefs and Quaternary shore-lines in the
Mascarene Archipelago. Proc. 2nd Internat. Symp. Coral reefs,
Brisbane (in press).

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31-45.

Bichon, Me. (LOTT) s Comparative study of the main features of some
coral reefs of Madagascar, La Réunion and Mauritius. In Stoddart
and Yonge : regional variation in Indian Ocean Coral reefs.

Gums 200L. Soc. London, 28": 185-216

‘

Stoddarme, Dek. (L973) - Coral reefs of the Indian Ocean. In Jones
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Works recording Octocorallia from Mascarene Archipelago

Briiggemann, F. (1879). Corals of Rodriguez. Phils Eran Si ROU 1SOC.
NGG (Extra volume) 2 569=57.9)-

Dana, J.D. (1846). Zoophytes. Philadelphia : 740 pp
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and uncommon zoophytes collected from various parts of the globe,
london 72) Gdka5—I7/3"

Gray wie tien GlLSS9))). Description of some new genera of litophytes, or
stony zoophytes. PROC ZOOL. SOGe LONGa, 2 ls z
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collection of the British Museum, London.

Kiltinzinger, (sb. (2877). Die Korallthiere des rothen Meeres I, Die
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Lamarck, J.B. de (1816). Histoire naturelle des animaux sans
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The

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Michelin, M. (1845). Zoophytes, Echinodermes et Stellerides. Essai
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Milne-Edwards, H. et Haime, J. (1857). Histoire naturelle des
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RBICULEN AA SOG (CMstsyxe) 6 Contribution to the knowledge of Alcyonaria
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Simpson, Jicdic, (L9TO):. A revision of the Gorgonellidae, I. The
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Stiasny, G.... (1940) . Alcyonaria und Gorgonaria von Mauritius Arch.
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Studer ils GLSnSNi. Uebersicht der Anthozoa Alcyonaria wdahrend der
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Templeton, R. (1835). Descriptions of a few invertebrated animals
obtained at Ile de France. PROC) 3 2ZOOl.§ SOC! Lond .7 NSS Sa-mseleles

Thomson, J.A. and MacKinnon, D. (1910). Alcyonarians collected by
the Percy Sladen Trust Expedition by M.J. Stanley-Gardiner. 2.
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Thomson, J.A. and Russel E.S. (1909). Alcyonarians collected by the
Percy Sladen Trust Expedition by M.J. Stanley-Gardiner. l.
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Tixier-Durivault, A. (1956). Les Alcyonnaires du Museum. I. Famille
des Alcyoniidae. 4. Genre Lobophytum. Bull. Hist. nat., Paris
(sexs 2) 28) (4)i- 3) -401=405-, > 28 (Seu: 476-482) 5 28) 1 (6)y_ 3541-5467

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NS)

Tixier-Durivault, A. (1972). Nouvel apport d'Octocoralliaires de
Madagascar et des Iles avoisinantes. TecGhys., SUppile, 2 = LL=68;

Tixier-Durivault, A. and d'Hondt, M.J. (1974). Nouvelles récoltes
d'Octocoralliaires 4 Madagascar. Tethys 5 : (2-3) 1973 : 251-266.

Wright, E.P. and Studer, Th. (1889). Report on the Alcyonaria
collected by H.M.S. "Challenger" during years 1873-1876. SCi.
Rep. Voy. H.M.S. Challenger, 31 : 341 pp.

ATOLL RESEARCH BULLETIN
NO. 205.

THE HOSTS OF THE CORAL-ASSOCIATED INDO.-
WEST-PACIFIC PONTONIINE SHRIMPS

by A. J. Bruce

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

February 1977

eee —————

Paratypton siebenrocki Balss

Uvigerous female in situ in cyst in acropora host coral,

Ba,

Kenya. vJrawn from colour transparency,

scale approximately 2 mm.

THE HOSTS OF THE CORAL-ASSOCIATED INDO.
WEST-PACIFIC PONTONIINE SHRIMPS

by

Beds Bruce!

Many of the shrimps belonging to the subfamily Pontoniinae
Kingsley live in permanent obligatory association with scleractinian
corals in the Indo-West Pacific region. These shrimps are abundant
on most coral reefs and represent a major component of the diversity
of the caridean fauna.

Borradaile (1917) mentioned briefly that shrimps of the genera
Harpiliopsis, Coralliocaris and Harpilius (>Periclimenes spp.,
Philarius spp.) are adapted to life amongst the branches of coral
colonies but provided no details. Kemp (1922) provided a list of
associations of the Pontoniinae with corals, including again
Coralliocaris, Harpilius, (in which he included Harpiliopsis), and
Periclimenes diversipes, together with P. spiniferus, a free-living
species frequently found in live or dead coral colonies. Holthuis
(1952), in the report on the Pontoniinae collected by the Siboga and
Snellius Expeditions, provided further details in tabular form of the
species then considered to be associates of corals. The decapod
fauna of the Queensland branching corals was analysed by Patton (1966),
who added particularly to knowledge of the shrimps associated with the
Pocilloporidae. The Proceedings of the Coral and Coral Reef Symposium,
1969, at Mandapam, included a review of the coral associations then
known of the Indo-West Pacific Pontoniinae, with a key for their |
identification, and with further details of the Acropora host-species
involved, (Bruce, 1972a).

The following report provides information on the coral species
known to act as hosts for the obligatory shrimp associates. A number
of other species may frequently be found when coral colonies are
examined, i.e. Palaemonella spp. specially P. rotumana (Borradaile)
and Periclimenes petitthouarsi (Audouin), and P. spiniferus De Man, and
probably other related species. These species are free-living micro-
predators or browsers. The exact identification of the hosts of some

1 Formerly East African Marine Fisheries Research Organisation (EAMFRO).
(Manuscript received September 1975 --Eds.)

rare species, for example Philarius lifuensis (Borradaile), have yet

to be ascertained, and it is probable that many more hosts of the known
shrimp species, as well as new shrimps associated with new hosts, remain
to be discovered.

SYSTEMATIC ACCOUNT
PALAEMONIDAE, Samouelle, 1819.
Pontoniinae, Kingsley, 1878..

Ae Vir orientalis (Dana)
Restricted synonymy:
Vir orientalis-— Bruce, 1972, 64, 65-67, fig.1.

Hosts. Pocillopora damicornis (L.). (Bruce, -1972a; 1972b).
Pocillopora verrucosa (Lam.) (Bruce, in press e).
Stylophora erythraea von Marenzeller (Bruce, 1972a).
Stylophora pistillata (Esper,) (Bruce, 1972a).

Acropora sp. (Bruce, in press, d, h).

Distribution. Sparsely recorded throughout the Indo-West Pacific region

from East Africa to Hawaii.

Remarks. A small species that is easily overlooked and usually only

present in small numbers in a host colony.

Pc Periclimenes amymone De Man
Restricted synonymy:
Periclimenes amymone- Holthuis, 1972, 10, 82-83, fig.32.

Hosts.. -Pocillopora’damicornis .(l..), (Patton, 19665, 1974)-.
Stylophora mordax (Dana) (Patton, 1966).
Stylophora pistillata (Esper.) (Patton, 1966).

Seriatopora hystrix (Dana) (Patton, 1966).

Acropora cymbicyathus (Brooks) (Bruce, 1972a).

Acropora digitifera (Dana) (Bruce, 1972a).

Acropora diversa (Dana) (Bruce, 1972a)

Acropora eurystoma (Klunz.) (Bruce, 1972a).

Acropora hyacinthus (Dana) (Bruce, 1972a).

Acropora kenti .(Brook) (Bruce, 1972a).

Acropora sarmentosa (Brook) (Bruce, 1972a).

Acropora syringodes (Brook) (Bruce, 1972a).

Acropora tenuis (Dana) (Bruce, 1972a).
Distribution. From Singapore and Djakarta in the west to Samoa in the
east; not recorded from the Indian Ocean.
Remarks. One of the few commensal shrimps that is found commonly in
both acroporid and pocilloprid coral hosts, where it is usually present
on the outer branches.

32 Periclimenes consobrinus De Man
Restricted synonymy:
Periclimenes (Harpilius) lutescens- Holthuis, 1952, 12,
88=9i, elgg. 35 (par tame:
Periclimenes consobrinus - Bruce, 1975, 27, fig.16.

Hosts. Pocillopora hemprichi (Ehrenberg) (Bruce, 1971, as P.lutescens).
Pocillopora damicornis (L.) (Bruce, in press e).
Distribution. Due to confusion with the closely related P. lutescens
auct. this species is known with certainty only from Ternate, Indonesia
the Comoro Islands, and Mombasa, Kenya.
Remarks. Bruce (1971) recorded the association of P. lutescens auct.
with Pocillopora hemprichi in the Comoro Islands. At that time
P. consobrinus was considered to be a synonym of P. lutescens, and the
subsequent re-examination of the specimen showed that it should be
correctly referred to P. consobrinus on account of the characteristic
morphology of the second maxilliped (Bruce, 1972a, fig. 1b). This
species is moderately common in the fringing lagoons of central East
Africa, in Pocillopora sp., often also in association with Harpiliopsis
beaupresi, H. depressus and H. spinigerus. It has not so far been
found on any other host genus.

4. Periclimenes lutescens auct.

Restricted synonymy:

Periclimenes (Harpilius) lutescens - Holthuis, 1952, 12,

S5—9i secs) a(parmt-um)r. Patton, L966 s27> 268, cals...

DOO w=eabis 2

Periclimenes lutescens - Bruce, 1971:2, 5; 1972a: 404, 405,

4065, 407.) 4409). 400 sarogwilia Yat? (key )ye SPOTS 22 7 Eig 21.5 ,.
Hosts. Seriatopora hystrix (Dana) (Patton, 1966).

Acropora convexa (Dana) (Bruce, 1972a).

Acropora humilis (Dana) (Bruce, 1972a).

Acropora kenti (Brooks) (Bruce, 1972a).

Acropora paniculata Verrill (Bruce, 1972a).

Acropora squarrosa (Ehrenberg) (Bruce, in press, qd).

Acropora surculosa (Dana) (Bruce, 1973).

Acropora tubicinaria (Dana) (Bruce, in press, e).
Distribution. Widespread throughout the Indo-West Pacific region from
the Red Sea to Madagascar, as far east as Tonga and Samoa, possibly
extending to Tahiti and the Marquesas Islands.

Remarks. As noted under P. consobrinus, the report of an association

of P. lutescens with Pocillopora in Bruce (1971) is erronous. aes:
species is commonly found in association with Jocaste and Coralliocaris
spp. The association of three specimens with Seriatopora hystrix

reported from Heron Island by Patton (1966) is most unusual.

Dis Periclimenes parvus Borradaile
Restricted synonymy:

Periclimenes parvus Borradaile, 1898:384; 1899:407, pl.36,

1PaMoi A SNe
Hosts. Unidentified.
Distribution. Known only from New Britain, the Makassar Straits and
Singapore.
Remarks. The association of this species appears uncertain. Johnson
(1961) notes that a single specimen from the Raffles Light, Singapore,
in the Bedford Lanchester collection, was obtained from "outer coral"
at very low tide. Periclimenes parvus possesses a biunguiculate
dactylus on the ambulatory pereiopods (Holthuis, 1952), a feature that

is not found in any other coral associated pontoniine shrimp, and
which suggests that the association may have been accidental. In
coral-associated pontoniine shrimps this dactylus is generally robust
and simple, except in Fennera, Jocaste and Coralliocaris, in which a
basal process is present.

6. Periclimenes diversipes Kemp
Restricted Synonymy:
Periclimenes (Ancylocaris) diversipes Kemp, 1922:117, 169
(key), 179-184, figs. 36-39 (partim).

Hosts. Psammocora togianensis Umbgrove (Bruce, 1972a).

Pocillopora damicornis (L.) (Bruce, in press, e).

Stylophora erythraea von Marenzeller (Bruce, 1972a).

Seriatopora sp. (Bruce, 1971).

Acropora tenuis (Dana) (Bruce, 1972a).

Acropora variabilis (Klunzinger) (Bruce, in press, e).

Montipora circumvallata (Ehrenberg) (Bruce, 1971).

Montipora prolifera Breuggemann

Pavona danai (M.-Edw. & Haime) (Bruce, 1972a).

Porites nigrescens (Dana) (Bruce, in press, e).

Porites iwayamanensis Eguchi (Bruce, 1972a).

Porites sp.nov. cf. andrewsi (Bruce, 1972a).

Galaxea clavus (Dana) (Bruce, 1972c).
Distribution. Common in the Indian Ocean; from the Red Sea to
Madagascar. Also recorded from the Australian Great Barrier Reef on
Acropora sp. (Patton, 1966).
Remarks. The least host-specific of the pontoniine coral associates,
with hosts belonging to nine different genera.

Te Periclimenes madreporae Bruce
Restricted synonymy:
Periclimenes madreporae Bruce, 1969:262-263, 264; 1972a:403,
404, 405:,..406,, 407:, 409), 410, -413: (key)i..-= Patton jlo awake
223—=226. taba, Zoli, oo pica Giles
Periclimenes (Harpilius) inornatus Patton, 1966:274-275, 288
tab.2, 291 tab.3, fig.2. - Miyake & Fujino, 1968 400,. 402
(key) 413-414, fig.3 g-h, 431. tab...
Hosts... Pocillopora, damicornis, (b.)s0 \batton, +1966) %
Pocillopora verrucosa (Ellis & Solander) (Patton, 1966).
Seriatopora hystrix (Dana), (Patton, 1966).
Stylophora mordax (Dana), (Bruce, 1972a).
Stylophora pistillata (Esper.), (Patton, 1966).
Acropora corymbosa (Lam.), (Bruce, 1972a).
Acropora cuneata (Dana), (Bruce, 1972a).
Acropora digitifera (Dana), (Bruce, 1972a).
Acropora nasuta (Dana), (Bruce, 1972a).
Acropora rotumana (Gardiner), (Bruce, 1972a).
TurbinaGlasSp 7; UB GUCe,. 97 Zal)e.
Distribution. NE Australia, and Palau Islands only.
Remarks. The only caridean so far found in association with corals of
the genus Turbinaria.

ie} 4 Periclimenes holthuisi Bruce
Restricted synonymy:
Periclimenes holthuisi Bruce, 1969:258-259.
Hosts. Fungia actiniformis Quoy & Gaimard.
Distribution. Zanzibar, Seychelle Is., Hong Kong, New Caledonia, and
N.E. Australia.
Remarks. This species has generally been found in association with
actiniarians, but it has also been found with scyphozoans (Bruce,
JUS T/PXS ee A pair of specimens were collected from the host at a depth
of 60 ft. at Krantet Is., Madang, New Guinea, on 13-9-73 by
J.E. Randall and R. Steene.

S) Periclimenes mahei Bruce
Restricted synonymy:
Periclimenes mahei Bruce, 1969:263-264; 1971:2, ll.
Hosts. Pocillopora elongata (Dana), host of type material, (Bruce,

1972a).
Seriatopora hystrix (Dana), (Bruce, 1971).
Acropora corymbosa (Lam.), (Bruce, in press, e).

Distribution. Known only from the Seychelle and Comoro Islands and
Zanzibar.

Remarks. This species appears to occupy the same niche in the western
Indian Ocean as P. madreporae does in the western Pacific Ocean.

Os Periclimenes difficilis Bruce

Periclimenes difficilis Bruce, (in press, e).
Hosts. Porites nigrescens Dana.
Distribution. Known only from the type locality on Praslin, Seychelle
Islands.
Remarks. Closely related to P. diversipes, which is the only other
species of this genus known to associate with corals of the genus
Porites.

Habe Periclimenes kororensis Bruce
Periclimenes kororensis Bruce, (in press, f).
Hosts. Fungia actiniformis Quoy & Gaimard.
Distribution. Known only from the type locality on Koror, Palau
Islands.
Remarks. A rather aberrant species of Periclimenes, known only from
the incomplete holotype specimen.

WBE Periclimenes gonioporae Bruce
Periclimenes gonioporae Bruce, (in press, b).
Hosts. Goniopora stokesi Milne Edwards & Haime.
Lobophyllia sp.
Distribution. Known only from fringing lagoons of southern Kenya.
Remarks. Closely related to P. mahei and P. diversipes, but appears
to be specially associated with the genus Goniopora.

WSs Philarius gerlachei (Nobili)
Restricted synonymy:

Harpilius gerlachei Kemp, 1922:299 (key), 238-239, figs.74-75.

Hosts. Acropora corymbosa (Lam.), (Bruce, in press, e).
Acropora cymbicyathus (Brooks)
Acropora digitifera (Dana), (Bruce, 1972a)
Acropora formosa (Dana), (Bruce, 1972a).
Acropora humilis (Dana), (Bruce, 1972a).
Acropora hyacinthus (Dana), (Bruce, 1972a).
Acropora nana (Studer), (Bruce, 1972a).
Acropora surculosa (Dana), (Bruce, 1973).

Distribution. From the Red Sea, East Africa and Madagascar to the
Marshall, Gilbert and Samoan Islands.

Remarks. Distinctly less common, in the Acropora hosts, than Jocaste
or Coralliocaris species.

14. Philarius imperialis (Kubo)
Restricted synonymy:
Harpilius amperialis, Kubo, 19402 1-4, figs: L-3:
Hosts. Acropora spp., (Bruce, 1972a).
Distribution. From the Red Sea to Zanzibar, as far east as the Palau
and Marshall Islands. E
Remarks. Generally uncommon; particularly in comparison with Jocaste
and Coralliocaris spp. Usually found right at the very innermost
bases of the branches of the coral hosts, often on the floor of the
grooves between branches. The colour pattern of this species shows
the closest resemblance to the illustration given by Dana (1855, pl.37
fig.4) of Harpilius lutescens. This species is apparently found in a
variety of Acropora host species, but none have been identified to
specific level.

Si Philarius lifuensis (Borradaile)
Restricted synonymy:

Periclimenes lifuensis Borradaile, 1898:384; 1899:405, pl.36

IEAL{O fey db,
Hosts. ‘Acropora sp. (Bruce, 1972a):.
Distribution. Recorded only from the type locality in the Loyalty
Islands.
Remarks. The rarest species of this genus. Additional material has
also been recently obtained from Heron Island and Erskine Island,
Queensland, Australia.

16. Hamopontonia corallicola Bruce
Restricted synonymy:
Hamopontonia corallicola Bruce, 1970: 38-48, figs. 1-4.

Hosts. Fungia actiniformis Quoy & Gaimard, (Bruce, in press, c).

Goniopora stokesi Milne-Edwards & Haime (Bruce, 1970).
Distribution. Known only from Hong Kong and Queensland Australia.
Remarks. The colouration of specimens found in association with the
Fungia is generally similar to that of the type material on Goniopora,
although the specimens are distinctly larger in size. Mortensen
(1925, in: Balss, 1956) refers to a shrimp from the Kei Islands, which
lives among the white tipped tentacles of a Fungia coral, and which is
colourless with two white spots on carapace, which almost certainly
belongs to this species.

TES Ischnopontonia lophos (Barnard)
Restricted synonymy:

Philarius lophos Barnard, 1962:242-243, fig.2.

Isehnopentonia lophos—Bruce, 1966:585-597, figs. 1-5.
Hosts. Galaxea fascicularis (L.), (Bruce, 1966).
Distribution. Common in the western Indian Ocean; also Singapore;
Pulau Perhentian Besar, Malaysia and Queensland, Australia.
Remarks. This species apparently strictly associated with this single
coral host type, where it is often found with Platycaris latirostris
and occasionally with Anapontonia denticauda. The species is
remarkable for its particularly strongly bilaterally compressed body
form, and it moves actively in the spaces between the host corallites.

18. Anapontonia denticauda Bruce

Restricted synonymy:
Anapontonia denticauda Bruce, 1967:8-12, figs.1-4.
Hosts. Galaxea fascicularis (L.), (Bruce, 1967)
Distribution. Known only from Zanzibar; Singapore; Pulau Perhentian
Besar, Malaysia and Queensland, Australia.
Remarks. Of the three species of pontoniine shrimps restricted to this
host, Anapontonia denticauda is the least common.

=

iS) Metapontonia fungiacola Bruce
Restricted synonymy:
Metapontonia fungiacola Bruce, 1967:24-30, figs.9-12.

Hosts. Fungia spp. (Bruce, 1967).

Halomitra sp. (Bruce, in press, h).

Hydnophora microconus Lam. (Bruce, 1972e).

Goniastrea pectinata (Ehrenberg) (Bruce, 1974b).
Distribution. Known only from the western Indian Ocean: Kenya,
Tanganyika and the Comoro and Seychelle Islands.
Remarks. One of the smallest pontoniine shrimp species. The species
of Fungia with which this shrimp has been found to associate have so
far not included Fungia actiniformis, which does not occur in the |
region where M. fungiacola has been found. i

—<—$—=

—<$—<———

—$—$—

BOR Platycaris latirostris Holthuis
Restricted synonymy:

PIAEGCAGES Iaerroseris, Holehuads, 952 2G), =l/3—-L7 6),

BEGSe 1So— Sor.
Hosts. Galaxea fascicularis (L.) (Bruce, 1966). |
Distribution. Common in the western Indian Ocean; Flores, Indonesia. |
Remarks. This species appears to be strictly associated with this |
single host species. It 1s remarkable for its strongly depressed
body form. It is sluggish in behaviour, resting in a head down
position on the corallites.

Babe Paratypton siebenrocki Balss
Restricted synonymy:
Paratypeon slebenrocki, — Bruce, LICO=/1—-l85, figs. L=5;,
jollodl Gee

Hosts. Acropora hyacinthus (Dana) (Bruce, 1972).

Acropora massawensis von Marenzeller (Bruce, in press, e).

Acropora palmerae Wells (Bruce, 1972a).

Acropora squamosa Brook (Bruce, 1972a).

Acropora squarrosa (Ehrenberg) (Bruce, 1972a).

Acropora variabilis (Klunzinger) (Bruce, 1974b) .
Distribution. Recorded from the Red Sea, Kenya, Tanganyika and
Zanzibar, the Seychelle Islands, Marshall and Samoan Islands and Heron
Island, Australia.

Remarks. This, species: is. remarkable, for 1s habi tof) living sine passas
in: small cysts completely enclosed by ‘the host !s,coral lum, withyonlayea
few minute apertures connecting the chamber with the exterior.

Dk Fennera chacei Holthuis
Restricted synonymy :
Fennera chacei. Holthuis,.1951: 171-178, pl.54 a-p. Bruce,
VIES 2 8O—82 7) eliguali wil OW Aloe. ki, sm talecien A
Hosts. Pocillopora verrucosa (Ellis & Solander), (Patton, 1966).
Pocillopora eydouxi Milne-Edwards & Haime, (Bruce, 1965).
Distribution. Kenya, Maldive and Seychelle Islands and Willis Island,
off NE Australia.
Remarks. One of the smallest pontoniine shrimps. It may be found on
the same host colonies with Harpiliopsis depressus and H. spinigerus.
Originally discovered in the eastern Pacific region on the shores of
Mexico, Costa Rica, Panama and Colombia, on Porites.

23. Harpilius beaupresi (Audouin)
Restricted synonymy:
Harpidliopsis: beaupresi Borradaile, 1917: .324, 379, spl.55,
fig .21., Kemp, L922. 229-231 Rugs...0/a00%

Hosts. “Pocillopora acuta A,amarck,', (Bruce, 19724).

PocilJlopora..damicornis, ‘(Uh.) (Patton, 1 L966).

Pocillopora danae (Verrill), (Bruce, 1972a).

Pocillopora elongata (Audouin), (Bruce, 1972a).

Pocillopora eydouxi Milne-Edwards & Haime, (Bruce, 1972a).

Pocillopora verrucosa (Ellis & Solander), (Patton, 1966).

Pocillopora woodjonesi Vaughan, (Bruce, 1972a).

Seriatopora. hystrix. (Dana), (Patton, 1966).

Stylophora erythraea von Marenzeller, (Bruce, 1972a).

Stylophora mordax (Dana).,, (Bruce, 1972a).

Stylophor palmata (Blainville), (Bruce, 1974b).

Stylophora: pistillata (Hsper..)5. (Brace, 19%2a)..

Acropora variabilis (Klunzinger), (Bruce, 1972a).
Distribution. Widespread throughout the Indo-West Pacific region from
the Red Sea to Mocambique and Madagascar as far east as Hawaii; also
extending into the East Pacific region.

Remarks. One of the commonest pontoniine coral commensals, frequently
found in association with H. depressus and H. spinigerus.

24. Harpiliopsis depressus (Ortmann)
Restricted synonymy:
Harpilius depressus Stimpson, 1860:38. Kemp, 1922:228
(key), 9231234, £igs 69-70.
Harpiliopsis depressus Borradaile, 1917: 379 (key), 380.
Poenius tos Oo, DlSast 22 NOS2 <6 l82—1eA ser a9O2
Hosts. Pocillopora damicornis (L.) (Bruce, 1972a)*
Pocillopora elongata Dana, (Bruce, 1972a).
Pocillopora eydouxi Milne-Edwards & Haime, (Bruce, 1972a).
Pocillopora ligulata (Dana), (Chace, 1937).
Pocillopora meandrina var. nobilis Verrill (Castro, 1971).
Pocillopora verrucosa (Ellis & Solander), (Patton, 1966) *
Pocillopora woodejonesii Vaughan (Bruce, 1972a).
Seriatopora angulata Klunzinger, (Bruce, 1972a).
Seriatopora hystrix (Dana), (Patton, 1966).
Stylophora mordax (Dana) (Patton, 1966).
Stylophora pistillata (Esper.), (Patton, 1966).
Acropora sp. (Bruce, 1972a).
Porites sp. (Holthuis, 1951).
Distribution. Widespread throughout the whole Indo-West Pacific region
from the Red Sea to Madagascar as far east as Hawaii and in the East
Pacific to the Galapagos Islands, California, Mexico, Costa Rica,
Panama and Colombia.
Remarks. One of the commonest pontoniine coral commensals, frequently
found in mixed association with H. beaupresi andH. spinigerus.
Juveniles of this species have also been collected on one occasion from
a colony of the hydrozoan Millepora sp. There has been considerable
confusion in the zoological literature between this species and
H. spinigerus and many of the records are of dubious value. Those
indicated with an asterisk have been recently confirmed. The
P. ligulata record is from the East Pacific region.

25 Harpiliopsis spinigerus (Ortmann)
Restricted synonymy:
Anchistus spinigerus Ortmann, 1890:511, pl.36 fig.23.
Harpilius depressus var. gracilis Kemp, 1922:228 (key),
A34—23 5 EUG. es
Harpiliopsis depressus var. spinigerus Holthuis, 1952:16

184-185.
Harpiliopsis spinigerus Bruce, 1974:224; (in press, d).
Hosts. Pocillopora eydouxi Milne-Edwards & Haime, (Bruce, in press, j).
Pocillopora verrucosa (Ellis & Solander), (Bruce, 1974).
Stylophora mordax (Dana), (Bruce, in press, d).
Stylophora palmata (Blainville), (Bruce, in press, e).

Stylophora pistillata (Esper.), (Bruce, in press, e).
Distribution. Known with certainty only from Kenya, Farquhar Island,
the Andaman Islands, Celebes and Samoa.

Remarks. Some of the host records given under H. depressus probably
refer to this species, which appears to be common and widespread
probably throughout the whole Indo-West Pacific region.

10

26. Cavicheles kempi Holthuis
Restricted synonymy:

Cavicheles kempi Holthuis, 1952:17, 205-208, figs. 99-101.

Bruce, 1966:266-269, figs.1-2.
Hosts. | Acropora. Sp.) (Byucel, 1966-0 jim jpress., in)
Distribution. Recorded only from Halmahera, the Molucca Archipelago;
Kenya, Zanzibar and Tanganyika; and the Comoro Islands.
Remarks. The East African and Comoro Islands material was found in
Acropora colonies generally in association with numerous specimens of
Jocaste spp. and it is considered probable that this species is a
post-larval or early juvenile of J. japonica. (Ortmann) -

Dylee Jocaste japonica (Ortmann)

Restricted synonymy:

Jocaste japonica—Patton,,.1966: 279-280, 288 tab. ll} 290ntabeey
IEAL(O fs S119)e lsabKore\ yy ADS) TMloye AUS SSI o)S) ee ae alio ey 9) a
Hosts. Pocijilopora-demicornis _Uu.);,, (Bruce, 19724).

Acropora abrotanoides (Lam.), (Bruce, in press, e).

Acropora assimilis (Brook), (Bruce, in press, e).

Acropora convexa (Quelch), (Bruce, 1972a).

Acropora corymbosa (Lam.), (Bruce, in press, e).

Acropora digitifera.(Dana),,..(Bruce;, 1972a) .

Acropora disticha (Brook), (Bruce, in press, e).

Acropora diversa (Dana), (Bruce, 1972a).

Acropora haimei (Milne-Edwards & Haime), (Bruce, 1972a).

Acropora humilis .(Dana), (Bruce, 1972a).

Acropora massawensis (von Marenzeller), (Bruce, in press, e).

Acropora nana (Studer), (Bruce, 1972a).

Acropora nasuta (Dana), (Bruce, 1972a).

Acropora pectinata (Brook), (Bruce, in press, h).

Acropora rotumana (Gardiner), (Bruce, 1972a).

Acropora spicifera (Dana), (Bruce, in press, h).

Acropora squamosa (Brook), (Bruce, 1972a).

Acropora squarrosa (Ehrenberg), (Bruce, in press h).

Acropora surculosa (Dana), (Bruce, 1973).

Acropora: valida. (Dana), .(Bruce, in press e).

Acropora variabilis (Klunzinger), (Bruce, 1972a, 1974b).
Distribution. Widespread throughout most of the Indo-West Pacific
region, but not recorded from the Red Sea, Arabian Sea or Bay of Bengal,
nor east of the Marshall Islands.

Remlarks. =,cOMme Lepoxnts Of J. lucina May Kener tO thas Species. ne
reference to.an association of this shrimp with Acropora tenuis,
(Bruce, 1972a) was erroneous due to misidentification. As noted above,

Cavicheles kempi is probably a juvenile stage of this species.

28.6 Jocaste lucina (Nobili)
Restricted synonymy:
Jocaste lucina - Patton, 1966:278-279, 288 tab.1, 290 tab.2,
2927) EG Sale Beuce, OAD OOP EgaS
Hosts. Pocillopora verrucosa (Ellis & Solander), (Patton, 1966).
Stylophora sp.
Acropora conferta (Quelch), (Bruce, 1972a).

11

Acropora corymbosa (Lam.), (Bruce, 1972a).

Acropora cuneata (Dana), (Bruce, 1972a).

Acropora eurystoma (Klunzinger), (Bruce, in press, j).

Acropora haimei (Milne-Edwards & Haime), (Bruce, in press, e).

Acropora hebes (Dana), (Bruce, 1972a).

Acropora humilis (Dana), (Bruce, 1972a).

Acropora palifera (Lam.), (Bruce, 1972a).

Acropora ramiculosa (Dana), (Bruce, 1972a).

Acropora Squamosa (Brook), (Bruce, 1972a).

Acropora tenuis (Dana), (Bruce, 1972a).

Acropora teres (Verrill).

Acropora variabilis (Klunzinger), (Bruce, 1972a).

Porites sp.
Distribution. Throughout the Indo-West Pacific region, except the
Hawaiian Islands.
Remarks. The associations of Jocaste spp. with Acropora spp. has been
reviewed by Bruce (1969c). Details were provided of eighteen host
species with J. jocaste exclusively associated with eleven species,
J. lucina with ten and three coral species, A. humilis, A. squamosa and
A. variabilis acting as hosts for both shrimp species. The number of
Acropora species now known to serve as hosts for Jocaste spp., iS now
increased to twenty nine, with fifteen exclusively for J. japonica,
nine for J. lucina, and with the addition of two species, A. corymbosa

and A. haimei, to those hosting both species of shrimps. The two
specimens found in association with Porites are juvenile, but those
with Stylophora were adult (11, 4 ovig. ) and were associated with

Coralliocaris and Harpiliopsis spp.

29. Coralliocaris brevirostris Borradaile
Restricted synonymy:
Coralliocaris brevirostris Borradaile, 1898:1006, pl.64,
PUCtie a. SRC peo antO> 406), 407 7 sao itkey)s, seal cel.

Hosts. Acropora corymbosa (Lam.), (Bruce, 1972a).
Acropora cymbicyathus (Brooks).
Acropora humilis (Dana), (Bruce, 1972a).

Acropora hyacinthus (Dana).
Distribution. Known only from the Ellice Islands, and Willis Island,
NE Australia.
Remarks. The specimen from Mauritius, referred to C. brevirostris by
Holthuis (1965), should be referred to C. nudirostris Heller. The
chelae of the second pereiopods in C. nudirostris show a close
resemblance to C. brevirostris, rather than C. venusta.

30. Coralliocaris graminea (Dana)
Restricted synonymy:
Coralliocaris graminea - Holthuis, 1952:17, 186-189, fig.91l.

Hosts. Pocillopora danae (Verrill), (Bruce, 1972a).
Pocillopora elongata Dana, (Bruce, 1972a).
Seriatopora hystrix (Dana), (Patton, 1966).
Stylophora erythraea von Marenzeller, (Bruce, 1972a).
Acropora corymbosa (Lam.), (Bruce, 1972a).

Acropora cymbicyathus (Brooks), (Bruce, 1972a).

WZ

Acropora digitifera (Dana), (Bruce, in press, e).

Acropora diversa (Dana), (Bruce, 1972a).

Acropora haimei (Milne-Edwards & Haime), (Bruce, 1972a).

Acropora hebes (Dana), (Bruce, 1972a).

Acropora humilis (Dana), (Bruce, 1972a).

Acropora nana (Studer), (Bruce, 1972a).

Acropora nasuta (Dana), (Bruce, 1972a).

Acropora ramiculosa (Dana), (Bruce, 1972a).

Acropora squamosa (Brooks), (Bruce, 1972a).

Acropora squarrosa (Ehrenberg), (Bruce, in press, e).

Acropora variabilis (Klunzinger), (Bruce, 1972a).
Distribution. Probably common throughout the whole Indo-West Pacific
region except for the Hawaiian Islands.
Remarks. Many of the records of C. graminea Probably refer to
C. viridis, and similarly some of the host records may also refer to
this species. Although these two closely related species may be
found in the same localities, they do not occur together on the same
coral host colonies, as may occur in the species-pair Harpiliopsis
depressus and H. spinigerus. Specimens from Seriatopora and Stylophora
are usually small juveniles and these genera must be regarded as
abnormal hosts.

Bile. Coralliocaris macrophthalma (Milne-Edwards)
Restricted synonymy:
Coralliocaris macrophthalma - Borradaile, 1898:385. Nobili,

ASTON ER eh
Hosts. Probably Acropora spp., but none so far recorded.
Distribution. Known only from "Mers d'Asie", the Red Sea and the
Seychelle Islands.
Remarks. This little known species is most closely related to
C. graminea and C. viridis, with sound-producing chelae on the second
pereiopods.

Bie Coralliocaris nudirostris (Heller)
Restricted synonymy:
Oedipus nudirostris Heller, 1862:279, pl.3 fig.25.
Coralliocaris nudirostris - Borradaile, 1971:382 (key), 384
Bruce, 197462262); Elge2.

Hosts. Acropora digitifera (Dana), (Bruce, 1972a).
Acropora humilis (Dana), (Bruce, 1972a).
Acropora hyacinthus (Dana), (Bruce, 1972a).
Acropora spicifera (Dana), (Bruce, press, h).
Acropora surculosa (Dana), (Bruce, 1973).

Distribution. Reported only from the Red Sea, Tanganyika, the Seychelle
and Maldive Islands and Tahiti.

Remarks. On East African fringing reefs, this species seems to prefer
hosts situated on the exposed side of the reef crest rather than the
lagoon side.

B38 Coralliocaris pavonae Bruce
Restricted synonymy:
Coralliocaris pavonae Bruce, 1972:63, 77-84, figs. 8-11.

183

Coralliocaris taiwamensis Fujino & Miyake, 1972:95-98,
iEaLeisig l=
Hosts. Pavona divaricata (Lam.), (Bruce, 1972a)
Pavona minor Breuggemann, (Bruce, 1972a).
Distribution. Known from Fiji and Taiwan only.
Remarks. The only species of the genus that is apparently normally
associated with a coral host not belonging to the genus Acropora.

34. Coralliocaris superba (Dana)

Restricted synonymy:

Coralivtocaris superba — Holthurs, LI52217, LS9-191, £ig.92.
Hosts.. Pocillopora danae (Verrill), (Bruce, 1972a)

Acropora africana (Brooks), (Bruce, in press, h).

Acropora convexa (Dana), (Bruce, in.press, e).

Acropora cymbicyathus (Brooks), (Bruce, 1972a).

Acropora digitifera (Dana), (Bruce, 1972a)

Acropora disticha (Brooks), (Bruce, in press, h).

Acropora diversa (Dana), (Bruce, 1972a).

Acropora eurystoma (Klunzinger), (Bruce, 1972a).

Acropora humilis (Dana), (Bruce, 1972a).

Acropora irregularis (Brooks), (Bruce, 1972a).

Acropora leptocyathus (Brooks), (Holthuis, 1953).

Acropora nana (Studer), (Bruce, 1972a).

Acropora nasuta (Dana), (Bruce, in press, h).

Acropora pectinata (Brooks), (Bruce, in press, h).

Acropora pulchra (Brooks), (Bruce, 1972a).

Acropora variabilis (Klunzinger), (Bruce, in press, h).
Distribution. Widespread throughout the Indo-West Pacific region from
the Red Sea to Madagascar as far east as Tahiti and the Society
Islands.

Remarks. Five of the host corals also serve as hosts for C. "graminea".
This species is one of the most beautifully coloured of pontoniine coral
commensals (Bruce, 1974c, fig.3), and was accurately illustrated by

Dama CkSS 57, spills sw! telcye2)).

S5k Coralliocaris venusta Kemp

Restricted synonymy:

Coralliocaris venusta Kemp, 1922:269, 274-276, figs.100-101.
Hosts. Acropora convexa (Dana), (Bruce, 1972a).

Acropora cymbicyathus (Brooks).

Acropora humilis (Dana), (Bruce, 1972a).

Acropora hyacinthus (Dana).

Acropora subulata (Dana), (Bruce, in press, h).

Acropora pectinata (Brooks), (Bruce, in press, h).
Distribution. Red Sea, central East Africa, the Seychelle Islands,
Mocambique Channel, Indonesia, the South China Sea, Samoa and the
Great Barrier Reef.

Remarks. It has been noted (Bruce, in press, h), that two distinct
species have probably been confused under this name. The two "species'
are readily separated by their distinct colour patterns in life, but no
satisfactory morphological distinction has yet been found.

14

Ors Coralliocaris viridis Bruce
Restricted synonymy:
Coralliocaris viridis, Bruce, 1974ac222-224)) figs livasb.

Hosts. Acropora disticha (Brooks), (Bruce, in press, h).

Acropora humilis (Dana), (Bruce, in press, h).

Acropora valida (Dana), (Bruce, in press, h).

Acropora variabilis (Klunzinger), (Bruce, in press, h).
Distribution. Known only from southern Kenya, the Mocambique Channel
and the Great Barrier Reef. Probably throughout the Indo-West Pacific
region.

Remarks. Many records of the closely related C. graminea probably

refer to this species.

Sine Pontonides maldivensis(Borradaile)
Restricted synonymy:
Pontonides maldivensis - Borradaile, 1917:387, pl.57 fig.28.
Host. Dendrophyllia micracantha (Ehrenberg), (Bruce, in press, h).
Distribution. Known only from the Maldive Islands and Mombasa, Kenya.
Remarks. This species and the next are the only known commensals of a-
hermatypic corals.

38. Pontonides unciger Calman
Restricted synonymy:

Pontonides unciger - Holthuis, 1952:219-223, fig.108-110.
Host. Dendrophyllia ijimae Yabe & Eguchi, (Fujino & Miyake, 1969).
Distribution. Reported from the Red Sea; Amakusa Islands, Japan, and
Indonesia.
Remarks. Other species of the genus Pontonides have been found to
associate with gorgonian or antipatharian hosts.

Discussion

Details are given of thirty seven species of Indo-West Pacific
pontoniine shrimps that are considered to be obligate commensals of
scleractinian corals. The criteria for assessing the obligate nature
of the association have been provided by Garth (in press), who indicated
that the shrimps concerned are not found away from their host, that the
association in recurrent, and that adult breeding individuals are
involved. One species, Periclimenes parvus, is considered to be a
doubtful coral-associate. Fourteen genera, out of a present total of
forty-three known Indo-West Pacific pontoniine genera, have now been
found to have some species living in association with corals and of
these genera, nine are monospecific. One of these monospecific genera,
Vir Holthuis, is one of the least specialized pontoniine genera, with
an unmodified set of palaemonid mouthparts, including a mandible with
ar small pala: The eight other nonospecific genera consist of highly
specialized species, showing an increasing degree of reduction of many
parts of the mouthparts.

The hosts are represented by seventy-six different species, of
seventeen genera, although forty-one of these belong to the genus

5

Acropora alone and sixteen to the branching corals of the families

Thamnasteridae (Pocillopora, Seriatopora and Stylophora). The
Acroporidae (Acropora spp.) clearly provide the most satisfactory
niches for these shrimps. It is certain that many hosts still remain

to be identified, especially those from the reef front and from deeper
water, as most of the records have been derived from shallow lagoons
OF pools on the reef flats easily accessible at low tide.

The most favoured host coral is Acropora humilis (Dana). This
species has been found to provide a niche for a Periclimenes, a
Philarius, two Jocaste and six Coralliocaris species. Although these

do not occur all on a single host colony, it is not uncommon to find
Periclimenes lutescens with two Jocaste and two Coralliocaris spp.
together in a single host. Similarly, Pocillopora damicornis (L.) may
act as host for Vir orientalis, three Periclimenes and two Harpiliopsis
spp. The number of species and individuals present in a coral colony
is largely dependant upon the size of the colony, small colonies often
having only a single male-female pair of one species. Large corals
may have over a hundred individual shrimps (Bruce, 1972a).

The abundance of coral-associated pontoniine shrimps in the Indo-
West Pacific region is in marked contrast with findings in other
tropical seas (Bruce, in press, g). Of the genera associated with
corals in the Indo-West Pacific region, only Periclimenes is found to
occur elsewhere and none of its extra-Indo-West Pacific species are
reported to be specifically associated with corals, although the hosts
of several species that are possibly commensals remain to be ascertained.
In the Eastern Pacific region two species of pontoniine shrimps,
Harpilius depressus and Fennera chacei, are known to live in association
with coral hosts. Both these species are of widespread distribution
and have probably extended their ranges across the East Pacific
barrier, via the Galapagos Islands, by means of the Equatorial Counter
Current. In the Caribbean region there are no reported associations
between pontoniine shrimps and corals and even the Acropora species,
which provide a niche for so many of the Indo-West Pacific species, are
without a Similar series of faunal associates.

Acknowledgements
For the identification of many of the coral hosts here reported,

I am particularly endebted to P.M.J. Woodhead, Dr. B.R. Rosen,
Dr. J.W. Wells and Dr. H. Utinomi.

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Nos. 206-207

VL
i a ae
AF ©
fag

May 1977

ATOLL RESEARCH
BULLETIN

206. THE NATURAL HISTORY OF NECKER ISLAND,
NORTHWESTERN HAWAIIAN ISLANDS

by Roger B. Clapp and Eugene Kridler

207. THE NATURAL HISTORY OF NIHOA ISLAND,
NORTHWESTERN HAWAIIAN ISLANDS

by Roger B. Clapp, Eugene Kridler and Robert R. Fleet

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

ATOLL RESEARCH BULLETIN
NOS. 206 - 207

206. THE NATURAL HISTORY OF NECKER ISLAND, |
NORTHWESTERN HAWAIIAN ISLANDS |

by Roger B. Clapp and Eugene Kridler .
4

207. THE NATURAL HISTORY OF NIHOA ISLAND,
NORTHWESTERN HAWAIIAN ISLANDS }

by Roger B. Clapp, Eugene Kridler and Robert R. Fleet

Issued by
THE SMITHSONIAN INSTITUTION
with the assistance of
The United States Fish and Wildlife Service
U.S. Department of the Interior

Washington, D. C., U.S.A.

May 1977

ACKNOWLEDGEMENT

The Atoll Research Bulletin is issued by the Smithsonian Institution
as a part of its Tropical Biology Program. It is co-sponsored by the
Museum of Natural History and the Smithsonian Press. The Press 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 de-
voted 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

ATOLL RESEARCH BULLETIN
NO. 206

THE NATURAL HISTORY OF NECKER ISLAND,
NORTHWESTERN HAWAIIAN ISLANDS

by Roger B. Clapp and Eugene Kridler

Issued by
THE SMITHSONIAN INSTITUTION
with the assistance of
The United States Fish and Wildlife Service
U.S. Department of the Interior
Washington, D. C., U.S.A.

May 1977

TABLE OF CONTENTS

Page
EATS Tega OFA HAT CURIE) Stews oncieanebaheichotapats stays csnctosteiosctievs: Sicmeteriotene: @ieus ehsce-s asauaee ara avets EAE
AT SHO PACTA TAD Gie ole cee Wes s oeeee ee Rete. Paslode oFetaUsbs, cde’Mer's laf: «.cuolsteltokioBels: «foie secbllel.» ve ttt
ARS O Fee APPEND IPNGCIAIB IGE: Gietewel stetroniey s totteticve its: telohiei elisuel oMetehe tile ‘etfebe: oMelierei lets ieveastene 1v
HUENGTSER DUT CATT’ O) Nitefnatorotrsiteiticebathaitewetrarent sthadia’tallelis lsiienararaileeNculleaiclciatets sede elewelidila cite rae ls il
DYESS CREE Teli ONicrokaeotions siete ciekatouetepecePelonetcrc e: «. eo. etovav0i06) ce elete acl ee elelisc ee: siecle (els) e's 4
(CHOICES diag 6d 6 0.0.0 0.010900 60.6 OOO OOO 6 0:0 OIG OIG. Olt CRO CIDIOLOIOTO Diol PII OR One JEN
HILSOR Ao obo oOo e coon oo OD Oo e OU CODD CUO MOOD COON DDO DO OO DN onU SC OnOOUC 2
AE CE ATSAC MNO) Nitetotet omomet cn alicla\iasis) Momma denen veramayia sicttel swell aclctolarel(cvies'es iiss leisite) ere ete} ofene: lias 25
THER RIE RED AT {ERG BVA HSiede tiatntise: ole altetcttelis, llsiselalicifelisyicitalle: ctfuilel ells) shofelieltcHalle voticlleite Sil
BS STECSiekkotelie! eneKeleNeiolsl(oMolexaalisielaisits! isiislie sielslielisifefelleiene! ois ie) cl eliclsl/sielevelle (ofelisirelefiols Bilt
Species NQCOWIMESB OSC OOS DODD OOOOOG DO OOO ODD OGOUOKO OO OOOO OO 3)5)
Dtomedea nigrtpes Sin ketohe Wanelicneblotiolats crnvecene detours epopeiemene:chevens 36
DEOMCACOR TTT E DAL GHo deter cnenenreiclicia/ ch tele ate: henelenetcee tenor. 38
Bia OVP OMLALIDC TD acteloc av elise ehetls: etebnliey shies eo sshepeneieney ©: eie/e\ elas 42
PUSFINUS! PACTFICUS 0.00 o eevee eesccceseccdesescees Ad
Puffinus NACTVICAETS +++ r reer ev cereecerveececees 46
OECANOGROMAMETLZEG FIGHT helstele elelcisicneleie ais/ ete) sielorelsicrsiosiote 46
PHACENO MPU LCC CGuord Med etel cho! ctel el cuelotele (ci dfeqeteneretie ts cll. 46
Sula dactylatra++.ereeceerecceeceerecececeeevcnee 48
Sula Leucogaster SGDOODO OOO OO OIC O OD OG6OO 0000 GOOD UC Syl |
Gyilte) BUILGI OAC CD ACO Dod RD OOD OOO OD COO ODOOLaGOD DONO GOGO 53
Fregata MBINOIDO OS DAR ood DOO OND OO DOD OOO oso Mood dDODdC 56 i
PLD LOMLUSUGLONIENLEC Gin tale) orctckoekehetel siete =| excl chiehoiteltohte del eieyoneue! 59
INDSMETTILUS WOM IPE DEL SO 000 Don bo DnOn COO dso udo on oOddS 60
HOGOCPOSGOLUE EHCONUGI: eke clerolele cleleroielersrs slle clue olptetate cre 61 {
APENAYTA INFEPPVES + sere ree rce eee ecnreererceccenes 62 f
COB WAGIELES GHOGID DO GO COCA COD DOO DOD OD ORO O10, 0,01cuGio.d Soc 63 i
Larus g Llaucescens SO Ob ORG ODOOOO.0006 DOO OOO.C OU OK 63 1
ECACC TITIELECINS cred cliotat ere ccbete alielic tel efeWots, oie belie ciel chelate ee ots 63 ;
StCPna FUSCATA++ +s reer ere re ere r eee eeererereecece 66 f
PHOCCUSECERCMCCHILLCG nie ctiubelatererele clea elie ic <itis}is et ael ells teits 69 |
AOU S COWES Merete ei istelone totetetenciavetets fetetcle otal letelerere ters UP ;
AHOUGMCCTIIEEO SELLS olebetetensl sie eleisteNetsiinfiarelieietsia)claheta erslicielels 75 )
Gygts DiLIOOLE.G 00000 010.0 60.0'0.0 0.0 60.0.0 006, 0.00 OOO DIG Gne Vi
Mimus polyglottos Sie a ee aatis aerate MeeveMe eRe: alisterroteivelie, evleMereievetio 79 f
MammalS--eeececeeeneereeererererererereererseseseseeesssvceseens 80 |
Monachus SChQUINSLANdT +++ eee eee revcnvesnnevesees 80 |
Reptiles DODO OOODGHO OOD OOD OCOOUOD OOD OD OOD OOD OC OOOOUOU UO UDO OOOO 81 i
Chetonta mydas 5.0,0,.010 0.0100 62001010010 010, 61010, 05610 OO 01010'0.0.0'0,0-0 82 ]
AGKNOWUIEED ChE NS temelaieretete letete re peteven -Neleiellehefoieiaisilclcielelic cletelerelclclsichelwlola fers creme 83 }
LigveRyNnis (tite dcocbgoubooboUo Gn dduoddoDOdoH OOO OU GOO UDOO Ob 85 I
EMRMIDID SNL R660 530060000000 00505000500 UE O00 DOGO OO OOCL 92

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14.

LIST OF FIGURES

The Hawaiian Islands.

Map of Necker Island. After Judd (7m Palmer, 1927) and Elschner
(1915). Names in brackets are those used by Elschner; the others
are those used by the Tanager Expedition and subsequent visitors.

Aerial photograph of Necker Island showing prominent features of
island. Official U.S. Navy photograph, January 1966.

Oblique aerial view of Necker Island from the northeast. BSFW
photograph by David B. Marshall, June 1962,

Shallow ridge between Flagpole (right) and Summit (left) Hills.
Photograph by Derral Herbst, 28 August 1968.

Shallow ridge between Bowl (middle) and Summit (left) Hills.
Photograph by Derral Herbst, 28 August 1968.

Low gap between Flagpole (left) and Annexation (right) Hills.
Photograph by Derral Herbst, 28 August 1968.

Gap between Northwest Cape and Annexation Hill as viewed from
Bowl Hill. Shark Bay in right mid-ground and looking into the
East Cove landing. Photograph by Derral Herbst, 28 August 1968.

Frequently used landing area in the West Cove looking at the
southeastern peak of Northwest Cape. Photograph by Derral Herbst,
28 August 1968.

Adult and immature Red-footed Boobies and Great Frigatebird roost-
ing on uprights of marae on Annexation Hill. Blue-faced Booby in
mid-ground and large nestling Great Frigatebirds in background.
Photograph by Derral Herbst, 28 August 1968.

Immature Red-footed Boobies roosting on uprights of marae on
Annexation Hill. Nestling Great Frigatebirds and immature Red-
footed Booby in foreground. Photograph by Derral Herbst,

28 August 1968.

Saddle between Summit and Flagpole Hills, August 1968. Note dense
growth of Chenopodtum. Photograph by Derral Herbst.

Saddle between Bowl and Summit Hills, August 1968. Photograph by
Derral Herbst.

Portulaca lutea Sol. growing on the Northwest Cape. Note nesting
Red-tailed Tropicbird in the left background. Photograph by
Derral Herbst, August 1968.

10

14

14

28

28

30

1336

16.

ttt

Page
Young Laysan Albatross among Red-footed Boobies and Great
Frigatebirds nesting in Chenopodium. Northwest Cape in the
background. BSFW photograph by David B. Marshall, 11 June 1962. 39
Sooty Tern nesting area in the saddle between Flagpole and
Annexation Hills. Looking toward Annexation Hill. Photograph 68
by Derral Herbst, 28 August 1968.

LIST OF TABLES

Recent surveys of Necker Island by the POBSP and BSFW. 3
The avifauna of Necker Island. 32
Observations of Black-footed Albatross on Necker Island. 3
Observations of Laysan Albatross on Necker Island. 40
Observations of Bulwer's Petrel on Necker Island. 43
Observations of Wedge-tailed Shearwaters on Necker Island. 45
Observations of Red-tailed Tropicbirds on Necker Island. 47
Observations of Blue-faced Boobies on Necker Island. 50
Observations of Brown Boobies on Necker Island. By
Observations of Red-footed Boobies on Necker Island. 54
Observations of Great Frigatebirds on Necker Island. 58
Observations of Golden Plovers on Necker Island. 60
Observations of Ruddy Turnstones.on Necker Island. 61
Observations of Wandering Tattlers on Necker Island. 62
Observations of Gray-backed Terns on Necker Island. 65
Observations of Sooty Terns on Necker Island. 67
Observations of Blue-gray Noddies on Necker Island. 7fal
Observations of Brown Noddies on Necker Island. 74
Observations of Black Noddies on Necker Island. 76

4v

20;

21.

22

Observations of White Terns on Necker Island.
Observations of Hawaiian Monk Seals at Necker Island.

Observations of Green Turtles at Necker Island.

LIST OF APPENDIX TABLES
Scientific visits to Necker Island, 1891-1973.
Results of scientific visits to Necker Island, 1891-1973.

Publications on collections and studies (with the exception
of birds) made on Necker Island, 1891-1973.

Page
78

80

82

GZ
96

98

THE NATURAL HISTORY OF NECKER ISLAND, NORTHWESTERN HAWATTAN ISLANDS!

by Roger B. Clapp and Eugene Kridler?

INTRODUCTION

Necker Island, now a part of the Hawaiian Islands National Wildlife
Refuge, is a small, precipitous, rocky island that lies at 23935' North,
164942' West (Off. of Geogr., 1956: 58) near the eastern end of the
northwestern Hawaiian Islands (Figure 1). Its nearest neighbors in the
chain are French Frigate Shoals, lying about 75 miles to the westward,
and Nihoa Island, some 155 miles to the east-south-east (Great. Brit.
Hydro. Dept., 1946: 296=297).

Relatively little is known about the fauna of the island, in part
perhaps due to the difficulty of making landings there. The principal
previous surveys of the island were made by the Albatross Expedition in
1902, by Carl Elschner in 1914, and by the Tanager Expeditions of 1923
and 1924. The various scientific reports resulting from these expedi-
tions, particularly those resulting from the Tanager Expeditions, and
Bryan's (1942) summary of information supply most of what was previously
known about the biota as well as the geology of the island.

Beginning in 1964 the Pacific Ocean Biological Survey Program
(hereafter POBSP) of the Smithsonian Institution and the Bureau of
Sport Fisheries and Wildlife (hereafter BSFW) began making periodic
surveys of the fauna of the island. From 1964 through 1973 13 visits
were made, all but five of them in the months of March and September,
which together totaled only 7.0 days of observation (Table 1). Of these
seven days, only three nights were spent on the island, which accounts,
in part, for the paucity of data on petrels.

This report, one of a series on the northwestern Hawaiian Islands,
is primarily intended as a summary of these and previous observations of

1/ Paper Number 75, Pacific Ocean Biological Survey Program, Smithsonian
Institution, Washington, D.C.

2/ Present address: National Fish and Wildlife Laboratory, Bureau
of Sport Fisheries and Wildlife, U.S. Fish and Wildlife Service,
National Museum of Natural History, Washington, D.C. 20560.

3/ Present address: Bureau of Sport Fisheries and Wildlife, U.S. Fish
and Wildlife Service, Kailua, Oahu, Hawaii 96734.

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the terrestrial vertebrates and vascular plants of the island. A brief
description and history of the island are also included. Secondarily,
this report should serve as a summary and reference (see Appendix Tables
1 and 2) to other biological and other information previously published
about the island. Through the courtesy of Dr. Alexander Wetmore, the
previously largely unreported observations on the birdlife made by him
in 1923 are also included.

The present report was largely in final draft form in late 1970
and includes only slight emendation or addition after that period.
Observations reported include those available through 1973. Except for
botanical observations, this largely excludes the August 1968 survey,
for which most data were not available.

BSFW and POBSP field notes and trip reports concerning Necker are,
respectively, stored in the Bureau of Sport Fisheries and Wildlife files,
Kailua, Oahu, Hawaii, and the Pacific Ocean Biological Survey Program
files, National Museum of Natural History, Washington, D.C.

DESCRIPTION

Necker, with an estimated area of 41 acres (Fig. 2), is a sharply
rising ridge’ of volcanic rock: (Figs. '3 and 4) which Palmer (1927; ;22)
reported to be the remnant of a volcanic cone that was formerly of much
greater extent. Palmer's report on the geology also gives one of the
best descriptions of Necker. He reported that the island "...consists
of two parts. The principal one is a ridge extending nearly due east
and west, 4,000 feet long and varying from 200 to 600 feet in width.

On this ridge are five peaks....The saddles between the peaks are
shallow [Figs. 5 and 6], except the most westerly [between Annexation
Peak and Flagpole Hill], which drops to only 76 feet in elevation [Fig.
7]. From the westernmost peak a peninsula extends 500 feet north-
northeast to a gap only a few feet above sea level [Fig. 8]." Landings
are generally made here and are often particularly difficult and dangerous
as the prevailing winds from the northeast often cause large breakers to
break across this gap. "The lesser part of Necker Island, Northwest
Cape [Fig. 9], extends about 800 feet northeast from this gap....At the
east end of the main part of Necker is a low islet about 75 feet wide
and 200 feet long. It is awash at high water and waves break over it
continually."

On the north side of the islands the rocks rise precipitously to
the peaks but on the north side are somewhat less steep and according to
Emory ((1928: 51)-"are easily climbed."

The steep slopes of the island have little or no vegetation and
are craggy and wave-sculpted where alternating layers of harder and more
soft rock have been differentially eroded and bird droppings often
whiten the face of the rock (see Fig. 9). Along the shallow valleys
and on the rounded crests of the hills a few species of plants are well
established in the thin soil (see Vegetation Section).

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Figure 5 Shallow ridge between Flagpole (right) and Summit (left)
Hills. Photograph by Derral Herbst, 26 August 1968.

Shallow ridge between Bowl (middle) and Summit (right )
Hills. Photograph by Derral Herbst, 28 August 1968.

Figure 6

Figure 7 Low gap between Flagpole (left) and Annexation (right)
Hills. Photograph by Derral Herbst, 28 August 1968.

Figure 8 Gap between Northwest Cape and Annexation Hill as 1
viewed from Bowl Hill. Shark Bat in right mid-ground and looking |
into the East Cove landing. Photograph by Derral Herbst, 28 August 1968. ii

231-338 O- 77 - 2 |
5 \y

hapure: 9 Frequently used landing area in the West Cove looking
at the Southeastern Peak of Northwest Cape. Photograph by Derral
Herbst, 28 August 1968,

aL

GEOLOGY

Palmer (1927) is the only one who has written to any extent on
the geology of Necker and our brief account here is largely taken from
that paper. Although the remnant of a volcanic cone, no part of the
original surface remains, all the present surfaces having resulted
from the action of wind, sea, and running water. Palmer suggested
that the deeper valley between Annexation and Flagpole Hills has re-
sulted from greater wind erosion caused there by the funneling of
the northwest trade winds by the main ridge of the island and by
Northwest Cape. He (op. ett.: 23) considered it likely that

at least a few hundred feet of overburden rocks
have been removed from all parts of the island.
There are now exposed at the surface various
dikes and sills, some of which are entirely free
of gas pores. This feature is usually taken to
mean that the lava cooled and solidified under
considerable pressure, which in turn implies a
considerable overburden.

Necker has no definite stream channels and all rainwater either
sinks into the rocks or runs off through unorganized channels. During
his visit Palmer found two seeps of ground water, both evidently much
contaminated by bird droppings. One was found near Bowl Cave (see
Fig. 2) on the north slope of the island; the other was about 30
feet above sea level on the north side of the saddle between Flagpole
and Annexation Hills.

Palmer also noted that the beds of lava composing the island
are in general four to six feet thick and that they extend laterally
for considerable distances. Two unusually thick beds, 15 to 20 feet
thick, are to be found at the east end of the main island. He noted
that in general the lava beds strike "N. 700 W., and dip 10° NE" and
that the lava beds are

cut by a number of nearly vertical dikes that

strike about N. 70° E....The dikes vary from one

to four feet in thickness. Some of them, notably
the one on the northeast face of [Annexation
Hill]...are dense and nearly free of vesicles,

but others are moderately porous. Most of the

dikes are fine grained at the margins and coarse
grained in the interior. Some of the dikes branch
one or more times, and one pair of dikes [is] con-
nected by an oblique dike running about east-

west. The dike on the southeast side of Northwest
Cape connects with and seems to have fed a sill that
outcrops on both sides of the cape. This sill meets
the shore near the middle of the cape and extends
upward toward the southeast end of the cape, cutting

2.

across several flows in one place. A sill

about two feet thick crops out for about 150 feet
on the summit of the main island near its east
end, and was also presumably fed by the dike
which is continuous with it.

Palmer concluded that the vent of the volcanic cone of which
Necker is a remnant lay to the southwest of the present position of
the island. He remarked, however, that his conclusion (1927: 24)
"disagrees with the fact that Necker lies to the southwest of the
axis of the ellipse outlined by the surrounding shoals," but
suggested that this might be explained by the fact that the shoal
"was composed of two or more volcanic cones....and that the windward
cones have been removed as they were most exposed to attack by waves
driven by the prevailing northeast trade winds. The area of the
shoal around Necker is about 650 square miles which is large enough
to have included two volcanic domes."

Palmer found no evidence of faulting in the beds composing
Necker and noted that the beds along the westernmost saddle are
continuous.

Analyses of the rock composition have been given by Palmer,
M¥hle (1902), Powers (1920) and Washington and Keyes (1926). A111
the rocks studied were basalts and within this classification con-
sisted of four types, andesine, olivine-labrodorite, picrite and
nephalitic basalts. Further details of chemical composition of
these rocks can be found by consulting the literature cited.

Palmer also found gypsum stalactites and stalagmites in some
of the shallow caves of the island. These differed from the usual
in that they had fairly good crystal outlines rather than being more
rounded in form. According to Palmer, these probably resulted from
the evaporation of salt spray.

HISTORY
Prehistoric Habitation

Evidence of prehistoric habitation of Necker Island was noted
during one of the earliest landings on Necker which occurred about
1879. Captain William C. Bruhn, then a young sailor, evidently on
the schooner JULIA, landed and explored the island.! When he was
interviewed by Atkinson many years later Bruhn remembered seeing
marae (stone platforms with a conventional arrangement of uprights),
four or five idols, a few stone adzes, and a stone "shaped like a
rough dumbbell, two rough ends with a handle to clutch with the
hand. The Hawaiians said that when sharks were led to shallow

enon (1928: 55) found no evidence of a visit in 1879 but discovered
that the JULIA had sailed to’ the South Pacific on 13 July 1882.

13)

places that this dumbbell was used to kill them by beating them on
the head."" None of the idols was collected because of the super-
stitious fear of the Hawaiians on Bruhn's ship, but the dumbbell-
shaped stone was taken and later transported to San Francisco (Emory,
WPS BYE)

The presence of idols and images was again noted on 27 May 1894
when the island was annexed by the Hawaiian government. Seven idols
were taken to Honolulu and a photograph of six of them was published
in the Journal of the Polynesian Society (Alexander, 1894: 153).

Subsequent visits and collections are listed in detail by Emory

(1928). Four images (two now in the British Museum) were collected

by a party from the HBMS CHAMPION on 24 September 1894, but no idols
were found on seven subsequent visits. The latter visits included |
one on 12 July 1895 by King, two of unknown date by George N. Wilcox,
visits in 1910 and 1913 by personnel from the Revenue Cutter THETIS, )
and two visits by H.L. Tucker and party in 1917. Photographs were |
taken of several marae on the latter visit. |

a re-shaped image. In the summer of 1923 a number of artifacts
(including a hammerstone, grindstones, adzes, a chisel, an awl and
bowls) and skeletal material were found by members of the Tanager
Expedition.

|
On 6 October 1919 Gerrit P. Wilder collected an image leg and |
|

Observations made during the Tanager Expedition and on earlier
visits were reported in great detail by Emory (1928) and need not
be repeated at length here. Some records, however, are worth noting.

Emory reported finding 33 marae. Most were rectangular struc-
tures with a narrow elevated platform usually found at the rear, |
and a lower broader terrace at the front (see Fig. 25 in Emory, 1928: |
60). Various stones standing upright were placed on the structure
(Figs. 10 and 11). These platforms ranged in length from 17.5 to
64 feet. The length in 16 of these averaged about 25 feet, and in
11 about 40 feet. In 25 maraes the width of the platform was four
to six feet, in two, three feet, and in three, eight to nine feet.

Emory also counted 25 terraces that were rectangular, unpaved,
low, and usually narrow. Two were just west of the summit of
Annexation Hill and a group of eight was south of Flagpole Hill.
Another group of six terraces was found between Flagpole and Summit
Hills, and seven were found just northwest of the crest of Summit
Hill where a double terrace also was discovered.

Despite a considerable search for areas where people might have
lived, only eight hollows were found in the bluffs of Necker that gave
evidence of occupancy. Emory reported that all eight grottos could
probably shelter no more than 24 persons. Only Bowl Cave, where
most artifacts were found, had evidently been continuously occupied.

immature HKed-footed Boobies and Great Frigatebird

roosting on uprights of marae on Annexation Hill. Blue-faced Booby
in mid-ground and large nestling Great Frigatebirds in background.

Photograph by Derral Herbst, 28 August 1968.

Figure 10 Adult and

Immature Red-footed Boobies roosting on uprights of

Figure: 11
Nestling Great Frigatebirds and immature

marae on Annexation Hill.
Red-footed Booby in foreground,

WS)

There human leg bones were found on the floor. Emory speculates
that the bones may have been taken to the cave for use in making
fish hooks, a common practice among Hawaiians.

Emory gives detailed descriptions of all images available to
him and stated that all were "male human figures carved in the con-
ventionalized form." The images, carved from vesicular basalt, varied
in color from light to dark gray, and ranged in height from eight to
18 inches. They weighed from four to 25 pounds, "inclining equally
to extremes."

After making a detailed comparison of the archaeology of Necker
and nearby Nihoa, and of their cultures in comparison with other
Pacific cultures, Emory concluded that

this review of the cultural affinities of Nihoa
and Necker, reveals the Necker culture as one
which had been introduced to the Hawaiian Islands
from southeast Polynesia, probably from Tahiti,
and which, on the islands excepting Necker had
become for the most part displaced, and for the
rest, modified by the historic Hawaiian culture.

He ended by saying "it seems reasonable to adopt the view that the
Necker culture is a pure sample of the culture prevailing in Hawaii
before the thirteenth century...."

About 15 years after the publication of Emory's paper, charcoal
and wood found in 1923 in Bowl Cave were carbon dated by Libby (1954:
742). The age of the charcoal was placed at 166 to 200 years and
that of the wood at 0 to 250 years. Although the material was pos-
sibly contaminated, the results suggest a fairly recent occupation
of Necker, but clearly do not rule out the possibility that the
structures themselves were erected six or seven centuries ago.
Certainly the charcoal and wood could have been brought to the island
at a later date by fishermen or wandering natives.

European Discovery and Early Visits

All authors agree that the first European to discover Necker
was the French navigator, Jean Frangois de Galaup, Comte de la Perouse.
On 4 November 1786, en route to Macao, la Pérouse's frigate, the
BOUSSOLE, and its companion the ASTROLABE sighted a small rocky island
(Buck, 1953: 54). The vessels stood offshore all night. The following
morning La Pérouse (1799) made the following notations:

This very small island is little more than a
rock of about five hundred toises [= 1,000
yards] in length, and sixty [= 120 yards] in
elevation at the most.

16

It does not exhibit a single tree, but there

is a great deal of grass near the summit. The
naked rock is covered with the dung of birds,

and its white appearance affords contrast to
various red spots, upon which the grass has

not sprung up. I approached within the distance
of a third of a league. The banks were perpen-
dicular, like a wall, and the sea broke so
violently against them, that it was impossible

to land. As we sailed almost entirely around it,
the plan of this island, as well as the different
views, are perfectly accurate. Its latitude and
longitude, as determined by Mr. Dagelet, are
23934" north, and 166952" west of Paris.2 I
called it Isle Necker [in honor of Monsieur
Jacques Necker, French Minister of Finance

under Louis XVI].

The exact date of the first historic landing on Necker is not
precisely known but apparently occurred at the beginning of the 19th
century. Bryan (1938: 22) reported that John Turnbull, who visited
Hawaii December 1802 to January 1803 in the British ship MARGARET,
learned that two Hawaiians had landed on Necker sometime previously
and had noted a "range of stones, placed with some regularity in the
manner of a wall, and about three feet high."

Necker was sighted at least four times in the mid-1800's but
no landings were made. The PORPOISE and the OREGON, part of the
U.S. Exploring Expedition, passed offshore on 1 December 1841.
Charles Wilkes (1845: 389-390), commander of the expedition, re-
marked that "birds, especially the white tern, had been seen in
numbers prior to [sighting the island]..."" The vessels, three miles
off the reef surrounding Necker, made soundings but did not attempt
to land because of a "furious surf beating on all sides of the
island."" Necker's position was calculated as 164937'00" W., 23°44" N.

Necker was again observed from offshore on 24 April 1857 by
Captain John Paty of the schooner MANUOKAWAI. Paty (1857: 40)
described Necker as "a precipitous rock, 300 feet high, 1 mile long
and half a mile broad, with small patches of grass on its surface."

Two years later, on the morning of 1 January 1859, Lt. John M.
Brooke (ms.) viewed the island from the U.S. schooner FENIMORE
COOPER. “Eater that year, on 29-April, ;Captain N.C. "Brooks of; the
GAMBIA sighted Necker and took its position. Of the island itself
Brooks (1860: 499) remarked only that, "it is rocky, and about 1-1/2
toe 2 matesmilong. 24

amie Greenwich longitude would be 164° 32'.

107/

In 1886 the fishing schooner GENERAL SIEGEL fished off Necker
(Farrell Pel928) 253s" ihis trip as notable in, that this was the
first time seals were recorded for the island.

On 28 May 1891 the island was viewed from the KAALOKATI by
Henry Palmer and George C. Munro, members of the Rothschild Expedi-
tion which was collecting bird specimens in the northwestern Hawaiian
Islands. Although the party wished to land, it was prevented from
doing so by heavy seas (Rothschild, 1893-1900: viii).

Hawaiian Annexation

The historical details of this period were carefully researched
and analyzed by Pauline N. King (ms.). Our account follows hers
closely, and we are deeply indebted to her. Her thesis should be
consulted for further details and for citations of unpublished
correspondence and official communications.

In the 1890's Necker became an island of international interest
since it was not clear which if any nation held title to it. Great
Britain was interested in laying claims to islands which would serve
as mid-ocean stations for a cable connecting Canada and the Austral-
Asian colonies. She already claimed Fanning in the Line Islands
and she possessed many islands between the Lines and Australia which
would serve adequately as stations. However, she needed an inter-
mediate station between Canada and Fanning and the only possibility
lay within the Hawaiian group.

In the fall of 1893 two Canadians, Mackenzie Bowell and Sanford
Fleming, began inquiring into the feasibility of a cable station in
the main Hawaiian Islands, and into the physical characteristics
and international status of Necker. They concluded that the main
Hawaiians would be unsuitable, partly because they felt that national
pride demanded national ownership of an island station, and partly
because they felt uncertain of the political future of the Hawaiian
government. Fleming strongly recommended that Necker Island be
secured as a British possession.

Some months of political maneuvering followed. During this
period, British officialdom attempted to discover whether the
Hawaiian government did, in fact, claim Necker, and, if not, whether
the United States might object to Britain making such a claim. The
United States government remained neutral and noncommittal on the
matter, only indicating that it considered trans—Pacific telegraphic
communication to be of benefit to the United States as well as to
Great Britain and her colonies. The Hawaiian government was initially
open to the idea but wanted to know more details about the British
plan. Hawaii wished to do nothing detrimental to the interests of
the United States.

On 24 May 1894 the presence in Honolulu of the CHAMPION, a
British warship, and the visit of inquiry to the survey office of

18

the Hawaiian government by the acting British Vice-Consul, led to
the rumor that the British were en route to claim Necker Island.

As a result the Executive Council of the Hawaiian government decided
that Captain James A. King, Minister of the Interior, should travel
to Necker immediately and claim it for Hawaii. On 25 May King's
ship, the chartered interisland steamer IWALANI, departed from
Honolulu at 1710. The CHAMPION departed 50 minutes later. Many
observers interpreted this as a race to claim Necker Island, a race
that the CHAMPION would probably win since it was the faster ship.
Subsequent events indicated, however, that the CHAMPION was on no
such mission.

On 27 May the IWALANI reached Necker, and King, accompanied by
members of the crew, went ashore to claim the island.

A quotation from the log of Captain Freeman (Emory, 1928: 55)
describes the landing:

At 11:00 AM arrived at the island and dropped
anchor in 18 fathoms of watér. We lowered a
boat and proceeded to land at once with His
Excellency, Capt. J.A. King, Capt. Freeman,
C.B. Norton and nine sailors, leaving the
vessel in charge of the second officer. After
considerable difficulty the party was safely
landed. A hard climb up a rugged cliff 260
feet high, was successfully accomplished, when
His Excellency Capt. King hoisted the Hawaiian
flag, read the Proclamation and took possession
of the island in the name of the Hawaiian
Government....after a stay of about four hours on
the island, we left at 5:30 PM for home, steer-
ing E. by S., arriving at Honolulu on Tuesday
evening [May 29].

While on the island, the party erected a flagpole and attached
to it, within a copper tube, a copy of the paper claiming annexation.
Tucker (ms.) reports the contents of this paper:

I, James A. King, Minister of the Interior

of the Provisional Government of the Hawaiian
Islands, in pursuance of a commission granted
to me by His Excellency, Sanford B. Dole,
President of the provisional government of
the Hawaiian Islands, do hereby in the name
of said government take possession of this
island, known as Necker Island as a part of
the Hawaiian Territory; the same being within
the Hawaiian archipelago’ in. Lat... 23935'18"
North and Long. 164°39'00" west, and having
been claimed by the Hawaiian territory since

19

the year 1845 when an expedition? under Capt.
W.M. Paty was sent to survey said island.

Done at Necker Island the 27th day of May in
the year of our Lord 1894.

Signed: J.A. King, Minister of the Interior
Wm. Freeman, Master of Hawaiian Str. Iwalani
Jas. Gregory, Chief Officer "Iwalani"
Albert Tullott, 2nd A y

Late 19th Century Visits

On 24 September 1894, the HBMS CHAMPION did pay a visit to
Necker and landed a survey party. At least six men went ashore, but
Captain Rooke was apparently not among them. Although the Hawaiian
government had asked that no images or artifacts be disturbed, four
were collected.

The British had not yet abandoned their intention of using
Necker Island for a cable station. The British government sent a
mission to Honolulu to engage in negotiations with the Hawaiian
government but they did not achieve their aims. They failed in part,
perhaps, because of anti-British sentiment in the Hawaiian Islands,
and in part because of the unwillingness of the Hawaiian government
to jeopardize possible ties with the United States.

Necker was again visited 12 July 1895 by Captain J.A. King (ms.)
and a survey party on the ship LEHUA. While mapping the island, they

landed and found that the flagpole...placed in
position in May, 1894, had blown down. We
replaced the staff in position. We found the
Copper Cylinder which had contained the
Proclamation open and the document on a rock
under the staff. The Proclamation had been
opened and a written memorandum in pencil was
on the back of it, signed by Officers of the
H.B.M.S. Champion. One name I was able to
decifer as that of Lieut. Nugent, the others

I could not make out. The memorandum stated
as near as I can remember as follows: 'We,

the undersigned officers of the H.B.M.S.
Champion, on 24th of September 1894, surveyed
Necker Island shoal; running S.E. 35 miles,
N.E. 15 miles, and ten miles in all directions.
We found no less than 15 fathoms of water one
mile distance from shore.'

ue have been unable to discover any other reference to this 1845
voyage.

—

20

We returned the Proclamation to the Cylinder and
made it fast to the base of the staff.

Tucker (ms.) in 1917 also saw this Proclamation and reported
the text as "this island known as Necker Island was surveyed by
the undermentioned officers of Her Hawaiian Majesty's ship Champion
Sept. 1894 and also the bank extending 35 miles SE, 15 miles NE,
? miles in the other direction."' The "Hawaiian Majesty's" part of
this quotation is undoubtedly incorrect but the absence of a date
for September 1894 throws some doubt on exactly when the island was
visited. Tucker also deciphered a number of other names indicating
that Lieut. Rowland Nugent, Lieut. Frederick, A.H. Walker, Richard
Markham and three others including two midshipmen had visited the
island.

George N. Wilcox evidently visited the island on at least two
occasions at about this period (Emory, 1928: 48), but we have no
information concerning his visits.

1900-1929

In 1902 Necker was visited by the U.S. Fish Commission Steamer
ALBATROSS which was engaged in deep-sea investigations around the
Hawaiian Islands. The ship arrived offshore on the afternoon of
30 May and anchored off the west end of the island. The following
day the four naturalists aboard (Charles H. Gilbert, Walter K. Fisher,
John 0. Snyder, and Charles C. Nutting) landed and spent a few hours
on the island (Thomas, ms.). Fisher later reported that 1/7 species
of birds were seen and described one of them (Blue-gray Noddy) as a
new species (Fisher, 1903a, b).

In response to considerable agitation over Japanese feather
gathering in the northwestern Hawaiian Islands in the early 1900's,
Necker was included in the Hawaiian Island Reservation. This
reservation, established 3 February 1909 by President Theodore
Roosevelt's Executive Order No. 1019, included all northwestern
Hawaiian Islands but Midway. They were placed under the jurisdiction
of the Department of Agriculture and set aside as a preserve for the
native: birds.

From 1909 to 1916 the U.S. Coast Guard Cutter THETIS voyaged
frequently along the northwestern Hawaiian Chain. The visits had
several objectives: apprehension of Japanese bird poachers who had
been destroying birds on various islands; inspection of the islands
to determine whether further poaching had occurred; transportation
of various scientific parties who wished to study the fauna of the
islands; and transportation of mail and supplies to Midway Atoll.
Parties attempted to land on Necker on several occasions.

As the THETIS passed Necker on the morning of 13 January 1910,
W. Jacobs, the vessel's commander, noted an absence of bird and
human life on the island which was "covered with a growth of grass

PAL

on the central ridge.'' No landing was made because of dangerous
surf (Jacobs, ms.). The THETIS visited Necker again on 22 May,
25 August, and 5 September but no evidence of molestation of the
birdlife was seen.

In 1912, while transporting a U.S. Biological Survey party to
Laysan, the THETIS passed offshore on 18 December. Heavy swells
prevented landing. On 9 March 1913, during the return trip of the
survey party, two of its members, George Willett and Alfred M. Bailey,
attempted to land in a ship's boat but were again prevented from
doing so by high surf. Willett, undaunted, swam ashore. Despite
the inconvenience of being naked, Willett spent about two hours on
the island and obtained a few notes on the kinds of birds present and
their nesting status (Bailey, 1956: 32; Willett, ms.).

On 8 September 1914 Carl Elschner and others went ashore from
the THETIS. Elschner (1915) made notes on minerals and geology and
named a number of the prominent features of the island (see Fig. 2).
Apparently these names were never in common usage and all were re-

placed by names subsequently applied to them by the Tanager Expedition.

In November 1914 W.A. Bryan (ms.), a noted ornithologist and
sometime Honolulu politician, made application to H.W. Henshaw of
the Biological Survey for the lease of Necker Island. His reasons
for so doing were that "the longing to be ‘monarch of all I survey'
has always been with me and a lease to Necker seems to be about as
near as I am liable to get to realization of the longing." Despite
Bryan's longing the lease was never granted.

Bryan was not the only one to attempt to lease Necker. The
Territory of Hawaii leased Necker to A.C. Lovekin on 2 June 1907
(Frear, ms.)).

The THETIS again passed Necker on 19 March 1915 but once more
heavy surf made a landing impossible.

In early 1916 the THETIS paid two visits to Necker and on both
visits men landed on the island. On the first visit, late in the
afternoon of 27 January, three officers landed by swimming to a shelf
of rock in the East Cove (now known as Shark Bay). During their brief
visit they explored only the westernmost peak of the island (Annexa-
tion Peak). An easier landing was effected on 11 February when slight
swells allowed the survey party to step from the dinghy to a rock
shelf edging Shark Bay. During the three hour visit most of the
eastern portion of the island was explored. Lt. W.H. Munter, who
visited the island on both occasions, later made a detailed report of
this visit and included a considerable amount of information on the
birdlife (Munter, ms.).

Necker was visited twice in 1917. H.L. Tucker and others landed
from the power sampan NAKAIWA on 25 October, and Tucker and seven
other members of the crew of the J.A. CUMMINS stopped there on

22

5 November (Tucker, ms.). They rediscovered the Annexation Documents
and noted the few birds present. Tucker and Eben P. Low took a num-
ber of photographs of the birds.

The following fall, on 3 September 1918, the USS HERMES steamed
around the island during an inspection trip of the northwestern
Hawaiian Islands. Ina report of this trip Diggs (ms.) briefly
noted the presence of six species of birds. On 6 October 1919,
Wilder, Warden of the Hawaiian Islands Bird Reservation, visited
Necker in the lighthouse tender KUKUI (Emory, 1928: 59).

The visits made to Necker by the Tanager Expedition in June 1923
and in July 1924 resulted in a greater accumulation of knowledge
about the island than did any previous or subsequent visit. Plans
for this expedition, which visited all the northwestern Hawaiian
Islands, Wake Island, and Johnston Atoll, were formulated in 1922
in conferences between the U.S. Navy National Research Council, the
U.S. Biological Survey, and the Bernice P. Bishop Museum. The Navy
provided the ship (the minesweeper TANAGER), a director of naval work
(Commander S.W. King), and agreed to chart hydrographic data; the
Biological Survey provided an ornithologist (Alexander Wetmore) to
lead the party, a rabbit-killing specialist (E.C. Reno), and a movie
camera. Other personnel were supplied by the Bishop Museum (Gregory,
1924: 19-22).

We are not certain of the exact itineraries of every individual
but believe the following account, derived primarily from an exami-
nation of Wetmore's (ms.) field notes, is accurate.

In all, five voyages were made which included three visits to
Necker (two on trip C and one on trip E). On trip C, while part of
the field party remained on Nihoa, others including Anderson,
Atkinson, Judd, Palmer, Caum, and Cartright, proceeded to Necker
where they set up camp on 12 June. On the 16th Judd, Palmer, Caunm,
and Cartright departed for Nihoa where they replaced Wetmore,
Schlemmer, Grant, Bryan, and Thaanum who sailed to Necker where they
remained from the afternoon of 17 June through the morning of 21 June.
Atkinson departed Necker on the afternoon of 17 June. After a survey
of French Frigate Shoals, part of that field party returned to
Necker for a day's survey on 29 June. Those going ashore included
Wetmore, Judd, Cartright, Anderson, Caum, Palmer, Bryan, and Schlemmer.

The following year, at the request of the National Research
Council, the Navy provided the TANAGER for a resurvey of Necker and
Nihoa. This survey placed most emphasis on archaeological work. The
field party (Appendix Table 1) camped on Necker from the morning of
15 July until the afternoon of 18 July (Gregory, 1925: 19-20).

753}
NEB OIE)

There are few records of landings on Necker during this period.
The island was often sighted in the 1930's and '40's by U.S. naval
vessels engaged in various fleet maneuvers (Amerson, ms.) or by U.S.
Coast Guard vessels cruising up the chain.

On 4 March 1936 a landing was made by A.D. Trempe (ms.), co-
operator for the Biological Survey, and members of the crew of the
Coast Guard vessel RELIANCE, B.L. Bassham commanding. Trempe later
“wrote a brief report on birds seen on other islands visited (Nihoa,
French Frigate Shoals, Laysan) during the cruise but for Necker
reported only that "much the same birds" were seen as were seen on
Nihoa.

The name of the Hawaiian Island Reservation was changed on
25 July 1940 to the Hawaiian Islands National Wildlife Refuge, and
its jurisdiction was transferred to the U.S. Fish and Wildlife
Service in the U.S. Department of the Interior. In December 1951
the Wildlife Service entered into an agreement with the Territory
of Hawaii; one of the provisions of the agreement was that patrol
of the refuge would be by Territorial personnel.

On 20 December 1953 Frank Richardson of the University of
Washington visited Necker briefly from the Coast Guard vessel
BUTTONWOOD. He covered only about half the island but recorded 12
species of birds (Richardson, pers. comm.). Some of his observations
were later incorporated in a paper on the breeding cycles of Hawaiian
seabirds (Richardson, 1957).

Dale W. Rice and Karl W. Kenyon made an aerial survey of the
island on 28 December 1957. Albatross populations were estimated
from counts in low-level photographs (Rice and Kenyon, 1962).

1960-1969

In 1961 Necker was visited 25 to 26 March by the USS DUVAL
COUNTY (LST 758) which was determining exact locations of the north-
western Hawaiian Islands. During this visit the project known as
HIRAN I was plotting first order astronomic strata and azimuth marks
by conventional methods (Roach, ms.).

On 11 June of the following year Necker was visited from the
USS STONE COUNTY by four biologists, two of them from the Hawaii
Division of Fish and Game (Appendix Table 1). During their very
brief visit a few notes were made on vegetation, birds, and the
debris left from the HIRAN I operation (Kramer and Beardsley, ms.).
At that time several military personnel were camping on the island
and were engaged in the HIRAN II project (Marshall, ms.).

Responsibility for patrol and inspection of Necker was assumed
in March 1964 by the U.S. Bureau of Sport Fisheries and Wildlife in

24

the U.S. Fish and Wildlife Service and a refuge manager was assigned
to Hawaii. From then through July 1973 13 landings were made on
Necker (see Table 1). Personnel from the Smithsonian Institution's
POBSP participated in four of the surveys, those made in March and
September 1964, March 1965, and March 1967.

Landings on this island have been exceedingly hazardous and
inclement weather has often prevented landings or, in some instances,
caused precipitate departures. Some idea of the difficulty and
hazards of landing on Necker can be gotten from two recent first-
hand accounts.

Concerning the departure on 22 March 1969, John Sincock noted
that

Viewing a 25 foot surge at the rock shelf [from
which departures are made] and frequent 3-4-foot-
deep waves crashing across [it] I suddenly agreed
wholeheartedly with [the] recent memorandum...
that hazardous duty pay was justified....[Eugene]
Kridler, [David] Olsen and I had our share of being
knocked off our feet, swept along the rock ledge,
and then being swept toward the edge when the sea
dropped 20-25 feet. Kridler's extended hand pre-
vented me from going over once, and I think I
reciprocated 2 or 3 waves later. We...lost track
of how many times we were knocked down, or which
way was up. Olsen jumped to the [rubber] boat in
good shape and was taken to the life boat. Two
knockdowns later the rubber boat returned for Gene
and me...we both had decided a free dive into the
rubber boat was preferable to another sluicing
across the rocks. We waved off the bowline...
because of the hazard of becoming entangled. I
jumped into space like a skydiver and felt like I
had forgotten my parachute, but I didn't catch up
with the boat until it was at the bottom of the
surge. Kridler...dove next, [and landed] spread-
eagle on top of the coast guardsman in the bow....
Only one or two bags of equipment were lost and
wounds were only superficial (BSFW).

On the next visit, 30 May 1969, Olsen described the perils of
landing on Necker. Since the usual landing place (the northwest side
of the low rock shelf connecting the main island and Northwest Cape)
had proven hazardous on previous visits, an alternative landing
place on Shark Bay was investigated and Olsen decided to swim ashore.
His account of the attempt points out another of the hazards around
the island.

I was about 75 ft. from the shoreline when six
sharks came rushing at me from my left front.

25

I turned to face them as they went behind me
--another 10 sharks came in from my right.
The water was simply infested with sharks
swimming round and round me in a...[frenzied]
-.-manner. Some were no more than a foot
from me as they sped by me. At that time I
was about 25 ft. from the ledge and I swam
like I never had before, expecting to be
attacked at any second. Fins [were] all
around me on the surface, and the water was
literally boiling with sharks...Finally, a
wave swept me up on the ledge, but I was
unable to hold on and I was washed into the
surge again--[into a] mass of swirling fins.
I caught the next swell and scampered to
safety. As I stood there I counted 45 sharks
swimming around in the waters at my feet
(BSFW).

The primary purpose of most of these recent surveys has been
inspection and patrol of the island. No survey has been longer than
two days. Nonetheless, a considerable amount of information on the
biota has been obtained and these data are the primary basis for the
accounts of the wildlife that are presented in the following sections
of the report. Itineraries, personnel, and a summary of the results
of the surveys are presented in Appendix Tables 1, 2, and 3.

In February 1967 Necker was designated a natural area within
the refuge system. This designation stipulates that the island's
ecology is to be kept as free as possible from outside influence
and disturbance. The island is dedicated to research. Visiting is
limited solely to scientists on approved research programs and i
entry is only by permit from the Bureau of Sport Fisheries and 1
Wildlife. ;

VEGETATION 1)
by Derral Herbst

Although numerous parties have stopped at Necker, visits were i
brief and few botanical collections were made. La Pérouse (1799), it
who was the first to mention the vegetation of the island, stated
that in 1786 "It [did] not exhibit a single tree, but there [was]
a great deal of grass near the summit." Both Captain John Paty
(1857: 40) and the annexation party who visited Necker in 1857 and
1894 respectively also noted patches of grass (Anon., 1894). Fisher |
(1903a: 777) of the Albatross Expedition gave a slightly more de- i
tailed description of the vegetation: it

The wider shelves of the island are sparsely i
covered with a flesh-stemmed, yellow-flowered i
portulaca (Portulaca lutea), and the summit is
rather plentifully grown over with Chenopodtum

|
231-338 O - 77 - 3

26

sandwutcheum bushes, on which large colonies
of Sula ptseator [= S. sula, Red-footed Booby]
and Fregata aqutla [= F. minor, Great Frigate-
bird] were nesting at the time of our visit.

Elschner (1915: 16) briefly mentions the vegetation as being
"slight...and this...[is found] in higher, more flat parts of the
island while the lower parts of the vertical walls and the shore
rocks are bare." While he alludes to plant collections made--
"my time being limited I was unable to gather many plants on this
island"--the disposition of the specimens is unknown.

The first compenhensive botanical collections were made by the
Tanager Expedition in 1923 and 1924. The resulting publication on
the vascular flora (Christophersen and Caum, 1931) concurs with all
earlier observations in that the vegetation of Necker is described
as sparse and inconspicuous. During the 1923 Tanager survey, C.S.
Judd, a forester, sowed seed of seven species of plants in the saddle
between Flagpole and Summit Hills (Christophersen and Caum, 1931:

7; cf. annotated list below). None of these plants was found growing
there subsequently.

Recent collections and observations were made by D. Herbst (ms.)
(UH), E. Kridler (BSFW), and C.R. Long (POBSP).

In comparing past accounts of the vegetation, it appears that
the composition has remained fairly constant over the years. Probably,
differences can be attributed to the amount of rainfall prior to the
visit. The vegetation cover was sparse on recent visits. Plants
were restricted primarily to the top of the island, with some inter-
mittently distributed on the natural terraces lower on the side.

Vascular plants have been collected on Necker by the following:
J.O. Snyder, May 1902; E.L. Caum, June 1923; E. Christophersen, July
1924; E. Kridler, July 1964; and C.R. Long and J.W. Beardsley,
September 1964. Specimens are deposited in the B.P. Bishop Museum
Herbarium (BPBM), the Herbarium of the University of Hawaii (UH), or
the U.S. National Herbarium of the National Museum of Natural History
(USNM) .

The following list notes all 13 species of vascular plants
collected from, introduced to, or observed on Necker Island. Three
of the five species now growing there, Panicum torridum, Chenopodium
oahuense, and Sesbanta tomentosa, are endemic to Hawaii, while the
other two, Sesuvium portulacastrum and Portulaca lutea, are wide-
spread throughout the Pacific islands. A list of lichens (eight
species) from this island may be found in Magnussen (1942); Tsuda
(1966) lists 17 species of marine algae.

27
Gramineae
Panteum torridum Gaud.

Caum 56 (BPBM), Christophersen 10 (BPBM), Kridler 3 (UH), Long
2445. 2A504 024550 CBE BM):.

As would be expected, the amount and distribution of this annual
grass have varied more than those of the other four phanerogams.
Christophersen and Caum (1931: 7) report that it was fairly common
on the north side of the main island in 1923, but one year later only
two clumps were seen. In June 1962 (Kramer and Beardsley, ms.) small
tufts were found everywhere on the island's crest, while in August
1968 the grass was found in moderate numbers only, and these pri-
marily on Bowl Hill. This grass was also thought to be most abundant
in the vicinity of Bowl Hill in September 1964 and 1966 (BSFW). The
short growing period, the rapid wearing of dead tufts by the seabirds
and the ease by which the wind can disperse the densely vestured
spikelets can explain the varied distribution patterns.

Palmae

[itvistonta australts Martius
Pritchardta pacitftca Wendl.
Prttehardta sp.

Seeds of all three sowed in 1923; not found subsequently.
Casuarinaceae
Casuartna equisettfolia L.

Seed sowed in 1923; not found subsequently.

Chenopodiaceae

Chenopodium oahuense (Meyen) Aellen
C. sandwichewn Moq. f. mtcrosperma Aellen

Snyder (Albatross Expedition, see Fisher 1903a: 807), Caum 58
(BPBM), Christophersen 14 (BPBM), Kridler 2, 5 (UH), Long 2447, 2452,
2454, 2458 (UH).

Since 1923, at least, it has been the most common plant on Necker.
Christophersen and Caum list it as being “abundant on the sloping
sides, but rare on the flat top.'' In August 1968 and on other recent
visits Chenopodium formed an almost pure stand in the saddle between
Flagpole and Summit Hills (Fig. 12); it was abundant on the portion
east of this region (Fig. 13) but somewhat less common on the tops
of the hills than on the sloping sides and the saddle between them.

It occurred in small amounts on the top of Flagpole Hill and was
rare on both Annexation Hill and Northwest Cape.

Saddle between Summit and Flagpole Hills, August 1968.

Figure 12
Photograph by Derral Herbst.

Note dense growth of Chenopodium.

Saddle between Bowl and Summit Hills, August 1968.
Photograph by Derral Herbst o

Figure al
Note Chenopodium and Blue-faced Boobies.

29
Aizoaceae
Sesuvtum portulacastrum L.
Caum 93 (BPBM), Christophersen 12 (BPBM), Long 2456 (UH).

The Tanager Expedition members found this species growing
within reach of the spray on the southern slopes of Annexation Hill.
In September 1966 Sesuvtum was most prevalent along the lower eleva-
tions where it would receive spray from the ocean (BSFW). In 1968
Herbst noticed one or two isolated plants on the northeast side of
Annexation Hill. The rest of the sparse population was limited
primarily to the southern slope near the top of the saddle between
Annexation and Flagpole Hills. A similar pattern of distribution
was noted by Long in September 1964 (POBSP).

Portulacaceae
Portulaca lutea Sol. i

Gilbert (Albatross Expedition, USNM 594972), Caum 59 (BPBM), i
Christophersen 13 (BPBM), Kridler 4 (UH), Long 2457 (UH). |

As in previous reports, Portulaca was common on the flat tops it
and ledges of the cliffs in August 1968. Except for a rare Cheno- |
podium shrub, it was the only plant growing on Northwest Cape--
in cracks and in shallow pockets of soil (Fig. 14). Some plants
on the summit of Flagpole Hill appeared intermediate between
P. lutea and P. oleracea.

Leguminosae |
Haematoxylen campechtanum L.

Seeds sowed in 1923; not found subsequently.
Sesbanta tomentosa H. & A.

Gilbert (USNM 594974), Caum 57 (BPBM), Christophersen 11 (BPBM), I
Kreadler sl (UE)). Longe 24490) 2450 2453. 2460 (UH): |

Christophersen and Caum (1931) observed a few plants along the
top of the main part of the island. Kramer (7” Kramer and Beardsley,
ms.) concurs and adds that it seemed "to be holding its own quite
well" in June 1962. Kridler (BSFW), Long (POBSP) and Herbst (UH)
found this shrub on the tops of all of the hills of the main island
and Kridler found a few plants on Northwest Cape in March 1967.

Sesbanta is evidently more common now than Christophersen and Caum
indicated.

g on the Northwest Cape. Note
tailed Tropicbird in the left background. Photograph by

Derral Herbst, August 1968.

Figure 14 Portulaca lutea Sol growin
nesting Red-

Sil
Euphorbiaceae

Aleurites moluceana L.

Seeds found on the shores of Shark Bay in 1923 (Christophersen
and) Gaun,* 193d": °7)*

Malvaceae
Thespesta populnea (L.) Sd.
Seeds sowed in 1923; not found subsequently.
Solanaceae
Solanum lycoperstecum L.
Seeds sowed in 1923; not found subsequently.

TERRESTRIAL VERTEBRATES

Birds

Twenty-four species of birds have been recorded from Necker it
Island (Table 2). Of these 24, 15 are species of seabirds that
breed at many localities in the central Pacific and two (Christmas
Shearwater, Sooty Storm Petrel) are seabirds that breed commonly at
other localities in the northwestern Hawaiian Islands but which as
yet have not been found breeding on Necker. It seems likely that
the absence of breeding Christmas Shearwaters may be largely due to ii
a lack of suitable nesting habitat but the lack of records for the \
Sooty Storm Petrel may result primarily from the infrequency with
which the island has been visited during the species’ breeding peak
in mid-winter. If the latter species does nest on Necker its
population is probably small.

Of the seven remaining species, five are migrant and possibly
winter resident shorebirds. Three of these five, Golden Plover,
Wandering Tattler and Ruddy Turnstone, probably occur at Necker
every year. The two others, the Sanderling and Bristle-thighed
Curlew, are regular visitors to most of the other northwestern
Hawaiian Islands but probably occur at Necker only sporadically.

The two remaining species are vagrants. One, the Glaucous-
winged Gull, wanders fairly frequently to the main and northwestern
Hawaiian Islands. The other, the Mockingbird, certainly wandered to
Necker from the main Hawaiian Islands where introduced populations
are well established on six islands (Berger, 1972: 215).

In addition to the 24 species known from the island, one other
species, the Bonin Petrel (Pterodroma hypoleuca), was seen flying
near Necker in September 1971. There is no evidence that this species

By.

utilizes the island for roosting or nesting. It seems likely that
these birds belonged to populations breeding on the northwestern
Hawaiian Islands to the west.

Examination of the available data suggests that the breeding
cycles of most of the birds of Necker Island are essentially similar
to those found on the other Northwestern Hawaiian Islands. Two
species, the Laysan and Black-footed Albatross, have clearly defined
winter breeding peaks. The Black Noddy, which has a late winter and
early spring breeding peak on other northwestern Hawaiian Islands,
may have such a peak on Necker but the evidence for this is far from
conclusive.

All other species, with the possible exception of the Sooty
Tern, have breeding peaks that occur either in spring or summer but
several species (e.g., Red-footed Booby, Brown Noddy) have such ex-
tended nesting seasons that some birds may be found nesting in every
month of the year.

One species, the Sooty Tern, apparently nests earlier on Necker
than on most of the other northwestern Hawaiian Islands. It appears
to have a clinal series of breeding peaks along the chain with popu-
lations laying progressively later as one moves to the westward. On
Necker the egg-laying peak apparently usually occurs in the period
from January through early March while on Kure at the westernmost

end of the chain, the laying peak usually occurs in May (Woodward,
HOTZ 2 50) a5

It is clear that much is yet to be learned about the status,
maximum populations and breeding cycles of the birds of Necker
Island. More surveys made during the mid-summer and mid-winter
months would greatly help to clarify these matters.

Table 2. The avifauna of Necker Island
Maximum Estimate
since 1960 and

Taxa Current Status when recorded

Order Procellariiformes
Family Diomedeidae

Diomedea ntgripes Uncommon breeder 350%
Black-footed Albatross Mar. 1969
Diomedea tmmutabtlts Common breeder 1,650-2,000
Laysan Albatross Mar. 1965

Family Procellariidae

Bulwerta bulwertt Common breeder 200+
Bulwer's Petrel Sept. 1966

Table 2. (continued)

Taxa

Pufftnus paetftcus
Wedge-tailed Shearwater

Puffinus nativitatis
Christmas Shearwater

Family Hydrobatidae

Oceanodroma tristramt
Sooty Storm Petrel

Order Pelecaniformes
Family Phaethontidae

Phaethon rubrteauda
Red-tailed Tropicbird

Family Sulidae

Sula dactylatra
Blue-faced Booby

Sula leucogaster
Brown Booby

Sula sula
Red-footed Booby

Family Fregatidae

Fregata minor
Great Frigatebird

Order Charadriiformes
Family Charadriidae

Pluvialts domitntea
Golden Plover

Family Scolopacidae

Numentus tahtttensts
Bristle-thighed Curlew

Heteroscelus tnecanus
Wandering Tattler

Current Status
Common breeder
IRewee walsat(eovre

(two records)

Rare visitor

(one record)

Common breeder

Common breeder

Uncommon breeder

Common breeder

Common breeder

Uncommon migrant

Rare migrant

Uncommon migrant

333)

Maximum Estimate
since 1960 and
when recorded

4,000-5,000
Sept. 1964

I
Malem O67 969

1

Mar. 1965
200

Sept. 1966
500

Mar. 1969
50

Sept. L972
1,400*
Mar. 1969
2,000

SeMieg WOOG5 AY7Al

6
Sept. 1967
1
Sept. 1972
3}
Sept. 1966

SS Ra ae Se a

es

Table 2. (continued)

Taxa Current Status

Arenaria tnterpres
Ruddy Turnstone

Calitdrtis alba
Sander ling

Family Laridae

Larus glaucescens
Glaucous-winged Gull

Sterna lunata
Gray-backed Tern

Sterna fuscata
Sooty Tern

Procelsterna cerulea

Uncommon migrant

Rare migrant
(one record)

Vagrant
(one record)
Abundant breeder

Abundant breeder

Common breeder

Maximum Estimate
Since 1960 and
when recorded

50
Mar. 1965

O*x*

O*x*
7 ,500*
Mar. 1965

50,000
Mar. 1965

2, 000-3 , 000

Blue-gray Noddy Mar. 1967

Anous stolidus Abundant breeder 50,000

Brown Noddy Sept. 1971

Anous tenutrostris Common breeder 1,000

Black Noddy Mar. 1967

Gygts alba Common breeder 600

White Tern Mar. 1967
Order Passeriformes

Family Mimidae
Mimus polygtlottos Vagrant 1
Mockingbird (one record) Sept. 1967

*Estimate is of the breeding population only.

t+This, one of very few numerical estimates available, almost certainly
understates the numbers of birds utilizing the island, as the estimate
was made in a month (September) when the species was nearing the end
of its breeding cycle. The maximum population inhabiting the island
at any one time may well be five to ten times as great.

**A single bird was recorded in June 1923.

5)5)

Species Accounts

Common names of seabirds follow King (1967) in the following
species accounts. Taxonomic order follows that of Peter's (1931,
1934) Checklist of Birds of the World, volumes I and II, with the
exception of Procellariformes which follow Alexander et al. (1965),
the Charadriidae and Scolopacidae which follow Jehl (1968), and the
Sulidae which follow the A.O.U. Checklist (1957). The scientific
names of two shorebirds, the Wandering Tattler and Sanderling, have
been modified following the latest supplement to the checklist
CAZO2U 5) 1973).

The following species accounts are set forth in a standard format
from which deviations are made only when warranted by lack of data.
The section of the species accounts under Status lists maximum recent
populations; delimits periods when birds occur most abundantly on the
island, and, if the bird breeds there, briefly indicates the principal
nesting habitat. It should be noted, except where otherwise indicated,
that the population estimates are of the number of flying birds
present on the island during any one visit; dependent non-flying young
are not included. The section listed under Populations briefly evalu-
ates recent numerical estimates, comparing them when possible with
estimates or statements of abundance made by earlier visitors. In
the Observation Tables a question mark indicates that the presence
of birds was noted but that no estimates were made.

The section headed Annual Cycle uses all available data in an
attempt to determine the regularity (or lack of it) in the breeding
regimes of each species and by interpolation to determine periods
during which birds breed. Data for Necker are less substantial than
for most of the other islands considered in this series of reports
and distinct conclusions sometimes could not be drawn. In the section
headed Breeding Habitat is given a general summary of both previous
and recent observations on nesting habitat. The section headed
Banding lists all known bandings by the BSFW and POBSP. If the
section is omitted, this indicates that no birds are known to have
been banded. Very little banding has been done on Necker as compared
with the other northwestern Hawaiian Islands. Only a total of 328
individuals of ten species was banded here by the POBSP and BSFW during
the study period. Two interisland movements of banded birds involving
Necker are known (see Great Frigatebird and Brown Noddy species
accounts). Finally, the section headed Specimens lists the location
of 91 specimens of 18 species, of which we are aware, that have been
collected on the island. Hopefully this section will be of use to
those interested in conducting taxonomic studies and will point out
where further collecting might be useful.

36

BLACK-FOOTED ALBATROSS Diomedea nigripes
Status

Uncommon breeder (maximum recent breeding population estimate:
350). Known to be present from late December through late June, but
probably present a month earlier and later; absent remainder of year.
Nests on the ground on higher slopes of the island.

Populations

Estimates of breeding Black-footed Albatross are among the best
obtained for any species for three reasons: the population is small
and easily counted, the young are conspicuous, and albatrosses have
been of particular interest to personnel visiting the island.

The estimate of June 1923 and most recent March estimates of
breeding populations are much smaller than the estimate derived by
an aerial survey in December 1957 (Table 3). On northwestern Hawaiian
Islands the number of fledging young may be at times considerably less
than the number of nests with eggs. This fact and variations in the
number of young present on recent March visits make it impossible to
be certain whether the 1957 estimate was erroneous, or whether the
breeding population was actually larger then than in other years. The
size of the albatross nesting population on Midway fluctuates from
year to year; there is no reason to believe that the population on
Necker does not vary also.

It is clear that the 1966-1967 population was either considerably
smaller or considerably less successful than the 1964-1965 or 1968-
1969 populations.

Annual Cycle

Too few data on breeding status are available to establish whether
the breeding cycle on Necker is different from the cycle on other
northwestern Hawaiian Islands. Available observations indicate a span
of breeding from at least late December through June or perhaps early
July. Young have been recorded from 10 March (1967) through 29 June
(1923) but are probably present for at least a month prior to that
period and for at least several weeks thereafter.

Breeding Habitat

In June 1923 Wetmore found young on the higher slopes among the
Laysan Albatross. In March 1965 and 1969 Kridler noted that most nests
were found on the upper slopes of Annexation and Flagpole Hills; no
nests were found on Northwest Cape.

Banding

One young albatross was banded by the BSFW in March 1965.

Bi/
Specimens
We have no record of any specimens of Black-footed Albatross
from Necker Island.
Table 3. Observations of Black-footed Albatross on Necker Island

Population

Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May least Seen sparingly...(Fisher, 1903a: 777,

abundant bird 790).

mgs, 9 Mar. ? Young present (Bailey, 1956: 32). A

few nesting (Willett, ms.).

UMS 27, Jan. only a few* Nesting (Munter, ms.).
11 Feb. several Island estimate, with eggs or very
hundred young chicks (Munter, ms.).
1923 17-20 June 200** No adults but ca. 100 nearly grown
young; a few young had down clinging |
to head and neck but many lacked any 1}
trace of down (Wetmore, ms.). .
29 June 200** Young somewhat more developed (Wetmore,
msec
LO53 2.0 Diee- 200-300*** Nesting (Richardson, pers. comm.). i
1957 28 Dec. 491(736)+ Nesting (Rice and Kenyon, 1962: 377). i
1962 11 June 2 A few young seen (Kramer and Beardsley,
ms.).
1964 8 Mar. 2 Numerous; ca. 130 seen in flight from
offshore (POBSP).
25-26 Sept. 0 (POBSP; BSFW). .
1965 15 Mar. 375-400 94 young counted; 100 estimated present |
(188-200** ) (POBSP; BSFW) .
1966 10-11 Sept. 0 (BSFW) .
1967) 20) Mar. 75 Ca. 25-30 1/4- to 1/2-grown young seen
(50-60** ) (POBSP; BSFW).
L969 822 sMarr 350** 175 young counted. No more than 10 or

15 missed (BSFW) .

38

Table 3. (continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References
LOT dA Sept. 0 (BSFW) .
1973 20 July 0 (BSFW) .

*Only westernmost peak examined.
**kEstimate is of number of breeding birds.
***Only about half of island seen during survey.

tData from aerial observations; figures are count of total birds
and an estimate of breeding birds, based on an allowance for 25
percent unoccupied birds (736 was-rounded to 740 by Rice and
Kenyon, op. ctt.).

LAYSAN ALBATROSS Dtomedea tmmutabilts

Status

Common breeder (maximum recent breeding population estimate:
1,650-2,000). Recorded present and breeding late December to late July,
but probably present from October through early August; not present
during remainder of year. Nests on ground, principally on summits of
hills and along topmost ridge of island.

Populations

Estimates of breeding populations of this species, like estimates
for the preceding species, are relatively accurate and show practically
the same degree of variation from survey to survey (Table 4).

It seems likely that current breeding populations are considerably
smaller than indicated by Rice and Kenyon. However, estimates from May
1902 and June 1923 compare quite favorably with the recent March esti-
mates. Thus we cannot conclude that there has been any real change in
population size during the period for which we have data. It seems
possible that Rice and Kenyon's count was biased upward by an inadvertent
inclusion of Blue-faced or Red-footed Boobies; at present both species
occur among the Laysan Albatross (Fig. 15).

Annual Cycle

Data on breeding status are few and poorly detailed but indicate
a breeding cycle no different from that found on other northwestern
Hawaiian Islands. June 1923, July 1964, and March 1967 and 1969

Figure 15 Young Laysan Albatross among Red-footed Boobies and
Great Frigatebirds nesting in Chenopodium. Northwest Cape in the
background, BSFW photograph by David B. Marshall, 11 June 1962.

40

observations reveal that this species has a later breeding cycle than
has the Black-footed Albatross, an observation often noted in other
areas where both breed.

Breeding has been recorded from late December through late July,
and young have been noted from 10 March (1967) through 26 July (1964).
Laysan Albatross are probably present from about late October through
August as on Laysan.

Breeding Habitat

Observers who recorded nesting areas indicated Laysan Albatross
nest primarily on the hilltops of the ridge running along the island.
Fisher (1903a: 789) also found young "over the shelves on the side of
the north point," which is evidently the area now called Northwest Cape.
In March 1967 Hackman similarly noted a few young on rock shelves on
cliff sides and slopes.

One observer specifically indicated where young were found. In
July 1964 Kridler noted five young on the top of Annexation Hill, 11 on
Flagpole Hill, 35 on Bowl Hill, and ten on the ridge extending east of
Bowl Hill. In March 1969 he noted that the majority of the young was
found on the upper slopes of Summit and Bowl Hills, with a few young
in various locations along the ridges of the main part of the island.
Less than 30 young were seen on the Northwest Cape.

Banding

BSFW personnel banded 100 young in March 1965.

Specimens

We have records of four specimens of Laysan Albatross from
Necker: two males (USNM 300852, 300854) and two females (USNM
300853, 300857) collected by Wetmore 18 June 1923.

Table 4. Observations of Laysan Albatross on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May 1,000-2,000 Young present (Fisher, 1903a: 789).
POTS) a 9s Mar. 800% Large downy young present (Bailey,
1956: 32)... (Willette, ms.)
O16 = 27 Jan; in large Most with eggs, a few with very young
number s** chicks (Munter, ms.).
11 Feb. most numerous Well incubated eggs and very young

bird chicks (Munter, ms.).

41

>

Table 4. (continued)
Population
Date of Survey Estimate Breeding Status, Remarks and References
1923" 17-20, 1,200 Nearly grown young present; many with
29 June much down lost from head and neck and
others with downy head and neck only
(Wetmore, ms.).
i538 420) Dec. 600-800*** Nesting (Richardson, pers. comm.).
1957 28 Dec. 3327) Nesting (Rice and Kenyon, 1962: 377).
(4,990)t
n962> 11 June 2 Abundant young (Kramer and Beardsley,
imiS}G)) ¢
1964 8 Mar. numerous From offshore ca. 800 seen on land and
Bor ey MO\Opaisay we Ihayeinie » (GEORISIE)) 4
26 July 140% Ca. 70 young counted (BSFW).
25-26 Sept. 0 (POBSP; BSFW).
965 e > Mar - 1,650-2, 200 Ca. 550 young present (POBSP; BSFW).
(1, 100%)
1966 10-11 Sept. 0 (BSFW) .
L967) 10) Mar . 600 Ca. 150-200 half-grown young seen (POBSP
(300-400* ) BSFW) .
Soe pit. 0) (BSFW) .
1969 22 Mar. 1,020-1,050* Count of 510 young, 10-15 perhaps over-
looked (BSFW).
1971 14 Sept. 0 (BSFW) .
a7 50) July: 2 Count of 40 fully-feathered young (BSFW).

*Estimate is of the number of breeding birds.

**Only westernmost peak examined.

***Only about half of island surveyed.

*Data from aerial observations; figures are a count of total birds
and an estimate of breeding birds, based on an allowance for 25
percent unoccupied birds (4,990 was rounded to 5,000 by Rice and
Kenyon, op. ctt.).

231-338 O- 77-4

42
BULWER'S PETREL Bulwerta bulwerit
Status

Common breeder (maximum recent estimate: 200). Breeds from at
least late May through mid-September. Nests in holes in the rocks.

Populations

Too little information is available (Table 5) to adequately
determine numbers present. The only numerical estimates probably do
not begin to represent maximum numbers, since estimates were not made
when birds would have been most abundant. We suspect that 1,000 or
more petrels occur on the island but considerably more work is needed
even to approximate numbers.

Annual Cycle

The few available observations on breeding status indicate that
the species breeds annually. We have no evidence that the cycle dif-
fers from that on other northwestern Hawaiian Islands. Eggs are
present from at least late May through late July, and young are
present from about mid-July through September.

Breeding Habitat

Few notes have been made about nesting sites on Necker. Fisher
(1903a: 794) gave the most detailed description: "Here the birds nest
in rather deep, bubble-like holes in the rocks, as far from the light
as possible....The favorite site...is a hole about .2 feet deep, with
a narrow entrance, and wider cavity at the rear....The nest, scarcely
worthy of the name, consists of a few old tern feathers gathered rudely
around the egg, as if merely to hold it in place. Sometimes there is no
trace of a nest, and again I found a few wing bones of a tern, as
though these had been used in place of sticks." More recent observers
found them nesting under stones and crevices on the sides of cliffs.

Banding
Two adults were banded by the BSFW in September 1964.

Specimens

We have records of nine specimens. Five males (USNM 189409,
189411-189413; SUI 18596) and three females (USNM 189407, 189408,
189410) were collected by Snyder, Fisher, and Nutting on 31 May 1902.
One female (USNM 300800) was collected by Wetmore on 18 June 1923.

43

Table 5. Observations of Bulwer's Petrel on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May considerable Eggs and pre-laying birds (Fisher, 1903a:
numbers 777-794).

pols 19 Mar. 0 (Baaleys V956 32).

1923 17-20 June common Nesting (Wetmore, ms.).

29 June abundant (Wetmore, ms.).

Ls 9 20 Dec.. O* None seen during diurnal survey (Rich-
ardson, pers. comm.).

1962 11 June ? (Kramer and Beardsley, ms.).

1964 26 July 2 Eggs and young found (BSFW).

25-26 Sept. 75-100 No eggs or young noted (POBSP; BSFW).

1965 15 Mar. 0 (POBSP;BSFW) .

1966 10-11 Sept. 200 Two downy young found, one of them
almost full grown (BSFW).

£967>" LO) Mar’. 0 (POBSP; BSFW) .

15 Sept. ? One young noted that was nearly fully
feathered but retained a small amount
of down on the head (BSFW).

1969 22 Mar. 0 None seen although a number of holes
were examined (BSFW).

Tote ies Sept. 2 Several young found, all fully feathered
except for large patches of down on the
breast (BSFW).

LOTS 30) Jaly ? Many heard calling deep in burrows

(BSFW).

*Only about half of island seen during survey.

44

WEDGE-TAILED SHEARWATER Pufftnus pactficus
Status

Common breeder (maximum recent estimate: 4,000-5,000). Present
from at least March through October; most breeding occurs from June
through October. Most nest in holes or crevices at ground level, but
a few dig burrows.

Populations

Since there are few numerical estimates (Table 6), and little
banding has been done, it is impossible to state how many birds occur
on the island. At least several thousand are present judging from the
two maximal estimates. It is entirely possible that 10,000 or more
may frequent the island.

Annual Cycle

Data on breeding are scanty but’ seem to conform to the general
pattern found on other northwestern Hawaiian Islands. As on Nihoa,
most birds evidently arrive in March but are not abundant until the
latter half of the month. Birds probably court and dig burrows, where
possible, for about two months before the first eggs are laid. Eggs
are laid in June (1923) and are probably present through July (1964).
The only observations made later in the breeding season were on three
September visits (1964, 1966, 1967) when only downy young, most of
them large, were seen. Most fledging presumably takes place in
October and November.

Breeding Habits

On Necker these shearwaters nest mostly on or near the surface of
the ground. Fisher (1903a: 792) stated that they nested in hollow
cavities in the rocks. Kridler recently found them nesting in holes
and depressions in rock rubble and along the upper cliff sides. He
noted a number nesting under Chenopodium on the surface of the ground
(particularly on the west slope of Flagpole Hill), and occasionally in
burrows beneath the Chenopodtum.

Banding

In March 1964 POBSP personnel banded 19 adult Wedge-tailed
Shearwaters. None was recaptured.

Specimens

We have records of seven specimens of Wedge-tailed Shearwaters
from Necker. Three males (USNM 240995, 250996, 240998) and one female
(USNM 240997), collected by Willett, and one male (UMMZ 120133) col-
lected by Bailey, were taken on 19 March 1913. A male (USNM 300722)
and a female (USNM 300721) were collected by Wetmore on 18 June 1923.

Table 6. Observations of Wedge-tailed Shearwaters on Necker Island
Population
Date of Survey Estimate Breeding Status, Remarks and References
nIO2s) 31 Maiy 2 Nesting but no eggs (Fisher, 1903a: 792).
EO Mar: abundant Mating and nest building but no eggs
found (Bailey, 1956: 32).
TONGISI2 7, Jan’. 2 Some seen flying about at dusk (Munter,
msi
1923 17-20, 29 6,000 Eggs present (Wetmore, ms.).
June
i95se 20) Dec. O* None seen during diurnal survey (Rich-
ardson, pers. comm.).
i962. June 2 Kramer and Beardsley, ms.).
1964 Sm Maitre ? Few seen from offshore (POBSP; BSFW).
26 July very common Many nests contained eggs (BSFW).
25-26' Sept. 4,000- An estimated 1,500 downy young present
5,000 (POBSP; BSFW).
i9Goe 15 Mar’ 2,000 No eggs or young found (POBSP; BSFW).
1966 10-11 Sept. abundant Only large downy young seen (BSFW).
SG yYiey eO) Mar. 10 Estimate represents number of birds
seen, not number of birds present.
No eggs found (POBSP; BSFW).
15 Sept. common Ca. 118 downy young counted, most of
them large; 8 eggs found (BSFW).
E969 e522) Mar. 4,000 Almost all birds observed were in pairs
(BSFW) .
LOE 4a Sept’ very common All chicks seen were large and downy
(BSFW) -
1G72 WS) Cepes 2 (BSFW) .
Uys) SO) Sey ? At least 275 nesting adults present

(BSFW) .

*Only about half of island seen during survey.

46
CHRISTMAS SHEARWATER Puffinus nattvttatis
Status
Rare visitor, two March sight records.
Observations

Richardson (1957: 17) suggested that this shearwater breeds on
Necker, but the near absence of records by several competent observers
at times when breeding should be at its peak indicates that such is
not the case. There are only two records, hitherto unpublished. On
10 March 1967 Kridler saw a single individual near the landing area;
on 22 March 1969 BSFW personnel observed another.

SOOTY STORM PETREL Oceanodroma trtstramt
Status

Rare visitor; one March sight record.
Observations

The only record is of an adult found by Kridler on 15 March 1965;
it was thought to have been dead about a week.

RED-TALLED TROPICBIRD Phaethon rubrtcauda
Status

Common breeder (maximum recent estimate: 200). Present through-
out most of year but probably much less abundant in late fall, winter
and early spring. Most breeding probably occurs between early April
and early October. Nests primarily in crevices and hollows in the rocks.

Populations

The maximum estimate given above is probably a poor indication
of maximal numbers since it was made late in the year when populations
should be smaller than during the probable breeding peak. Wetmore's
estimate of 1,000 birds made in June 1923 probably better represents
maximal numbers, both then and in recent years.

Annual Cycle

Tropicbirds on Necker evidently have an extended nesting season
(Table 7) similar to that on other northwestern Hawaiian Islands. The
presence of near-fledging young (at least two months.old) in late July
1964 indicates that egg laying can occur by mid-April; it is likely
that some birds lay even earlier. Eggs are presumably present from
then at least through September. Young are present at least from late
May through December or January.

47

Various March observations imply that populations and breeding
decrease considerably late in the year and suggest that little breed-
ing occurs until early April.

Breeding Habitat

On Necker these tropicbirds nest principally in rock crevices.
Fisher (1903a: 796) noted that tropicbirds "accomodated [themselves]
to the rocks and [laid eggs] in any rounded cavity' and Wetmore (ms.)
noted that several pairs could be found nesting eight to 20 feet apart
under a single over-hanging ledge. Kridler noted that shallow small
caves on ledges along the upper portions of Bowl, Summit, and Flagpole
Hills were favored as nest sites, as were crevices on the upper western
slopes of Northwest Cape.

Banding

The POBSP banded three adults and 18 young in September 1964;
none was subsequently recaptured.

Specimens

Two females (USNM 300986, 300987) were collected by Wetmore on
17 June 1923, and are now in the National Museum of Natural History. ~ |

Table 7. Observations of Red-tailed Tropicbirds on Necker Island

Population

Date of Survey Estimate Breeding Status, Remarks and References }
1902 31 May rather Nesting, at least some eggs (Fisher, i
common 1903a: 796).
HOS LOwMar\ 1 (@Badiliey, L956) 32).
1923" 17-20 June 1,000 Fresh eggs to recently hatched young
(Wetmore, ms.).
[95se0 2ONDec: 2% Still unfledged young present (Rich-
slravelisvonnis Ik) shy ALS) ee
1962 11 June ? (Kramer and Beardsley, ms.). '
1964 8 Mar. 2 Four seen in flight from offshore

(Waiker, ms.).

26 July 2 Eggs to nearly full-grown young noted in
ten nests found (BSFW).

25-26 Sept. 125 25 nests with near-fledging young
counted and a few eggs seen (POBSP;
BSFW).

4

Table 7. (continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References
1965) /ielbom Man: 0 (POBSP; BSFW).
1966 10-11 Sept. 200 72 nests with young, mostly large, and
1 nest with egg counted (BSFW).
967 iO} Marr 15 Birds in courtship flight; no nests
found (POBSP; BSFW).

15 Sept. 2 23 young from small downy chicks to
near-fledging young seen, 15 adults
seen (BSFW).

1969 22 Mar. 20 No eggs or young found (BSFW).

971 14° Sept. ? All young seen were very large (BSFW).
O72 sal Sepier ? (BSFW).

973s 30) July Q5*x* Young seen (BSFW).

*Only about half of island seen during survey.

**Estimate is of adults and young.

BLUE-FACED BOOBY Sula dactylatra
Status

Common breeder (maximum recent estimate: 500). Present and breed-
ing in all months, but most breeding occurs from February through late
September or early October. Nests on the ground on the higher slopes.

Populations

Recent estimates from March and September surveys consistently
indicate populations of 200 to 250 birds (Table 8). There are no recent
summer estimates for Blue-faced Booby populations on Necker; however,
comparisons of March, summer, and fall estimates on other northwestern
Hawaiian Islands indicate that summer populations are not much larger
than spring or fall populations. Thus we suspect that Necker mid-
summer populations seldom exceed 500 to 600 birds. This figure is
considerably less than that reported for June 1923 by Wetmore and may
indicate that populations were formerly larger.

49

Annual Cycle

Data on the breeding cycle were primarily obtained during March
and September but they afford us some idea of its duration and nature.
Egg laying began as early as late November (1964) or early December
(1913). Data on breeding status from March visits indicate that much
greater numbers of eggs are laid from mid- or late January through
March than in the preceding months.

The smallest young present on all recent September visits were
probably no less than two or three months old. This suggests that
most laying ceases by about mid-June, although eggs are possibly
present through early August. This laying span is similar to that
found on other northwestern Hawaiian Islands such as Nihoa and Laysan
and implies that other features of the breeding season are also similar.
If this is true, then we would expect that the hatching peak occurs
about May and June and that the fledging peak occurs from about late
August to perhaps early October.

The length of the periods during which hatching and fledging
occur is considerably longer. Hatching has occurred from early or
mid-January (1965) to as late as August. Fledging has probably oc-
curred as early as May and certainly continues through October, with
a few chicks possibly fledging in November.

Thus, on Necker, as on other northwestern Hawaiian Islands, the
Blue-faced Booby has nested in all months of the year. It is possible,
however, that breeding may not occur in all months every year and that
there are yearly variations in the breeding cycle which cannot be de-
tected from the available data.

Breeding Habitat

All observers who noted where these birds nested indicated they
nested on the higher slopes of the island. Fisher (1903a: 797) stated
that the Blue-faced Boobies were "nesting among the bushes on the top i
of the island, and also out on the bare rocks. They chose often a
jutting crag, where they could obtain a gyod prospect of the surround- it
ing island and sea.'' On several subsequent visits (June 1923, July i
1964, March 1965, 1967, 1969) other observers reported these birds '
nesting on the higher slopes, the summit of the island, or along the
ridge in more bare areas.

On Necker Blue-faced Boobies usually have well-separated nest
sites.

Band ing

Fifty adults were banded by the BSFW in March 1965.

50

Specimens

We have records of only two specimens from Necker, both of which
are in the USNM. Both are adult females (USNM 300940, 300941) collected
by Wetmore on 18 and 19 June 1923.

Table 8. Observations of Blue-faced Boobies on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May rather The few nests examined had young (Fisher,
abundant 1903a: 797).
LOIS; ; elt. Mary. common Eggs to large young (Bailey, 1956: 32).
TOVG 276 Jan: small No eggs or young (Munter, ms.).
numbers*
11 Feb. large A very few with eggs; two nearly full
numbers** grown young (Munter, ms.).
1923 17-20 June 1,000 Young from half to nearly grown
(Wetmore, ms.).
1953 5,20 Dee. 100-150**%* (Richardson, pers. comm.).
1962 11 June present (Kramer and Beardsley, ms.).
1964 8 Mar. 25 Seen from offshore (POBSP; BSFW).
26 July ? Eggs to flying young (BSFW).
25-26 Sept. 250 Count of 58 adults, 57 immatures and

60 locals (POBSP; BSFW).

1965: 15 Mar. 230 Count of 110 nests; all with eggs
except for one with an egg and recently
hatched young and one with a large
downy young (POBSP; BSFW).

1966 10-11 Sept. 200 Some young still dependent but all of
fledging size (BSFW).

1967 10 Mar. 21:0 All on eggs except for three pairs
with recently hatched young (out of a
count of 103 nests) (POBSP; BSFW).

15 Sept. 240 Count of 204 adults and 32 flying im-
matures; 6 downy chicks counted (BSFW).

Sal

Table 8. (continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References
9G9 -22) Mar: 500 Count of 230 nests. Sample count of
101 nests: 6% without eggs; 63% with
eggs; 16% with an egg and one young,
and 15% with young (BSFW).
197E 14 Sept. 200 Estimate includes 40 young (BSFW).
Poel “Sept. 2 (BSFW).
1973. 30 July 190 Estimate includes young (BSFW).

*Only westernmost peak examined.
**Only the eastern part of the island surveyed.

*k*kOnly about half of island seen during survey.

BROWN BOOBY Sula leucogaster
Status
Uncommon breeder, but poorly known (maximum recent estimate: 50).

Present and probably has bred in all months of the year. Nests on
ground along the higher elevations of the island or on cliff ledges.

Populations

Recent estimates consistently suggest that 50 or fewer Brown
Boobies occur on Necker (Table 9). These estimates are possibly too
low since at least some proportion of the population nests in in-
accessible and seldom-observed areas on the rock cliffs. Wetmore's
estimate (100 birds in June 1923) is considerably larger than recent
estimates and may best represent current populations.

Annual Cycle

We have too few data on breeding to accurately delineate the
major features of the breeding season. Available data suggest that
breeding can occur in all months of the year and that eggs and young
have been present in all months, although not necessarily during any
one year. The data also suggest that most nesting occurs in spring
and summer, as on other northwestern Hawaiian Islands.

Sy

Breeding Habitat

Very little information on nesting habitat has been recorded
here. Fisher (1903a: 798) merely noted that "The two eggs [were]
laid on a level place, where there happens to be a little soil, upon
a shelf of the rock.'' Wetmore and more recent visitors observed
them nesting on rock ledges and along the top of the island. Ob-
servations from two recent surveys (March 1965 and September 1967)
indicate that these boobies nested more commonly on the Northwest Cape
than on the main portion of the island.

Specimens

We have found records of three specimens of Brown Boobies from
Necker: a female (USNM 189414), collected by Fisher on 31 May 1902,
and a male and female (USNM 300858, 300859), collected by Wetmore on
19 and 20 June 1923, respectively.

Table 9. Observations of Brown Boobies on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May not vat “all Fresh eggs to large downy young (Fisher,
abundant 1903a: 798).
POLS) oe Max*: 2 (Willett, ms.).
1923 1/=20_ June 100 One or two half-grown young seen
(Wetmore, ms.).
19537 220) Dec. O* (Richardson, pers. comm.).
1962 11 June present (Kramer and Beardsley, ms.).
1964 8 Mar. ? Ca. 50 seen from offshore (POBSP;
BSFW).
26 July ? A nest with two eggs found (BSFW).
25-26 Sept. 15 Count of eight adults and three flying
immatures. No nests seen (POBSP; BSFW).
L965). 5) Mar.. 25 Ten nests counted, most with eggs. One

nest with a near-fledging chick and
one with a small downy young observed
(POBSP; BSFW).

1966 , 10-11 Sept. 2 No young or eggs found (BSFW).

53
Table 9. (continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References
1967 10 Mar. 10 Not breeding (POBSP; BSFW).

15 Sept. 10 Count of six adult and three flying
immatures. A large downy young and a
nest or nests with eggs seen (BSFW).

LOG A22-Mar. 40*% Count of 20 nests, all containing
eggs (BSFW).

Loy Ee Le Sept. 10 Four adults and two flying immatures
seen (BSFW).

1972 15 Sept. 50 (BSFW) .

*Only about half of island seen during survey.

**Estimate is of the number of breeding birds.

RED-FOOTED BOOBY Sula sula
Status

Common breeder (maximum recent estimate: 1,400). Present and
may breed throughout the year, but most nesting usually occurs from
February or March through September or October. Builds bulky nests
in the tops of Chenopodium bushes.

Populations

The few recent numerical estimates (Table 10) are reasonably
consistent except for the differences in the March estimates for 1965
and 1967. The difference in these estimates is partly a result of the
difference in the breeding cycle in these tro vears (see Annual Cycle |
below).

A single early estimate (2,000 in June 1923) is larger than any
recent estimate (1,400 in March 1969) but is not enough so that we
can conclude that there has been any change in numbers. Since the
largest recent estimate was made relatively early in the breeding
season, when all birds might not have returned to the island, we
suspect that maximal populations at the present time may be as much
as a third or a half again larger (to perhaps as many as 2,000 birds).

54
Annual Cycle

This species has a very extended breeding season on Necker and,
in some years, may nest in all months. Most of the population, however,
nests from March (or perhaps February) through about mid-October.

Laying has begun at least as early as early January (1913) and
has probably occurred in all subsequent months through September.
Since nothing larger than small young was seen on any of four March
visits (1913, 1965, 1967, 1969), it is likely that eggs are usually
not laid from October through December. Observations from those four
March visits also indicate that the timing of the initiation of laying
varies from year to year. Initiation of breeding appears to have been
earlier in 1965 and 1969 than in 1967 (see also Great Frigatebird
account).

In most years most fledging probably occurs from August through
October. If the eggs seen in September 1964 and 1967 were viable and
produced young, however, fledging could have occurred in December and
January.

Breeding Habitat

Various observers agree that most Red-footed Boobies on Necker
nest in the tops of Chenopodium bushes. A few nests have been found
on the stone ruins and in Sesbania. Several observers (Fisher, 1903a:
797; BSFW) indicated that this species nests principally on the higher
slopes of the island. On various March visits and in September 1966
Kridler noted that most nests were found on the north slopes of the
main portion of the island. Red-footed Boobies have not been recorded
nesting on Northwest Cape where no Chenopodium grows.

Banding

A single adult was banded by the BSFW in March 1965.

Specimens

We know of only three specimens (USNM 300906-08); they were
collected 19 June 1923 by Wetmore.

Table 10. Observations of Red-footed Boobies on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May abundant Eggs and young common (Fisher, 1903a:
797).
1913 19 Mar. common New nests to small young (Bailey, 1956:

32).

55)
Table 10. (continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References

1916 11 Feb. quite None found nesting. About 30 immatures
common* seen (Munter, ms.).

1923 17-20 June 2,000 Most nests with 1/3 to 1/2 grown young,
a few with eggs, and some recently
built nests (Wetmore, ms.).

1953-20) Dec. 200-3 00** (Richardson, pers. comm.).

1962 11 June 2 (Kramer and Beardsley, ms.). A photo-
graph by Marshall shows birds on nests
and a near-fledging young.

1964 8 Mar. Few seen from offshore (POBSP).
26 July 2 Eggs to downy young (BSFW).

25-26 Sept. 650 Most nests contained young. 97 nests
were counted but contents are known
for only 72: 27 (38%) contained eggs;
22 (31%) contained small or medium-
sized young, and 23 (32%) contained
large young. 64 immatures capable of
flight were also counted (POBSP; BSFW).

LI65) 15 Mar. 1,000*** Count of 412 nests and an estimated
500 present. All contained eggs
(POBSP; BSFW).

1966 10-11 Sept. a More than 95% of young had fledged.
Some dependent immatures and less than
ten downy young observed (BSFW).

1967 10 Mar. 350 Mostly pre-breeding birds, three nests
of 100 checked had eggs (POBSP; BSFW).

15 Sept. 550 Count of 420 adults and 95 flying im-
matures... 7/6) nests counted: 2 17 "(2/)
with eggs and 59 (782%) with young. An
additional 17 new nests as yet lacking
eggs counted (BSFW).

PIGS 22 Mate. 1,400*** Count of 700 nests with perhaps a few
having been overlooked. Of 118 nests
inspected, 30% were new, but empty, and
70% contained eggs (BSFW).

56

Table 10. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References
NOFA Ea Sept. 425 47 nests contained chicks, most of them
1/2 grown (BSFW).
1972 15- Sept. 700 (BSFW).
LO7S 230 duly 108 Estimate is of adults and dependent

young (BSFW).

*Only the eastern part of the island explored.
**Only about half of island seen during survey.

*kkEstimate is of the number of breeding birds present.

GREAT FRIGATEBIRD Fregata minor
Status

Common breeder (maximum recent estimate: 2,000). Present
throughout the year but less abundant in winter. Breeding birds
present in all months but primary nesting period is from March
through October or November. Builds bulky nests in Chenopodtum
bushes, primarily at higher elevations.

Populations

Recent estimates consistently indicate populations in the low
thousands (Table 11). The single December estimate suggests that
populations may decrease by as much as three-quarters when breeding
is mostly completed. The remarkable similarity in the nest counts
of March 1965 and 1969 (832 vs. 850) suggests a similar correspondence
in the breeding cycles these years. The distinct difference between
the 1965, 1969 March estimates and the 1967 March estimate parallels
the difference in Red-footed Booby estimates for those months and
probably has the same cause, an earlier initiation of breeding in 1965
and 1969 ‘than: ns 1967;

The single early numerical estimate (600 in June 1923) is lower
than most recent estimates, enough so that it seems likely that either
the size of the population has increased or possibly that 1923 was a
particularly unfavorable year for breeding.

S)//

Annual Cycle

Populations breed on an annual basis but exhibit an extended
breeding season in which nesting probably occurs in all months. The
near absence of observations from winter months limits our analysis of
the nesting season but observations from March and September indicate
that only a very small proportion of the nesting population has un-
fledged young in January and February and that no eggs are present
from September through about February.

A few eggs may be laid in late February but most egg laying
occurs in March and April. The presence of eggs in late July (1964)
implies that egg laying continues from March at least through May.
Hatching occurs primarily in May or June but may occur as late as
early August.

The first young probably fledge in early or mid-September but
May remain dependent on the adults for at least several more months.
Most young fledge from about late October through November but
Richardson's observations indicate that some young may not have
fledged by January or even February. The immatures seen by Kridler
in July 1964 were young from the preceding nesting seasons.

Breeding Habitat

On Necker Great Frigatebirds nest in the same areas and habitat
as do Red-footed Boobies. Most build bulky nests about a foot toa
foot and a half above the ground in dense Chenopodium bushes and in
almost every Sesbanta bush found along the ridges and higher slopes.
Fisher (1903a: 799) reported that "A few had nests on the rocks,
generally on jutting crags."’ On recent March visits frigatebirds
were found nesting all over the north slopes of Summit and Flagpole
Hills among the Red-footed Boobies and other nests were found scattered
over the ridges and tops of the other parts of the main island. In
March 1965 Kridler noted that the majority nested on the north slopes
of Summit Hill but that none nested on the Northwest Cape.

Banding and Movements

No frigatebirds have been banded on Necker. However, on 10 and
11 September 1966 Kridler saw an immature bird with an orange streamer
on its left leg. Orange plastic leg streamers were placed on frigate-
birds by the POBSP only on Sand Island, Johnston Atoll, about 560
nautical miles to the south-southwest.

Specimens

We found records of three Great Frigatebird specimens from Necker.
An immature male (USNM 189416) was collected 31 May 1902 by Fisher and
Snyder, and a male and a female (USNM 465205, 300971) were collected
18 June 1923 by Wetmore.

231-338 O- 77-5

58

Table 11. Observations of Great Frigatebirds on Necker Island

Population

Date of Survey Estimate Breeding Status, Remarks and References

1891 28 May 2 Seen offshore (Munro, 1941b: 51).

1902 31 May large Nesting, displaying males seen (Fisher,

numbers 190383 777 50799)!

POS ho eMar. several small Eggs present (Bailey, 1956: 32).

colonies

1916); +27) Jan. 2% No nests found. Only seen roosting
(Munter, ms.).

11 Feb. very None found nesting. Females apparently
numerous** more common than males (Munter, ms.).

1923 17-20 June 600*** Eggs to one-third grown young (Wetmore,
MSc).

1953 °20 Dec. 400-600+ All young flying except for two half-
grown young (Richardson, 1957: 23; and
pers. comm.).

1962 11 June present Nesting (Kramer and Beardsley, 1962).

1964 | 8 Mar. ? Males with inflated throat pouches seen
from offshore. Ca. 2,000 seen on land
and ea. 1,500 in flight (POBSP).

26 July Ca. 500** Eggs to half-grown young. Many imma-
tures seen flying (BSFW).
25-26 Sept. 1,500- Eight small downy nestlings; 85 large
1,800 young and 375 immatures counted. Ob-
servations indicate a ca. 50Z nest
failure (POBSP; BSFW).

1965< 15 Mar? 2,000 832 nests counted and an estimated 850
present (POBSP; BSFW).

1966 10-11 Sept. 2,000 Ca. 200 dependent young still on nests,
a few of which were downy young (BSFW).

1967: .10;Mar. 500 Most birds building or with recently
completed nests. Thirty-seven of 100
active nests with eggs. An estimated

200 nests present (POBSP; BSFW).

59

Table il. (Ccontinued)

Population

Date of Survey Estimate Breeding Status, Remarks and References

1967) 15 Sept. 1,200 Many half- to nearly full-grown young
(BSFW).

ugG9 22 Mar. 1,900- Count of 850 nests. Of 112 nests whose

2,000 contents were checked, 42% were new but
empty; 56% contained one egg and 2%
contained two eggs. One almost full-
grown young with a trace of down on the
head and more than two hundred flying
immatures also seen (BSFW).

Eon 4 Sept: 2,000 A minimum of 478 nests present con-
taining two-thirds to almost full-
grown young (BSFW).

EO72 U5"Sept. 835 Estimate believed to be low. Most
birds seen were adults and flying im-
matures (BSFW).

TofS 30 aul y 374 Partial count of chicks and adults

(BSFW).

*Only westernmost peak examined.
*&*kOnly the eastern part of the island visited.
**k*kEstimate is of the number of breeding birds.

tonly about half of island seen during survey.

GOLDEN PLOVER Pluvtalts domintica
Status

Uncommon migrant (maximum recent estimate: 6). Recorded in
March, June, and September.

Observations

All observations are listed in Table 12.

Specimens

The only specimen (USNM 301057) is a female collected 20 June
1923 by Wetmore.

60

Table 12. Observations of Golden Plovers on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902° 31 May 0 (Fisher, 1903a).
TONS 9 SMart 0 (Bailey, 1956: 32).
1923...1/-20 June 2 One female collected on 20 June (Wetmore,
mMSey)s

1953) 20° Dec. oO* (Richardson, pers. comm.).
1964 26 July ? (BSFW).

25-26 Sept. 3 (POBSP; BSFW).
i965 35 MaG. 0 (POBSP; BSFW).
1966, 10-11 Sept. 4 (BSFW).
1967 10 Mar. 2 (POBSP; BSFW).

15 Sept. 6 (BSFW) .
1969 22 Mar. 0 (BSFW).
1971 14 Sept. 0 (BSFW) .
O72) 15) Sept. 0 (BSFW) .
Los SON sully 0 (BSFW) .

*Only about half of island seen during survey.

BRISTLE-THIGHED CURLEW Numenius tahtttensts
Status

Rare migrant (maximum recent estimate: 1). Recorded in
September.

Observations
A single bird seen near the east end by Kridler on 15 September

1972 constitutes the only record for Necker Island. The species is
of regular occurrence on the other Northwestern Hawaiian Islands.

61

WANDERING TATTLER Heteroscelus tneanus
Status
Uncommon migrant (maximum recent estimate: 3). Recorded in

March, May, June, and September.
Observations
All observations are listed in Table 13.

Specimens

The only specimen (USNM 301028) is a male collected 19 June 1923
by Wetmore.

Table 13. Observations of Wandering Tattlers on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May 2 (Fisher, 1903a: 778).
IL@bs} ALS) WiEnee 0 @aditley, 19562.32))-
1923 17-20 June 1-2 Seen daily; a male collected 19 June
(Wetmore, ms.).

955).,.20: Dec. O* (Richardson, pers. comm.).
1964 26 July 2 (BSFW) .

25-26 Sept. it (POBSP; BSFW).
1965 15 Mar. 0 (POBSP; BSFW).
1966 10-11 Sept. 3 (BSFW).
L967 55 0) Mar. i (POBSP; BSFW).

15 Sept. 0 (BSFW) .
1969 22 Mar. 0 (BSFW).
1971 14 Sept. 0 (BSFW).
M72 LoeSept. 0) (BSFW).
O73 eS On Tully, 0 (BSFW) .

*Only about half of island seen during survey.

62

RUDDY TURNSTONE Arenarta interpres

Status

Uncommon migrant (maximum recent estimate: 50). Recorded in
March, May to July, September, and December.

Observations

Although never occurring in large numbers, turnstones are the most
abundant and most frequently seen shorebird that visits Necker (Table 14).

Specimens

The only specimen (USNM 393503) is a male collected 20 June 1923
by Wetmore.

Table 14. Observations of Ruddy Turnstones on Necker Island

Population

Date of Survey Estimate Observations, Remarks and References

1902 31 May a few (Fisher, 1903a: 803).

1913 19 Mar. 1-2 (Willett, ms.).

1916 11 Feb. fairly Scattered about island (Munter, ms.).

common*

1923 17-20 June 20 A male collected 20 June (Wetmore, ms.).

1953-20 Dec. 20-30** (Richardson, pers. comm.).

1964 26 July several Seen at the top of the easternmost
peak (BSFW).

25-26 Sept. 23% (POBSP; BSFW).

1965 15 Mar. 50 30 seen in one flock (POBSP; BSFW).

1966 10-11 Sept. 11* (BSFW).

1967 10 Mar. 22 One flock of 12 seen flying over Shark
Bay. Solitary birds observed along
the ridge (POBSP; BSFW).

15 Sept. 37%&kK A flock of 30 seen. Another flock of
seven seen on Northwest Cape (BSFW).

1969 22 Mar. 2 (BSFW) .

63

Table 14. (continued)

Population
Date of Survey Estimate Observations, Remarks and References
Loy 14 Sept. <10 (BSFW).
L722) L5i'Sepe. 2 (BSFW).
oy Sie SO} uly. 30 Seen in one flock (BSFW).

*Only eastern part of the island explored.
**Only about half of island seen during survey.

**K*KCount.

SANDERLING Caltdris alba
Status

Rare migrant; one June sight record.
Observations

Wetmore (ms.) noted that one was reported by Grant on 19 June
1923. This record, hitherto unpublished, constitutes the only known
occurrence of the Sanderling on Necker Island.
GLAUCOUS-WINGED GULL Larus glaucescens
Status

Vagrant; a single June specimen record.
Observations and Specimens

A dead immature, found on the beach 18 June 1923, and subsequently
collected by Wetmore (USNM 489329), was reported by Clapp and Woodward
(968s 27)
GRAY-BACKED TERN Sterna lunata

Status

Abundant breeder (maximum recent estimate: 7,500). Present from
at least February through September or October, probably absent or

64

occurring only as a visitor during the rest of the year. Breeds from
at least March (and occasionally February) through September or October.
Nests on the ground, on rock cliffs, or open slopes.

Populations

No recent numerical estimates (Table 15) are available for months
when the population is presumably near its peak (April-June). The
largest recent numerical estimate (March 1965) was very similar to
that obtained in June 1923 suggesting that present populations are as
large if not larger than in 1923. Since few birds were seen in
September (1964, 1966, 1967) and December (1953), it seems likely
that the entire population departs the island after breeding.

Annual Cycle

No observations are available that indicate when birds first
arrive at the island but data from March surveys indicate that egg
laying usually begins in March but may occasionally occur in February
(1916, 1967?). Some egg laying occurs during succeeding months
through about July but the peak laying period occurs from about late
March through mid-May. Hatching young are present from about early
April through about late August and young fledge from about early or
mid-June through late September.

Breeding Habitat

Gray-backed Terns nest primarily on ledges on the cliffs and
upper slopes of the island. Fisher (1903a: 781) found them nesting
"in shallow cavities and hollows of the rock on more exposed portions
of the island, and only very sparingly on the broad shelves with
Sterna fultginosa [=fuscata]." Wetmore (ms.) stated "they were nesting
on ledges of cliffs and open slopes of the island.'' More recently,
observers have found them nesting commonly on broad rock shelves,
particularly on the Shark Bay side of Northwest Cape. Others nested
on the inaccessible cliffs and on the various slopes of the main island
as well as the upper portion of Northwest Cape. On Necker, the Gray-
backed Terns seem to favor more elevated areas for nesting to a
greater degree than does their congener, the Sooty Tern.

Specimens

Wetmore is apparently the only visitor who collected Gray-backed
Terns on Necker Island. On 18 and 20 June 1923 he collected five
specimens, three males (USNM 300630, 300631, 300645) and two females
(USNM 300632, 300646). Two of these (USNM 300645, 300646) were young
birds.

65

Table 15. Observations of Gray-backed Terns on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
WOO 25 3 May, many Nesting (Fisher, 1903a: 781).
observed
Ons) 19 Mar. common Eggs (Bailey, 1956: 32).
UGE Neb. not very Only eggs found (Munter, ms.).
numerous*
923 17-20 June 8,000 From eggs to near fledging young
(Wetmore, ms.).
Loose 20 Dec. 2** Not breeding (Richardson, pers. comm.). j
1962 11 June present (Kramer and Beardsley, ms.).
1964 8 Mar. ? Ca. 200 seen in flight from offshore i
(POBSP; BSFW).
26 July at least sev- Many near-fledging young present

eral thousand (BSFW).

2Z5—26 ‘Sept. 3 Not breeding; 2 immatures still
present (POBSP; BSFW).

LOGS ewelo Mar. 7,500**%* Eggs (POBSP; BSFW).
1966 10-11 Sept. 2 Few adults seen. Over 25 flightless

but nearly full-grown young observed.
Several 2/3-grown young seen (BSFW).

1967 10 Mar. 500 Fresh to lightly incubated eggs (POBSP;
BSFW).
LS) Sq 20 17 adults counted. Three large near-

fledging young seen (BSFW).

G9 22) Mar. 1,300 About 25% on eggs (BSFW). |

LOE Ta Sept. ? No adults seen. A number of large |
young, some fledged, present (BSFW).

LOT Za Sept « 2 (BSFW).

LOWS WSO sawilky 37 0*** 160 chicks of varying sizes seen and

about 25 birds were on eggs. Estimate
probably too low (BSFW).

66

Table 15. (continued)

*Only eastern part of the island explored.
**Only about half of island seen during brief survey.

**k*Estimate is of the breeding population.

SOOTY TERN Sterna fuscata
Status

Abundant breeder (maximum recent estimate: 50,000). Present
from about December or January through about late September; probably
absent during remainder of year. Has bred from early December through
September but most breeding probably occurs from February through
August. Nests on the ground in bare areas on slopes and cliff ledges
or under dense Chenopodtum bushes.

Populations

Recent estimates (Table 16) indicate that the Sooty Tern is the
most abundant breeder on Necker. As with the Gray-backed Tern, the
smaller September and December estimates indicate this species is
absent from the island during the non-breeding season.

Annual Cycle

Sooty Terns breed annually on Necker Island, but their nesting
season is significant in that here it apparently regularly begins
earlier than on almost any of the other northwestern Hawaiian Islands.
The March 1913 observations indicate that laying must have occurred
at least as early as mid-February while the presence of near-fledging
young on two recent March visits (1965, 1967) indicates some laying
began at least by early December (1964) or early January (1965). The
beginning of laying may vary considerably from year to year. The
size of the young reported by Fisher suggests that laying did not
begin until about late April in 1902.

Laying continues from January or February through June (1923)
and in some years (1966) possibly occurs into early July. Hatching
has occurred from about mid-January to early February through about
late July or early August and fledging young have occurred from early
March through about mid-September. The-small numbers of fledged and
near-fledging young seen on September visits, however, suggest that
most fledging was completed at least a month earlier.

67

Breeding Habitat

Sooty Terns have been recorded nesting in many different areas
of the island (Fig. 16). Fisher (1903a: 780) reported that these terns
laid "their eggs on the shelves of rocks where there [were] some soil
and matted succulent portulacas....A few eggs...were laid in cavities
in the face of the rock...many eggs were laid out on the bare rock in
the full glare of the sun...a few perilously near high-water mark, in
fact wet with spray."' On recent visits colonies were found all over
the island. Birds nested on bare rocks on Northwest Cape and the
main part of the island and under Chenopodium from above the cliffs to
the tops of the ridges. In September 1966, when the breeding season
was nearly over, a few young were found on the west slope of Flagpole
and Summit Hills and on the north slope of Annexation Hill.

Specimens

We have found records of six specimens collected on Necker
Island: Fisher collected a juvenile male (USNM 189430) on 31 May
1902 and Wetmore collected three males (USNM 300541, 300545, 300547)
and few females (USNM 300542, 300546) on 18 and 20 June 1923. Two of
these birds (USNM 300546-547) are juveniles.

Table 16. Observations of Sooty Terns on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May most abundant Heavily incubated eggs and recently
bird hatched young, some chicks 7-10 days
old (Fisher, 1903a: 780).
LOTS 9) Mar. thousands Eggs and small young (Bailey, 1956:
32).
1916 11 Feb. very Fresh eggs to near-fledging young
abundant* (Munter, ms.).
1923" 17=20' June 30,000 Fresh eggs to grown young; fledged
young present on 29 June (Wetmore, ms.).
953 20) Dec. 1*% Not breeding (Richardson, pers. comm.).
1962 11 June most abundant (Kramer and Beardsley, ms.).
bird
1964 8 Mar. ts Hundreds seen from offshore (POBSP;

BSFW) .

26 July u Many young chicks seen (BSFW).

Figure 16 Sooty Tern nesting area in the saddle between Flagpole
and Annexation Hills, Looking toward Annexation Hill. Photograph by

Derral Herbst, 28 August 1968.

69
Table 16. (continued)

Population

Date of Survey Estimate Breeding Status, Remarks and References

1964 25-26 Sept. 4 Not breeding. Four birds flew over
island in the evening (POBSP; BSFW).

Soo 5 Mar’ 50,000 80% with eggs, 20% with chicks overall;
about 50% with young on the Northwest
Cape and 15-20% with young elsewhere;
Ca. three near-fledging young seen
(POBSP; BSFW).

1966 10-11 Sept. 300 Present by day, more present at night.
A few unfledged chicks observed (BSFW).

67>" £0) Mar. 15,000 All stages from eggs to fledged young
(POBSP; BSFW).

SE Oepit. 100 Ca. 100 adults flying about with a few

immatures among them (BSFW).

ICIS 22. Mar « 16,600 About 75% on eggs which appeared to
have been newly laid. No young noted
(BSFW).

oe AES ep . ¢100 Less than five adults seen. All
others seen were immatures (BSFW).

LOT 2 tS Sept. 2 (BSFW) .

Se 0) July: several 190 fully feathered young still

thousand present (BSFW).

*Only eastern part of island explored.

**Only about half of island seen during brief survey.

BLUE-GRAY NODDY Procelsterna cerulea
Status

Common breeder (maximum recent estimate: 2,000 to 3.000),.
Present and breeding throughout the year but periods of peak breeding
either non-existent or not yet established. Nests primarily in holes
in cliffs and rocks.

7S
Populations

Estimates (Table 17) are too few and too variable to draw any
valid conclusions about changes in numbers from season to season or
even to be certain about maximal numbers present. All that may be
safely concluded is that the species is a common resident and that
populations fluctuate considerably.

Annual Cycle

Various observations on breeding status are only complete enough
to indicate that Necker Blue-gray Noddies have a very extended breeding
season. Eggs were found during four of the months that the island has
been surveyed (March, May, July, and September) and possibly in a
fifth (December, 1953). Wetmore found only fledged young in June, sug-
gesting that the breeding season had been completed but some recent
September observations (1964, 1967) show that birds may be found
breeding much later in the year.

Breeding Habitat

All observers reported or suggested that these birds nest over
much of the island, and, in particular, on the steeper slopes and
cliffs. Fisher (1903a: 781) reported that "The single egg is laid in
a shallow bowl-like cavity or recess in the rock with no nest, but
occasionally a few stray quills and rubbish scattered about." More
recent observers frequently noted that they nested in holes in the
cliffs; on at least two occasions (September 1966 and 1967) these
noddies were considered more abundant on Northwest Cape than on the
main part of the island.

Banding

Six adults were banded by the BSFW: three adults and a local in
September 1964 and two adults in March 1965.

Specimens

Blue-gray Noddies were originally described from Necker Island
as a new species (Procelsterna saxatilis) by Fisher (1903b). The
species was since reduced to synonymy with P. cerulea but retained
subspecific identity as P. ec. saxatilts. Consequently, since Necker
is the type locality, a large series of specimens has been collected
by many observers.

In all, we have been able to find records of 35 specimens, most
of them (29) now located in the National Museum of Natural History
(USNM). Four specimens, the type, an adult male (USNM 188651), and
three cotypes, an adult female, an immature female, and a juvenile
male (USNM 188652-654), were collected by Fisher on 31 May 1902.

Tak

Four females and two males were subsequently collected by Bailey
and Willett on 19 March 1913. Two of these, a male and a female, are
located in the University of Michigan Museum of Zoology (UMMZ 121850-
851) and the rest are located in the USNM (USNM 239016, 239996, 240015,
240017).

The largest series, consisting of eight males and 14 females,
was collected by Wetmore on 19 June 1923. Twenty-one of these specimens
(USNM 300363-367, 300369-382, 300448, 300449) are now located in the
USNM but one (USNM 300362) was subsequently exchanged with the Paris
Museum of Natural History.

Three additional specimens (BPBM 4853-4855) are located in the

B.P. Bishop Museum, Honolulu, but we do not know when or by whom they
were collected.

Table 17. Observations of Blue-gray Noddies on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May fairly Eggs, all heavily incubated; downy young
common including one recently hatched, and ju-
veniles (Fisher, 1903a: 777, 781-782).
1913 19 Mar. common Only one egg found (Bailey, 1956: 32).
on cliffs
LONG 27, San. a few* (Munter, ms.).
11 Feb. quite Only eggs found (Munter, ms.).
common**
1923 17-20 June 800 Apparently all young fledged (Wetmore,
ms.).
£953)> 30) Dec. 4-8*x* Apparently beginning to lay (Richard-
son, 1957: 25; and pers. com.).
1962 11 June ? (Kramer and Beardsley, ms.).
1964 8 Mar. 2? Hundreds seen from offshore (POBSP;
BSFW).
26 July 200 Some nests with eggs found (BSFW).
25-26 Sept. 450-500 From eggs to immatures present (POBSP;
BSFW).
1965 15 Mar. 100+ One nest with a fresh egg found (POBSP;
BSFW).

shinee I ec

TZ

Table 217 ., }-(continued)

Population

Date of Survey Estimate Breeding Status, Remarks and References

1966 10-11 Sept. 2 No eggs or young found (BSFW).

1967 10 Mar,. 2, 000- Mostly eggs and small downy chicks;

3,000 one chick about a week from fledging
seen (POBSP; BSFW).
5 Sept. >254 254 counted. Population stated to
be probably larger than this figure.
One bird found incubating an egg
(BSFW).

HOG. 220 Mare 750 375 counted. Estimated based on other
available but inaccessible nesting
habitat. In most nests birds were in-
cubating eggs (BSFW).

197 24 Sept: <50 No nests found (BSFW).

1973:-°30° July 40 (BSFW).

*Only westernmost peak of island examined.
**kOnly eastern part of island explored.
**kkOnly about half of island seen during brief survey.
+Kridler considers that this estimate was low in view of observa-
tions made on subsequent visits.
BROWN NODDY Anous stolidus
Status
Abundant breeder (maximum recent estimate: 50,000). Probably
present throughout the year but evidently much less common from about
late December through at least late March. Breeding may occur in all

months but in most years it apparently occurs principally between
April or May and October or November. Nests on ground on open slopes.

Populations

Recent numerical estimates (Table 18) are usually somewhat larger
than the estimate made by Wetmore in 1923, but not enough so that we

73

conclude that there has been any change in the size of the population.
The very low numbers seen in December 1953 and on recent March visits
indicate great variation in the numbers of birds present at different
times of year.

Annual Cycle

Observations from three recent March visits (1965, 1967, 1969)
indicate that few birds had returned to the island or begun to breed
by early and mid-March, but observations from December 1953 and January
and February 1916 indicate that breeding began earlier than March. Ob-
servations from July 1964 show that laying began by at least mid-May
and suggest an egg peak in late June. The September 1964 observations Hi |
and those from September 1966 indicate laying occurred into August or
September and suggest a fledging peak about late September or early
October. On the other hand, observations from September 1967 indicate
that a larger proportion of the breeding population laid eggs in
August and September that year than in 1964 and 1966. If, as the data
suggest, there was a late August-early September laying peak in 1967,
a fledging peak probably occurred in October and November.

The data available indicate that breeding can occur in ald months
but also suggest that the number breeding in some periods (December
through March) is considerably reduced. On Necker this species
evidently nests later in the year in larger numbers than on islands
toward the northwestern end of the Hawaiian Chain (é.g., Laysan and
Lisianski Islands).

Breeding Habitat

Brown Noddies were found nesting in most areas on Necker but
seemed to prefer open slopes. On three occasions (July 1964, Septem- i
ber 1964 and 1966) Kridler noted that Brown Noddies nested in greatest
abundance on Northwest Cape. Conversely, in September 1967 more than
twice as many nests were seen on the main part of the island as on
Northwest Cape.

No observer reported this species nesting in bushes and only
Fisher (1903a: 783) described the nest. He reported that "the nest
was smaller than on Laysan, the material being restricted from
necessity to the fleshy stems of Portulaca lutea, which grows abund-
antly on the shelves of the rocks."

Banding

The POBSP banded 97 Brown Noddy chicks in September 1964. One
of these (USFW band number 793-29149) banded 25 September 1964, not
24 September 1965 as reported by Amerson (1971: 381), was recaptured :
as an adult at Trig Island, French Frigate Shoals, on 8 June 1967 by
POBSP personnel.

231-338 O- 77-6

74

Specimens

We know of two specimens from Necker, both adult females, col-
lected by Wetmore on 18 and 19 June 1923 (USNM 300520, 300501).

Table 18. Observations of Brown Noddies on Necker Island
Population

Date of Survey Estimate Breeding Status, Remarks and References

1902 31 May fairly Nests and eggs (Fisher, 1903a: 783).

common

1913 19 Mar. fairly Nesting (Bailey, 1956: 32; Willett,

common ms.).

1916 27 Jan. very Only eggs present (Munter, ms.).

numerous*
11 Feb. quite Only eggs found (Munter, ms.).
common**

1923 17-20 June 6,000 Eggs found (Wetmore, ms.).

1953 20 Dec. 100-150*** A definite breeding season beginning
(Richardson, 1957: 26; and pers. comm.).

1962 11 June present (Kramer and Beardsley, ms.).

1964 8 Mar. Q Ca. 10-25 seen from offshore (POBSP;
BSFW).

26 July 920,000 At least 10,000 nests on the Northwest
Cape. Most contained eggs but chicks
in all stages of growth seen (BSFW).

25-26 Sept. 8 ,000- Eggs to flying young. An estimated

10,000 3,000 chicks present (POBSP; BSFW).

1965 15 Mar. CQ.+25 Only three birds actually seen. No
evidence of breeding noted (POBSP;
BSFW).

1966 10-11 Sept. 10,000 From eggs to fledged chicks. Only
about one percent of breeding birds
with eggs. An estimated 4,000+ young
present. Chicks in all stages of
growth seen but most 1/3 to 1/2
grown. Very few had fledged (BSFW).

1967 10 Mar. 0 None seen (POBSP; BSFW).

79

Table 18. (continued)

Population

Date of Survey Estimate Breeding Status, Remarks and References

1967 15 Sept. 9,100 Ca. 2,200 nests with eggs; 950 with
downy chicks counted and/or estimated.
Notes state that these figures are less
than actual totals present (BSFW).

1969 22 Mar. 25 No nests found (BSFW).

1971 14 Sept. 50,000 Most abundant bird on island. Nests
contained eggs to full grown young
(BSFW) .

nS7Se 30) July 11,000 Eggs and an estimated 800 young present

Estimate possibly low (BSFW).

*Only westernmost peak examined.
**Only eastern part of the island explored.

*kkOnly about half of island seen during survey.

BLACK NODDY Anous tenutrostris
Status

Common breeder (maximum recent estimate: 1,000). Present and
has bred in all months, but probably does not breed in all months every
year.

Populations

Recent estimates (Table 19) consistently indicate that only small
numbers of Black Noddies occur on Necker. Maximal populations are
evidently on the order of 1,000 birds but considerably fewer birds
were recorded on visits when no active nests were found. The largest
recent estimates are about three to five times as large as the single
early estimate (June 1923) but we cannot be certain whether there has
been any real increase in numbers.

Annual Cycle

Available data on breeding status indicate that breeding has
occurred in all months but not necessarily in all months every year.
No well-defined pattern of annual breeding can be deduced from the data.

76

Eggs have been laid in November or December (1953), January (1967),
February (1965, probably 1967), June or July (1964), and August or
September (1964). However, data from two of three September surveys
suggest that breeding was largely completed by then. Clearly, more
detailed data on breeding status are needed from many months to
determine breeding peaks and the breeding schedule.

Breeding Habitat

Wetmore (ms.) noted that Black Noddies nested in clefts in the
rocks in June 1923. Recent observers have found these birds nesting
most commonly in small holes in the cliffs on Northwest Cape with
smaller numbers nesting on steep, inaccessible slopes of the main part
of the island. Others probably nest in the steep cliffs of the north
side of the island.

Specimens
Apparently only three specimens have been collected. Wetmore

collected two adult males (USNM 300446, 300459) on 19 June 1923 and
an adult female (USNM 300460) on 20 June 1923.

Table 19. Observations of Black Noddies on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1902 31 May ? No nests found (Fisher, 1903: 784).
TOUS ar kOe Manes fairly (Bailey, 1956: 32).
common
1923 17-20 June 200 Nesting (Wetmore, ms.).
1953 ZO eDec. 400-500* Eggs and young present (Richardson,
1957: 26; and pers. comm.).
1964 8 Mar. ? Ca. 25-50 seen from offshore (POBSP;
BSFW).
26 July 600** Ca. 300 nests found; all contained
eggs (BSFW).
25-26 Sept. 300-400 Eggs to flying young in seven nests
found on the Northwest Cape (POBSP;
BSFW).
1965 15. Mar. 250-300 Six of ten nests checked on Northwest

Cape contained eggs, one a small downy
chick (POBSP; BSFW).

Ha

Table 19. (continued)

Population
Date of Survey Estimate Breeding Status, Remarks and References
L9G6- 10-11 Sept. 20 No eggs or young noted (BSFW).
HOGiey LO Mar. 1,000 Some nests with eggs and at least one
half-grown nestling seen on Northwest
Cape (POBSP; BSFW).
1S Sept. 150 Seen in one flock at the end of the
Northwest Cape. No active nests found
(BSFW).
IGS 22 Mar. 500 About 80% of birds present on eggs 1
(BSFW) . i
1971 14 Sept. not too No nesting noted (BSFW).
common

*Only about half of island seen during brief survey.

**kEstimate is only for the number of breeding birds present.

WHITE TERN Gygts alba

Status |
Common breeder (maximum recent estimate: 600). Present and

breeding throughout the year. Lays single egg in holes in rocks or

on rock ledges, particularly on the steeper cliffs.

Populations

Numerical estimates (Table 20) indicate neither seasonal change in
numbers nor that numbers have changed since 1923. This apparent lack )
of seasonal variation is quite different from that observed on other
northwestern Hawaiian Islands such as Laysan and Lisianski. However, |
estimates may be less accurate than those obtained on Laysan and ;
Lisianski, because Necker has much nesting habitat that is inaccessible
and which is observed with difficulty, if at all. Few numerical esti-
mates have been made in mid-summer or winter; it is possible that
there are seasonal differences in size of populations which are not
revealed by the available data.

Annual Cycle

Our data indicate that breeding may occur throughout the year
and that eggs and young may be found in all months. Since proportions

| dj

78

of nests containing eggs and various sizes of young were almost never
recorded, we cannot determine whether there is a regular, seasonal
breeding peak. Data from the only survey where such information was.
recorded (September 1966) suggest a late summer breeding peak.

Breeding Habitat

All observers who mentioned nest sites agree that White Terns
nested principally on the steep rocky cliffs of the island. Some
have also been found nesting along the higher ridges (March 1967).
On at least two surveys (March 1965, September 1966), this species
was thought to be more abundant on the Northwest Cape than on the main
part of the island.

Banding

In September 1964 the BSFW banded 31 White Terns (nine adults, two
immatures, and 20 locals).

Specimens
Three specimens, all collected by Wetmore, are in the USNM. One

(USNM 300390) is a juvenile female; the other two (USNM 300408, 300419)
are adult males.

Table 20. Observations of White Terns on Necker Island

Population
Date of Survey Estimate Breeding Status, Remarks and References
1891 28 May ? Seen offshore (Munro, 194la: 2).
1902 31 May one of the Eggs, a hatching young, and many small
commonest terns nestlings (Fisher, 1903a: 785).

1913) 219) Mar. abundant Nesting (Bailey, 1956: 32).
LOG e27yi dan. a few* (Munter, ms.).

11 Feb. a few** None found nesting (Munter, ms.).
1923 17-20 June 800 Eggs and young present (Wetmore, ms.).
1953 220 Dec. 300-400*** Eggs and few newly fledged young seen

(Richardson, 1957: 27; and pers. comm.).

1962. 11 June present (Kramer and Beardsley, ms.).
1964 8 Mar. u Hundreds seen from offshore (POBSP;

BSFW).

ie)

Table 20. (continued)
Population
Date of Survey Estimate Breeding Status, Remarks and References
1964 25 July 500-600 Eggs to nearly fledged young. An
estimate of at least 50 nests in the
vicinity of the top of Bow Hill (BSFW).

25-26 Sept. 400-500 Eggs to immatures (POBSP; BSFW).

1965 15 Mar. 200 Nests with eggs found (POBSP; BSFW).

1966 10-11 Sept. 500-600 Eggs to fledged young seen. 75-80% of
nests contained young (BSFW).

1967 10 Mar. 600 Eggs and small chicks present (POBSP;
BSFW) .

15 Sept. ? 363 Count of 363 stated to be a minimum
figure. A few seen on eggs. Both
small downy young and large chicks
also seen (BSFW).

1969 22 Mar. 500 150 birds counted. Estimate based on
additional nesting habitat which
could not be censused. Eggs to nearly
fledged young (BSFW).

1971 14 Sept. 400-500 (BSFW).

LOn2 15 Sepe. ? (BSFW) .

1WS)7/So= SiO) eae Ay 200 An estimated 50 young present. Esti-

mate almost certainly too low (BSFW).

*Only westernmost peak of island examined.

**kOnly eastern part of island explored.

**k*kOnly about half of island seen during survey.

MOCKINGBIRD Mimus polyglottos
Status

Vagrant; one September sight record.

80

Observations

Kridler and others saw a Mockingbird fly over the lower southeast
slopes of Flagpole Hill on 15 September 1967. Although seen but briefly,
and not observed subsequently that day, all relevant field characters
were clearly seen.

This species has not been recorded previously from Necker but has
been recorded at least three times previously in the northwestern
Hawaiian Islands, all at French Frigate Shoals (Clapp and Woodward,
1968: 33; Amerson, 1971: 302). All these records were presumably of
birds that wandered from resident populations in the main Hawaiian
Islands.

Mammals
The only mammal ever recorded from Necker Island is the Hawaiian

Monk Seal which occurs and breeds on most of the northwestern Hawaiian
Islands.

HAWAIIAN MONK SEAL Monachus schautnslandt
Status

Regularly present in small numbers. Maximum count: 20.
Observations

Observations of seals at Necker (Table 21) suggest that the
species has occurred there regularly for almost a century. While no
pups have yet been recorded, the consistency with which animals have
been seen there in recent years, as well as the observation of four

pregnant females, suggest the possibility that the seal may breed there.

Table 21. Observations of Hawaiian Monk Seals at Necker Island

Number

Date of Survey Seen Remarks and References
1886 late summer a few One or more killed for shark-bait

or early fall CFaxrel il. 1928 2253) 4
1894 27-29 May is A few seals seen by annexation party

(Enory,. 1928: 56):

1964 26 July 6 Swimming in Shark Bay (BSFW).

25-26 Sept. 12 Adults hauled out on rocky shelf 25

September: six females, sex of rest
not determined (BSFW; POBSP).

81

Table 21. (continued)

Number

Date of Survey Seen Remarks and References

1965) 15. Mar. 7 Basking on Shark Bay side of shelf con-
necting the two parts of the island:
six adults (two sexed were male and fe-
male), one subadult female (BSFW, POBSP).

1966 10-11 Sept. 6 In same locality as previous observation:
four adults (two females, one male), two
subadults; one female molting (BSFW).

L967. 10 Mar. 12 Six adults (at least one male; two
females), six yearlings (at least one
male; one female) (BSFW).

> epi. 15 Basking in same locality as above.
Thirteen adults (two males, five fe-
males), one yearling male, one yearling
female (BSFW).

1968 28-29 Aug. ? Several seen (BSFW).

969m 22>Mar. 20 On Shark Bay side of junction between
Northwest Cape and mainland: seven
adult males, four pregnant females,
three subadults (one male, two fe- 1
males), six not aged or sexed (BSFW).

1971 18 Aug. 10 Counted in Shark Bay (BSFW).

Sept. 16 All adults. Twelve were on the flat
shelf on the Shark Bay side of the cut
separating Northwest Cape from the main
island; the other four were on a ledge
on Shark Bay below Summit Hill (BSFW).

IQS) SHO Analy, 18 All adults (BSFW).

Reptiles

Only one species of reptile, a sea turtle, is known to occur at
Necker. The inhospitable habitat and infrequency of landings on
Necker presumably account for the absence of lizards, which are found
on a number of the other northwestern Hawaiian Islands.

82

GREEN TURTLE Chelonta mydas [=agasst2t]
Status

Regular visitor in small numbers. Maximum count: 6.
Observations

Recent observations of turtles at Necker (Table 22) suggest that
the dearth of early records is probably more the result of lack of
interest in recording their presence than the result of a lack of tur-
tles. These turtles do not breed at Necker but the remarkable frequency
with which they have been seen on recent visits suggests that the area
offshore is a much used feeding area.

Taggin

BSFW personnel tagged ten turtles at Necker (Table 22). The tag
return obtained by the BSFW also suggests that some turtles may remain
for awhile in the vicinity before continuing their pelagic wandering or
returning to their natal island, which, for most Necker turtles, is
probably French Frigate Shoals.

Table 22. Observations of Green Turtles at Necker Island

Number
Date of Survey Seen Remarks and References
1923. 12-21 June a few Found hauled out on rock shelves
(Wetmore in Mellen, 1925: 161).

1964 26 July il Seen near the island in Shark Bay (BSFW).
25-26 Sept. 2 One cq. 35" and one eq. 18" (BSFW;
POBSP).

1965° “15° Mar’ 5 Ca. 2.5-3' long; resting on low ledge
on Shark Bay side of island (BSFW:
POBSP).

1966 10-11 Sept. 6 Five noted, ca. 2.5-3' turtles seen
(and tagged) on same ledge as above.
Another ca. 3' seen subsequently in west
cove (BSFW).

1967 10 Mar. 3 Ca. 3'3 two tagged; in same location as
above (BSFW; POBSP).

15 Sept. 3 Same size and locality as above; two

tagged, one a recovery of a turtle
tagged the previous visit (BSFW).

83

Table 22. (continued)

Number

Date of Survey Seen Remarks and References

1969 22 Mar. 4 In same area as above; one tagged
(BSFW).

ry 21 Aug. 1 Seen swimming offshore (BSFW).

14 Sept. 3 Two large females and one smaller
turtle were hauled up on the ledge on
the Shark Bay side of the cut separat-
ing Northwest Cape from the main
Island (BSFW).

POTS 0) July 3 A dead turtle found on shelf adjacent
to Shark Bay. Three other turtles
observed at a depth of about 45 feet
(BSFW).

ACKNOWLEDGMENTS

Field work on Necker Island was made possible by a co-operative
agreement between the Department of the Interior, Bureau of Sport
Fisheries and Wildlife, and the Smithsonian Institution. We are in-
debted to the U.S. Coast Guard whose logistic support made possible the
visits to Necker.

Mr. Edwin H. Bryan, Jr., B.P. Bishop Museum, Honolulu, Hawaii,
generously allowed us access to his reference files which were of
particular value in our preparation of the history section of this
report. We are also particularly indebted to Dr. Alexander Wetmore,
Smithsonian Institution, Washington, D.C., who allowed us full use of
his previously unpublished observations made during the Tanager expedi-
tion in 1923. We also thank Dr. Frank Richardson, University of
Washington, Seattle, Washington, who communicated to us avian popula-
tion estimates that he made during his visit to Necker in December 1953.

Dr. Philip S. Humphrey, University of Kansas, Lawrence, Kansas,
principal investigator of the POBSP, offered constant encouragement
which led to the completion of this manuscript. We also thank the
POBSP and BSFW personnel, listed in Appendix Table 1, who did the field
work on which the bulk of this report is based. Derral Herbst kindly
wrote the vegetation section. A. Binion Amerson, Jr., and Phillip C.
Shelton made many helpful comments during various stages of manuscript
preparation and Dr. F. Raymond Fosberg was kind enough to read and
criticize the vegetation section. We are also grateful to Tina C.
Clapp who aided in the preparation of several of the figures. A. Binion
Amerson, Jr., edited the final manuscript.

EES ee

84

The camera copy was typed by Barbara B. Anderson with funding
through a contract with the Bureau of Sport Fisheries and Wildlife,
Department of the Interior (contract number 14-16-008-596, February
35 E97 AS) i.

85
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92

Appendix Table 1. Scientific visits to Necker Isiand,;, 1891-1973

Date Personnel Vessel
1891 28 May Rothschild Expedition* KAALOKAT

Henry C. Palmer
George C. Munro

1894 27 May Capt. James A. King IWALANI
Capt. William K. Freeman
lst Off. James Gregory
Eng. Benjamin H. Norton
Eight Hawaiian sailors

24 Sept. Members of the crew CHAMP LON

G9 5a. de2euilsy: James A. King LEHUA
William D. Alexander
William T. Brigham (BPBM)
Frank Dodge

1902” 31 May, Albatross Expedition ALBATROSS
Charles H. Gilbert (SU)
Walter K. Fisher (SU)
Charles C. Nutting =(SUL)
John O. Snyder (SU)

POs Osa: A.M. Bailey (BBS)* THERES
George Willett (BBS)

1914 Sm Sepit. Carl Elschner THETIS
Members of the crew

1916 -27 Jan , William H. Munter’ (CG) THELIS
Members of the crew

11 Feb. William H. Munter (CG)
Members of the crew

1918 3 Sept. Members of the crew* HERMES
1919 OF OCcE. Gerrit P.. Wilder KUKUL
[92733 ABV Tanager Expedition TANAGER
21,29 June Alexander Wetmore (BBS)
(ornithologist)
William G. Anderson
(collector)

A.l.C; Atkinson: (HBAF)

Edwin H.! Bryan, *J7c.. (BPBM)
(entomologist)

Bruce Cartwright (BPBM)
(assistant in hydrographic work)

Appendix Table l.

Date

1924 15-18 July

O55 20) Dec.

i957. 928) Dec.

1962 11 June**
(0630-0900)

1964 8 Mar.

(continued)

Personnel

Tanager Expedition (cont.)
Chapman Grant (BBS)

(naturalist)

Charles S. Judd
(forester)

Edward L. Caum (BPBM)
(botanist)

Harold S. Palmer (BPBM)
(geologist)

Eric L. Schlemmer
(assistant to Wetmore)

David L. Thaanum (BPBM)
(conchologist)

Tanager Expedition
Harold S. Palmer (BPBM)

(geologist)

William G. Anderson
(collector)

William Bush
(collector)

Erling Christophersen (BPBI)
(botanist)

Theodore T. Dranga (BPBM)
(conchologist)

Kenneth P. Emory (BPBM)
(ethnologist)

Kenneth I. Hobson
(collector)

A. Landgraf
(topologist)

Frank Richardson (UW)

Karl W. Kenyon (BSFW)
Dale W. Rice (BSFW)

Raymond J. Kramer (HDFG)
David H. Woodside (HDFG)
David B. Marshall (BSFW)
John W. Beardsley (HSPA)

Eugene Kridler (BSFW)

A. Binion Amerson, Jr. (POBSP)
Loren Kroenke (UH)

Edward O'Neill (BSFW)

Ronald L. Walker (HDFG)

George S. Wislocki (POBSP)

93

Vessel

TANAGER

BUTTONWOOD

Aerial Survey

STONE COUNTY

PLANETREE

94

Appendix Table 1.

Date

1964

1965

1966

1967

1968

1969

26 July
(0900-1845)

25-26 Sept.

(0830-1300)

15 Mar.
(0900-1800)

10-11 Sept.

(0700-1520)

10), Maa.
(1040-1715)

{5 Sept.
(0910-1840)

28-29 Aug.
(0830-1845)

22 Mar.
(0900-1740)

30 May
(0900-1200)

(continued)
Personnel

Eugene Kridler (BSFW)

Eugene Kridler (BSFW)
John W. Beardsley (UH)
Robert R. Fleet (POBSP)
Charles R. Long (POBSP)
Ronald L. Walker (HDFG)

Eugene Kridler (BSFW)
Winston Banko (POBSP)
Chandler S. Robbins (BSFW)
Ronald L. Walker (HDFG)

Eugene Kridler (BSFW)
Sherwin Carlquist (CC)
Karl W. Kenyon (BSFW)
Warren Roll (HSB)
Ronald L. Walker (HDFG)

Eugene Kridler (BSFW)

C. Douglas Hackman (POBSP)
Ernest Kosaka (HDFG)

John Maciolek (BSFW)
Richard Wass (UH)

Eugene Kridler (BSFW)
Robert Ballou (BSFW)
John L. Sincock (BSFW)
Ronald L. Walker (HDFG)

Eugene Kridler (BSFW)

G. Brent Dalyrymple (CGS)
Richard R. Doell (CGS)
Robert Eddinger (UH)
Derral, Herbst (UH)

John L. Sincock (BSFW)

Eugene Kridler (BSFW)
Karl W. Kenyon (BSFW)
George Laycock (NAS)

David L. Olsen (BSFW)
John L. Sincock (BSFW)

David L. Olsen (BSFW)

Vessel

CHARLES H. GILBERT

BASSWOOD

BLACKHAW

IRONWOOD

BUTTONWOOD

BUTTONWOOD

BUTTONWOOD

BUTTONWOOD

MAHI

Appendix Table 1. (continued)
Date Personnel

2971 21. Aug. David L. Olsen (BSFW)
(1230-1500) David Childs (SI)
Richard Grigg (HIMB)
Robert J. Shallenberger (01)
James Vansant (UH)
William Worcester (UH)

14 Sept. Eugene Kridler (BSFW)
(0900-1500) Erwin A. Bauer

Kenneth S. Norris (OI)

John L. Sincock (BSFW)

1972 15 Sept. Eugene Kridler (BSFW)
(1000-1500) Bruce Benson (HA)
Ernest Kosaka .(HDFG)
David L. Olsen (BSFW)
John L. Sincock (BSFW)

1973) 30) July David L. Olsen (BSFW)
(1100-1830) John L. Sincock (BSFW)
Leighton Taylor (BSFW)
Thomas Telfer (HDFG)

*No landing made on island.

**Times of arrival and departure, where known, are listed under the

Vessel

TERR) os

BUTTONWOOD

BUTT ONWOOD* -

BUTTONWOOD

dates of visit for surveys made during the 1960's and 1970's.

Glossary of Abbreviations:

BBS Bureau of Biological Survey
BPBM Bernice P. Bishop Museum

BSFW United States Bureau of Sport Fisheries and Wildlife

cc Claremont College, Claremont, California
CG United States Coast Guard

CGS United States Coast and Geodetic Survey
HA Honolulu Adviser

HBAF Hawaiian Board of Agriculture and Forestry
HDFG Hawaii Division of Fish and Game

HIMB Hawaii Institute of Marine Biology

HSB Honolulu Star Bulletin

HSPA Hawaiian Sugar Planters Association

NAS National Audubon Society

OI Oceanic Institute, Waimanalo, Hawaii

POBSP Pacific Ocean Biological Survey Program

95

96

Appendix Table 1. (continued)

SI Smithsonian Institution

SU Stanford University

SUL State University of Lowa
UH University of Hawaii

UW University of Washington

Appendix Table 2. Results of scientific visits to Necker Island, 1891-

1973
Date Results
1891 28 May Bird observations made from offshore (Munro, 1941la,b).
1894 27 May Hawaiian domain proclaimed and documents left on
island. Seven stone images and a stone bowl col-
lected.
24 Sept. Four images collected.
18953 12 duly Topographic survey made by Frank Dodge, and a number
of specimens of birds and their eggs collected by
W.T. Brigham.*
1902 30 May Observations of birds (Fisher, 1903a), collection of
plants. Blue-gray Noddy described as new species
(Fisher, 1903b). Extensive offshore marine survey.
Twenty bird specimens of five species collected.**
1913) 189 (Mar. Observations on birds and their breeding status
(Bailey, 1956). Eleven specimens of two species
collected .**
1914 8 Sept. Observations on geology and guano; map drawn.
196: 272) Jans Observations of birdlife.
Ji Feb. Observations of birdlife.
IS His: 3 Sept. Scanty observations of birds.
EAI) 60cE. Image leg and a reshaped image collected.
1923 12-045. Ornithological observations (Wetmore, 1925) ;***
29 June archaeological investigations; observations and col-
1924 .15=18 July lections of: vascular plants, crustacea, echinoderms,

annelids, foraminifera, fish, chilopods, insects,
molluscs, lichens and sponges. Geological and
petrological observations and a topographic map made.

Appendix Table 2.

Date
1923
1924 (cont. )
O>3 20) Dee.
LOS Zoe DEC.

1962 11 June

1964 8 Mar.

26 July

25-26 Sept.

LIGSmel oa Mar .

1966 10-11 Sept.

1967 10 Mar.

LSSept

1968 28-29 Aug.

6922 Mar.

97

(continued)
Results

Ca. 58 bird specimens of 17 species collected;
seven species of plants sowed by Judd.

Observations on birds and their breeding status
(Richardson, 1957).

Aerial photographs taken to determine size of
albatross populations (Rice and Kenyon, 1962).

Brief notes on vegetation, birdlife, effects of
military occupation of island; insects and
arachnids collected.

Observations of birds made from offshore. Patrol
of refuge.

Observations of birds, turtles, seals; erection
of refuge sign. Patrol of refuge.

Observations of birds, census of turtles and
seals. Collected: plants, crustacea (isopods),
arachnids, and insects. 1/74 birds of six species
banded.*** Patrol of refuge.

Survey of birdlife; censuses of turtles and seals;
153 birds of four species banded. Patrol of refuge.

Observations of birds; censuses of turtles and
seals; five turtles tagged; refuge sign erected.
Patrol of refuge.

Studies and censuses of vertebrates, vegetation
and marine life; patrol of refuge; two turtles
tagged.

Patrol of refuge; observations of birds; two
turtles tagged.

Studies and censuses of vertebrates, vegetation and
marine life. Botanical survey for purpose of study-
ing Portulaca; rock samples collected for a study

of their magnetic properties; refuge sign erected;
patrol of refuge.

Studies and censuses of vertebrates, vegetation and
marine life; cover map made of island vegetation;
current meters established offshore; patrol of
refuge.

98

Appendix Table 2.
Date

30 May

1971 21 Aug.

14 Sept.

1972 15 Sept.

1973. S0n July

(continued)
Results

Marine investigations around island; retrieval of
current meters. No observations made of wildlife
as only one observer landed on the island for a
very brief period.

Coralline algae collected. Seals and turtles cen-
sused. No observations made of birds. Patrol of
refuge.

Studies and censuses of vertebrates, vegetation
and marine life; patrol of refuge.

Cursory observations of birds; seals censused;
patrol of refuge.

Studies and censuses of vertebrates, vegetation
and marine life; patrol of refuge.

*Present location of these specimens unknown.

**kWe cannot be certain if this total is correct since the original
papers reporting the visit did not report all collections.

x*x*kA new bird distributional record from this trip was reported by
Clapp and Woodward (1968).

Appendix Table 3.

Publications on collections and studies (with the
exception of birds) made on Necker Island, 1891-
19 73%

Protozoa

Cushman tm Edmondson Records 27 species of foraminifera

Cradle. ¥ £925

Pilsbrys. 1927

from offshore.
Mollusca

Reports two species of barnacles col-
lected by the Tanager Expedition

Annelida

Treadwell tm Edmondson Reports a polychaete collected by the

Ct. abx 1925

Tanager Expedition

99

Appendix Table 3. (continued)

Arthropoda

Arachnomorpha (Arachnida)

Bryan et al., 1926 Mentions occurrence of spiders and
bird ticks.
Beardsley, 1966 Records four species of Araneida and

one of Acarina from collections made
in June 1962 and September 1964.

Amerson, 1968 Reports the distribution of hosts of
ticks from collections made by the
POBSP.
Crustacea
Rathbun, 1906 Reports two species of brachyuran

crabs collected at Necker and offshore
by the Albatross Expedition.

Edmondson 717” Reports nine species of decapods col-
Edmondson et al., lected by the Tanager Expedition.
925

Bryan et al., 1926 Mentions that isopods were collected.

Beardsley, 1966 Reports one isopod from a collection

made in September 1964.

Labiata
Myriapoda
Bryan et al., 1926 Describes a new species of Chilopoda
from collections made by the Tanager
Expedition.
Hexapoda - Insecta
Kellogg and Paine, Reports one species of mallophaga col-
1910 lected by the Albatross Expedition.
Bryan et al., 1926 Reports ca. 39 species of insects col-

lected by the Tanager Expedition.

Aldniechs i t93i1 Describes a new ephydrid fly from
specimens collected by the Tanager
Expedition.

100

Appendix Table 3. (continued)

Hexapoda - Insecta (cont.)

Lopes, 1938

Usinger, 1942

Zimmerman,** 1948a

Zimmerman, 1948b

Zimmerman, 1958a

Zimmerman, 1958b
Hardy, 1964

Hardwick, 1965

Maa, 1962

Beardsley, 1966

Describes a new species of sarcophagid
fly from material collected by the 1923
Tanager Expedition.

Describes two new species of Nystus
(Hemiptera, Lygaeidae) from specimens
collected by the Tanager Expedition.

Lists six species (one thysanuran, a
cockroach, an embiompteran, an earwig,
and two mallophaga).

Lists two species of Hemiptera
(Lygaeidae).

Lists a noctuid moth and a lycaenid
butterfly.

Lists a pyralid moth.
Lists a dolichopodid fly.

Describes a noctuid moth from collec-
tions made by the Tanager Expedition.

Reports specimens of hippoboscid col-
lected by the Tanager Expedition.

Records 28 new records from collections
made in June 1962 and September 1964,
and summarizes earlier records exclud-
ing Mallophaga.

Echinodermata

Fisher, 1906

Fisher, 1907

Agassiz and Clark,
1907 =19 12

Reports one species of starfish
(Asteroidea) collected by the Alba-
tross Expedition.

Reports four species of sea cucumbers
(Holothuroidea) collected by the Alba-
tross Expedition.

Reports five species of Echinoidea
collected by the Albatross Expedition.

Appendix Table 3. (continued)

Clark tn Edmondson
Bie Obs « MWS

Clark, 1949
Vertebrata
Pisces

Snyder, 1904

Fowler and Ball,
925

Fowler, 1927

Strasberg, 1956

Reptilia

Mellen, 1925

Emory, 1928

Mammalia

Emory, 1928

Farrell, 1928

Tomich, 1969

Laycock, 1970

101

Echinodermata (cont.)

Reports two species of Ophiuroidea,
three species of Echinoidea, and two
species of Holothuroidea collected by
the Tanager Expedition.

Summarizes records of echinoderms.

Chordata

Reports five species of fishes col-
lected by the Albatross Expedition;
most had been carried to the island
by birds.

Reports 17 species collected by the
Tanager Expedition.

Lists two species collected by the
Tanager Expedition and previously
reported in Fowler, 1925.

Revises taxonomy of Hawaiian blen-
nioid fishes, recording one species
from Necker.

Reports that Green Turtles were found
by the Tanager Expedition in 1923.

Erroneously stated that ‘Turtles...
abound."

States that "The Hawaiian seal has
been shot on Necker."

Reports that seals were seen at Necker
in 1886.

Reports occurrence of monk seal in
March 1965.

Mentions occurrence of monk seals in
March 1969.

102

Appendix Table 3. (continued)
Flora

Fisher, 1903a Reports three species of vascular plants
collected by the Albatross Expedition.

Christophersen and Reports five species of vascular plants
Gaum 371931 collected by the Tanager Expedition and
lists seven species planted in 1923
which were not present in 1924.

Magnusson, 1942 Lists eight species of lichens, six of
which are newly described, from collec-
tions by the Tanager Expedition of 1924.

Tsuda, 1966 Reports collection of 17 species of
marine benthic algae collected in July
1924 and September 1964.

Geophysical
M8hle, 1902 Describes three specimens of lava.
Elschner, 1915 Gives comments on the geology and

chemical composition of guano from
observations made in September 1914.

Powers, 1920 Describes a specimen of lava collected
im 194.

Washington and Keyes, Reports results of studies of rocks
1926 collected by the Tanager Expedition.
Palmer, 1927 Gives geological and topographical ob-

servations made by the Tanager Expedi-
tion.
Archaeology
Alexander, 1894 Gives a photograph of idols collected
TTS 94:
Emory, 1928 Reports work done by the Tanager Expe-

dition and summarizes all available
earlier information.

*Authors are in chronological order within taxa.

**Zimmerman and Hardy, in the Insects of Hawaii series, present dis-
tributional records derived primarily from the Tanager collections,
but extensively revise taxonomy, re-identify specimens, and identify
to species hitherto unidentified specimens.

ATOLL RESEARCH BULLETIN
NO. 207

THE NATURAL HISTORY OF NIHOA ISLAND,
NORTHWESTERN HAWAIIAN ISLANDS

by Roger B. Clapp, Eugene Kridler and Robert R. Fleet

Issued by
THE SMITHSONIAN INSTITUTION
with the assistance of
The United States Fish and Wildlife Service
U.S. Department of the Interior
Washington, D. C., U.S.A.

May 1977

TABLE OF CONTENTS

Page
INST. ON VELCURHS « ./cyais she. aes.o.e 43 Steen i ER tad D wprinde ets Be
TALS TMORMUABIGH Siavs-oi sists srt. sine o oe Bessie Sac Ie coe een ieee Dy
LIST OF APPENDIX TABLES........ 550/0 tac C0 OxD CODD sitelellotetetevel olatele ele (alle sieteters +2)
PN TROD UCT UON sicicve.e.s eye. «01,6 aeooasd we ccee siolersilele)ocs] sreilejisleie KonhoAoUnS eisiele)s it
DNS GIRTE ET ELON Gidea GIS GGG oycacuncic wile elie) els) ee eliel nle!e) alee o°6) wielvils se e's olla teveicl ow evelene ele e 3
GE OL OG Yaya reireile liar llaionelssiial/aiisilejele)lalielielsiissieils/ el elleis eHaleliseKclielisheiioliel siiccel ers sieve) eifelrelete relate oftctene 9
ESTES LORY cvs ici wylc)s)eliclietoyel ofc) oie) © s/s) ise) 0. 0 oe) olie) 0s OF 6d CIO CIO CROKE OO OIG CID CRC OOO OICIOICLD. 14
WEG EIDAST TON ciittetette s/t vraiia taro) al elislta}isl eitalletter’oiiel siete! slaxcllels eile] afetieberelleils, oeleveliel’s oh ovejeielledel elie 96
UP EU PEs le Roe eu Ps Role Ey VACA EL Oy aralslfalloliaitetaial olelellellokcNedlsiielioie te) elfs) e/el(siiells)e\lelelle!« (o)lel olielis) e)ie) siie.'e 38
Species Accounts....... o.0.0069 4.0.000 6 OIb.6 FeSO OG Od ado AdaAdoBOOOCONS 38
BASH (cporeie) bevel, si'etete\'e siielte oMfe Tolls; ehabeke cece. Bicileialsiele alsteite ofisia « nese 39

Ditomedea ntgripes....... jlo o. biol VoD EOOe Id.C DO Dom OOK OD Ld

Diomedea immutabilis..... Mie ee Neu so oaddoDOgd 47

BUCA WUNWICIAL Ven ate) clotehe heloke) stele eo iete: ete eerste erettaie ie 50

EU aE ML TAL CL IU CLUCT Mel etatialelala/ atelier el eliodeveker ele 'a/eKore/ ei anelele) «icles 58

PUREMUS HACE LLQCES ami anne ante. Meret Ree, ewe, 57
OCCAMOGROMAMELES TRO dave s/aic) s/e)s) « 01s) « 01 + >\6ks)s FdooDoooCOd 60

Phaethon aethereus MESONAUTA.......evevevee 5.56 Growc.o 6il

PUACEROM, RUDRCAUGA waste voltioisis c.lelsyel> ae ietolencies ere cheilelere sale 62

Si hes IBILY UORER Gh 0.610, Coit CHICO CIO. Oni HOG DODO NI0.0.0.00 65

SUL GMILCULCOGG SILC) Nel alelavos -lanaioye ciocelciaya ie) o atetercter cl otciere tere ero 69

SULA SULA. Jerse sacobonéoodouudne aneNekot stielcltelet ovat letelelee- 72

Fregata minor...... 0 O10 DiCnOnOsOtd Osc. WO OOO 00.00 BODO OO Gn 78

AOC GUC Menetteretenenneketetetsteheieialsieietehele cial oteielsveleieretelst sich siete 81

PLUDEGEES "AOMUNECO st etate et lareretshers WOO caterers. Ati So Ob O.0G 82

NWUMCHIVUS SCONE CECT SUG vile ate cshtey s vivise siete sae aie tetenten sNowars 83

HEE CHOSCCVNG <UNCANUS ath tate! tatotehelctsiete etetatale etl eateratole boon YA

Arenarta tnterpres.......+.- eEnCHALD 0 On Habel toc. OOK 550 1G

LAYUS APGENTATUS.»..saeeee elie) © ie1e © (ein) alle ese cetiehershaletieloclalevette 88

SuCciies Wiehe. Ab66.0 Cb0nb OOD QO ObOO OO SSO OOU OOOO D OGG 88

DUCCENAMUSCACA yc. haste oe eels we OO. G Oe UOGOb Hon OtOOOO0.40 92

PROCOLSLCUIG NCCTULOA Aelaretaher tel ee ote tetatetele) tyrone hclolels o¥eletal 96

AROUGHSGOU LAU Sin. AMM TNH. aie eel. Nelo oierehstehs eVotadol. oxcteholiers 101
AVOUSMECIUGLOS LICL Shin ate lela a leusitele/a)e\(sleis\\e So acdouaadn OO ONT 105

GUGES “AUDAa ee eheis JoouSoccoy. ooueoBoooodo bos SGC ONS 107

Mimus nolyglottos. ieteel ei cteloystciates- favaenevens fexeierelcr oie Soc odoOn 110
ACROCCDONALUS*FOMULTAP US TKENG Era « clele er slciel olele whele orale SS TO

PSU ELAVOSEMANCAMLANIS) WLU LING oleic s we) iele)oiele\ eee) 0 ere (ses \° 113

Memmma is reretece leper otenMera er ater efeiterela, stots any « cheks aici e Moe) Ml sLIDS)

MONACHUS SCRAUTNGLANAT Nols soe eee. oe SQD 0060000 -119

ISN SIOE ISI S) eNey sc rhics ty-0° CW CRORE WOE ERE RE MPS OLE aI LCR ERE ISR 120
Lepidodacty lus (DIG MARER6 Gols 580 stodaeon5 cooO0 ts S0-0.0.ie 120

CRELONGA MYAAS xo aicle es « Jadoudsoceos0UroODuOaC ae eletevenel ges
ACKNOWLEDGMENTS.......ceee000- RAR RR GOR one tes cS Be ARIE 122
LITERATURE sCEDED Rte. sia tis aioe REM EO CeO Oe a eee 124
APPEND PXGMUABIEE St. serainig ates. c esterase rere Be atece: cilaratapa Sac heeeee FEA: IER rok 4

231-338 O- 77 - 8

tt

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ate

ie

13:

14.

15.

16.

Ld

ats

LIST OF FIGURES

Page

The Hawaiian Islands. Facing 1
Map of Nihoa Island. fh
Aerial photograph of Nihoa Island showing prominent features of

the island. 5
Looking at Nihoa Island from the south, December 1912. 6
Viewing Miller Peak from the ridge west of Middle Valley. The

dots in the right foreground are nesting Red-footed Boobies. 6
Viewing Tanager Peak from the top of Middie Valley. Note dense
Chenopodtum on slopes and dry Stda in left foreground. 7

Low point along the north rim between Miller's and Tanager Peaks. 7
Tanager Peak as seen from the air. Note steepness of north cliffs

Miller's Peak as seen from the northeastern rim of the island,
September 1964. 8

The steep west clif of Nihoa Island. 10

Western portion of island showing sandy beach in westernmost cove.
Miller's Peak and Miller's Valley are to the right. In the center
is Pinnacle Peak and West Palm Valley. To the extreme left is

Dog's Head Peak. Just to the west of Dog's Head Peak is the smal110
West Valley.

Best landing area is in the right foreground. Note the extent of
rock terrace along the edge of the island. Camp is on a reasonably

level spot near the base of Miller's Valley. a:
Three of the many dikes cutting the western cliffs. — ial
Looking north down the "Devil's Slide." 14
Native structure on the northeast corner of Miller's Plateau. 15

Remains of ancient house terrace in East Palm Valley, August 1968.
Vegetation is chiefly Chenopodium shrubs with some Sida. 27

Middle Valley, August 1968; vegetative cover--typical of the sides
of the valleys--consists primarily of 2 to 3 foot tall Stda and
Chenopodtum shrubs. 27

Low scrambling Euphorbia shrubs around rock outcroppings along
northern cliffs, August 1968. In the foreground are Chenopodtwn
and Solanum shrubs. 30

9

20.

2,

ZZ),

23}

24.

25h

ZO).

OAS

23).

29%.

30.

31.

S2%

Small grove of Pritchardta remota at base of cliff in upper East
Palm Valley, August 1968.

Grove of Pritchardia (right foreground) in West Palm Valley, 10
June 1962.

Portulaca villosa plant growing from crack in face of stone ledge,
August 1968.

Euphorbta celastrotdes, August 1968. A variety with the same habit
growing in a somewhat similar environment at Kaena Point, Oahu,

was in flower at this time but was leafless.

The Solanum nelsont flowers on Nihoa in August 1968 had a white
corolla and purple anthers; those of Moomi Beach, Molokai, have
light blue petels.

Blue-faced Booby at nest on rocky outcropping.

White-phase plumage morph of the Red-footed Booby, 8 March 1968.
Dark-phase plumage morph of the Red-footed Booby, 8 March 1968.

Nesting Great Frigatebird with downy young in July 1966.

Bristle-thighed Curlew foraging among the rocks in lower Miller
Valley, 8 March 1968.

Sooty Tern incubating egg in lower part of Middle Valley, 9 March
1968.

Blue-gray Noddy chick at nest site in niche in rock wall, 9 March
1968.

Brown Noddy at nest with egg, March 1967.

Nihoa Finch on rocky outcropping, 16 September 1972.

CEL

Page
30

34

34

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3)
66
7h)
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85

85

97

MTL
102

114

Lv

Ta

24.

LIST OF TABLES

Recent surveys of Nihoa Island by the POBSP and BSFW.

The avifauna of

Periods of peak
Islands.

Observations of
Observations of
Observations of
Bulwer's Petrel

Observations of

Nihoa Island.

egg-laying on three northwestern Hawaiian

Black-footed Albatross on Nihoa Island.
Laysan Albatross on Nihoa Island.
Bulwer's Petrels on Nihoa Island.
specimens from Nihoa Island.

Wedge-tailed Shearwaters on Nihoa Island.

Wedge-tailed Shearwater specimens from Nihoa Island.

Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Observations of
Blue-gray Noddy
Observations of

Observations of

Christmas Shearwaters on Nihoa Island.
Red-tailed Tropicbirds on Nihoa Island.
Blue-faced Boobies on Nihoa Island.
Brown Boobies on Nihoa Island.
Red-footed Boobies on Nihoa Island.
Great Frigatebirds on Nihoa Island.
Golden Plovers on Nihoa Island.
Wandering Tattlers on Nihoa Island.
Ruddy Turnstones on Nihoa Island.
Gray-backed Terns on Nihoa Island.
Sooty Terns on Nihoa Island.

Blue-gray Noddies on Nihoa Island.
specimens from Nihoa Island.

Brown Noddies on Nihoa Island.

Black Noddies on Nihoa Island.

725) ¢

26;

Observations of White Terns on Nihoa Island.
Observations of Nihoa Millerbirds on Nihoa Island.
Nihoa Millerbird specimens from Nihoa Island.
Observations of Nihoa Finches on Nihoa Island.

Nihoa Finch specimens from Nihoa Island.
Observations of Hawaiian Monk Seals at Nihoa Island.

Observations of Green Turtles at Nihoa Island.

LIST OF APPENDIX TABLES
Scientific visits to Nihoa Island, 1885-1973.
Results of scientific visits to Nihoa Island, 1885-1973.

Publications on collections and studies (with the exception
of birds) made on Nihoa Island, 1885-1973.

108

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118

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AVMOIW ©

THE NATURAL HISTORY OF NIHOA ISLAND,
NORTHWESTERN HAWAIIAN ISLANDS}!

by Roger B. eres Eugene Reider and Robert R. Pieces

INTRODUCTION

Nihoa Island, sometimes known also as Bird Island, is a
precipitous remnant of a volcanic peak and is the easternmost of a
chain of islands comprising the Hawaiian Islands National Wildlife
Refuge (Figure 1). Approximately 156 acres in extent, it lies at
23°06'N, 161°58'W (Off. of Geogr., 1956: 58), about 250 miles from
Honolulu (Bryan, 1942: 167). Its nearest neighbor in the northwestern
Hawaiian Islands is Necker Island lying about 155 miles to the west-
north-west.

Few reports have been made about the biota of the island. Most of
our knowledge of the bird life stems from a report by Vanderbilt and
de Schauensee (1941) as well as from several papers dealing with the
two endemic forms of passerines found on the island (Bryan, 1916,
1917; Wetmore, 1924, Richardson, 1954). A considerable amount of
information was also obtained by the Tanager Expeditions of 1923
and 1924, but the material dealing with the birds was never published.

Beginning in 1964 the Pacific Ocean Biological Survey Program
(hereafter POBSP) of the Smithsonian Institution and the Bureau of
Sport Fisheries and Wildlife (hereafter BSFW) began making periodic
surveys of Nihoa. From 1964 through 1973, 16 visits were made,
together totaling 30.3 days of observation (Table 1 and Appendix Table
1). In the tables the unpublished survey material obtained by the
BSFW and POBSP can be located through reference to the date of the
survey. Dates are listed in the Literature Cited section under BSFW
and POBSP.

1/

Paper Number 76, Pacific Ocean Biological Survey Program, Smith-
sonian Institution, Washington, D.C.

National Fish and Wildlife Laboratory, U.S. Fish and Wildlife
Service, Department of the Interior, National Museum of Natural
History, Washington, D.C. 20560.

Office of Endangered Species and International Activities, U.S.
Fish and Wildlife Service, Department of the Interior, 1311
Kapiolani Boulevard, Honolulu, Hawaii 96814.

Department of Wildlife Sciences, College of Agriculture, Texas
A & M University, College Station, Texas 77840.

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The primary purpose of this report, one of a series on the
northwestern Hawaiian Islands, is to summarize present knowledge
of the vertebrate fauna and vascular flora of Nihoa Island. A
considerable effort has been made to thoroughly document previous
information and many of these unpublished notes, particularly those
of Dr. Alexander Wetmore, who led the 1923 Tanager Expedition, add
considerably to our knowledge of the island's biota.

Secondarily, this report should serve as a reference to papers
(see Appendix Tables 2 and 3) dealing with other aspects of the
island's biota.

The present report was largely in final draft form in late 1970
and includes only slight emendations and additions after that period.
The tables of observations have been emended to include BSFW informa-
tion available through 1973 but only seldom has this additional
information made changes in the text necessary.

BSFW and POBSP field notes and trip reports concerning Nihoa
are, respectively, stored in the Bureau of Sport Fisheries and
Wildlife files, Kailua, Oahu, Hawaii and the Pacific Ocean Biological
Survey Program files, National Museum of Natural History, Washington,
DG.

DESCRIPTION

Nihoa, remnant of a volcanic cone, is characterized by steep
slopes, rocky outcroppings, well developed valleys, and precipitous
cliffs. Figures 2 and 3 present a vertical overview of this island
which measures about 1,500 yards east to west and from 300 to 1,000
yards wide (Bryan, 1942: 167).

From the south the island present a distinct saddle-shaped
appearance (Fig. 4) with the highest peaks being found on the north-
eastern and northwestern corners of the island. The maximum elevation
of 895 feet is to the northwest at Miller's Peak (Fig. 5). In the
vicinity of Miller's Peak are several acres of reasonably level land
which is much favored by albatrosses for nesting. At the north-
eastern corner of the island is Tanager Peak (Fig. 6), its 852 foot
elevation only slightly lower than that of the peak to the west.
Between the peaks the elevation of the ridge drops to 360 feet near
the head of Middle Valley (Fig. 7). This area often has small nesting
concentrations of Blue-faced Boobies.

Cliffs dominate the perimeter of the island. The north, west,
and east cliffs (Figs. 8-10) are nearly perpendicular to the surface
of the ocean and at times may exhibit a small overhang. The cliffs
of the western perimeter decline rather irregularly in height and in
a few places rise to small peaks. The decline in height of the eastern
cliffs is far more uniform. The cliffs of the south side are only 50
to 100 feet high and may be relatively easily scaled. Six valleys,
varying considerably in the steepness of the slopes of the bordering
ridges, fan out radially to the center or south of a point in Adam's Bay.

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Figure 4. Looking at Nihoa Island from the south, December 1912.

Photo courtesy of Virginia Frear Wild.

Figurey or

Viewing Miller Peak from the ridge west of Middle Valley.

The dots in the right foreground are nesting Red-footed

Boobies. POBSP photograph, 8 March 1968, by Roger B.
Clapp.

He

oa
G
58
oO
ON

Figure 7.

Viewing Tanager Peak from the top of Middle Valley. Note

dense Chenopodium on slopes and dry Sida in left foreground.
POBSP photograph, 8 March 1968, by Roger B. Clapp

{ Se '
he oe Tet
-_

“ee

Low point along the north rim between Miller's and Tanager
POBSP photograph, 8 March 1968, by Roger B. Clapp.

Peaks.

Figure 8.

Tanager Peak as seen from the air. Note steepness of north
cliffs. BSFW photograph, July 1962, by David B. Marshall.

Figure 9.

Miller's Peak as seen from the northeastern rim of the
island, September 1964. BSFW photograph by Eugene Kridler.

The irregular southern perimeter of the island, enclosing
Adam's Bay, is divided into three coves, the westernmost of which
has a small sandy beach (Fig. 11) that is often called "Derby's
Landing."' It received its name in 1914 when a Lieutenant Derby
swam ashore there to obtain some rock specimens for Carl Elschner
(Bryan, 1942: 170). This is a poor landing spot because of the
presence of submerged rocks.

The coves to the west are edged by rock terraces and the best
landing spot is on the eastern side of the middle cove (Fig. 12).
Nonetheless, if the weather is not favorable, landing here can be
extremely hazardous.

The island is well vegetated but occasionally subject to
drought which changes its green appearance to a sere yellowish-
brown. Grasses tend to dominate the ridges with the valleys
covered by dense scrub.

GEOLOGY

The earliest geologic observations of Nihoa were made by
Sereno E. Bishop during a one day visit on 22 July 1885. His ob-
servations were of some value but were very brief and essentially
similar to those made later and in greater detail by Palmer (1927).

Carl Elschner, aboard the USCG ship THETIS in 1914, was unable
to land on the island but a Lieutenant Derby swam ashore and obtained
some rock specimens. Elschner (1915) described the island in some
detail but his observations were much improved upon by Harold S.
Palmer.

Palmer, geologist on the Tanager Expedition, visited the island
in 1923 and 1924 and later wrote the only detailed account of the
geology of Nihoa (Palmer, 1927). Our brief account here, unless
otherwise indicated, derives largely from his report.

Utilizing observations on the strike and dip of the dikes and
of the positioning of the some 25 dikes cutting the west cliffs
(Eger), Palmer’ (19272 11) concluded that

...Nihoa is a portion of the southwest quadrant

of the original cone. This is further corroborated
by the fact that Nihoa lies near the southwesterly
end of a submarine bank with depths of 20 to 40
fathoms. [The U.S. Coast and Geodetic Survey

Chart No. 4181 indicates that the bank has a depth
of 25 to 35 fathoms and extends 13 nautical miles
from Nihoa in a northwesterly direction.] The
survival of a part of the southwest quadrant may

be the result of protection from the attack of waves
driven by the northeast trades. If no other action
interfered, a homogenous [sic] island should be cut

10

Figure 10.

Figure J1.

The steep west cliff of Nihoa Island. BSFW photograph,
September 1971, by Eugene Kridler.

Western portion of island showing sandy beach in western-
most cove. Miller's Peak and Miller's Valley are to the

right. In the center is Pinnacle Peak and West Palm

Valley. To the extreme left is Dog's Head Peak. Just to

the West of Dog's Head Peak is the small West Valley. BSFW
photograph, June 1962, by David B. Marshall.

Figure 12. Best landing area is in the right foreground. Note the
extent of rock terrace along the edge of the island. Camp
is on a reasonably level spot near the base of Miller's
Valley. POBSP photograph, 8 March 1968, by Roger B. Clapp.

Figure 13. Three of the many dikes cutting the western cliffs. BSFW

nhotograph, 15 September 1971, by Eugene Kridler.

231-338 O-77-9

ay

back most on its windward side both by wave attack
and by stream erosion, and the last remnant should
be a part of the original leeward quadrant.

Palmer noted that none of the original lava surface remained
on Nihoa and stated that the flows were from 1 to 20 feet in thickness
and exhibited considerable lateral extent.

Perpendicular or extremely steep cliffs are found on the east,
west, and north sides of the island. Along the south shore cliffs
are 50 to 100 feet high. These cliffs were caused by wave erosion
which undermined higher layers of lava. Two other kinds of cliffs
may be found on Nihoa. The more abundant type is found where massive,
thick lava flows have resisted weathering and erosion more success-—
fully than have adjacent flows, thus forming low cliffs. A third
type of cliff is found where heavy dikes have protected the rock
behind them while that in front was worn away. Palmer (1927: 13)
described these as follows.

In this way two great dikes northwest of the sand
beach have formed a cliff 200 feet high, faced for
nearly its full length by the dike material. The
steep, chute like gash [the Devil's Slide, Fig. 14]
which debouches through the north cliff near the
northwest corner of the island has a steeply inclined
floor and cliffs for side walls. The gash is due to
the removal of the weaker flow lavas from the region
between two dikes....A shallower chute on the west
cliffs is due to the same processes working between
the western extensions of the same dikes.

Nihoa has no permanent streams but Palmer thought that inter-
mittent streams might be of considerable power, partly because of
the scarcity of small boulders and gravel in the streamways and
partly because the average slope of the island, excluding the cliffs,
is about 23 degrees. Runoff is mostly to the south side of the
island through the base of the valleys but the northwestern plateau
drains mostly to the north. Palmer found three small seeps at the
bottoms of streamways.

[These seeps] represent ground water which is held

up by relatively impervious layers of basalt, or which
circulates through fissures in the basalt. The smallest
seep is a little west of the head of the middle cove,

and issues from crevices in the cliff behind the terrace.
The second is at 270 feet elevation in the large valley
to the east....The water here appears to be brought out
by the overlying soil and rock by a massive and relatively
impervious lava flow. The largest discharge of water is
by seepage from the conglomerate body at the head of

the west cove, and is brought to the surface in the same
way. The water of all these seeps is heavily charged

with acrid-tasting matter, presumably derived from
the dejecta of the multitudinous birds and is
nitrogenous and phosphatic in character (Palmer, 1927: 15).

Along the south shore lies a nearly continuous wave cut terrace,
broken in three places, 4 to 8 feet above sea level and varying in
width from 10 to 50 feet. Fragments of terrace also remain along
other sides of the island.

At present wave action is cutting caves along the base of the
north, northeast, southeast and particularly the west cliffs of the
island. At the eastern end of the island a tunnel, through which
small boats may pass, cuts through the 300 foot promontory (Palmer,
ope edt.: 14).

Palmer found no slickensides, offset beds, or other signs of
faulting.

The rocks of the island are mostly olivine basalts and occur
either as dikes or flows. A few are olivine free basalts. No ash,
bombs or tuff was found. Most of the rocks are medium gray except for
a few which are dark brick red, due to oxide of iron. Oxidation is
not restricted to the surface of the rocks and presumably occurred
while the rock was still hot.

Olivine was found in at least 18 to 21 specimens examined by
Palmer and occurred particularly commonly in the dike rocks. Feld-
spar (labradorite) phenocrysts were found in four sections. The
ground mass of the rock consists primarily of elongated feldspar
grains. Among the feldspar grains are augite grains, a few olivine
grains and a little glass (Palmer, op. cit.: 16). Powers (1920)
also briefly described a rock collected prior to the visits by the
Tanager Expedition.

A detailed account of these rock specimens may be found in
Washington and Keyes (1926). All the rocks examined by them were
basaltic and occurred in four varieties: andesine basalt, olivine
basalt, labradorite basalt and picrite basalt. In their chemical
analysis of four specimens (one of each type of basalt) from 55 to
65 percent of the rocks consisted of Si0, and Al,0O, but appreciable
quantities of FeO, MgO, and CaO were also sHesenl (ieoninecon and
Keyes, 1926: 344).

The rocks of Nihoa are of high specific gravity. Palmer weighed
11 specimens of dike rock and found specific gravities ranging from
2.53 to 3.06, with an average of 2.88. The varying abundance of
olivine, a heavy mineral, accounts for the varying specific gravities.

Palmer also analyzed the sand from the beach in some detail.
The principal components were grains of calcium carbonate, olivine
and magnetite. Also found, but in much smaller amounts, were grains
of feldspar, augite and lava. Details on the size of the grains,

13

14

their relative frequency of occurrence, and discussions of relative
wear and appearance may be found by consulting Palmer.

Magnetite in the rocks of Nihoa causes wide variation in local
magnetic declination. The average declination on Nihoa, like that
of the surrounding ocean, is about 11 degrees E of N. However,
declination at 10 points on Nihoa ranged from 2 to 28.5 degrees
E of N, these two extremes coming from stations only 800 feet apart.

HISTORY
Prehistoric Habitation

Nihoa, like nearby Necker Island, was once inhabited by
Polynesians. Emory (1928) 2/ reported that of the many stone struc-
tures (Fig. 15) at least 25 were foundations of houses and about 15
were ceremonial structures. Fifteen bluff shelters were discovered.
He thought that the island could have sheltered a population of up
to 174 persons but believed that probably no more than 100 ever lived
there.

A large number of artifacts from terraces, house sites, and
bluff shelters was collected by the Tanager Expedition. Artifacts
of rock included grindstones, hammerstones, bowls, jars, and a
mortar.

Artifacts of plant origin included a bed and a pillow made of
bunchgrass, a wooden netting shuttle, a piece of breadfruit wood
which had been shaped into a crude tiller of European form, and ashes
and charcoal.

Artifacts of animal origin included a coral rubbing stone and
file, awls or needles made of bird bones, fragments of turtle shells,
and a fishhook of bone, probably from a human femur. Cowries
(Cypraea mauritiana) had been used as squid lures.

Human skeletal material was found at two sites. At one, ina
recess on the east side of Dog's Head, the remains of four adults
were found with petrels nesting among the skulls. At the other, on
a ledge facing the sea, were partial remains of an adult and two
children.

After comparing the archaeological materials collected on Nihoa
and Necker, inter se, and with those from other areas of Polynesia,
Emory (1928) concludes that at one time Nihoa supported a permanent

Dy)

— Emory (1928) gives an exhaustive account of the archaeology of
Nihoa derived primarily from the results of the field work done
by the Tanager Expeditions of 1923 and 1924.

Figure 14. Looking north down the "Devil's Slide". BSFW photograph,
15 August 1970, by Eugene Kridler.

Figure 15. Native structure on the northeast corner of Miller's
Plateau. BSFW photograph, 15 August 1970, by Eugene
Kridler.

16

population which was part of the culture that also sprang up on
Necker. He suggests that the permanent population of Nihoa arose
when Nihoa was cut off from communication with the larger Hawaiian
Islands.§/ He hypothesizes that the people of the Necker culture
were forced out of the main Hawaiians and settled for a time on
Nihoa. Emory concludes that the Necker culture was ''a pure sample
of the culture prevailing in Hawaii before the thirteenth century,
and that prehistoric as well as the historic Hawaiian culture may
be considered Tahitian in origin" (Emory, 1928: 122).

European Discovery and Early Visits

Europeans discovered the existence of Nihoa when the island was
sighted by Captain William Douglas of the H.M.S. IPHIGENIA. The
IPHIGENIA, sailing in company with the schooner NORTH WEST AMERICA,
sighted Nihoa at three in the morning of 19 March 1779 and hove to /
until daybreak. Meares (1790: 212), who evidently owned the ship,<
later reported that :

This island or rock, bears the form of a saddle,
high at each end, and low in the middle. To the
South it is covered with verdure; but on the North,
West, and East sides, it is a barren rock, perpen-
dicularly steep, and did not appear to be accessible
but to the feathered race, with which it abounds.
It was therefore named Bird Island.8/ It lies in
the latitude of 23 degrees 07' North, and in the
longitude of 198 degrees 10' East [161 degrees 50'
West], by a medium of several observed distances of
the sun and the moon.

During ensuing years a number of ships passed offshore without
making a landing: in March 1794 the British ships the DISCOVERY and
the CHATHAM, and on 1 August 1795 the British sloop PROVIDENCE (Buck,
1953: 44, 46); on 17 April 1817 the bark COLUMBIA (Corney 1896: 73);
and on 1 July 1825 the TARTAR (Morrell, 1841: 216).

In 1822, Queen Kaahumanu, having heard about Nihoa, sent two or
three vessels commanded by Captain William Sumner to find the island

6/

— Nihoa, but not Necker, was by tradition believed to have been

known to the Hawaiians prior to the arrival of Europeans (Emory,
O28 3. kL Oi

Tf Rack. 1953" 96.

oy) A name by which Nihoa was best known through the first half of
the 20th Century.

a7,

and annex it to the Hawaiian Kingdom (Thrum, 1893: 7; Emory, 1928:
8;9/ Bryan, 1942: 169). Adams, himself, states that he visited the
island twice but never landed and further notes that ''Kaahumanu
visited the island in the Summer of 1822 and some of her natives
landed on it...."10/

Another early landing was made by the crew of the U.S. Schooner
DOLPHIN (Paulding, 1831: 192-195). Like many subsequent visitors
they had considerable difficulty landing and returning from the
island. Paulding's description follows.

On the ninth of January, [1826] at eight in the
morning...[Nihoa] was discovered close to us.

We tacked and stood back close in with the south-
west side, where was a small sand-beach, fifty to
a hundred yards long.

The captain, taking the Globe's whale-boat went

in shore to fish, but seeing a few seal upon the
sand beach was induced to land. It soon afterwards
became squally and blew with great violence. The

surf...rose with the wind, and,...the captain,
after a short examination of the island...found it
impossible to launch his boat...[and was forced to

pass] the night on the island. It blew a gale and
rained in torrents all night. The captain and his
boat's crew took shelter in a cavern upon the sea-
shore, where they had not been long by a comfortai:le
fire they had made, when, by the rising of the tide
the sea broke in upon them, and they with difficulty
escaped to the side of the rocks, and thence upon
the sand-beach. The island was high and almost
perpendicular, and with the floods that fell and
rushed down its steep sides, rocks of a large size
were disengaged from their beds, and came tumbling
down in every direction, to the great peril of the
captain and his boat's crew....After a little search,
they found an asylum in a cave at the side of the
mountain, where they passed the night....

9/ Emory states also that Thrum's information about the 1822 visit
was obtained from the journal of Captain Alexander Adams. This
is impossible as the Adam's journal cited (Thrum, 1905: 66-74)
covers only the period 16 January 1816 to 26 December 1817.

10/ Adams to L. Kamehameha, 18 March 1857. State of Hawaii, State
Archives, Honolulu.

18

[The following morning] When the captain saw the
schooner approaching close in with the island, he
made a last effort to launch his boat. They suc-
ceeded in getting her into the breakers but the
first heavy roller that broke under them severed
the boat amid-ships, and the captain upon end of
her and a man that could not swim on the other,
were hove up safely on the beach by the succeeding
wave. The rest of the boat's crew were good
swimmers and also landed in safety. Our boat was
not far off when this occurred, and anchoring as
near as possible to the shore, the men, all but
one, swam off to her through the surf. The only
way we could devise to get the captain and seaman
off, was to float a cork-jacket on shore, at the
end of a line, which being put on by the captain
and seaman, afternately, and a rope tied around
them, they were hauled through the surf without
any other injury than swallowing a quantity of
salt-water.

Bird's Island is an uninhabited rock, about a league
in circumference, and the highest part from five to
eight hundred feet above the ocean. Where our boat
landed, is the only spot where a landing could be
effected, and upon that side alone it has an
inclination by which it may be ascended. Every
where else it is perpendicular, and at a distance,
looks like the work of art. It has a scanty vegeta-
tion.

In the late 1850's many of the central Pacific islands became
objects of commercial interest because of guano which was treated to
make fertilizer. Most interest centered on the dry equatorial islands,
many of which had substantial deposits, but the strong demand for guano
and the potential for quick profits resulted in almost all central
Pacific islands being investigated.

In December 1856 the Brisith warship HAVANA sailed from Honolulu
to determine whether guano deposits were present on Nihoa. The ship
arrived offshore on the morning of the eighth and remained offshore
through the eleventh, but heavy seas prevented a landing. Captain
Harvey went as close to shore as he dared but saw no guano, concluding
that "from the formation of the rock and the large amount of heavy
rain that falls in...[the island's] vicinity, I do not imagine it
possible that any quantity could accumulate; --nor were birds seen
in such quantities as to warrant the expectation" (Harvey, 1860: 423).

This expedition by a foreign ship in Hawaiian territory evi-
dently alarmed the Hawaiian government, for shortly thereafter a

iL)

11/

circular appeared claiming Hawaiian dominion over Nihoa.—

The following spring an expedition was dispatched to Nihoa to
annex formally the island to the Hawaiian Kingdom.

On 23 April 1857 Captain John Paty of the schooner MANUKAWAI,
with King Kamehameha IV on board, landed and again annexed the island
to the Hawaiian Kingdom. Paty's log describes the visit.

At 10 a.m. went ashore (got upset in landing).
The King and Governor landed at the same time
Naa CANOC sais).

I deposited a bottle at the foot of the pole

near the landing place, containing notes agreeable
to my instructions...also a plate of copper on
which I scratched 23rd April A.D. 1857. King
Kamehameha IV visited this Island, and took
Possession.

Not seeing anything to warrant my longer stay
here, I got under way at 3 p.m. (Paty, 1857).

The EURYDICE, a French man-of-war, visited Nihoa at the same time and
returned to Honolulu with the King and his party on board.

One of the primary purposes of the visit was to determine whether
guano was present in sufficient quantity and quality for profitable
mining. To Kamehameha's disappointment, Wm. Hillebrand, who analysed
the samples brought back, concluded that the material was not valuable
enough for shipping and export but that it might be of some use as a
fertilizer in areas of the main Hawaiian Islands.12/

On 30 December 1858, Nihoa was viewed offshore by Lt. J.N.
Brooke of the U.S. Schooner FENIMORE COOPER while en route to French
Frigate Shoals.23/

In the early morning of 22 July 1885 an excursion party of over
200 people, including Princess Liliuokalani, landed on Nihoa from
the steamer IWALANI. Others in the party included Sereno E. Bishop

au D.L. Gregg to the U.S. Secretary of State, 17, 24 December 1856.

State of Hawaii, State Archives, Honolulu.

Report from W. Hillebrand to L. Kamehameha, Minister of the
Interior, 26 August 1856. State of Hawaii, State Archives,
Honolulu.

2) Log of the U.S. Schooner FENIMORE COOPER. Rec. Group 37, U.S.

Nat. Archives, Washington.

and his assistants who made topographic and geological observations,
Sanford B. Dole, who made ornithological observations, Mr. Jaeger, a
botanist, and Mrs. E.M. Beckley, representative of the Hawaiian
Government Museum (Emory, 1928: 10). Mr. Williamsl4/ and W.E.H.
Deverill took photographs and the latter collected several archaeo-
logical specimens.

Bishop's statement that "the island had been ransacked for birds,
skins, eggs, and feathers" (Bishop, 1885b: 5) suggests that quite a
number of ornithological specimens were collected but the present
whereabouts of these specimens is unknown. Possibly most were lost,
as were most photographs and the cameras, when two boats swamped
during the departure of the excursion party.

Toward the end of the visit a fire had broken out that evidently
consumed much of the island's vegetation (Bishop, 1885a: 4).

The schooner KAALOKAI carrying the Rothschild Expedition was off
Nihoa on 26 and 27 May 1891. No landing could be made because of
heavy seas, but expedition personnel Henry C. Palmer and George C.
Munro made observations of birdlife offshore. Seventeen different
species were seen (Palmer 7” Rothschild, 1893-1900: vii; Munro 1941a:
41, 493: 1941b: 16)’.

Bryan (1942: 170) reported a visit to Nihoa in September 1894
by the H.M.S. HYACINTH; soundings were taken.

Nihoa was acquired by the United States as a part of the Territory
of Hawaii on 7 July 1898.

In 1902 the U.S. Fish Commission ship ALBATROSS visited Nihoa
twice, 1 to 3 June and 5 to 9 August, while engaged in deep-sea
explorations off the Hawaiian Islands. No landing could be made on
either visit. Walter K. Fisher, one of the biologists, subsequently
reported observations of 19 species of birds that were seen on or around
the island (Fisher, 1903). On both visits the ALBATROSS dredged off-
shore to collect marine organisms (see Appendix Tables 2 and 3).

In April 1909, by Executive Order 1019, Nihoa was included in
the Hawaiian Islands Bird Reservation.

Visits by the Thetis - 1904-1916

During the early 1900's Nihoa was visited often by the U.S.
Revenue Cutter THETIS. Most of these visits were casual inspections
which were made to discover whether birds had been molested by feather
poachers, a recurring problem on several of the other northwestern
Hawaiian Islands. No evidence was ever found that poachers had
visited Nihoa. Lack of poaching, in all probability, was not due to
insufficient numbers of birds but rather to the difficulty of landing
and working on the steep slopes.

14
ab) Possibly J.J. Williams who took bird photographs on Laysan Island
in the 1890's.

21

Nihoa was sighted on 9 June 1904 but no landing was made since
the ship was en route to Lisianski to apprehend Japanese who were
killing birds there (Hamlet, ms.) .5/

Another visit was made by the THETIS on 12 January 1910 but
this time heavy seas prevented a landing. W.V.E. Jacobs (ms.),
the vessel's commander, noted that he saw no birds except the Sooty
[Black-footed] Albatross and no evidence of human habitation.

The THETIS passed offshore on numerous subsequent occasions
between 1910 and 1912. On one of these offshore visits (17 December
1912), George Willett (ms.), who was engaged in surveying the north-
western Hawaaian Islands for the Bureau of Biological Survey, listed
eight species of birds.

The THETIS sighted Nihoa twice in 1913 but no landing was made
until 7 September 191416/ when crewmen swam ashore to collect rock
specimens for Carl Elschner. One of those who swam ashore was Lt.
W.N. Derby. The small sand beach on which they landed was named for
him, "Derby's Landing" (Bryan, 1942: 170).

On 18 March 1915 six persons went ashore, landing on a bit of
sandy beach on the south side of the island near the western end.
During the day the officers made a bird survey, noting species and
numbers and taking photographs.

One of the officers, Lt. W.H. Munter, later made a report in
which he listed 14 species, nine of which were recorded as breeding
(Munter, 1916). Two other species, the Christmas Shearwater and the
Bristle-thighed Curlew, were seen by the ship's captain, J.H. Brown
(ms.). Munter's obscure and little-known report was the first fairly
comprehensive list of birds of Nihoa. His report also includes the
earliest mention of the Nihoa Finch, later described by W.A. Bryan
(1917). Although some of Munter's estimates seem high in view of
more recent, detailed observations, others seem quite reasonable. On
the whole the report was one of the most valuable early surveys of the
birdlife of Nihoa, particularly when one considers the brevity of the
vaisit.

While the landing party was making observations on the high
slopes, the seas began to rise. The long-boat's crew lost control

) Log of the U.S. Revenue Cutter THETIS. Rec. Group 26, U.S.
Nat. Archives, Washington.

Elschner (1915: 9) stated that sailors swam ashore at Nihoa in
May 1910, but this is evidently incorrect as an examination of
the logs of the THETIS for that period reveals no such landing.
Probably Elschner confused Nihoa with Necker where a landing was
made on 23 May 1910.

Ze

of the boat and the coxswain severely injured his leg while trying
to hold the boat in position. By the time the survey party returned
the long-boat had been partially stove in and could not be used for
transportation. Consequently the ship's officers had to jump into
the sea from a low cliff and swim for a second long-boat.

The following year on 12 February a party from the THETIS again
landed. Several officers, including Munter, again explored the island.
Munter (ms.) recorded 13 species of birds, seven of which were breed-
ing. He also collected five specimens of the then undescribed Nihoa
Finch and made a collection of plant specimens.

In response to a request by the Department of Agriculture, the
U.S.S. HERMES made a survey of the birdlife of the leeward chain in
September 1918. Nihoa was visited on the second but, as had often
happened before, heavy seas prevented a landing. White Terns, Sooty
Terns, Wedge-tailed Shearwaters, albatross, and tropicbirds were noted
from offshore, but no other bird observations were recorded (Diggs,
MSi)

In 1919 Nihoa was visited by Gerrit P. Wilder, Warden of the
Hawaiian Islands. He arrived offshore on 7 October aboard the
lighthouse service tender KUKUI. The ship anchored in Adams Bay
and a landing was made about 150 yards east of Derby's Landing
(Anon 29.20'2 (560).

Visits by the Tanager Expedition

Nihoa was visited in 1923 and 1924 as a result of a cooperative
scientific venture sponsored by the U.S. Navy, U.S. Biological Survey,
and Bernice P. Bishop Museum. These visits, known collectively as the
Tanager Expedition,1// are by far the most important early scientific
visits.

On 24 May 1923 the TANAGER sighted Nihoa. Commander King,
Alexander Wetmore, leader of the scientific party, David L. Thaanum,
Theodore Dranga, Chapman Grant, and Eric L. Schlemmer took a long-
boat in to look over landing possibilities but found that high surf
precluded landing the whole party (Wetmore, ms.). Grant, Thaanum,
and Dranga got ashore for a brief period on the rock ledge and
managed to collect some molluscs and echinoids. The following day
Wetmore and others tried to get ashore but sea conditions had not
improved. Finally on 26 May the attempt to land was abandoned and
the TANAGER sailed for Honolulu (Ball, ms.).

The following month, on another of her several cruises along
the chain, the TANAGER arrived the afternoon of 10 June and the field

au The expedition received its name from the support vessel, the

Navy minesweeper TANAGER, commanded by Samuel W. King.

party landed and established camp. The following day camp was set

up on nearby Necker and radio communicationl8/ between the two
islands was established. Part of the party subsequently sailed for
Honolulu, another group remained on Necker, and a third crew stayed
on Nihoa (Edward L. Caum, Harold S. Palmer, Charles S. Judd, and
Bruce Cartwright). On 20 and 21 June the TANAGER removed the parties
from Nihoa and Necker, respectively (Gregory, 1924: 21-22). During
the protracted visit hydrographic work was done, plane table topo-
graphic surveys taken, and extensive collections and studies were
made of the archaeology and biota (see Appendix Table 3).

The following year the TANAGER paid another visit to Nihoa (and
Necker), primarily to follow up on the archaeological work begun the
previous year. The party (see Appendix Table 1) was under the leader-
ship of Harold S. Palmer, but archaeological work was directed by
Kenneth P. Emory. Although the work done during the 4-day visit
(9-13 July) was primarily archaeological, data were also obtained on
topography, geology, and botany (Gregory, 1925: 19-20).

Visits in the Late 1920's and Early 1930's

The U.S. Coast and Geodetic Survey Ship GUIDE, commanded by
Thomas Maher, visited Nihoa in April, May, and June 1928 19
Most of the survey party's work was hydrographic in nature but at
least one archaeological specimen was collected (Emory, 1928: 45).

On 19 August 1932 Nihoa was visited by the U.S.S. MONTGOMERY
(DM 17) commanded by L.E. Clifford. The landing party saw many birds
and noted a small lean-to facing south on the slope about 200 feet
above Adams Bay. Remains of a radio receiver were found in the shack
but no other evidence of human habitation was seen.29/ This shack
may have been built by personnel from the GUIDE during its survey of
the island a few years previously.

In 1934 the U.S. Coast Guard Cutter ITASCA paid a brief visit
to Nihoa to observe conditions on the island. The ship arrived late
in the afternoonof 8 February but, like many preceding visitors, the
crew was unable to effect a landing. After visiting Necker and French
Frigate Shoals, the ship returned to Nihoa on 11 February; an even

ey The exact itineraries of each individual are difficult to
establish as several trips were made between the islands.
Appendix Table I lists personnel known to have spent some time
on Nihoa.

19/ Log of the U.S.S. GUIDE. Rec. Group 37, U.S. Nat. Archives,
Washington.

20/

Report from the commanding officer, L.E. Clifford, Rec. Group
37, U.S. Nat. Archives, Washington.

78)

24

heavier surf than before discouraged any attempt at landing. On
both visits the ITASCA circled the island and neither time found
any evidence of recent landings (Baylis, ms.).

The ITASCA circled Nihoa again on 19 June during another in-
spection of the northwestern Hawaiian Islands. Later that year
(10 December), while searching Hawaiian waters for a missing plane,
the STAR OF AUSTRALIA, the ITASCA revisited Nihoa. A private vessel,
the LANIKAI, also visited Nihoa during the search for the downed
Plane (Honolulu Star Bulletin, 6, 9 Feb. 1935).

On 3 March 1936, A.D. Trempe, in cooperation with the Biological
Survey and five crew members of the Coast Guard vessel RELIANCE, made
a three-hour inspection of the birdlife of the island. Trempe (ms.)
reported seeing 14 species of birds, six of them breeding.

Visits During the 1940's and 1950's

George Vanderbilt, his wife, and Clifton Weaver paid a visit to
Nihoa from a private yacht in 1940. The party arrived at Nihoa on
7 August and completed landing their gear by the afternoon of that
day. The ship then departed, returning to remove the party from the
island on the 16th. The party was primarily interested in ornitho-
logical observations and collections (Vanderbilt and de Schauensee,
JUCHAa by

In July 1951 Vanderbilt revisited Nihoa on the George Vanderbilt
Pacific Equatorial Expedition that was collecting fish and observing
seabirds. With him were: Vernon E. Brock of the Hawaii Division of
Fish and Game and Robert R. Harvy of Stanford University who served
as naturalists; his wife Anita and daughter Lucille; Jack Lance, a
sport-fishing guide; and T. Ivar Vatland and Baba Green, the captain
and crew, respectively, of the PIONEER. Little has been reported
about this visit and we know of no zoological publications resulting
from it (see Herald, 1952; Harry, 1953).

Nihoa was visited twice in the early 1950's by Frank Richardson
of the University of Washington. Richardson made observations on the
occurrence and breeding activities of seabirds. Both visits, one 21
to 22 December 1953, the other on 18 March 1954, were made from the

Coast Guard vessel BUTTONWOOD. Thirteen species of birds were recorded

on the first visit and 11 on the second (Richardson, pers. comm.).

The AUKAKA, owned by a Honolulu insurance man, David G. Nottage,
paid a visit to Nihoa in 1955. Aboard were Nottage and his brother
Peter; George Carter, a Pearl Harbor mechanic; Ed Sheehan, a Honolulu
radio announcer; and Ivan T. Rainwater, an airport supervisor with
the plant quarantine branch of the U.S. Department of Agriculture.
Rainwater, an amateur archaeologist, had been detailed to collect
specimens of charcoal for radioactive carbon dating by Dr. K.P. Emory

of the Bishop Museum. Other objectives were the collection of plants,
and the capture of frigatebirds for the Honolulu Zoo. The party ar-
rived early on 21 August and visited the island daily until 24 August
when a final departure was made at 1100 (Honolulu Advertiser, 28 Aug.,
1955; Sheehan, 1966).

Dale W. Rice and Karl W. Kenyon made a low-level photographic
aerial survey of Nihoa on 28 December 1957. Their purpose was an
accurate census of the albatross and seal populations.

Visits During the 1960's

Nihoa was visited 5 to 15 March 1961 by the U.S.S. DUVAL COUNTY
which was conducting the first phase of a military project to establish
first order astronomic stations, HIRAN, and azimuth marks on the
northwestern Hawaiian Islands in connection with the Hawaiian geo-
detic survey. Transportation between the ship and the island was
by helicopter (Roach, ms.).

One of the more extensive surveys of Nihoa was made the follow-
ing December by Raymond J, Kramer and Gerald Swedberg, biologists
with the Hawaii Division of Fish and Game. They landed on the island
by helicopter from the U.S.S..FLOYD COUNTY which was engaged in
Phase II of the HIRAN project. They landed about noon on 9 December
and left the island early in the morning of the 15th (Kramer, ms.).

During their stay detailed notes were made on the status of the
two endemic landbirds, the Nihoa Finch and the Nihoa Millerbird, and
on the status and distribution of the various species of vascular
plants occurring on the island.

Several of their botanical observations are of particular
interest since they indicate distinct changes in the vegetation since
the detailed survey made by the Tanager Expedition in 1923. Kramer
and Swedberg's careful count of the endemic palms (Prttechardia remota)
revealed a total of 283 palms as compared with 515 in 1923.

Brief notes were taken on seabirds and shorebirds. Eleven
species of seabirds and two of shorebirds were recorded.

At the time of the biologists' visit, they found four military
personnel, two from the Air Force and two from the Army, who had been
living on the island for 2 weeks and who left Nihoa with the biologists.

Kramer, accompanied by David H. Woodside, John W. Beardsley,
and David B. Marshall, revisited Nihoa for about 7 hours on 10 June
1962 during the HIRAN II project. During their visit the biologists
made brief notes on vegetation and birdlife, collected arthropods,
and examined the island to determine what effect the HIRAN operations
had had on its ecology. They found two species of plants (Cenchrus
sp., Paspalum sp.), evidently introduced by the military, growing at
the HIRAN antennae sites.

25

26

The Bureau of Sport Fisheries and Wildlife (BSFW), now no longer
a separate entity but subsumed within the U.S. Fish and Wildlife Service,
assumed responsibility for the management, inspection, and patrol of the
Hawaiian Islands National Wildlife Refuge in 1964 when a refuge manager
was assigned to Honolulu. Since then at least one landing, and usually
two, were made each year.21/ On six of these recent visits (Table 1)
BSFW survey parties were composed in part of personnel from the Smith-
sonian Pacific Ocean Biological Survey Program (POBSP) which investigated
the biota of the central Pacific from 1963 until mid-1969.

During these often brief visits Refuge personnel were primarily
concerned with administrative and management duties, and with studies
of the populations and breeding status of albatrosses, the Nihoa
Millerbird, and the Nihoa Finch. POBSP activities were primarily
directed towards obtaining data on occurrence, numbers, and breeding
status of seabird populations. Data gathered during these visits, and
much hitherto unpublished data from earlier visits, form the basis for
the faunistic accounts presented in-following sections of this report.

In 1967 Nihoa and other islands of the Refuge were designated a
"natural area" by the Bureau of Sport Fisheries and Wildlife. This
means that the Refuge management is seeking, as far as is possible, to
prevent any disturbance of the ecology of the islands. Landings may
be made only by permit from the Bureau, and visits are restricted to
personnel involved in scientific studies.

VEGETATION
by Derral Herbst22/

Low shrubs, seldom above 1 meter high, cover the sides and much
of the floors of the valleys (Figs. 16-17). These consist primarily
of Solanum nelsont, Chenopodium oahuense and Sida fallax, and may be
mixed or in almost pure stands. Less common elements include Sesbanta
tomentosa, Euphorbia celastrotdes and Eragrostis vartabilits. Of the
very few plants of the annual grass Pantewn torridum sighted in
August 1968, all but one were young seedlings. The shrubs are sparser
and the tufts of Hragrostis more common on the ridges. Intertwining
branches of the scrambling Euphorbia shrubs form dense mats around
rock outcroppings along the edges of the north cliffs (Fig. 18). In
August 1968 Herbst did not notice well defined vegetative associa-
tions, although these are found on other northwestern Hawaiian Islands.

2A),

— ' Appendix Tables 1 to 3 summarize personnel, accomplishment, and
papers resulting from these and earlier visits.

22)

— Department of Botany, University of Hawaii, Honolulu, Oahu,
Hawaii 96822.

Figure 16.

Figure 17.

231-338 O- 77 - 10

Remains of ancient house terrace in East Palm Valley,

August 1968. Vegetation is chiefly Chenopodiwn shrubs
with some Stda. Photograph by Derral Herbst.

Middle Valley, August 1968; vegetative cover--typical

of the sides of the valleys--consists primarily of 2 to

3 foot tall Stda and Chenopodium shrubs. Photograph by
Derral Herbst.

27

28

The first recorded botanical collection was made by a Dr. Rooke
in 1858 when he brought specimens of the Nihoa loulu, Prttchardta
remota, to Honolulu (Hillebrand, 1888: 451). Later collections
were made by the following: Capt. J.H. Brown of the U.S. Revenue
Cutter THETIS, September 1914; E.L. Caum, E.H. Bryan, Jr., C.M.
Cooke, Jr., and C.S. Judd, June 1923; E. Christophersen and T.T.
Dranga, July 1924; R.J. Kramer and G. Swedberg, December 1961;
J.W. Beardsley, June 1962; C.R. Long, September 1964; E. Kridler,
J.L. Sincock, and D. Herbst, August 1968; and D. Yen, May-June 1969.23/

The following annotated list includes all species of vascular
plants known from Nihoa Island. Twenty species were collected during
the Tanager Expedition visits of 1923-1924--three species and three
varieties subsequently being described as new (Christophersen and
Caum, 1931). Additional species collected subsequently and not
previously reported were Portulaca oleracea, collected by Long in
1964, Cenchrus echinatus var. htllebrandtanus, collected by Herbst
in 1968, and Setarta vertitctllata and Ipomoea pes-caprae, collected
by Yen in 1969. These, together with a sighting of a grass (Paspalum
sp.), that has not been collected, bring the total number of species
of vascular plants known to have grown on Nihoa to 25.

Magnusson (1942) published a list of lichens and Tsuda (1966)
included two algae from Nihoa in his enumeration of the marine benthic
algae from the Northwestern Hawaiian Islands. The specimens cited
below are deposited in the B.P. Bishop Museum Herbarium (BPBM) or in
the Herbarium of the University of Hawaii (UH).

Annotated Species List

Gramineae

Cenchrus echinatus var. hillebrandtanus (Hitche.) F.B.H. Brown
C. htllebrandianus Hitchcock

Herbst 1206 (UH), Yen 1010 (BPBM). Kramer (ms.) reported find-
ing and burning, in 1961, six Cenchrus spikelets which were stuck to
a towel of one of the HIRAN personnel. Herbst found two plants
growing in a pocket of soil on the floor of Miller Valley in 1968.
Both were removed and the area searched for seeds. Apparently these
efforts were in vain, as this plant is included in the 1969 collections
of Douglas Yen.

23/

— ' Plants were also collected in February 1916, August 1955, and in
March 1964 (see Appendix Table 2) but we do not know the present
disposition of these collections.

7g)

Eragrostis vartabilts (Gaud.) Steud.

Caum 61 (BPBM), Long 2417 (UH), Herbst 1208 (UH), Yen 1009
(BPBM). Found widespread over the island as in 1923-1924 but
observers in both December 1961 and September 1964 believed this
species was less abundant than indicated by Christophersen and Caum.

Panteum torrtdum Gaud.

Caum 60 (BPBM), Christophersen 5 (BPBM), Kramer and Swedberg 7
(BPBM) , Long 2408 2432 (UH), Herbst 1209 (UH), Yen 1011 (BPBM).
Widely distributed in 1923. Observers in December 1961 and September
1964 believed this species to be more abundant than indicated by
Christophersen and Caum. Only a few plants were seen in August 1968,
however.

Paspalum sp.

Found growing in the vicinity of the HIRAN area of operations in
June 1962 (Kramer and Beardsley, ms.). Not seen subsequently.

Setarta vertictllata (L.) Beauv.
Yen 1012 (BPBM). Near south coast, May, 1969.
Palmae
Pritchardta remota Becc.

Dr. Rooke in 1858 (mentioned in Hillebrand, 1888: 451; not
seen), Brown s.n. (BPBM), two sheets, one with Rock's no. 10347,
Caum 72 (BPBM), Cooke 299 (BPBM), Christophersen 9a (BPBM), Long
2412, 2440, 2443 (UH). This endemic species of palm, found only in
East and West Palm Valleys (Figs. 19-20), is the most conspicuous
component of the flora. A few trees--most of which are mature--grow
at the bases of basaltic cliffs on the steep outer slopes of each
valley. Most of the population, however, is crowded into small,
dense groves on the terraces lower in the valleys. Kramer (ms.)
suggests that their distribution may be determined by soil depth and
availability of water. The faces of the cliffs may act as natural
catchment areas which would add to the moisture at their bases, while
part of the run-off during rains would be held by the deep accumula-
tions of debris in the ancient, man-made terraces.

The palms have been counted four times in the last 45 years.
In June 1923 C.S. Judd counted 347 trees in West Palm Valley and 168
in East Palm Valley. The count did not include small seedlings
(Christophersen and Caum, 1931: 6). In December 1961 Kramer and
Swedberg counted 229 trees in West Palm Valley and 54 in the east
valley. Seedlings under one meter were omitted, and immature fruits
were noted on some of the plants (Kramer, ms.). C.R. Long divided

30

Figure 18. Low scrambling Euphorbta shrubs around rock outcroppings
along northern cliffs, August 1968. In the foreground
are Chenopodium and Solanwn shrubs. Photograph by Derral
Herbst.

Figure 19. Small grove of Prttchardta remota at base of cliff in
upper East Palm Valley, August 1968. Photograph by
Derral Herbst.

Sit

his count of September 1964 into six localities: West Palm Valley
contained 107 seedlings, 148 non-flowering or non-fruiting trees,

and 127 with flowers or fruit; East Palm Valley had 32 seedlings,

69 non-flowering or non-fruiting, and 46 with flowers or fruits.
Long's total is 390 mature trees and 139 seedlings. Herbst attempted
to follow Long's system when he counted the trees in August 1968.

His census follows: in the west valley, 142 seedlings, 106 non-
flowering or non-fruiting and 139 with flowers or fruits; in the east
valley, 238 seedlings, 11 non-flowering or non-fruiting plants and

43 with flowers or fruits. The total of 516 palms does not include
the very young seedlings with less than five leaves.

Kramer attributed the decline in the palm population to the ex-
cessive crowding of the trees in the terraces and to a natural
thinning of a long-range cycle of the population. Herbst's observa-
tions of this species and of the growth rate of closely allied species
on Oahu left little doubt in his mind that the discrepancies between
the counts lay primarily in the lack of a uniform definition of the
word "seedling." In addition, the difficulty of counting the palms
is aggravated by the density of the population and by the numerous,
vociferous seabirds that nest in the trees. The population probably
has remained fairly static over the last 45 years.

In August 1968 mature fruits were collected from both valleys
and were distributed to the Lyon Arboretum of the University of
Hawaii, Honolulu, and the Foster Botanic Garden, also located in
Honolulu.

Polygonaceae
Rumex gtganteus Ait.
Caum 71 (BPBM), Christophersen 8 (BPBM), Long 2411 (UH), Yen
1015 (BPBM). Has been found only in the Devil's Slide-Miller Peak
area. Sterile in September 1964 and August 1968.

Chenopodiaceae

Chenopodtum oahuense (Mey.) Aellen
C. sandwicheum Moq. f£. mterospermum Aellen

Caum 58, 67 (BPBM), Judd 1 (BPBM), Christophersen 7 (BPBM),
Kramer and Swedberg 6 (BPBM), Long 2410, 2413, 2418, 2444 (UH),

Yen 1019 (BPBM). Widespread; one of the most common, if not the
most common, plant on the island (Fig. 6).

Amaranthaceae
Amaranthus browntt Christophersen and Caum

Caum 73 (BPBM), Judd 2 (BPBM), Yen 1013 (BPBM). Caum and Judd
made the first collections of this plant in June 1923, observing that

it was 'most common on the ridge leading to Miller Peak, but [was]
abundant also on the ridges to the east" (Christophersen and Caum,
1931: 6). This endemic species was not seen again until May 1969,
when Yen found four plants growing near the summit of Miller's Peak.
The region around Miller's Peak was traversed by most, if not all,

of the parties recently visiting Nihoa. It does not seem likely

that other botanists would have missed the plant. However, Amaranthus
brountt is an annual and the ridges are the driest part of the

island. If it has the delayed germination period and the rapid life
cycle that are common in many annuals, it could easily have been missed
by the botanists who collected during August, September and December.

Nyctaginaceae

Eoerhavta repens ly
B. diffusa UL

Bryan 3a (BPBM), Caum 79 (BPBM). In 1923 it was found only on
the sandy beach in Adams Bay and may have washed away. Not collected
or seen since 1923.

Aizoaceae

Tetragonta tetragontotdes (Pallas) 0. Kuntze
Tetragonta expansa Murray

Caum 80 (BPBM), Christophersen 2 (BPBM). Christophersen and
Caum (op. cit.) listed it as "rare, found only just above the sand
beach and on steep rock faces west of it." Possibly the corky
buoyant fruits floated to Nihoa from one of the main islands and
although at least one generation grew there, the species apparently
failed to become established. Not collected since 1924.

Portulacaceae
Portulaca lutea Sol
Caum 65 (BPBM), Kramer and Swedberg 1 (BPBM), Yen 1002 (BPBM).
Portulaca oleracea L.
Long 2431 (UH), Herbst 1204, 1205 (UH), Yen 1001 (BPBM).

Portulaca villosa Cham.
P. caumit F.Br.

Caum 66 (BPBM), Kramer and Swedberg 3 (BPBM), Long 2414, 2433
(UH), Herbst 1207 (UH), Yen 1003 (BPBM). Portulaca caumtt was
described as a new species by F. Brown in 1931 (in Christophersen
and Caum, 1931: 29) with Caum 66 designated as the type. Brown states
that P. cawnit is "allied to P. villosa Chamisso, from which it differs
in the color of the flower, the commonly flat leaves and the smaller,

33

minutely rugous seeds."" Egler (1938: 265)--after studying living
plants of P. villosa from Koko Head on Oahu and the islet of
Kauhikaipu near Oahu, and from the collections of the Bishop Museum
--states that "with the possible exception of a statement of petal
color,'' all characters listed from the Nihoa species existed in the
portulacas of Oahu. The Nihoa specimen collected by Caum had white
petals, while those of P. villosa generally have the outer third to
outer half pink. However, Egler did find plants with entirely white
petals in the colony on Kaohikaupu. Stone (1963) concurs with Egler's
conclusions.

After observing plants from Nihoa and Kaohikaipu, grown in the
botany greenhouse of the University of Hawaii, Herbst concurs with
Egler's conclusions and notes that a number of plants on Nihoa had
deep pink or pink-tipped petals in August 1968. These plants were
found at an elevation of 165 meters in West Palm and Miller Valleys.

On Nihoa, P. lutea grows in shallow, rocky soil and in cracks
along the lower south side; a large colony was found in August 1968
near the remains of a heiau (temple) at the northeast end of Miller
Plateau. Portulaca villosa was found in pockets of soil and in
cracks throughout the island (Fig. 21). An introduced species,

P. oleracea, occurs in two places, the largest colony being on
Miller Plateau. A smaller colony, probably established within a year
of the August 1968 visit, is located at the base of Miller Valley on
a small rise usually used as a campsite by recent survey parties.

Seeds of P, oleracea probably were introduced accidentally
during the HIRAN operation in 1961 when the Miller Plateau area was
used as a helicopter landing pad and campsite.

Caryophyllaceae
Sehtedea verttctllata F. Br.

Bryan 2 (BPBM), Caum 70 (BPBM), Christophersen 2 (BPBM), Kridler
and Sincock, s.n. (UH), Herbst 1210a (UH), Herbst 1401 (BPBM, UH),
Yen 1005 (BPBM). In 1923-1924 "seen only on the cliff west of the
sand beach, and north of Millers Peak, just below the summit"
(Christophersen and Caum, 1931: 6). Two fleshy roots just beginning
to send out new leaves were collected by Kridler and Sincock in
August 1968, about half-way down Devil's Slide; specimens from one of
the roots, grown in the botany greenhouse, Manoa Campus, University
of Hawaii, were listed under Herbst's no. 1401. Seeds from Herbst
no. 1401 were later distributed to the Foster Botanic Garden and the
Lyon Arboretum. Herbst 1210a, consisting of a dried fragment of a
plant, was found at about 165 meters elevation in West Palm Valley.
Yen 1005 was collected on the south face of the cliffs of Dog's
Head (west end), adjacent to Emory's site No. 2 (Emory, 1928: 18).

34

Figure 20.

Figure 21.

Grove of Prttchardia (right foreground) in West Palm
Valley, 10 June 1962. BSFW photograph by David B.
Marshall.

Portulaca villosa plant growing from crack in face of
stone ledge, August 1968. Photograph by Derral Herbst.

35)

Leguminosae
Sesbanta tomentosa H. and A.

Bryan 5 (BPBM), Caum 63 (BPBM), Judd 3 (BPBM), Christophersen 4
(BPBM), Kramer and Swedberg 2 (BPBM), Long 2409, 2428 (UH), Yen 1016
(BPBM). As in 1923-1924, Sesbanta is now distributed widely over the
island. Found flowering in December 1961 and June 1962.

Zygophyllaceae
Trtbulus cetstotdes L.

Bryan 2a (BPBM), Caum 78 (BPBM), Long 2420 (UH). In 1923
"observed only at the foot of the ridge leading up to Miller's Peak,
just above the cliff" (Christophersen and Caum, 1931: 6). In 1962
found in flower at the bottom of West Palm Valley (Kramer and
Beardsley, ms.). Last collected in September 1964 when it was
found on the south side of the cliffs on the upper slopes.

Euphorbiaceae
Euphorbta celastroides Boiss.

Bryan 6 (BPBM), Caum 64 (BPBM), Judd 5 (BPBM), Kramer and
Swedberg 10 (BPBM), Long 2405, 2430 (UH), Yen 1014 (BPBM). Common
on the edge of the cliffs in 1923-1924 and on recent visits (Fig. 22)

Malvaceae
Stda fallax Walp.

Caum 69 (BPBM), Bryan la (BPBM), Judd 4 (BPBM), Kramer and
Swedberg 8, 11 (BPBM), Long 2421, 2429, 2437 (UH), Yen 1004 (BPBM).
Common over entire island.

Convolvulaceae

Ipomoea indica (Burm.) Merr.
I. insularts (Choisy) Steud.
I. congesta R. Br.

Caum 83 (BPBM), Christophersen 1 (BPBM), Kramer and Swedberg 9
(BPBM), Long 2427 (UH), Yen 1017 (BPBM). In 1923-1924 “observed in
the gorge just below Millers Peak, at the base of the pinnacle on
the west cliff, and above the sand beach" (Christophersen and Caum,
OSI Oo) ie Seen anmthe ronrmervareas am: 19 6il:

Ipomoea pes-caprae (L.) Sw.

Yen 1007 (BPBM). Collected from stony south face adjacent to
the east slope of the mouth of West Palm Valley; elevation about 35

36

meters. A littoral species cosmopolitan in the tropics and sub-
tropics. Its large, buoyant seeds were doubtless carried to the
island by ocean currents and deposited at the mouth of West Palm
Valley--an area to which several strand species have been restricted.

Boraginaceae
Heltotropium curassavicum L.

Bryan 4 (BPBM), Caum 77 (BPBM), Christophersen 6 (BPBM), Long
2436 (UH), Yen 1008 (BPBM). Collected from the sand beach at the
base of West Palm Valley and the immediately adjacent area in 1923,
1924, and 1964. In September 1964, 27 plants were counted mainly
on the slopes above the beach.

Solanaceae

Solanum nelsoni Dunal
S. nelsoni Var. cqumit F.Br.
S. nelsoni var. acuminatum F.Br.

Bryan 3 (BPBM), Caum 68, 84 (BPBM), Judd 6, 7, 8 (BPBM), Kramer
and Swedberg 5, 12 (BPBM), Long 2424, 2434, 2339 (UH), Herbst 1210
(UH), Yen 1018 (BPBM). Found commonly over the island (Fig. 23).

Solanum nigrun L. var. nthoense F. Br.

Caum 62 (BPBM). Two plants were seen in 1923 "one on the edge
of the southern cliff, about the middle of the island, and the other
in the pocket of a stream bed just above the sand beach" (Christophersen
and Caum, 1931: 6-7). Not reported subsequently.

Cucurbitaceae
Steyos nthoaensts St. John

Christophersen and Dranga 9 (BPBM), Kramer and Swedberg 4
(BPBM), Beardsley s.n. (BPBM), Yen 1006 (BPBM). This species, pre-
viously identified as Steyos pachyecarpus H. and A., has recently been
described as a new species by St. John (1970). It was found only
below Tanager Peak in 1924. Several patches found just below and
south of the highest Pritchardias in West Palm Valley in 1961. In
1962 found in the HIRAN camp area on Miller Plateau.

Plant Affinities

Of the 25 taxa known from Nihoa, four species and one variety
are endemic to the island:

Pritehardta remota Sehtedea vertictllata
Amaranthus browntt Solanum nigrum var. nthoense
Steyos nthoanensts

Figure 22. Euphorbia celastroides, August 1968.

A variety with the
same habit growing in a somewhat similar environment at

Kaena Point, Oahu, was in flower at this time but was
leafless. Photograph by Derral Herbst.

Figure 23.

The Solanwn nelsont flowers on Nihoa in August 1968 had

a white corolla and purple anthers; those of Moomi Beach,
Molokai, have light blue petels.

37

Among the remaining taxa, eight are restricted to the Hawaiian
Islands:

Eragrostts vartabtlts Sesbanta tomentosa
Panteum torrtdum Euphorbta celastrotdes
Rumex gtganteus Solanum nelsont
Chenopodtum oahuense Portulaca vtllosa

Seven taxa are native to Hawaii, but are distributed widely
throughout the Pacific:

Boerhavia repens Ipomoea tndtea
Portulaca lutea Ipomoea pes-caprae
Tribulus etstoides Heltotroptum curassavicum

Stda fallax

The remaining taxa are not native to Nihoa, to Hawaii or the
Pacihics ;

Cenehrus echtnatus 24 / Portulaca oleracea
var. htllebrandtanus— Setarta verticillata
Paspalum sp.

Tetragonia tetragontotodes

To summarize: of the 25 taxa of vascular plants known from
Nihoa, 20 are native to the Hawaiian archipelago; of the 13 taxa
endemic to the Hawaiian archipelago, 5 are restricted to Nihoa.

TERRESTRIAL VERTEBRATES

species Accounts

Common names of seabirds follow King (1967) in the following
species accounts. Taxonomic order follows that of Peter's (1931,
1934) Checklist of Birds of the World, volumes I and II, with the
exception of the Procellariiformes which follow Alexander et al.
(1965), the Charadriidae and Scolopacidae which follow Jehl (1968),
and the Sulidae which follow the A.0O.U. Checklist (1957). The
scientific name of the Wandering Tattler has been modified to follow
the latest supplement to the A.O.U. Checklist (A.0.U., 1973).

The species accounts which follow are set forth in a standard
format which is used throughout except in those instances (e.g.,
accounts of vagrants) where its use would be inappropriate. The
section headed Status gives the maximum recent population estimate,
delimits periods of occurrence and breeding, and briefly indicates
areas of the island used for nesting. The maximum recent estimate
is the maximum estimate obtained since 1960. These estimates, as well
as those others listed in the tables of observations are meant to

24 ; ; Ae
a Variety native to Hawaii.

39

represent the maximum number of flying birds present during any one
Survey. Such estimates include breeding and non-breeding adults, sub-
adults, and flying immatures but do not include dependent non-flying
young.

The section entitled Populations discusses numerical estimates in
part attempting to discover whether there have been any historical
changes in population levels and in part to analyze seasonal popula-
tion changes. Most population numbers were based on visual estimates
supplemented by nest counts and banding data. For the two endemic
subspecies of birds, the Nihoa Millerbird and Nihoa Finch, recent
estimates were usually based on randomly chosen transect censuses.

In these transects, usually 16.5 feet wide and 200 to 250 feet long,
all birds of either species seen were counted. Usually about 50
transects were covered and a total area of about 4 to 5 acres was
sampled. Standard deviation was calculated from the results of the
transect counts to obtain 95 percent confidence levels.

The section entitled Annual Cycle contains an attempt, based on
all available data, to determine the reproductive cycles and, where
possible, to pinpoint peaks of breeding activity. The section headed
Breeding Habitat specifies areas or habitats which seem particularly
favored by the species under discussion.

The section headed Banding summarizes all known bandings by the
POBSP and BSFW and the section headed Specimens lists the locations
of all specimens from Nihoa of which we are aware. The specimens
cited are deposited in the American Museum of Natural History (AMNH),
Bernice P. Bishop Museum (BPBM), Los Angeles County Museum (LACM),
Museum of Comparative Zoology (MCZ), Philadelphia Academy of Natural
Sciences (PAS), State University of Iowa (SUI), and United States
National Museum of Natural History (USNM). These two latter sections
are omitted if we have no records indicating that the species has ever
been banded or collected on Nihoa.

Birds

In all, 27 species of birds have been recorded from Nihoa Island
(Table 2). Eighteen of these are seabirds, all but one of which (the
vagrant Red-billed Tropicbird) breed on Nihoa. The remaining avifauna
consists of four migrant shorebirds, three vagrants (the Herring Gull,
Pintail, and Mockingbird) and two endemic passerines, the Nihoa Finch
and the Nihoa Millerbird.

Most species exhibit a distinct annual breeding cycle but in
some species at least a few individuals can be found breeding in every
month of the year. It seems likely that the birds of Nihoa, like
those on other northwestern Hawaiian Islands more intensively studied,
vary initiation of nesting by up to several weeks from year to year.
This certainly appears to be the case for the Gray-backed Terns and
Sooty Terns on Nihoa and may well apply to other species as well.

40

Comparison (Table 3) of peak egg laying periods on Nihoa with
those on Laysan, a more westerly island in the leeward chain, and
with those on Kure Atoll, westernmost of the northwestern Hawaiian
Islands, reveals that some species show distinct geographical varia-
tion in their reproductive cycles. In general, peak laying periods
on Laysan and Kure are in rough accord but it appears that on Nihoa
at least five species breed earlier than on the outlying islands. At

present we have no adequate explanation for these differences in

breeding regimes.

Table 2. The avifauna of Nihoa Island.

Taxa

ORDER PROCELLARIIFORMES
FAMILY DIOMEDEIDAE

Diomedea nigripes
Black-footed Albatross

Diomedea immutabilts
Laysan Albatross

FAMILY PROCELLARIIDAE

Bulwerta bulwertt
Bulwer's Petrel

Puffinus pactficus
Wedge-tailed Shear-
water

Puffinus nativttatts
Christmas Shearwater

FAMILY HYDROBATIDAE

Oceanodroma trtstramt
Sooty Storm Petrel

ORDER PELECANIFORMES
FAMILY PHAETHONTIDAE

Phaethon aethereus
mesonauta
Red-tailed Tropicbird

Phaethon rubricauda
Red-tailed Tropicbird

Current Status

Ustcommon breeder

Rare breeder

Abundant breeder

Abundant breeder

Common breeder

Uncertain; almost
certainly breeds

in at least small
numbers

Vagrant

Common breeder

Maximum Estimate
since 1960 and
when recorded

120-130;
March 1964

40;
July 1973

250,000;
July-August 1966

20, 000-31, 000;
July-August 1966

800;
March 1965

150%
March 1965

375-625;
July-August 1966

fable 2: (Continued)
Taxa
FAMILY SULIDAE

Sula daetylatra
Blue-faced Booby

Sula leueogaster
Brown Booby

Sula sula
Red-footed Booby

FAMILY FREGATIDAE

Fregata minor
Great Frigatebird

ORDER ANSERIFORMES
FAMILY ANATIDAE

Anas acuta
Pintail

ORDER CHARADRIIFORMES
FAMILY CHARADRIIDAE

Pluvialts domintea
Golden Plover

FAMILY SCOLOPACIDAE

Numentus tahttiensis
Bristle-thighed Curlew

Heteroseelus tneanus
Wandering Tattler

Arenarta interpres
Ruddy Turnstone
FAMILY LARIDAE

Larus argentatus
Herring Gull

Sterna lunata
Gray-backed Tern

Current Status

Common breeder

Common breeder

Common breeder

Abundant breeder

Vagrant

Uncommon migrant

Uncommon migrant

Uncommon migrant

Common migrant

Vagrant

Abundant breeder

41

Maximum Estimate
since 1960 and
when recorded

350;
March 1968

225;
March 1968 i

3,500;
March 1965

10,000;
March 1964

25
September 1971

50s
March 1965

a
September 1972

2;
September 1972

200;
March 1968

1s
March 1965

10,000;
March 1967

42

Table 2. (Continued)
Taxa

Sterna fuscata
Sooty Tern

Procelsterna cerulea
Blue-gray Noddy

Anous stoltdus
Brown Noddy

Anous tenutrostrts
Black Noddy

Gygts alba
White Tern

ORDER PASSERIFORMES
FAMILY SYLVIIDAE

Acrocephalus familtaris
kingt
Nihoa Millerbird
FAMILY DREPANIDIDAE
Psittatrostra eantans
ultima
Nihoa Finch
FAMILY MIMIDAE

Mimus polyglottos
Mockingbird

* Recorded once in 1923.

Current Status

Abundant breeder

Common breeder

Abundant breeder

Common breeder

Common breeder

Common endemic
breeder

Abundant endemic
breeder

Vagrant

Maximum Estimate
since 1960 and
when recorded

100,000;
March 1965

2,500:
July-August 1966

20,000;
July-August 1966

several or low
thousands;

March 1967, September
1971

3,000;
August 1971

625%
September 1967

6,686;
August 1968

1;
August 1971

43

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44

BLACK-FOOTED ALBATROSS Dtomedea nigritpes
Status

Uncommon breeder; maximum recent breeding population estimate:
120 to 130. Present and breeding from at least December25/ through
June; probably absent from July through at least the middle of October.
Nests on the ground in the Miller Plateau area.

Populations

Estimates from the 1915 survey (Table 4) are so large in com-
parison to all others that we believe they are probably erroneous.
Wetmore's 1923 estimate, and all but one of the more recent estimates
that have incorporated careful observations of the Miller Plateau
area, have consistently indicated a breeding population of about 100

yYvv vasa ec:

Reports from a number of March visits indicate that the number
of young present mav vary considerably. This may indicate either
that the number of breeding birds varies from year to year or that
mortality may be considerably greater in some years. No evidence of
mortality was found in July 1964, July-August 1966 and August 1968
so it seems more likely that the former hypothesis is correct.

Annual Cycle

The scant data indicate that birds lay at least by early December
and that all young hatch by early March. Young apparently fledge by
mid-July since none has been seen at the end of that month or later.

Breeding Habitat

All observers who noted the location of Black-footed Albatross
nests stated or indicated that this species was confined to areas of
little vegetation on Miller Plateau, north and northwest of Miller's
Peak.

Munter (1915: 132) described an area which was likely Miller
Plateau. He noted that the colony was located on "a plateau
several acres in extent...near the highest part of the island."
Similarly, Wetmore (ms.) noted that the colony was "on a small flat
below the pinnacle point of Miller's Peak at a point about 850 feet
above the sea.'’ The Miller Plateau area was the only one where these
birds were found nesting in December 1961 and March 1964, 1965, and 1968.

25 : ;
ce) These albatross almost certainly return earlier than December,

judging from observations on other northwestern Hawaiian Islands,
but direct observational evidence is unavailable for the late fall
and early winter on Nihoa.

45

Banding

The BSFW banded 186 Black-footed Albatross on recent visits to
Nihoa: 50 young in March 1964, 35 young and 41 adults in March 1965,
and 60 young in March 1968. None has been recaptured.

Specimens

Only one Black-footed Albatross, a male (USNM 300832), has been
collected on Nihoa. It was collected by Wetmore on 12 June 1923.

Table 4. Observations of Black-footed Albatross on Nihoa Island

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1891 26-27 May ? Presence noted from offshore (Munro,
1941a: 49).
1902 1-3 June u A number seen some miles west of
island (Fisher, d903% 7:78):
5-9 Aug. 4 None seen from offshore (Fisher,
US OSs 779).
LOTS) os: Mar’. 500 Young were still in "pin feathers"
(2,000) [probably means birds were still
downy] (Munter, 1915: 132). Alternate
estimate by Brown (ms.).
L9LG .12..Feb. 2 Nesting on Miller Plateau. About
as abundant as Laysan Albatross
(Munter, ms.).
1923 BVADT« ? Seen offshore (Wetmore, ms.).

11-16 June 120* Only 3 adults seen but 60 well grown
young found, some with wings fully
feathered (Wetmore, ms.).

LOSGe. 3) May. 2 Evidently nesting (Trempe, ms.).

1940 7-15 Aug. 0 (Vanderbilt and de Schauensee, 1941: 8).

1953 21-22 Dec. 0(?) Only part of island surveyed (Rich-
ardson, pers. comm.).

O54) 9S) Max 0(?) Only part of island surveyed (Rich-
ardson, pers. comm.).

1957 28 Dec. 100* ,+ An estimated 50 nests present (Rice

and Kenyon, 1962: 377).

46

Table 4. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1961 2 Mar. 2 Not noted from offshore (Woodside and
Kramer, ms.).
9-16 Dec. 50 Many incubating eggs (Kramer, ms.).
1962 10 June u Only 5 young seen, downy on head but
capable of flight (Kramer and
Beardsley, ms.).
1963 5-6 June ? One seen offshore (POBSP).
1964 6-7 Mar. 120-130% 50. young banded, about 10-15 more
present (BSFW, POBSP).
Zo: eduliy, 0 (BSFW).
23-24 Sept. 0 (BSFW, POBSP).
1965 13-14 Mar. 80-90% 35 young banded. Possibly 5 or 10
more present (BSFW, POBSP).
1966 28 July- 0 (BSFW, POBSP).
1 Aug.
1967 8-9 Mar. ? 5 birds seen about ship; breeding
area on Miller Plateau not censused
(BSFW, POBSP).
13-14 Sept. 0 (BSFW) .
1968 7-9 Mar. 120-124* 60 young banded on Miller Plateau.
1 or 2 more may have been present
(BSFW, POBSP).
24-27 Aug. 0 (BSFW) .
1969 21 Mar. 2 None observed, but primary nesting
area on Miller Peak not visited (BSFW).
1970°715 Aug. 0 (BSFW).
1971 18-19 Aug. 0 (BSFW).

15. Sept. 0 (BSFW).

Table 4. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1972 16 Sept. 0 (BSFW).
ojo Si July 0 (BSFW) .

* Number of breeding birds utilizing island.

+ Aerial observations only.

LAYSAN ALBATROSS Dtomedea tmmutabilis
Status

Rare breeder; maximum recent breeding population estimate: 40.
Present from at least December (see footnote 25 in the Black-footed
Albatross species account) through May or June, but may not breed in
all years. Nests on the ground, primarily in the Miller Plateau area.

Populations

Laysan Albatross apparently were never common on Nihoa (see
Table 5). The only large estimate--1,000 birds in December 1957
(Kenyon and Rice, 1962: 377)--is certainly erroneous in view of the
numbers recorded on other surveys. Evidently Kenyon and Rice con-
fused this species with either Red-footed or Blue-faced Boobies when
they made their aerial count, a possibility they themselves suggested.

Direct evidence of breeding has been noted only in March 1915,
December 1953, March 1968, and July 1973. The lack of nests on the
three other March visits indicates that during some years Laysan
Albatross nest unsuccessfully, if at all. No more than 40 birds are
known to have nested in any one year but it is possible that a few
more may have initiated nests. Rice and Kenyon's estimate of numbers
breeding in 1956-1957 is much larger but is erroneous (see Table 5).

Annual Cycle

The data are insufficient to document adequately the periods
of arrival, laying, hatching, and fledging. Existing data do not
indicate that Laysan Albatross on Nihoa have a cycle any different
from that on other northwestern Hawaiian Islands.

47

48

Breeding Habitat

Only Munter (1915: 132) and Kridler have given any indication
of where nests were found. Munter found a half-grown young about
500 feet up one of the slopes, and in March 1968 Kridler banded four
chicks within the Black-footed Albatross colony on Miller Plateau.

Other Areas of Occurrence

Other observers reported pre-breeding or non-breeding birds
from several areas. In December 1961, Kramer noted that all Laysan
Albatross were on Miller Plateau. In March 1964, BSFW personnel
saw most Laysan Albatross on Miller Plateau, but a few were as much
as a hundred feet down the south slope of Miller's Peak. In March
1965 Kridler noted that most were on the high ridge between Miller's
and Tanager Peaks; in March 1968 these albatross were seen only on
Miller Plateau and Tanager Peak.

Banding

Ten Laysan Albatross were banded by the BSFW on recent visits:
6 adults in March 1964 and 4 young in March 1968. No returns have
been obtained for these birds.

Table 5. Observations of Laysan Albatross on Nihoa Island

Population
Date of Survey Estimate Breeding Status, Remarks, and References
18919" 26=27° May kK Presence noted from offshore (Munro,
1941a: 49).
1902 1-3 June ? Only 1 or 2 seen some miles west of
island (Fisher, 1903: 778).
5-9 Aug. ? None seen from offshore (Fisher,
MOOS 7/9).
TOUS: SiS Mars Cas 120 At least 1 half-grown young seen
(100) (Munter, 1915: 132). Alternate
estimate by Brown (ms.).
1916 12 Feb. 2 Nesting on Miller Plateau. About
as abundant as Black-footed Albatross;
2 pairs seen "near the top of the
middle peak" (Munter, ms.).
19230") Go" Apy ; ? Many seen in ship's wake offshore
(Wetmore, ms.).
14 June 1 1 adult seen with Black-footed

Albatross (Wetmore, ms.).

49

Table 5. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1936 SMa. ? Evidently nesting (Trempe, ms.).
1940 7-15 Aug. 0 (Vanderbilt and de Schauensee,
LOAN 8).
1955), (21-22) Dec. 6-8 Breeding in small numbers (Richardson,
1957: 16; pers. comm.).
1954 18 Mar. 0 Only part of island surveyed (Rich-
ardson, pers. comm.).
Lay. 28 Dec. COR. "Nesting'' (Rice and Kenyon, 1962:
1,000+ Sits)
1961 9-16 Dec. Cay 30 "No eggs. Much bachelor dancing"
(Kramer, ms.).
1963") 45-6 June ? None noted offshore (POBSP).
1964 6-7 Mar. 38 Only adults seen. 1 seen on empty
nest (BSFW, POBSP).
25 July 0 (BSFW) .
23-24 Sept. 0 (BSFW, POBSP).
1965 8-9 Mar. ? One seen from offshore; Miller Plateau
not censused (BSFW, POBSP).
1966 28 July- 0 (BSFW, POBSP).
1 Aug.
1967 8-9 Mar. 1 No young seen; Miller Plateau not
censused (BSFW, POBSP).
13-14 Sept. 0 (BSFW) .
1968 7-9 Mar. 10* 4 young banded on Miller Plateau.
6 adults seen on Tanager Peak; no
young found there (BSFW, POBSP).
24-27 Aug. 0 (BSFW).
1969 24 Mar. i Counted. Miller Plateau area not

visited (BSFW).

50

Table 5. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1970° 25> Aug; 0 (BSFW).
1971 18-19 Aug. 0. (BSFW).
U5 \Septe 0 (BSFW) .
LOZe Leesept. 0 (BSFW).
L973) Sinduly, 4O0** 20 nearly fledging young counted (BSFW).

* Probable number of breeding birds utilizing island.

** Figure represents a minimum estimate of numbers of birds that
bred on the island.

Aerial observations only.

BULWER'S PETREL Bulwerta bulwertt
Status

Abundant breeder; maximum recent estimate: 250,000. Present
from March through September or October; probably absent the rest of
the year. Breeds from at least June through September. Nests under
rocks, in holes or crevices in rock ledges or occasionally in shallow
depressions under dense vegetation.

Populations

None of the numerical estimates (Table 6) is very accurate
because of the difficulty of censusing populations of this very
numerous petrel. Clearly, however, many thousands are present during
July and August.

Annual Cycle

March 1964 and 1965 observations suggest that Bulwer's Petrels
begin to return to Nihoa in March but none was observed on a number
of other March visits. It seems likely that their reported "absence"
in a number of cases may have been the result of simply overlooking
small numbers that were present. Descriptions of a bird call heard
in March 1967 suggest that Sooty Storm Petrels rather than Bulwer's
Petrels were heard on that visit.

eye

Although detailed data on the nesting cycle are scanty, those
available suggest that these petrels have much the same cycle on
Nihoa as on Laysan and other northwestern Hawaiian Islands. Most
birds probably do not arrive at Nihoa until April and eggs probably
are not laid until a month or so later. Only eggs were found on two
June visits, and only eggs and newly hatched young were found on the
two visits in late July and early August. Only young were found
from mid-August through mid-September but it seems likely that a few
late fledging birds may be present as late as October.

We tentatively assume that the hatching peak occurs in July and
early August and that most young fledge about late September,

Breeding Habitat

Several observers noted that this species was found throughout
the island but few described the nest sites. Nests were found under
rocks, in small holes in the rocks, and particularly in crevices in
rock ledges. A few nests were also found in shallow depressions
under dense vegetation.

Banding

The POBSP and BSFW banded 808 adults on recent visits: 8 by
the BSFW in September 1964 and 800 by the POBSP in July and August
1966. None has been recaptured.

Specimens

Ten Bulwer's Petrel specimens have been collected on Nihoa
(Table 7).

Table 6. Observations of Bulwer's Petrels on Nihoa Island

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1902 1-3 June e. Abundant offshore. "Birds had been
feeding on fish eggs? and ctenophores
or comb-jelly"' (Fisher, 1903: 795).
5-9 Aug. ? Presence noted from offshore (Fisher,
SOS 7.79).
1923 11-16 June abundant Incubating eggs. No nests with young
found (Wetmore, ms.).
940 7-15 Aug. extremely "No eggs were found, all nests ex-
common amined containing downy, black young"
(Vanderbilt and de Schauensee, 1941: 9).
Lobe eacl=22e pec. 0 Only part of island surveyed (Rich-

ardson, pers. comm.).

bz

Table 6. (Continued)
Population

Date of Survey Estimate Breeding Status, Remarks, and References

1954 18 Mar. 0 Only part of island surveyed (Rich-
ardson, pers. comm.).

1961 2 Mar. 2 Not noted from offshore (Woodside and
Kramer, ms.).

9-16 Dec. 0 (Kramer, ms.).

1962 10 June common* On eggs (Kramer and Beardsley, ms.).

1963 5-6 June ? Thousands seen offshore (POBSP).

1964 6-7 Mar. 10 Oné photographed by Kridler (BSFW,
POBSP).

25. July very Many nests found. Only eggs and

abundant newly hatched young seen (BSFW).

23-24 Sept. 5,00 Nearly full grown young and ea.25
dead fledglings seen (BSFW, POBSP).

1966 28 July- 250, 000** Mostly on heavily incubated eggs or

1 Aug. (225,000) with ca. 1 week old nestlings (BSFW,
POBSP). Alternate estimate by
Berger (197227 33)y.

1967 8-9 Mar. 2 Heard calling in burrows (BSFW,
POBSP); see text.

13-14 Sept. li A number of adults and half-feathered
young seen but most young evidently
departed from island (BSFW).

1968 7-9 Mar. 0? None seen on island but a few small
petrels seen offshore may have been
this species (BSFW, POBSP).

24-27 Aug. thousands Hundreds of adults seen during day
with numbers increasing to thousands
at night. Several downy chicks
found (BSFW).

1969 21 Mar. 0 (BSFW) .

L970) ESV Aus. 75,000- Burrows examined contained pairs

100,000 and small downy chicks. Much calling

by adults (BSFW).

53
Table 6, (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1971 18-19 Aug. 4,000 Half-grown young seen (BSFW).
15esept. ? Nearly fully-grown young were common

(BSFW) .

O20 lbs Sepit. ? At least 300 adults seen during day
(BSFW) .

Lo7see31 i July. 2 At least 400 adults seen during day
(BSFW).

* In their report Kramer and Beardsley noted that ''Bonin Petrels"
were common and on eggs. Since no other observer has found that
species on Nihoa, and, since Kramer and Beardsley did not mention
the presence of Bulwer's Petrel, we assume that ''Bonin Petrel"
was a lapsus calimts for Bulwer's Petrel. Marshall (1964: 160),
in a popular account of the visit, also indicates, we think er-
roneously, that Bonin Petrels were present.

** Most abundant species on island.

Table 7. Bulwer's Petrel Specimens from Nihoa Island.

2?
Museum Museum and Museum Date
Museum Males Nos. Females Nos. yg: Nos. Collected Collector
SUL at 18597 3 June Nutting
1902
USNM 2 300799, ul 530876 11-15 June Wetmore
530875 1923
PAS 3 146160, 1 146161 Aug. 1940 Vanderbilt
146168-
69
USNM tL 493022* 1 493023* 6 June POBSP
1963
* Collected offshore.
WEDGE-TAILED SHEARWATER Puffinus pactficus

Status

Abundant breeder; maximum recent estimate: 20,000 to 31,000.
Present from March through at least mid-November; most birds probably

54

absent during remainder of year. Most nesting occurs from June
through November. Nests in burrows, on the surface of the ground
under thick vegetation, and in cavities and crevices in rock falls
and ledges. Well distributed over entire island.

Populations

The few numerical estimates (Table 8) suggest that maximal
populations are on the order of tens of thousands rather than
hundreds of thousands. In August and September many thousands are
present; at dawn the steady chorus of calling results in a dull roar
that, at lower elevations, is muffled only by the surf.

Annual Cycle

March observations suggest that these shearwaters return to
Nihoa in great numbers within a very short period. In 1964, 1967,
and 1968 only a few birds were present in early March, while in 1915
and 1965 thousands were present by the middle of the month. Rich-
ardson's lower mid-March estimate does not fit this pattern. However,
he did not spend the night on the island and as a result possibly
did not see many birds that may have come to the island to roost at
night.

The nesting cycle appears to be similar to that found on Laysan
and other northwestern Hawaiian Islands. Most egg laying evidently
occurs in June although a few eggs may be laid in May. A few young
hatch in July but by far the greater proportion hatch in August. It
seems likely that most fledge within 5 to 5 1/2 months26/ after the
peak of laying. Thus, a few possible fledge as early as late October
and as late as early December with the peak being in November. No
surveys were made on the island during the presumed fledging period.
Kramer observed none in mid-December 1961 (Table 8).

Breeding Habitat

Nests have been found in a wide variety of situations on the
slopes of the island. A small proportion of the population digs
burrows in the ground but few such burrows are possible because of
the shallow soil cover and rocky nature of the island. Wetmore (ms.)
noted a number of burrows as much as 3 or 4 feet long but most were
only deep enough to provide shelter. A much larger proportion nest
in natural cavities in the rocks and on rock ledges protected by
overhanging rock. Many others nest on the surface, particularly in
areas covered by dense vegetation such as Chenopodium or Sida.

26 Based on the range of fledging periods (99-111 days) obtained in
a small sample (n = 10) on Kure Atoll (Woodward, 1972: 125) and
on an estimated 52 to 54 day incubation period (Woodward, pers.
comm.).

55

Color-Phases

Dark, light, and intermediate plumaged morphs have all been
seen on the island. No quantitative data exist on the occurrence of
the different color phases but one POBSP observer estimated that 95
percent of the birds were light-phase morphs. Wetmore (ms.) saw
gray-breasted birds occasionally, but the great majority was white-
breasted.

Banding

The BSFW and POBSP banded 288 adults on recent visits: 3 in
March 1964 and 85 in March 1965 by the BSFW, and 200 by the POBSP
in July and August 1966. None has been recaptured.

Specimens
We have found records of 8 study skins from Nihoa (Table 9).

The USNM houses in addition an embryo alcoholic (USNM 289301)
collected by Wetmore on 14 June 1923.

Table 8. Observations of Wedge-tailed Shearwaters on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May ? Presence noted from offshore (Munro,
1941a: 49).

1902 1-3 June 2 A number flew aboard. Stomachs
contained the "hard parts of small
cephalopods (squid, octopus, and
the like)" (Fisher, 1903: 792).

5-9 Aug. ? Presence noted (Fisher, 1903: 779).

1915 18 Mar. "very None. found nesting (Munter, 1915:

common''* 131). Alternate estimate by Brown

(50,000) (ms.).

1923 24-26 May ? Thousands seen circling ship
(Wetmore, ms.).

11-16 June 30, 000% Birds laying. Many fresh eggs
(Wetmore, ms.).

1936 3 Mar. ? 25 birds, found under clumps of grass
and rock, had red celluloid bands
placed on their legs (Trempe, ms.).

1940 7-15 Aug. 2% "Breeding was in its early stages,
both eggs and small young being ob-
served" (Vanderbilt and de Schauensee,
1941: 9).

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1953 21-22 Dec. 0 Only part of island surveyed (Rich-
ardson, pers. comm.).
1954 18 Mar, 40-60 ? (Richardson, pers. comm.).
1961 2 Mar. ? Not noted offshore (Woodside and
Kramer, ms.).
9-16 Dec. 0 (Kramer, ms.).
1962 10 June "common" On eggs (Kramer and Beardsley, ms.).
1963 5-6 June ? Thousands seen offshore (POBSP).
1964 6-7 Mar. 2 Courting behavior observed (BSFW,
POBSP):.
25 July very Several nests with eggs found (BSFW).
common
23-24 Sept. 2 Ca. 2,000 downy nestlings present
(BSFW, POBSP).
1965 13-14 Mar. 18,000 No eggs or young seen (BSFW, POBSP).
1966 28 July- 20,000- » Ca. 10,000 nests present, almost
1 Aug. 31,000 all with incubated eggs. One ca.
3 week old nestling observed (BSFW,
POBSP).
1967 8-9 Mar. 5 (seen) Apparently vanguard of breeding
population (BSFW, POBSP).
13-14 Sept. many Small and large downy chicks (BSFW).
thousands
1968 7-9 Mar. less than No evidence of breeding. Only 2
50 individuals actually noted (BSFW,
POBSP)’:
24-27 Aug. many So many calling in early dawn hours
thousands that combined sound made a low steady
roar (BSFW).
196929 21) Mar: 875 Nesting had evidently begun (BSFW).
1970 15 Aug. 20, 000- (BSFW) .
25,000
1971-. 18=19 Aug. 2 At least 2,000 birds were present
(BSFW).

57

Table 8. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
po7t 15° Sept. 2 Active burrows were widespread. Oc-
casional calling was heard (BSFW).
E9O72> L6<Sept.« 10,000 (BSFW).
1973 ¥31xJuly ? At least 500 birds were seen (BSFW).

* Stated to be most common bird on island.

Table 9. Wedge-tailed Shearwater Specimens from Nihoa Island

Ng
Museum Museum and Museum Date
Museum Males _ Nos. Females Nos. yg: Nos. Collected Collector
USNM iL 189399 Z 189400- 2 June Fisher
401 1902
SUI i 18592 3 June Nutting
1902
USNM it 300723 2 300724- 1 146153 Aug. 1940 Vanderbilt
725
CHRISTMAS SHEARWATER Puffinus nattvitatts

Status

Common breeder; maximum recent estimate: 800. Present from at
least early March through at least early October; absent remainder of
year. Most nesting probably occurs from April through September.
Nests most frequently at higher elevations and on the ground.

Populations

Recent numerical estimates made in March and July and August
(Table 10) suggest that less than a thousand birds nest on the
island. These estimates are of low reliability, however, and it is
possible that intensive censusing might reveal greater numbers present.

Annual Cycle

Information that would permit an analysis of the annual cycle
is sparse but what is available suggests that this species' cycle

58

closely parallels, but is earlier than, that of the Wedge-tailed
Shearwater. It is not known when these birds begin to return to
the island, but most evidently return in early and mid-March. Eggs
have been laid as early as late March (1969), but observations made
in June and July suggest that most laying probably takes place in
April or May. Considerably more detailed information is needed

on the nesting status in April and May in order to document ade-
quately the period of peak egg laying. Most eggs evidently hatch
in late June and July but at least a few hatch by late May. Some
fledging may occur in late August but most probably occurs in
September. Judging from the available observations, the number of
adults greatly decreases by mid-September, and it seems likely that
most have departed by late September.

Breeding Habitat

A number of observers have indicated that this species nests on
the higher elevations of the island. Many have recorded nests from
such areas. Wetmore (ms.) found a number of pairs on the slopes of
Miller's Peak at about 800 to 900 feet. Vanderbilt and de Schauensee
(1941: 9) recorded their presence "at the peak of Middle Valley and
along the cliff leading to Devil's Slide." Kridler found young
among the rocks just below Miller's Peak in July 1964, on the ridge
west of Miller's Peak in September 1967, and near Miller's Peak in
August 1968. He noted in 1965 that this species was "commonly found
at the higher elevations.'' In July and August 1966 most young were
found on the ridges and peaks and along the ledges on the northern
rim of the island between Miller's and Tanager Peaks. None was
observed on the vegetated slopes of the valleys.

Few data are available on the nature of the nest sites.
Wetmore (ms.) reported that Christmas Shearwaters nested "in little
shelters beneath overhanging stones or tussocks of grass" where
they might or might not be covered.

Banding

Eight adults, none of which has been recaptured, were banded
by the BSFW: 1 in March 1964 and 7 in March 1965.

Specimens

Six specimens have been collected on Nihoa. Wetmore collected
an adult female (USNM 300699) on 12 June 1923, a juvenile male (USNM
300700) on 14 June 1923, and an embryonic alcoholic (USNM 289399) the
same day. The other three specimens (PAS 146165-167), not certainly
sexed, were collected in August 1940 by the Vanderbilt Expedition.

Table 10. Observations of Christmas Shearwaters on Nihoa Island
Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May ? Presence noted offshore (Munro, 194la:
49).

1902 1-3 June ? None seen from offshore (Fisher, 1903:
793).

5-9 Aug. A few seen offshore (Fisher, 1903:
723)

1915 18 Mar. (100,000) Estimate by Brown (ms.). is most
likely erroneous,

1923 11-16 June 0? Ca. 20 pairs found building nests;
most birds prelaying or with eggs;
one ca. 1 week old young found
(Wetmore, ms.).

1940 7-15 Aug. rarest bird "No nests or eggs were observed but

on island one nestling young was seen" (Van-
derbilt and de Schauensee, 1941: 9).
£953 | 1S. Mar. 0? Only part of island surveyed (Rich-
ardson, pers. comm.).
21-22 Dec. 0 Only part of island surveyed (Rich-
ardson, pers. comm.).

1961 2 Mar. ? Not noted from offshore (Woodside

and Kramer, ms.).
9-16 Dec. 0 (Kramer, ms.).

1962 10 June very "On eggs and with downy chicks"

abundant (Kramer and Beardsley, ms.).

1963 5-6 June 2 50-100 seen offshore (POBSP).

1964 6-7 Mar. 2 Courting behavior observed. No
nests with eggs found (BSFW, POBSP).

25 July 2 2 young noted. Both were nearly
feathered with down remaining only
on nape of neck (BSFW).

23-24 Sept. 0 (BSFW, POBSP).

1965 13-14 Mar. 800 No eggs or young seen (BSFW, POBSP).

231-338 O - 77 - 12

59

60

Table 10. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
L966" 28) *July= 500 Ca. 250 young, all older than 3 weeks
1 Aug. (BSFW, POBSP).
1967 8-9 Mar. 1 seen Most of breeding population not
yet present (BSFW, POBSP).
13-14 Sept. 10 One young, feathered with down re-
maining on neck, noted (BSFW).
1968), -7=9 Mar. 100 No nests found but 1 pair seen
copulating by Kridler (BSFW, POBSP).
24-27 Aug. 6 3 adults and 3 chicks seen near
Miller's Peak. Chicks in juvenile
plumage except for scant down on
top of head and nape of neck (BSFW).
1969) 21) Mars 50 3 found incubating eggs (BSFW).
197:0.% 15.'‘Aug’. 50-100 (BSFW) .
1971 18-19 Aug. 40 Downy to near-fledging young seen
(BSFW).
15)Septy ? Several seen (BSFW).
1972 .16.Sept. 30 (BSFW).
SOOTY STORM PETREL Oceanodroma tristramt

Status

Poorly known. Occurs on Nihoa, at least in small numbers, and
apparently breeds there.

Observations

The Sooty Storm Petrel has been recorded thrice on Nihoa and
once from offshore. Fisher (1903: 795) obtained an immature that
retained a trace of down which flew aboard his ship on 1 June 1902.
The present location of this specimen is unknown (Vanderbilt and
de Schauensee, 1941: 9-10).

On 6 and 7 March 1964 POBSP personnel saw none but heard one
calling from a burrow. In addition, they collected two very young

61

petrel chicks that were found dead. These specimens cannot be
located but they were probably young Sooty Storm Petrels since no
other species of petrel known to occur on Nihoa would be expected
to have chicks at that time of year.

During the visit of 13 to 14 March 1965 Kridler banded one
petrel and collected another. In March 1969 Kridler captured and
banded two adults which were taken from a burrow about half-way up
one of the canyons. Several other petrels were seen in crevices
and hollows during this visit.

There are no other definite sightings but observations at-
tributed to Bulwer's Petrel (see that species account) in March 1967
may have referred to this species. In March 1968 several small
petrels which could not be positively identified were seen offshore.

This species is the least conspicuous of the petrels breeding
in the northwestern Hawaiian Islands. Its call does not carry well
and it tends to form localized colonies. Little attempt was made
to make nocturnal surveys on some of the recent visits. Thus, if
Nihoa has only a small population of these petrels, they could have
been easily overlooked.

Banding

The BSFW banded 1 adult in March 1965 and 2 adults in March 1969.

Specimens

We have been unable to discover the location of any of the four
specimens mentioned above.

RED-BILLED TROPICBIRD Phaethon aethereus mesonauta
Status

Vagrant; one specimen record from June 1923.
Observations

Wetmore collected an immature female (USNM 300997) as it flew
over Nihoa on 15 June 1923. The specimen was identified as belonging
to the race P. qa. mesonauta (Clapp and Woodward, 1968: 10-11). (The
specimen number was erroneously reported as 300977 in Clapp and Woodward
(1968) ). This race breeds in the Pacific from the Gulf of California
and the Revilla Gigedo Islands to the Galapagos and islands near the
coast of Ecuador (AOU, 1957: 27).

The only other specimen record for the central Pacific is one
taken in 1968 at French Frigate Shoals (Amerson, 1971: 184). Recent
sight records also exist for Johnston Atoll (Moynihan, 1957: 36;
Amerson and Shelton, in press).

es

62

RED-TAILED TROPICBIRD Phaethon rubricauda

Status

Common breeder; maximum recent estimate: 375 to 625. At least
small numbers (tens) present in all months, but more numerous from
March through September. Known to nest from at least mid-March
through at least early October. Nests on ground in shelter or
rocks or vegetation.

Populations

Although of low reliability, the various population estimates
(Table 11) indicate that less than a thousand tropicbirds breed on
Nihoa. Wetmore's May and June 1923 estimate of 800 birds is probably
not significantly different from the estimate of 375 to 625 birds made
in July and August 1966 due to the low reliability of estimates,

The March 1965 estimate of 400 seems unusually large for that
time of year--so large, in fact, that the difference from other
years may be significant; possibly more birds initiated nesting
earlier than usual. The March 1965 survey was the only recent March
survey on which active nests were found.

Annual Cycle

Data are too scanty for detailed analysis of seasonal variation
in numbers, but available observations indicate that maximal numbers
are present in mid-summer and minimal numbers in mid-winter.

The earliest that eggs were recorded was 3 March and the latest
they were recorded was 24 to 25 August. The largest number of eggs
are probably present from mid- or late April through June. The
presence of eggs in March 1936 and 1965 indicates that hatching those
years could have occurred by early May. Most hatching probably occurs
in late June and early July.

If the nests initiated in March were successful, young could have
fledged by late July. Most fledging likely occurs from late August
through September, with at least a few birds fledging in October.

Breeding Habitat

According to most observers, Red-tailed Tropicbirds are widely
distributed over the slopes of the island. They are common in rocky
crevices at the base of rimrock outcroppings at the west end of the
island and in West Palm Canyon, and have been reported as abundant
in the vicinity of the cliff face between Miller's and Tanager Peaks.
Most nests are in rocky cavities but others are found under over-
hanging rock ledges. Some nest on the surface of the ground beneath
dense vegetation such as Chenopodium and Sida.

63

Banding

Seven birds were banded by the POBSP and BSFW in 1964: 4
adults in March by the POBSP and 3 nestlings in September by the
BSFW. No returns have been obtained.

Specimens

Two study skins collected by Wetmore in 1923 are apparently all
that have been collected on Nihoa. Adult males (USNM 300998-999) were
collected on 11 and 12 June; and a skull (USNM 289154) was collected
on 15 June.

Table 11. Observations of Red-tailed Tropicbirds on Nihoa

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1891 26-27 May ? Several seen displaying from offshore
(Munro, 194la: 50; Palmer tn Roth-
schild, 1893-1900: vii-viii).
1902 1-3 June ? ' Possibly seen from offshore (Fisher,
LQOS: (78.5 77.90)
1915 18 Mar. several 1 pair found nesting (Munter, 1915:
132).
bg23 5 Apr. ? Occasionally seen offshore (Wetmore,
ms.).
24-26 May ? Seen from offshore (Wetmore, ms.).
11-16 June 800 "Nesting'' (Wetmore, ms.).
OB Go Mar. 2 Nests with eggs found (Trempe, ms.).
1940 7-15 Aug. fairly Mostly young nearly full grown but
numerous 2 nests with eggs found (Vanderbilt
and de Schauensee, 1941: 10).
1953 21-22 Dec. 0) Only part of island surveyed (Rich-
ardson, pers. comm.).
1954 18 Mar. 0 Only part of island surveyed (Rich-
ardson, pers. comm.).
1961 2 Mar. 2 Not noted from offshore (Woodside and
Kramer, ms.).
9-16 Dec. if Seen flying high over island (Kramer,

mS.) <

Table 11. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1962 10 June common Many with eggs (Kramer and Beardsley,
TS\s)) 13

1963 5-6 June 2 Ca. 20 seen from offshore (POBSP).

1964 6-7 Mar. 30 Birds at nest sites but no nests with
contents (BSFW, POBSP).

20 July ? Eggs to nearly grown young (BSFW).
23-24 Sept. 50 4 large downy young and 5 near-
fledging young found (BSFW, POBSP).

1965 13-14 Mar. 400 Eggs but no young present (BSFW, POBSP).

1966 28 July- Ca. Ca. 100 nests with young; ca. 202

i; Aug. 375-625 about 2-3 weeks old; ca. 80% older
than 3 weeks (BSFW, POBSP).

1967. 8-9 Mar. 20 No eggs or young found (BSFW, POBSP).

13-14 Sept. common A few very large chicks observed
(BSFW).

1968 7-9 Mar. 150-200 No nests with contents found. Ca.

50 seen in flight over the island at
once but no more than 10 found on the
ground (BSFW, POBSP).

1968 24-27 Aug. 200-300 From eggs to nearly fledged young.
Over 50 nests recorded during course
of other activities (BSFW).

1970" (15*Aug: 100 Large feathered young noted (BSFW).

1971 18-19 Aug. 200 Young noted were from about 3 weeks
old to nearly fledged (BSFW).

15 Sept. 2 Near-fledging chicks seen (BSFW).

1972. 16 Sept. 100 (BSFW).

1973 31 July 150 (BSFW) .

BLUE-FACED BOOBY Sula dactylatra
Status

Common breeder; maximum recent estimate: 350. Probably present
throughout year, but most nesting occurs from February through September
or October. Nests on the ground, primarily in areas of higher elevation.

Populations

Recent estimates (Table 12) suggest a maximal population level of
300 to 400 birds. Wetmore's 1923 estimate is reasonably consistent
with these estimates but Munter's 1915 estimate is inexplicably higher
than recent estimates. Several recent estimates made during summer
(1966, 1971) are lower than would be expected, judging from most of
the estimates made during March, early in the breeding season. The
variability in March and late summer estimates is apparently greater
than for estimates from most other northwestern Hawaiian Islands at
these times of year, and suggests that the population levels on this
island may be more variable from year to year than on other north-
western Hawaiian Islands.

Annual Cycle

Numbers present and numbers breeding are probably considerably
lower in late fall and winter but the absence of numerical estimates
from these months makes it impossible to determine the degree to which
the population varies throughout the year.

Eggs are laid primarily from February through March and at least
a few are laid from April through July. In most years apparently
little laying occurs in late summer or early fall but the presence of
recently fledged young in mid-June 1923 indicates that some egg-laying
took place 5 1/2 to 6 months earlier--November or early December.
Kramer's observations in December 1961 and June 1962 indicate that
some eggs were laid in January or February, and BSFW observations on
21 March 1969 suggest a laying peak about 2 months earlier (late
January). Vanderbilt and de Schauensee's observations similarly
suggest that laying took place very early in 1940.

Peak numbers of young are probably present from early May through
July. Most young evidently fledge from late July through September
with a few fledging in October and November.

Breeding Habitat

All observers who noted the location of Blue-faced Booby nests
agree that most nests were in open areas (Fig. 24) on the ridges
and slopes of the island. On many surveys (1923, 1964, 1965, 1966,
1970, September 1971) the largest concentration of nests or birds

66

Figure 24. Blue-faced Booby at nest on rocky outcroppings. POBSP
photograph, 8 March 1968, by Roger B. Clapp.

was on Miller Plateau. On several surveys (July 1964, July-August
1966, March 1968) a small concentration of nests was located just
above the cliff face on the ridge along the north side of the island
between Miller's and Tanager Peaks.

In March 1968 a more detailed appraisal of the location of the
nests was made. Of the 122 nests counted, 51 (42 percent) were
found on Miller Plateau and the upper west slopes of the island; 12
(10 percent) were found on the lower west portion of the island (the
area around West and West Palm Valleys); 11 (9 percent) were found on
the upper central slopes from the ridge east of Miller Canyon across
Middle Valley to the lower slopes of Tanager Peak; 3 (2 percent)
were found on the upper east portion of the island (the area around
Tanager Peak and the upper portions of East and East Palm Valleys);
and 45 (37 percent) were found on the lower eastern slopes of the
island (the area around East and East Palm Valleys). In summary, 53
percent were on upper slopes and 47 percent were on lower slopes.

Banding

Nine adults were banded in March 1964 by the POBSP (6) and the
BSFW (3), but none has been recaptured. A Blue-faced Booby, banded
(USFWS 568-71825) as a "local" at Trig Island, French Frigate Shoals,
on 9 June 1967 by POBSP personnel, was found dead (age and sex un-
known) at Nihoa on 24 August 1968 (Amerson, 1971: 143, 197, 356).

67

Specimens

Two adults, a male (USNM 300949) and a female (USNM 300948), |
were collected by Wetmore 13 June 1923,

Table 12. Observations of Blue-faced Boobies on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May ? Presence noted from offshore (Munro,
L9ATNas 49).

1902 1-3 June ? Noted as "common" from offshore
(Fisher, 1903: 7/97).

5-9 Aug. ? Common; numerous birds in juvenal
plumage seen from offshore (Fisher,
1903) 797).

1915 18 Mar. 5,000 Eggs present (Munter, 1915: 131-132).
Brown (ms.) gives an identical
alternative estimate.

1916 12 Feb. [apparently not Some found nesting (Munter, ms.).

very common]

1923 5 Apr. ? Occasionally seen offshore (Wetmore,
ms.).
11-16 June 250 Newly-hatched to recently fledged
young observed (Wetmore, ms.).
1936)... Mar. 4 Nesting; eggs noted (Trempe, ms.).
1940, 7-15 Aug. 2 "The breeding season had just been

completed" (Vanderbilt and de
Schauensee, 1941: 10).

1961 2 Mar. ? Seen on the island from offshore
(Woodside and Kramer, ms.).

9-16 Dec. very Birds paired but no nests noted
abundant (Kramer, ms.).
1962 10 June z Large downy young observed (Kramer

and Beardsley, ms.).

1963 5-6 June ? Ca. 20 seen from offshore (POBSP).

Table 12. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1964 6-7 Mar. 100 Ca. 40 active nests found: 90%
with eggs, none with young (BSFW,
POBSP).

25) da ly, ? Nesting (BSFW).

23-24 Sept. 50 14 immatures counted. No nests with
eggs or downy young seen (BSFW, POBSP).

1965 13-14 Mar. 300 Eggs but no young observed. An
estimated 150 nests present (BSFW,
POBSP)).

1966 28 July- 115-140 Sample count of 17 nests: 2 (12%)

1 Aug. with heavily incubated eggs, 15 (88%)
with 10-21 day old.young. 24 im-
matures also counted (BSFW, POBSP).

1967. 8-9 Mar. 2 None found nesting (BSFW, POBSP).

13-14 Sept. very Downy chicks observed (BSFW).
common

1968 7-9 Mar. 350 Only eggs present, most probably fresh
or slightly incubated. 122 nests
counted. Of 77 nests of which the
contents are known, 3 (4%) were empty
but active and 74 (96%) contained eggs.
Ca. 300 nesting birds present (BSFW,
POBSP).

24-27 Aug. 200-250 Most abundant on Miller Plateau. Nests
there contained eggs to nearly fledged
young. Many young fledged (BSFW).

1969 21 Mar. 90* Eggs to flying young. Most nests with
small young (BSFW).
£970 15Aug:. 160 Estimate includes 30 flying young. No

dependent young or eggs seen (BSFW).

Table 12. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1971" 18-19 Aug. 80* Most young seen were full grown (BSFW).
15 Sept. q Most birds were on Miller Plateau
(BSFW).

Loe 16. Sept. 300 At least 75 young were present (BSFW).

mosis July 300 (BSFW).

* Estimate is of the number of nesting birds only.

BROWN BOOBY Sula leucogaster

Status

Common breeder; maximum recent estimate: 225. Present through-
out year with fewer present in winter. Most nesting evidently occurs
from February through August but some may occur outside this period.
Nests on the ground.

Populations

Recent numerical estimates indicate that peak populations comprise
150 to 225 birds (see Table 13). The estimates from September and
December suggest that populations usually decrease during fall and
winter. The only early estimate (1923) does not appear significantly
different from most recent estimates. The March 1967 estimate is
inexplicably small, which may be the result of observer error or
possibly indicates a very late nesting season.

Annual Cycle

On Nihoa this species appears to nest earlier than its congener,
the Blue-faced Booby. Observations of variously-sized young in
March, anda Kramer's observations in December 1961, indicate that
laying may begin as early as November or December. Most eggs, however,
are probably laid in February or March.

69

70

The occurrence of a nest with eggs in June (Wetmore, ms.)
suggests that an occasional nest is started in May or June and a
few nests are probably begun in April. These are possibly renesters.

Observations of the size of young made during the summer suggest
that most hatching occurs from about April through May. Most fledg-
ing probably occurs from late July through August, with a few birds
fledging in September or later; judging from recent observations
fledging is largely completed by the end of August.

Breeding Habitat

Brown Booby nests are widely scattered over the slopes of the
island and usually are found in locations overlooking rather sharp
drops in elevation. Although Brown Booby nests are as restricted
to the slopes of the island as are those of the Blue-faced Boobies,
nests may often be found along the ridge above the cliff face on
the north edge of the island. Some nest sites are possibly on the
cliff face itself, but we lack information in this area. In March
1965 most nests were near Tanager Peak and along the western ridge
from Miller's Peak to Dog's Head Peak.

In March 1968 we attempted to determine what proportion of the
nests was found on Miller Plateau and the upper west slopes of the
island; 14 (18 percent) were found on the lower west slopes of the
island (in the area around Vest and West Palm Valleys); 6 (8 percent)
were found on the upper west slopes (including Miller Plateau); 6
(8 percent) were found on the upper central slopes and the ridges
east of Miller Canyon across Middle Valley to the lower slopes of
Tanager Peak; 31 (40 percent) were found on the upper east portion
of the island (the area around Tanager Peak and the upper slopes of
East and East Palm Valleys; and 20 (26 percent) were found on the
lower eastern slopes of East and East Palm Valleys and the surround-
ing area. In summary, 56 percent were located on upper slopes and
44 percent were on lower slopes.

Banding

The POBSP banded two nestlings, one in March 1964, the other
in July 1966. Neither has been recaptured.

Specimens

Two males were collected by Wetmore: USNM 300878 on 14 June
1923, USNM 300880 the following day.

Table 13. Observations of Brown Boobies on Nihoa Island
Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May B Presence noted from offshore (Munro,
1940a=" 49) .

1902 1-3 June 2? Presence noted from offshore (Fisher,
HES OS 777.9) he

5-9 Aug. Q Numbers of young birds seen from off-
shore (Fisher, 1903: 798).

1923 11-16 June 100 Mostly young, many half-grown, but at
least 1 nest with eggs present (Wet-
more, ms.).

1940 7-15 Aug. u "Breeding was in a late stage, and large
young were seen on the nests" (Vanderbilt
and de schauensee, 1941: 10).

1953. = 21-22" Dec. 8-10 ? (Richardson, pers. comm.).

Si ceeo dU Ly ? Two dozen birds followed ship as it
passed island (Labrecque, 1957: 19).

1961 2 Mar. ? Not noted from offshore (Woodside and
Kramer, ms.).

9-16 Dec. relatively "Females were on eggs" (Kramer, ms.).
rare

1962 10 June 2 Large downy young observed (Kramer and
Beardsley, ms.).

1963 5-6 June ? Ca. 40 seen from offshore (POBSP).

1964 6-7 Mar. 150-200 Ca. 75 nests present, all with eggs
except for 1 with a very large nest-
ling (BSFW, POBSP).

ZO ry ? Several nearly fledged and fledged
young observed (BSFW).
23-24 Sept. 20 4 immature birds seen. No nests with

eggs or downy young observed (BSFW,
POBSE)

72

Table 13. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1965 13-14 Mar. 15.0 Eggs and young present. Most nests con-
tained 2 eggs (BSFW, POBSP).

1966,):23; July— 40-65 Near end of breeding cycle. Cq. 20

1 Aug. immatures on island (BSFW, POBSP).
1967 8-9 Mar. 15 "Nesting'' (BSFW, POBSP).
13-14 Sept. 100 About 50% seen were fledged immatures

(BSFW).

1968 7-9: Mar. 225 Fresh eggs to medium-sized downy young.

24-27 Aug.
O69 eZ leat.
LOT ON Sr Aug:

1971 18-19 Aug.

1972 16 Sept.

LOPS ol July,

minimum of
50

110%

135

80%*

150

ils)

77> nests counted. Of 38 nests whose
contents were checked, 1 (3%) was
empty but active; 32 (84%) contained
eggs; 1 (3%) held an egg and a naked
young; 4 (11%) contained a medium-
downy young. Ca. 200 nesting birds
present (BSFW, POBSP).

No nests with eggs or downy young found.
Flying immatures seen. 14 seen in one

group on Tanager Peak (BSFW).

Eggs to downy young. Some flying im-
matures seen.

Very large and fledged young seen (BSFW).

Young from about a month old to fully-
fledged birds were present (BSFW).

(BSFW).

(BSFW).

* Estimate is of the number of breeding birds only.

RED-FOOTED BOOBY

Status

Sula sula

Common breeder; maximum recent estimate: 3,500. Present through-
out the year but most abundant in spring and summer. Breeding occurs

throughout year but most of the population breeds from February
through October. Builds bulky nests in low bushes or Pritchardta
palms.

Populations

Recent population estimates (Table 14) consistently indicate
maximal populations of 3,000 to 4,000 birds. Surveys made towards the
end of the year suggest a consistent decrease in the population, and
the one available December estimate suggests that populations decrease
markedly, to perhaps one-tenth of the peak.

Wetmore's 1923 estimate is almost twice as large as any recent -
estimate, but the absence of recent numerical estimates from June, a
month when populations may well reach their zenith, precludes any
conclusion that populations are smaller today.

Annual Cycle

Our data clearly indicate that laying usually begins in February,
with the peak occurring in late February or March. A small downy
young about 2 weeks old seen in March 1968 must have come from an egg
laid in January, and observations from March 1969 suggest that fairly
large numbers of birds laid eggs in January that year. Thus, a few
eggs may be laid earlier in the year but the number nesting at this time
must represent only a small proportion of the breeding population.
Some laying also occurs in May and June but laying thereafter is ap-
parently quite variable from year to year. Observations in July and
August 1966 suggest that laying had been largely completed by early
July, but observations in September 1967 and 1971 indicate that some
laying had occurred in July or August. In September 1964 and August 1970,
on the other hand, observations indicate that laying had been completed
by late June.

Young are known to have hatched as early as late February and, if
eggs observed in the fall were fertile, may have hatched as late as
September or October. However, by far the largest proportion of the
young probably hatch from May through June or early July.

Little data are available on fledging but a few young may fledge
as early as June. (If eggs present in September were fresh, fledging
could occur even earlier in the year.) Most probably fledge in August
or September, and a very small proportion fledge in October or November.

Thus, breeding probably occurs in all months, but only a few birds
breed from late September through early February.

Breeding Habitat

On Nihoa Red-footed Boobies nest on the slopes of the island in
small colonies and in widely dispersed individual nests. A large pro-
portion of the nests is found at middle and lower rather than at
higher elevations.

73

74

A nesting concentration has several times been noted on Miller
Plateau (= Albatross Plateau of Venderbilt and de Schauensee, 1941:
10) and in nearby areas. Vanderbilt and de Schauensee noted that they
were most abundant there in August 1940 and Kramer stated that the
main colony was found on the upper slopes of Miller Valley in December
1961. Kridler found a colony of about 100 nests on the Plateau in July
1964, and considerable numbers were found nesting in that general area
in March 1965, July-August 1966, and March 1968. On the latter visit
perhaps half the nests were found in the Miller's Peak-Miller Plateau
area.

Considerable numbers also nest on slopes of the various valleys,
and smaller numbers nest in groves of Pritchardia palms. Most nests
are built from 3 to 4 feet above the ground in low Chenopodtum, Sida
and Sesbanta.

Color Phases

No quantitative data are available on the proportion of the dif-
ferent color phases in the breeding population but observational data
suggest that over 99 percent of the population consists of the white-
plumaged morph (Fig. 25). Dark-plumaged morphs have been seen on three
occasions. Wetmore (ms.) flushed a "bird in grey plumage with a pure
white tail" from a nest with a newly hatched young in June 1923.
Another dark-phase bird was noted in March 1964 and still another,
evidently paired with a white-phase bird, was noted in March 1968
GEiig’s 3 2'6))

Banding

Fifty-one adults and 7 nestlings were banded by the BSFW and POBSP
on recent visits. Fifty nesting adults were banded by the BSFW in
March 1965 and 1 adult and 7 nestlings were banded by the POBSP in
July 1966. None has been recaptured.

Specimens

Three specimens were collected by Wetmore: two males (USNM 300912,
300915) on 12 June 1923, and an embryonic alcoholic (USNM 289297) on 14
June 1923.

Table 14. Observations of Red-footed Boobies on Nihoa

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1891 26-27 May 2 Presence noted from offshore. Several
captured as they roosted on the ship.
An immature seen (Munro, 1941la: 49-50).
1902 1-3 June plentiful Seen sitting on nests from offshore

(Fisher, 1903: 797).

75

Figure 25. White-phase plumage morph of the Red-footed Booby,
8 March 1968. POBSP photograph by Roger B. Clapp.

Figure 26. Dark-phase plumage morph of the Red-footed Booby, 8 March
1968. POBSP photograph by Roger B. Clapp.

231-338 O - 77 - 13

Table 14. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References |

1902 5-9 Aug. ? No adults certainly noted but immatures
seen from offshore (Fisher, 1903: 797).

19d 5 yalS Marc 800 Eggs present (Munter, 1915: 131). Al-

(20,000) ternative estimate by Brown (ms.) seems
much larger than credible.

1916-12 Feb. ? Mating and building nests. Several
fledged young seen (Munter, ms.).

1923 11-16 June 6,000 From recently completed nests through
fresh and incubated eggs to recently-
hatched through half-grown young
(Wetmore, ms.).

1936 3,Mar. 2 Nesting, eggs noted (Trempe, ms.).

1940 7-15 Aug. ? "Breeding seemed to be in all stages"
(Vanderbilt and de Schauensee, 1941: 10).

1953 21-22 Dec. 200-300 ? (Richardson, pers. comm.).

1954 18 Mar. 30-40 22? (Richardson, pers. comm.).

1961 2 Mar. 2 Seen on the island from offshore (Wood-
side and Kramer, ms.).

9-16 Dec. ? "All young were nearly full grown and
capable of flying" (Kramer, ms.).

1962 10 June 2 Young in various growth stages. No
eggs observed (Kramer and Beardsley,
mss).

1963 5-6 June 2 Ca. 80 seen from offshore (POBSP).

1964 6-7 Mar. 2,600 Ca. 1,000 nests counted. Of those whose
contents were checked, all contained eggs,
none young (BSFW, POBSP).

2a ley, 2 Eggs to well feathered young but most
nests with young (BSFW).

23-24 Sept. 900 15 large nestlings and 68 immatures
counted. No nests with eggs or small
young observed (BSFW, POBSP).

77

Table 14. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1965 13-14 Mar. 3,500* Ca. 1,700 nests counted. All investigated
contained eggs. None contained young
(BSFW, POBSP).

1966 28 July- 2,800- Ca. 1,400 nests counted: ca. 3-7% with

1 Aug. 3, 000% incubated eggs, the rest with young. Of
the young about 10% were recently hatched,
40% were about 2-3 weeks old, and 50%
were more than 3 weeks old (BSFW, POBSP).

1967 8-9 Mar. 3,000 Only about 5% of nests with eggs, none
with young (BSFW, POBSP).

13-14 Sept. very Eggs to near fledging young (BSFW).

abundant

1968 7-9 Mar. 3,000 Ca. 1,200 nests counted. Eggs to small
downy young but most nests with contents
had eggs. Sample count of 115 nests: 60
(52%) empty but active; 54 (47%) with
eggs; 1(1%) with a small downy young
(POBSP, BSFW).

24-27 Aug. hundreds Eggs to flying young. Scattered through-
out vegetated slopes of island. Large
colony of 150 nests on Miller Plateau
(BSFW).

1969 21 Mar. 1,050* Most nests contained small young but a
few held eggs (BSFW).

1970 15: Aug. if 500 An estimated 500 young present, ranging
in size from 3/4 grown to fully fledged.
Most young had not yet fledged and no
nests with eggs were seen (BSFW).

1971 18-19 Aug. 1,000 Most nests contained near-fledging young
but a few nests with eggs were also
present (BSFW).

1S Sept. b A few nests contained eggs but most con-
tained young of varying ages (BSFW).

1O7 2° onSepier ea 00 Chicks seen (BSFW).

LOTS 3 suly 2 A minimum of 350 birds was present (BSFW).

* Estimate is for the number of nesting birds.

78

GREAT FRIGATEBIRD Fregata minor
Status

Abundant breeder; maximum recent estimate: 10,000. Present
throughout year but decidedly less abundant in winter. Some breeding
birds present throughout the year but most breeding occurs from Febru-
ary or March through September. Builds bulky nests in low bushes.

Populations

Recent estimates place maximal population levels at about 10,000
(Table 15) but populations may be considerably smaller during late fall
and winter. The only two early numerical estimates are not consistent
with recent observations. Munter's estimate of 50,000 is much higher
than any recent estimate, and Wetmore's estimate of 1,200 is considerably
lower than any recent estimate made when a large proportion of the popu-
lation was breeding. Recent estimates are subject to a degree of error
but do not vary from one another enough to support either Munter or
Wetmore's estimates.

Annual Cycle

Displaying males have been noted from early December to early June
but the earliest that eggs have been known to be laid is about mid-
January. The presence of recently hatched young in early March 1965
indicates that some eggs were laid as early as the second week in
January. However, the egg-laying peak apparently usually occurs from
late February through March with an undetermined number of eggs laid
in April, May and perhaps June. Observations from early June 1923
indicate that some egg-laying must have occurred in April, May and
June, and the presence of young about a week old in late July 1966 (Fig.
27) predicates laying in late May.

Most hatching probably occurs from April through June and the
majority of the young probably fledge from September through October.
An occasional bird may fledge as early as late August, and a few as late
as November. Dependent immatures have been present in mid-March well
into the succeeding nesting season.

Breeding Habitat

Munter (1915: 132) indicated that frigatebirds nested less commonly
at high elevations, but most recent observers have found them common on
most of the upper two-thirds of the island's slopes.

Vanderbilt and de Schauensee (1941: 10) stated that the "colonies
appeared to be limited to localities half-way up most of the valleys."
In March 1964 Kridler noted that nests were most abundant near the top
of East Palm Valley, Middle Valley, and Miller Canyon, and in September
1967 found them most abundant in the former two areas. In March 1965,

YS

1967, and 1968 these frigatebirds nested most densely on the upper
slopes of Miller Valley up to Miller Peak.

On Nihoa these birds nest mostly in many small colonies containing
a dozen to 30 nesting pairs, but are also found nesting in widely scat-
tered individual sites. Most nests are placed in low, thick Chenopodium
and Sida.

Banding

The POBSP banded 1 adult and 490 nestlings in July 1966 but none
has been recaptured.

Specimens

Two specimens, an adult male (USNM 464440) and an adult female
(USNM 464441) were collected on 14 June 1923 by Wetmore.

Table 15. Observations of Great Frigatebirds on Nihoa Island.

Population

Date of Survey Estimate Breeding Status, Remarks, and References

Uses) 22 July 2 Large downy young (Bishop, 1885a: 2).

1891 26-27 May 2 Presence noted from offshore (Munro,
1941a: 49).

1902 1-3 June ? Abundant, seen nesting in bushes from
offshore (Fisher, 1903: 799).

5-9 Aug. ? "Still to be seen in considerable

numbers'' [from offshore] (Fisher, 1903:
VIDE

gS) LS Max. 50,000 Eggs and males with inflated throat
pouches reported (Munter, 1915: 132).
Brown (ms.) gives an identical alterna-
tive estimate.

ROG 2) heb. very Breeding season apparently just begun

common but no eggs or young present (Munter,

ms.)

O23 5) None ? Occasionally seen offshore (Wetmore, ms.).

11-16 June ZOO, Incubating eggs or with young from

recently hatched to a few half-grown.
Only 1 male with inflated throat pouch
seen (Wetmore, ms.).

EGSGiew So) Maltare many Many nests with eggs (Trempe, ms.).

80

Table 15. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1940 7-15 Aug. "Small colon- ''The nests contained almost mature
ies were quite young" (Vanderbilt and de Schauensee,
common" POAT TO):

1953 21-22 Dec. 700-900 ? (Richardson, pers. comm.).

1954 18 Mar. 500-1, 000 ? (Richardson, pers. comm.).

1961 2 Mar. 2 Seen on the island from offshore
(Woodside and Kramer, ms.).

9-16 Dec. very Large flying young but no nests with
abundant eggs seen (Kramer, ms.).

1962 10 June 4 "Half grown downy young on nests. No
males seen in nuptial displays"
(Kramer and Beardsley, ms.).

1963 5-6 June 2 Ca. 20 seen from offshore (POBSP).

1964 6-7 Mar. 10,000 Ca. 4,800 active nests counted, most with
eggs. 10 nests found with recently
hatched young; courting birds observed
(BSFW, POBSP).

250 duly, & Downy to well-feathered young but most
nests with downy young (BSFW).

23-24 Sept. 6,000 167 large young counted. Ca. 1,000
already fledged young present (BSFW,
POBSP).

1965 13-14 Mar. 5,200* Ca. 2,500 nests counted about 70% con-
taining eggs (BSFW, POBSP).

1966 28 July- 4,200- Ca. 2,500 young present, 10% recently

1 Aug. 6,900 hatched, 30% from 2-3 weeks old, and
60% older than 3 weeks old. No nests
with eggs observed (BSFW, POBSP).

1967 8-9 Mar. 9,000 Initiating nesting. Of 31 nests checked,
10 (32%) were empty and 21 (68%) con-
tained eggs, 43% of which were fresh
and 57% were incubated (BSFW, POBSP).

13-14 Sept. very A few small downy young and many im-

abundant matures notes (BSFW).

Table 15. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References
1968 7-9 Mar. 7, 000- Only eggs present. A considerable pro-
8,000 portion of population still in prelaying

stages (30-40% of males with inflated
throat pouches in some areas; 70-80% in
others). Sample count of 55 nests: 26

(47%) empty but active, 29 (53%) with

eggs. Ca. 2,000 nests counted and an

estimated 4,000 nesting birds present

(BSFW, POBSP).

24-27 Aug. ? From half-grown to fledged young (BSFW).

969 21 *Mar : 3,450* Most nesting birds just beginning to in-
cubate eggs (BSFW).

1970 15 Aug. 4,600 More than 1,100 young were counted. Most
were very large and well-feathered but
were not yet fledged (BSFW).

1971 18-19 Aug. ? At least 1,200 present. Most young were
about 3/4 grown (BSFW).

15 Sept. 2 Young were from 1/2 grown to near-
fledging size (BSFW).

Lo72r Lor Sept: 3, 000* An estimated 1,500 young were present
(BSFW).

1973.31, July 2 At least 2,000 present (BSFW).

* Estimate is for the number of breeding birds only.

PINTAIL Anas acuta
Status

Accidental; one record of two birds from September 1971.
Observations

On 15 September 1971 the BSFW survey party saw a female Pintail

near a very small puddle of water about one-third of the way up East
Palm Valley. The rotting carcass of a drake that had probably been

dead for 2 to 3 weeks was found in the same puddle and a wing (USNM
567292) was collected. Pintails have not previously been recorded
from Nihoa but are of fairly regular occurrence on Laysan and Midway
Atolls where much more extensive ponds or lagoons are to be found.
The species has also been recorded from French Frigate Shoals
(Amerson, 1971: 228), Pearl and Hermes Reef (Amerson, Clapp and
Wirtz, 1974: 182) and from Lisianski Island (Kridler, unpub. observ.)
in the northwestern Hawaiian Islands, as well as from Johnston Atoll
(Amerson and Shelton, in press).

GOLDEN PLOVER Pluvtalts domintca

Status

Uncommon but regular migrant; maximum recent estimate: 50.
Recorded in February, March, June, August, September, and December.
Largest numbers occurred in March. .
Observations

All observations are listed in Table 16.

Specimens

Evidently only one specimen has been collected. The Vanderbilt
Expedition obtained a female (PAS 146156) in August 1940.

Table 16. Observations of Golden Plovers on Nihoa Island

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1902 1-3 June a few Seen from offshore (Fisher, 1903: 778).
5-9 Aug. a few Seen from offshore (Fisher, 1903: 779).

1915), 8 Mair. several Seen "on the plateau'' (Munter, 1915:

seen li 3))
1916 12 Feb. quite (Munter, ms.).

common
19237 Vhi=16, June 8) Seen on rocks of Adams Bay 11 June

(Wetmore, ms.).

1940 7-15 Aug. a few "not present in any numbers." Number

of dead birds found (Vanderbilt and
de Schauensee, 1941: 10).

1953 21-22 Dec. 0 (Richardson, pers. comm.).

83

Table 16. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References
1954 18 Mar. 0 (Richardson, pers. comm.).
1961 9-16 Dec. iW (Kramer, ms.).
1963 5-6 June itt Seen from offshore (POBSP).
1964 6-7 Mar. 75) (BSFW, POBSP).

23-24 Sept. 5 (BSFW, POBSP).
1965 13-14 Mar. 50 5 seen near Dog Head Peak (BSFW, POBSP).
1966 28 July- 0 (BSFW, POBSP).

1 Aug.

1967 8-9 Mar. 5) (BSFW, POBSP).

13-14 Sept. 20 (BSFW) .
1968 7-9 Mar. 50 Seen in small flocks but more often as

individuals. Scattered all over island
(BSFW, POBSP).

1968 24-27 Aug. 10-15 Individuals scattered over island (BSFW).
L969) 2ihiMar’. 36 (BSFW).
COTO Se Au eS 2 (BSFW).
1971 18-19 Aug. 0 (BSFW).

15 Sept. ? Rare (BSFW).
Lo 2) OmSept. iL (BSFW).
BRISTLE-THIGHED CURLEW Numentus tahittensis

Status

Uncommon migrant; five sight records; maximum recent estimate:
5. Recorded in March, August and September.

Observations

Vanderbilt and de Schauensee (1941: 10) reported a small flock of
curlews seen "Shortly after landing on Nihoa,' presumably on 7 August

84

1940. There are four more recent records. On 13 and 14 September
1967 Kridler saw a single Bristle-thighed Curlew. Two others were
seen on rocks about 150 to 200 feet from the base of Miller Canyon

7 to 9 March 1968 (Fig. 28). Another was seen 18-19 August 1971

and five were present on 16 September 1972.

WANDERING TATTLER Heteroscelus tneanus
Status

Uncommon migrant; maximum recent estimate: 2. Recorded in
March, May, June, and August.

Observations
All observations are listed in Table 17.

Specimens

A single specimen was collected by Vanderbilt but we do not know
the specific collection data.

Table 17. Observations of Wandering Tattlers on Nihoa Island

Population

Date of Survey Estimate Observations, Remarks, and References
1923 24 May 2-3 (Wetmore, ms.).

11-16 June 1-2 Seen daily on rock ledges of Adams

Bay (Wetmore, ms.).

1940 7-15 Aug. very few (Vanderbilt and de Schauensee, 1941: 11).
1953 21-22 Dec. al (Richardson, pers. comm.).
1954 18 Mar. 0 (Richardson, pers. comm.).
1961 9-16 Dec. 0 (Kramer, ms’. ).
1964 6-7 Mar. 0 (BSFW, POBSP).

23-24 Sept. 0 (BSFW, POBSP).
1965 13-14 Mar. 0 (BSFW, POBSP).
1966 28; duly= 0 (BSFW, POBSP).

1 Aug.

1967 8-9 Mar. Z- Along ledges on south side of island

(BSFW).

13-14 Sept. 0 (BSFW).

85

%
~ > Spee e-
<*,: Si 3h
Se RW on,

Figure 27. Nesting Great Frigatebird with downy young in July 1966.
BSFW photograph by Eugene Kridler.

Figure 28. Bristle-thighed Curlew foraging among the rocks in lower
Miller Valley, 8 March 1968. POBSP photograph by Roger
B. Clapp.

86

Table 17. (Continued)

Population

Date of Survey Estimate Observations, Remarks, and References
1968 7-9 Mar. 0 (BSFW, POBSP).

24-27 Aug. 0 (BSFW).
1969 — 20Mar. 0 (BSFW).
1970 15@Auer il (BSFW).
1971 18-19 Aug. i (BSFW).

15 Sept. 0 (BSFW).
1972 Ver Sept. 0) (BSFW) .
RUDDY TURNSTONE Arenarta tnterpres

Status

Common migrant, present in small numbers; maximum recent estimate:
200. Recorded in March, May, June, August, September, and December.
Largest numbers occurred in spring (March) and fall (September).
Observations

All observations are listed in Table 18.
Specimens

We know of but one specimen, a female (PAS 146157), collected by
the Vanderbilt Expedition in August 1940.

Table 18. Observations of Ruddy Turnstones on Nihoa Island

Population
Date of Survey Estimate Observations, Remarks, and References
1891 26-27 May 2 2 small flocks seen from offshore
(Munro, .1941a: 50)’;

1902 1-3 June ? Seen from offshore (Fisher, 1903: 778).

4-9 Aug. 2 Seen from offshore (Fisher, 1903: 779).
1916 12 Feb. a few seen (Munters ms..)'.
1923 24 May 2 Seen on ledges below western cliffs

(Wetmore, ms.).

87

Table 18. (Continued)

Population

Date of Survey Estimate Observations, Remarks, and References

1923 11-16 June ? "One or two seen daily on the rocks of
Adams Bay. Two seen resting on a cairn
on Millers Peak 900 feet above the sea"
(Wetmore, ms.).

1940 7-15 Aug. "Probably... ‘Observed in small numbers everywhere,

most common even on the highest peaks." Seen feed-
shorebird." ing on beetles as well as marine life
(Vanderbilt and de Schauensee, 1941: 11).

1953 21-22 Dec. a few (Richardson, pers. comm.).

1954 6-7 Mar. ik (Richardson, pers. comm.).

1961 9-16 Dec. about 5 (Kramer, ms.).

seen

1963 5-6 June i Seen from offshore (POBSP).

1964 6-7 Mar. 10 Seen in gulches and sea cliffs (BSFW,
POBSP).

23-24 Sept. 30-75 (BSFW, POBSP).

1965 13-14 Mar. 100 (BSFW, POBSP).

1966 28 July- 0 (BSFW, POBSP).

1 Aug.

1967 8-9 Mar. Ys Flocks of 17, 23, and 32 flying over
water on south side of island (BSFW,
POBSP).

13-14 Sept. 75 Scattered small flocks from sea level
to the top of the ridge between the
peaks (BSFW).

1968 7-9 Mar. 200 Primarily in small flocks or scattered
individuals in the lower areas of the
island. Common along the rocks at the
edge of the surf where a flock of about
28 birds seen bathing. Small flocks of
10-20 birds seen on the crests of some
lower ridges (BSFW, POBSP).

24-27 Aug. 30 Scattered. Most on rock shelf at sea

level (BSFW).

88

Table 18. (Continued)

Population
Date of Survey Estimate Observations, Remarks, and References
969 Pe 2s Marry 82 (BSFW) .
1970 lS Auor 8 One seen near the top of Miller's Peak
(BSFW) .
1971 18-19 Aug. 8 (BSFW).
15 Sept. ? Rare (BSFW).
1972 :\16 Sept. 2 (BSFW) .
HERRING GULL ; Larus argentatus

Status

Vagrant; one sight record in March 1965.

Observations

Kridler observed an adult near Derby's Landing on 24 March 1965.
Subsequently, while attempting to photograph the bird, Kridler got close
enough to make a positive identification. Herring Gulls are one of the
several species of gull that occur fairly frequently in the northwestern
Hawaiian Islands (Clapp and Woodward, 1968: 26; Sibley and McFarlane,
1968: 314-318). None has been reported previously from Nihoa.

GRAY-BACKED TERN Sterna lunata

Status

Abundant breeder; maximum recent estimate: 10,000. Present from
February or earlier through September or October; probably absent during
remainder of year. Breeds from at least March through September. Nests
on the ground, usually in areas of sparse vegetation.

Populations

Recent population estimates and those from 1915 and 1923 agree
reasonably well (Table 19) and suggest that maximal populations are
about 10,000 birds. Estimates from various March visits are variable,
but this may reflect differences in the timing of the breeding cycle
from year to year as much as it does any inaccuracies in the estimates
themselves.

Annual Cycle

The paucity of observations from January and February makes it
impossible to tell when the first birds arrive at the island but the
numbers and stage of breeding observed on various March surveys make
it obvious that these terns usually arrive at least as early as
February.

The initiation of laying apparently varies from year to year.
Observations from March 1915, 1964, 1967, and 1968 indicate that the
population was just beginning to breed and would probably reach an
egg-laying peak later in the month or in April. In March 1965,
however, at least several thousand eggs were present by mid-March,
suggesting that some laying occurred in February and that the egg
laying peak was earlier than on the other March visits. Laying may
also occur in May or June but it is not known whether the Gray-backed
Terns usually lay in large numbers during these months. Wetmore's
1923 observations indicate a laying peak in early June but the presence
of half-grown young at that time shows that at least some eggs had been
laid in April.

Hatching may occur from late March or early April through late
July but most young probably hatch from about May through early June.
The presence of slightly incubated eggs in early March suggests that
some young could fledge by early or mid-June but the earliest report
of flying young is early August.

Despite the lack of positive observations, it seems likely that
most fledging occurs from mid-July through August. Small numbers of
young are present on the island in September but by the end of the month
almost all young have fledged and most adults have left the island.

Breeding Habitat

Gray-backed Terns are found in much the same areas as are Sooty
Terns, but apparently prefer less densely vegetated areas and are much
more prone to nest on rock ledges and ridges. Colonies were widely
distributed over the slopes of the island (March 1965, July-August 1966,
March 1967, 1968), but seem to reach maximum nesting densities in the
various canyons and gulches (July-August, 1966; March 1968). On the
latter visit they were apparently most abundant in the lower portions
of Miller Canyon, Middle Valley, and the lower eastern slopes of the
island. Vanderbilt and de Schauensee reported that in August 1940 they
occurred in two small colonies, both of them in the upper portion of
Middle Valley.

Banding

The BSFW and POBSP banded 119 Gray-backed Terns on recent visits.
The BSFW banded 10 adults in March 1964 and 79 adults in March 1965.
The POBSP banded 6 adults, 13 flying young, and 11 nestlings in July
and August 1966. None has been recaptured.

89

90

Specimens

We know of four specimens from Nihoa: Wetmore in 1923 collected
an adult male (USNM 300628) on 15 June, a female (USNM 300629) on
12 June, and an embryonic alcoholic (USNM 289312) on 13 June; another
male (PAS 145155) was collected by the Vanderbilt Expedition in August

1940.

Table 19. Observations of Gray-backed Terns on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May 2 Presence noted from offshore (as "Bridled
Tern") (Munro, 194la: 49).

1902 1-3 June & Common; seen from offshore (Fisher, 1903:
781).

5-9 Aug. i Common; seen from offshore. Numbers of
flying young seen (Fisher, 1903: 781).

1915: Se Mar. 10,000 ",..seen in scattered pairs....Two only
were flushed from eggs" (Munter, 1915:
133). Brown (ms.) gives an identical
alternative estimate.

1916 12 Feb. not very No nests found (Munter, ms.).

numerous
1923 5 Apr. ? Occasionally seen offshore (Wetmore, ms.).
11-16 June 10,000 Majority with fresh eggs but some re-
cently hatched young and a few half-
grown young seen (Wetmore, mS.) .

1936 3a Marr ? No nests found (Trempe, ms.).

1940 7-15 Aug. ? Found in two small colonies. 'Breeding
was in its last stages and only young
were seen'' (Vanderbilt and de Schauensee,
LO Gslee de) ye

1953 21-22 Dec. 0 Only part of island surveyed (Rich-
ardson, pers. comm.).

1954 -18°Mar. 12-14 Only part of island surveyed (Rich-
ardson, pers. comm.).

1961 2e Mar. ? Abundant, seen from offshore (Woodside

and Kramer, ms.).

Table 19. (Continued)
Population
Date of Survey Estimate
1961 9-16 Dec. 0
1962 10 June ?
1963 5-6 June 2
1964 6-7 Mar. 100
25 July very
common
23-24 Sept. 50
1965 13-14 Mar. 5 ,000-
6,500
1966 28 July- 8,000*
1 Aug.
1967 8-9 Mar. 10,000
13-14 Sept. very
common
1968 7-9 Mar. 2,000
24-27 Aug. low
thousands
< 3,000
1969" 21° Mar’. 350
USTO) a5 Aug. 6,009
971 18-19 Aug. 7,000
15 Sept. b

231-338 O- 77-14

OF

Breeding Status, Remarks, and References
(Kramer, ms.).

With eggs and downy young (Kramer and
Beardsley, ms.).

Ca. 40 seen from offshore (POBSP).

Ca. 25 nests with eggs observed (BSFW,
POBSP).

Very small chicks to 2/3-grown young
(BSFW).

6 immatures remaining on the island
from the preceding season (BSFW, POBSP).

Nests contained only eggs. An estimated
2,500 nests present (BSFW, POBSP).

Ca. 4,000 young; 5% recently hatched,
25% from 2-3 weeks old, and 70% older
than 3 weeks (BSFW, POBSP).

On fresh to slightly incubated eggs
(BSFW, POBSP).

Some near fledging young noted (BSFW).
Only 1 nest with egg found. Most birds
evidently pre-nesting (BSFW, POBSP).

A very few 3/4 feathered young but most
young flying (BSFW).

2? (BSFW).

Young ranged in size from about 1/4
grown to fledging. The great majority

of birds was near fledging (BSFW).

Young were from about 1/2 grown to
fledging (BSFW).

Small numbers of near-fledging young
were present (BSFW).

92

Table 19. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1972 16 Sept. 4 ,000* Young were present (BSFW).
Lise, 3) July f At least 1,500 birds were present (BSFW).

* Estimate is of the number of breeding birds only.

SOOTY TERN Sterna fuscata

Status

Abundant breeder; maximum recent estimate: 100,000. Usually
present from at least mid-February through late September or October
but breeds primarily from late February through July or August;
probably absent during much of remainder of year. Nests on the
ground (Fig. 29), often in areas of dense vegetation.

Populations

Recent population estimates fairly consistently suggest popula-
tion levels in the low tens of thousands, and do not indicate any
difference from estimates made earlier in the 20th century (Table 20).
The estimate from March 1965 is considerably higher than any other but
it is based on better data. On that visit the portion of the population
breeding near Miller's Peak was censused by making a nest density count
and applying the density figure to the approximate area covered by nesting
birds. (No details on methods employed are available.) That area was
estimated to contain about 27,000 breeding birds. This figure, repre-
senting only about one-fourth of the total breeding population, is still
higher than any other population estimate that has been made on Nihoa.
We believe that this datum indicates that many of the other population
estimates may have been too low and that total breeding populations
are usually 50,000 to 100,000 birds. (Date on the reliability of visual
estimates of Sooty Terns, obtained on other islands by the POBSP, sug-
gest that populations of this species are more often underestimated
than overestimated.)

Annual Cycle

The initiation of the breeding season may vary by several months
from year to year. In at least three years (1915, 1967, 1969) laying
evidently began in late February or early March. In 1964, however,
the presence of week-old chicks in early March indicates that laying
began as early as late January or early February. The presence of young
(of unspecified size) in mid-March 1965 and Munter's observations in

93

February 1916 indicate laying occurred by mid-February in those years.
The size of young seen in March 1968 (Table 19) indicates that some
egg-laying must have begun by the second or third week of December

and continued until the first or second week of January. On the latter
visit the distinct hiatus between stages of breeding in different
colonies suggested that no eggs were laid in late January or early
February. In 1964, 1966, and 1967 only a very small porportion of the
breeding population laid before late February, however. Thus, data from
five years indicate that the majority of the breeding population begins
to lay in late February and early March...

The winter breeding in 1967-1968 probably was an exceptional occur-
rence as winter breeding populations have not been reported from any of
the other northwestern Hawaiian Islands. However, a winter breeding
population was reported from Moku Manu in the main Hawaiian Islands
(Richardson, 1957<% 24).

Some laying occurs in April and May and fresh eggs were reported
in June (Wetmore, ms.). Observations made in mid-August 1940 indicate
that some eggs were laid as late as mid-July. On Nihoa Sooty Terns
have thus exhibited a laying span of about seven months (mid-December
to mid-July).

Hatching has occurred from at least January through late August
but probably occurs primarily from late March through May. Fledging
May occur as early as early March but in most years most fledging
probably occurs from late May through July. Fledging possibly occurs
as late as October (1940) but is evidently usually completed by August
or mid-September.

The absence of breeding birds in late July and early August 1966
is puzzling and may indicate that the population did not breed or that
early breeding attempts were entirely unsuccessful. It is also pos-
sible that laying began as early as in 1967-1968 and that the breeding
season had been completed prior to the survey.

Also puzzling is the paucity of birds in March 1954. Richardson's
observations suggest that not only had nesting not begun but that the
birds had not begun to return to the island. Clearly, much may yet be
discovered about the nature and variability of the Sooty Tern breeding
cycle on Nihoa.

Breeding Habitat

Sooty Terns nest in a wide variety of situations on Nihoa but a
number of observers noted that they were more common at higher elevations.
Munter (1915: 132) found the birds "in ever increasing numbers" as he
ascended the slopes. Wetmore (ms.) found them nesting "from the lower
rock cliffs clear to the higher summits.'"' He also noted that they
occurred in a number of small colonies on the lower slopes but formed
larger colonies on the highest slopes. The two colonies reported by

94

Vanderbilt and de Schauensee (1941: 11) were also found at higher
elevations, the larger colony near the top of Middle Valley, the
smaller high up in East Palm Valley.

Recent observations confirm that these terns tend to nest in a
number of colonies and in larger numbers at higher elevations. In
March 1964 BSFW personnel found large colonies on the east slopes of
Middle Valley and on Miller Plateau and noted that birds were found
nesting elsewhere as well. In July 1964 a large colony was found along
the top of the ridge between Miller's and Tanager Peaks, and in March
1965 birds nested there, on Miller Plateau, and on the east slope
below Tanager Peak. In March 1968 these birds nested on the slopes
at the base of Miller Canyon, near the top of Middle Valley, on the
slopes of East Palm Valley, on Miller Plateau, and in a number of other
areas. In August 1970 the last few nesting birds were present on the
upper slope of the saddle between Miller's and Tanager Peaks.

Most eggs are laid in small depressions on bare ground but are
occasionally found on small amounts of dead vegetation. Typically
nests are placed beside tufts of grass or under dense vegetation,
such as Chenopodtum, Stda, or Solanum.

Banding

Two adults were banded by the BSFW in March 1964, Neither has
been recaptured.

Specimens

We have found records of five specimens from Nihoa. Four, two
adult males (USNM 300548, 300550), an adult female (USNM 300549) and
a juvenile female (USNM 300551), were collected by Wetmore between 12
and 15 June 1923. A female (PAS 146154) was collected by the Vanderbilt
Expedition in August 1940.

Table 20. Observations of Sooty Terns on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May ? Presence noted from offshore (Munro,
1941a: 49).

1902 1-3 June ? Abundant, seen from offshore (Fisher,
1903i 2F780) 2

5-9 Aug. ? Abundant, seen from offshore. Many

flying young seen (Fisher, 1903: 780).

1915) Si Mars 20,000 Only eggs found (Munter, 1915). Alter-

(10,000) native estimate by Brown (ms.).

95

Table 20. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
LOrG) 12 Feb. in large Only eggs found (Munter, ms.).

numbers

1923 11-16 June 12,000 Majority of nests with fresh eggs but
fledged young present in some parts of
colony (Wetmore, ms.).

9S 6a oS) Mar. Z No nests found (Trempe, ms.).

1940 7-15 Aug. 2 2 colonies, 1 large, 1 small, present.
"Nesting was in all stages from un-
hatched eggs to fully fledged young"
(Vanderbilt and de Schauensee, 1941:
iM ys

LO53e21—22 Dec. 15-20 ? (Richardson, pers. comm.).

1954 18 Mar. 6-8 ? (Richarcson, pers. comm,).

1961 2 Mar. ? Abundant, seen from offshore (Woodside
and Kramer, ms.).

9-16 Dec. 2 Seen soaring over cliffs (Kramer, ms.).

1962 10 June ? With eggs and fledged young (Kramer and
Beardsley, ms.).

1963 5-6 June ? Hundreds seen from offshore (POBSP).

1964 6-7 Mar. 10,000- Ca. 6,000 nests with eggs, ca. 100

15,000 week-old chicks found (BSFW, POBSP).
25) July, very Downy chicks to fledged young (BSFW).
common
23-24 Sept. i Heard flying overhead. Not breeding
(BSFW, POBSO).

1965 iS —il4 | Mar. 100,000 Ca. 90% with eggs, 10% with young
(BSFW, POBSP).

1966 28 July- 500 Not breeding. 2 groups of birds ob-

eA on. served on ground (BSFW, POBSP).
1967 8-9 Mar. 25,000 On eggs, no young found (BSFW, POBSP).
13-14 Sept. U Only a few adults and flying young

noted (BSFW).

Table:20. (Continwed))

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1968 7-9 Mar. ca. 20,000 Ca. 2,000 to 4,000 nests with fresh to
slightly incubated eggs. 2 small colonies
with large young also present. 1 contained
several hundred young from 1/2 grown (ea.
4 weeks old) to near fledging young (ea.
6-7 weeks old). The other contained about
50 young (ca. 4-5 weeks old). The ma-
jority present was not yet nesting
(BSFW, POBSP).

24-27 Aug. ? ? (BSFW).

LOGO 20 Many. 6,800 Most incubating eggs; no young seen
(BSFW).

1O7Oe 5) Awe. 2, 000- A few very large young were still present

3,000 (BSFW) .

1971 18-19 Aug. 1,000 Young were fully feathered (BSFW).

L972 16" Sept. 20 No breeding birds were found (BSFW).

1.973% 330s July 3,000 (BSFW).

BLUE-GRAY NODDY Procelsterna cerulea

Status

Common breeder; maximum recent estimate: 2,500. Present through-
out year but evidently more common in spring and summer. Breeds
throughout year but majority of birds apparently breeds during spring
and early summer. Lays single egg in holes and niches under ledges in
cliff faces and rock outcroppings (Fig. 30).

Populations

Most recent estimates and the one made by Wetmore in 1923 suggest
that the maximal population level is in the low thousands (Table 21).
Considerable variability is found in estimates made at the same time of
year but this probably only indicates the low level of reliability of the
estimates.

Annual Cycle

Too few detailed observations are available to document completely
the breeding cycle; available observations indicate an extended breeding

;
.

i

wie se

Figure 29. Sooty Tern incubating egg in lower part of Middle Valley,
9 March 1968. POBSP photograph by Roger B. Clapp.

— — . ‘3 f
oe <a : 1 antl “S a? Jor

a

Figure 30. Blue-gray Noddy chick at nest site in niche in rock wall,
9 March 1968. POBSP photograph by Roger B. Clapp.

98

season from at least December through October or November in some
years (1964, perhaps 1940). In other years (1967, 1968, 1971, 1972)
observations from late summer and early fall indicate that nesting
had probably been completed by August or September. In March 1964,
1965, and 1968 an estimated 50, 90, and 88 percent, respectively, of
the nests contained eggs which suggests a laying peak then or in Feb-
ruary. Observations made in December 1953, and reports of variously
sized young on a number of March visits (1964, 1965, 1967, 1968),
indicate that some laying normally occurs as early as December or
January.

Most estimates from late summer and fall (mid-August 1970 and
1971; September 1964, 1967, 1971, 1972) indicate a population decrease
at that time which probably means a decrease in the number of breeding
birds. More detailed sample nest counts are needed from most seasons
to better document the laying peaks and the degree to which breeding
activity may diminish during the latter part of the year.

Breeding Habitat

Blue-gray Noddies, like White Terns, nest in considerable abundance
on the north rock cliffs of the island but unlike that species nest
commonly in various areas on the south slopes of the island. Nests are
particularly abundant in niches and cavities in the rock outcroppings
just above the shoreline, but many nests are also present on the
rock faces and under ledges along the sides of the valleys and in rock
outcroppings near the tops of the ridges.

Banding

The BSFW banded 32 Blue-gray Noddies on recent visits: 6 adults
in March 1964, 5 adults in March 1965, 2 adults in July 1966, 17 adults
and 1 local in March 1968,and 1 adult in August 1968. None has been
recaptured.

Specimens

More Blue-gray Noddies have been collected on Nihoa than almost
all other species combined--at least 42 specimens. Of these, 33 are
located in the National Museum of Natural History, 2 in the Philadelphia
Academy of Science, and 7 in the Bernice P. Bishop Museum. A list of
these specimens is presented in Table 22.

Table 21. Observations of Blue-gray Noddies on Nihoa Island

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1891 26-27 May ? Presence noted from offshore (Munro,
1941a: 49).
1903 1-3 June 2 Abundant; seen from offshore (Fisher,

1903:,- 781).

99

Table 21. (Continued)
Population

Date of Survey Estimate Breeding Status, Remarks, and References

1902 5-9 Aug. ? Abundant; seen from offshore (Fisher,
903: 779).

PONS 118 Mar’. "Several" 2 young birds found (Munter, 1915).

196) 12 Feb. 5-6 seen A nest with an egg found (Munter, ms.).

1923 5 Apr. 2 Occasionally seen offshore (Wetmore,
NGS

24-26 May 100 noted Associated in pairs (Wetmore, ms.).
11-16 June 1,600 From slightly incubated to heavily in-

cubated eggs and from recently hatched
to fledged young (Wetmore, ms.).

19356: o- 3°Mar. ? Presence noted (Trempe, ms.).

1940 7-15 Aug. rather "Nesting" (Vanderbilt and de Schauensee,

rare TOA. 11).

95a) 2-22 Dec: 150-200 "Nesting and laying were apparently just
starting'' (Richardson, 1957, pers. comm.).

1954 18 Mar. 100-150 ? (Richardson, pers. comm.).

1961 2 Mar. q Abundant; seen from offshore (Woodside
and Kramer, ms.).

9-16 Dec. not too ? "Paired" (Kramer, ms.).
common

1962 10 June abundant "Breeding...one dead chick seen"
(Kramer and Beardsley, ms.).

1963 5-6 June 2 Thousands seen from offshore (POBSP).
1964 6-7 Mar. thousands From eggs to fledged young found (BSFW,
POBSP).

25) July fairly ? (BSFW).
common
23-24 Sept. 150-300 Eggs to fledged young (BSFW, POBSP).
1965 13-14 Mar. 500-600 Ca. 250 nests, 5% with young (BSFW,

POBSP).

100

Table 21. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1966 28 July- 2,500 2) (BSEW,. POBSP).
1 Aug.
1967 8-9 Mar. several From eggs to 1/2 grown young (BSFW,

thousand POBSP).

13-14 Sept. very common No eggs or young found (BSFW).
along cliffs

1968 7-9 Mar. 2,000 Fresh eggs to near fledging young.
Sample count of 34 nests: 30 (88%) with
eges; 3 (9%) with small downy chicks;
and 1 (3%) with a near-fledging young
(BSFW, POBSP).

24-27 Aug. high A special effort made to find nests but
hundreds no nesting activity noted (BSFW).
<1, 000
1969 21 Mar. 85 2? (BSFW).
1970 15 Aug. 75-100 No nests were found (BSFW).
1971 18-19 Aug. 20 (BSFW).
15 Sept. 0? None was seen (BSFW).
1972; 16 Sept. 50 (BSFW).
LO Se ole Udy, 150 (BSFW).

Table 22. Blue-gray Noddy Specimens from Nihoa Island.

Gh

Museum Museum and Museum Date
Museum Males Nos. Females Nos. yg. Nos. Collected Collector
USNM 13) 300383, 6 300384- 3 300451- 12-15 June Wetmore
434,436, 386, 453 1923
437,440, 438,442
441,443-
450
USNM i]! 289216%* 2 289214- 12-15 June Wetmore
215% 1923
USNM 7 289305- 11,15 June Wetmore
308** 1923

323-325%%**

101

Table 22. (Continued)

29
Museum Museum and Museum Date
Museum Males Nos. Females Nos. yg. Nos. Collected Collector
PAS 2 146162, Aug. 1940 Vanderbilt
164
BPBM 7 4850- Be RY
Sa)
7846-
850
USNM 1 4932757 6 June POBSP
1963
* Skeletons.
** Embryonic alcoholics.
*&KK Older alcoholics.
es Shot from offshore.
BROWN NODDY Anous stoltdus

Status

Abundant breeder; maximum recent estimate: 20,000. Present
throughout the year but most numerous in summer and fall. Breeds
throughout the year but in greater numbers from spring through early
fall. Builds nest on the ground in both vegetated and open areas
(asters | SUD

Populations

The general trend of estimates (Table 23) shows that birds number
at least in the low thousands in summer and fall and that considerably
fewer birds are present in March.

The early estimates (1915 and 1923) do not appear to be signifi-
cantly different from recent estimates, although Wetmore's estimate of
4,000 for June 1923 is a little lower than we would expect, and Brown's
estimate for March 1915 is probably erroneously large. The population
estimate for July and August 1966 seems particularly large compared with
other numerical estimates, but no other estimates have been made at
that time of year.

102

Figure 31. Brown Noddy at nest with egg, March 1967. BSFW photograph
by Eugene Kridler.

Annual Cycle

This species shows a less clearly defined breeding cycle than
that of any other species breeding in the northwestern Hawaiian Islands.
On Nihoa eggs have been laid in all months from December through August
and some may have been laid in the other months as well. The more quan-
titative data suggest that the numbers of laying birds increase con-
siderably during March and several succeeding months, with little laying
occurring after mid-August. Observations indicate periods of peak laying
as: 1923-June, 1964-July, 1966-June or July, and 1967-late July or
early August.

Greatest numbers probably hatch from late June through mid- or
late August, and largest numbers probably fledge from late August
through October. Judging from data gathered on other northwestern
Hawaiian Islands, probably relatively little breeding occurs from
November through February.

103

Breeding Habitat

Brown Noddy nests are widely dispersed over the slopes of the
island but tend to be most numerous in ravines. Nests consist pri-
marily of loose aggregations of sticks, weed stems, and straws, and
occasionally contain feathers, bones, or pebbles. All nests whose
actual locations were reported were found on the ground but the sites
varied considerably. Many of the nests were under thick shrubs such
as Chenopodium, and others were on rock outcroppings, ledges, and in
shallow holes in the cliffs.

Banding

In all, 37 Brown Noddies were banded on recent visits to Nihoa:
3 adults by the BSFW in March 1964 and 34 large young by the POBSP in
September 1964. None has been recaptured.

Specimens

We have found records of six specimens. Wetmore collected two
adult females (USNM 300499-500) on 12 June 1923 and an embryonic alco-
holic (USNM 289399) on 14 June. In August 1940 the Vanderbilt Expedition
collected two others, a male (PAS 146159) and a doubtfully sexed female
(PAS 146158). A third specimen from 1940 is listed by Vanderbilt and
de Schauensee (1941: 12) but we do not know its present disposition.

Table 23. Observations of Brown Noddies on Nihoa Island

Population
Date of Surve Estimate Breeding Status, Remarks, and References
1891 26-27 May ? Presence noted from offshore (Munro,
1941a: 49).
1902 1-3 June ? Seen flying offshore and on island
(Fisher, 1903: 778-783).
5-9 Aug. ? Seen flying offshore and on island
(Fisher, 1903: 779-783).
UV ao sMar. 500 2 no nests noted (Munter, 19152 133).
(5,000) Alternative estimate by Brown (ms.).
Ole, 12) Feb: very Fresh eggs found. Many young from
numerous preceding breeding season seen (Munter,

TSiey)

104

Table 23. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1923 11-16 June 4,000 Mostly with fresh or partly incubated
eggs (Wetmore, ms.).
1936 3) Mari. ? No nests found (Trempe, ms.).
1940 7-15 Aug. extremely "Breeding was in all stages" (Vander-

plentiful bilt and de Schauensee, 1941: 12).

1953. 21-22 Dec. 200-250 "a dedinite breeding season had started"
(Richardson, 1957 and pers. comm.).
19540 S18 Man's 200-300 ? (Richardson, pers. comm.).
1961 9-16 Dec. t Eggs and downy chicks (Kramer, ms.).
1962 10 June ? From eggs to fledged young (Kramer and
Beardsley, ms.).
1963 5-6 June 2 Thousands seen from offshore (POBSP).
1964 6-7 Mar. 600 1 nest with eggs and 2 with young found.
Courtship behavior observed (BSFW, POBSP).
2 ally, very Eggs to near-fledging young but ea.
abundant 99% of nests with eggs (BSFW).
23-24 Sept. 7 ,000% Ca. 3,500 young, mostly large chicks or

flying immatures. A few nests with eggs
or smaller young (BSFW, POBSP).

1965 13-14 Mar. 1,000 90% with eggs, 10% with young (BSFW,
POBSP).
1966 28 July- 20, 000* Ca. 10,000 nests, 90% with eggs, 10%
1 Aug. with young. Ca. 75% of young recently

hatched, 25% from 2-3 weeks old (BSFW,
POBSP).

105

Mable 23. (Continued)
Population
Date of Surve Estimate Breeding Status, Remarks, and References
1967 8-9 Mar. several From eggs to 3/4 grown young. Most
thousand chicks from 1/3 to 3/4 grown (BSFW,
POBSP).
13-14 Sept. common Eggs to flying young but eq. 75-80%
of nests with small downy chicks (BSFW).

1968 7-9 Mar. 1,000 Fresh eggs to young about 3-4 weeks
old. Most of population not nesting
(BSFW, POBSP).

24-27 Aug. low Eggs to flying young (BSFW).
thousands
€5,000

1969 21 Mar. 2,000 Eggs to flying young (BSFW).

1970 15 Aug. 5,000 In a sample of 100 nests, 84 contained
eges, 14 had small downy chicks, and 2
had large near-fledging young. Recently
fledged young were also seen (BSFW).

1971 18-19 Aug. 5,000 From eggs to fledged young (BSFW).

15 Sept. thousands (BSFW).
inon2.” 16 Sept . 15,000 (BSFW).
aaa, Sil uly >» 4,000 (BSFW).

* Estimate is of number of breeding birds only.

BLACK NODDY

Status

Poorly known.
changes in population level unknown.
thousand or low thousands.

Anous tenutrostris

Birds probably present throughout the year but

Maximum recent estimate: several

Only once found breeding (August 1940) but

almost certainly breeds on Nihoa every year.

106

Populations

Virtually nothing is known of the population size since these
birds occur primarily on the nearly inaccessible north cliff face,
and to a lesser extent on the eastern and western cliff faces. Since
the north cliff face cannot be carefully observed, differences in
population estimates from survey to survey are nearly meaningless.
Estimates (Table 24) indicate that this species is fairly common during
the spring.

Annual Cycle

Vanderbilt and de Schauensee (1941: 12) were the only observers
who found nests of this species. They noted that this species "nested
particularly in Middle Valley." The validity of this observation seems
questionabie since no other observer has found them nesting in this area.
They also noted that there were "no distinct colonies and the bird
seemed to mix freely with the...[Brown] Noddy."

Table 24. Observations of Black Noddies on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1891 26-27 May ? Presence noted from offshore (Munro,
1941a: 49).

1902 1-3 June 3 Presence noted from offshore (Fisher,

j 1903: 778).
5-9 Aug. U Presence noted from offshore (Fisher,

1903: 779) but the comment that they
were seen in August (loc. cit., p. 784)
might suggest they were not seen in June.

1923 24 May 2 ? (Wetmore, ms.).

11-16 June small Seen in small numbers in the rock shelves
numbers below the cliffs on the western side of

the island (Wetmore, ms.).

1936 3 Mar. ? No nests found (Trempe, ms.).

1940 77-15 Aug. fairly "Breeding was in all stages" (Vanderbilt

abundant and de Schauensee, 1941: 12).

1953 21-22 Dec. 4-8 ? (Richardson, pers. comm.).

1954 18 Mar. 2 ? (Richardson, pers. comm.).

1961 2 Mar. te Not noted from offshore (Woodside and

Kramer, ms.).

Table 24. (Continued)
Population
Date of Survey Estimate
EG 9-16 Dee. 0
1962 10 June ?
1963 5-6 June ?
wes, 6-7 Mar... Cas 1,000
25 July 2
23024 Sept. Ca. 250
1965 13-14 Mar. 700
1966 28 July- 6
1 Aug.
1967 8-9 Mar. several
thousand
1968 7-9 Mar. at least sev-
eral hundreds
24-27 Aug. 2
LGTO- 15 “Aug 1,000
ioe cE Sept. low
thousands
EST 16 Sept. 1,000
WHITE TERN
Status

Common breeder; maximum recent estimate:

107

Breeding Status, Remarks, and References
(Kramer, 1961).

? Seen only on cliffs (Kramer and
Beardsley, ms.).

Ca. 100 seen from offshore (POBSP).
2? (BSFW, POBSP).

(BSFW).

? (BSFW, POBSP).

No eggs or young seen (BSFW, POBSP).

? (BSFW, POBSP).

2? (BSFW, POBSP).

? No nests found on accessible part of
island (BSFW, POBSP).

2? Seen in area of northern cliff face
(BSFW).

(BSFW).
(BSFW).
(BSFW).
Gygts alba
3,000. Present through-

out the year but apparently less abundant in fall and winter; probably
breeds throughout the year but data are too few to establish when

breeding peaks occur.

holes

Inmet acel.

231-338 O-77 = 15

Lays single egg on rock outcroppings and in

108

Populations

Most recent estimates of the population place it in the low
hundreds (Table 25) but these estimates are quite subjective. Since
many surveys covered only part of the area wherethese birds occur most
densely, we suspect that many of the estimates are low and that the
maximal populations are in the low thousands. Both Munter's and
Wetmore's early estimates are considerably larger than any recent
estimate, the former so much so that we suspect it was highly inaccu-
rate. Wetmore's estimate is also larger than recent estimates but he
may have seen more of the northern cliff face than did observers on
more recent trips.

Annual Cycle

Many, if not most, White Terns, nest on the nearly inaccessible
north face of the island and many surveys have been unable to find any
nests although they were probably present. Consequently we know little
about the breeding cycle of this species on Nihoa.

Eggs have been found in March, June, August, and September and
pre-fledging young have been found in March, May, June, and August,
indicating that laying has occurred in at least February, April, and
July. It is likely that at least a small proportion of the population
breeds throughout the year.

Breeding Habitat

The great majority of White Terns nests on the sheer cliffs of
the north, east, and west sides of the island. A much smaller number
nests on the rocky outcroppings of the south slopes of the island.
Wetmore (ms.) noted that those on the north cliff face tended to nest
below 500 feet and that most nested between 20 and 250 feet above the
sea.

Specimens

Four specimens were collected by Wetmore on 13 June 1923: an
adult female (USNM 300417), an adult male (USNM 300418) and 2 alco-
holics (USNM 289327-328).

Table 25. Observations of White Terns on Nihoa Island

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1891 26-27 May 2 Presence noted from offshore (Munro,
1941a: 49).
1902 1-3 June ? Common; seen from offshore (Fisher,

1903: 785-786).

Table 25. (Continued)
Population
Date of Survey Estimate Breeding Status, Remarks, and References
1902 5-9 Aug. ? Common; seen from offshore (Fisher,
1903: 785-786).
1915/18 Mar. 50,000 ? (Munter, 1915: 133). Alternative
(100,000) estimate by Brown (ms.).
1916 12 Feb. seen Stated to be not as numerous as on the
occasionally March, 1915, survey (Munter, ms.).
923 5 Apr. 2 Occasionally seen offshore (Wetmore,
ms):
24-27 May abundant Eggs and young found (Wetmore, ms.).
11-16 June 8,000 Some beginning to breed; "other seemed
to have young" (Wetmore, ms.).
1936) -3-Mar. 2 No nests found (Trempe, ms.).
1940 7-15 Aug. not particu- "Breeding was in an early stage, the
larly abundant eggs just commencing to hatch" (Van-
derbilt and de Schauensee, 1941: 12).
1953 s921=22 Dec. 20-30 ? (Richardson, pers. comm.).
1954 18 Mar. 20-30 ? (Richardson, pers. comm.).
1961 2 Mar. i Abundant; seen from offshore (Woodside
and Kramer, ms.).
9-16 Dec. 2 "Many flying immatures seen" (Kramer,
ms.).
1963 5-6 June 2 Thousands seen from offshore, including
many young birds (POBSP).
1964 6-7 Mar. thousands Eggs to fledged young (BSFW, POBSP).
25 July ? (BSEW) .
23-24 Sept. 350 Eggs to fledged young (BSFW, POBSP).
1965 13-14 Mar. 800 ? No attempt made to look for nests
(BSFW, POBSP).
1966 28 July- 500 7 5CBSEW, BOBSE)).

1 Aug.

110

Table 25. (Continued)
Population
Date of Survey Estimate Breeding Status, Remarks, and References
1967 8-9 Mar. 600 ? (BSFW, POBSP).
13-14 Sept. 2 One found incubating an egg (BSFW).
1968 7-9 Mar. many ? No nests found on accessible part of
hundreds island (BSFW, POBSP).
24-27 Aug. many Eggs to flying young (BSFW).
hundreds
1961 1212Mar« 30 2? (BSFW).
1970 15 Aug. 100-200 (BSFW).
1971 18-19 Aug. 3,000 Near fledging young seen (BSFW).
19720 LO. Sept . 300 (BSFW).
19/3 31 sally, 2,000 (BSFW) .
MOCKINGBIRD
Status Mimus polyglottos

Vagrant; one sight record in August 1971.
Observations

Robert J. Shallenberger observed and photographed a Mockingbird
that he saw on Miller Plateau during the BSFW survey of 18-19 August
1971. The species has not been recorded previously from Nihoa but is
well established in the main Hawaiian Islands and has previously wan-
dered to both French Frigate Shoals and Necker Island in the north-
western Hawaiian Islands (Amerson, 1971: 302; Berger, 1972: 215).

NIHOA MILLERBIRD Aerocephalus familiarts kingt

Status

Common endemic breeder; maximum recent population estimate: 625.
Present throughout the year but infrequently seen because of its tendency
to skulk in dense shrubbery. Nests in low bushes and probably breeds
from at least February through late August or early September.

This

Populations

Recent population estimates, based primarily on transect censuses,
have varied widely (Table 26) but this variation is more likely attributed
to inadequacies of the censuses than to very marked changes in the popu-
lations of Millerbirds. In any case it seems likely that the population
consists of at. least several hundred birds.

Annual Cycle

Little is known of the reproductive habits of the Nihoa Millerbird
since only four active nests had been found through July 1973 (Table 26).
What little evidence is available suggests that the species nests pri-
marily during the northern hemisphere spring and summer.

Breeding Habitat

This species occurs widely throughout the island but seems to prefer
areas of dense Stda and Chenopodium. Berger (1972: 110) has pointed out
that of all nests found to date, two were in Sida and the rest were in
Chenopodium. Berger (op. ctt.: 110) has described old nests seen by
him as averaging "about 3 by 4 inches in maximum diameter and were com-
posed primarily of strips and pieces of grass stems and blades, with
varying amounts of rootlets. All of the nests contained some feathers
of other species of birds, white being the predominant color used."

Banding

Thirty-two Millerbirds have been banded on Nihoa by the BSFW: 8
in September 1964, 1 in March 1965, March 1967, March 1968, 4 in August
1968 and 17 in June 1969.

Specimens

We know of 26 study skins of the Millerbird from Nihoa Island.
These are listed in Table 27. There are, in addition, a skeleton (USNM
289276) and an alcoholic (USNM 289299), both collected by Alexander
Wetmore in June 1923.

Table 26. Observations of Nihoa Millerbirds on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1923 11-16 June near 100 Described as a new species Conopoderas
kingt by Wetmore (1924). The breeding
season had ended and young were fully
grown.

1940 7-15 Aug. ? No nests or young birds seen (Vander-
bilt and de Schauensee, 1941: 13).

953° 21-22) Dee. ? Only two individuals seen (Richardson,

O54 224;

112

Table 26. (Continued)

Population
Date of Survey Estimate Breeding Status, Remarks, and References
1961 9-16 Dec. 200 "Pairing appeared to be taking place"
(Kramer, ms.).
1962 10 June ? Many males singing. One nest contain-
ing one egg was found (Kramer and
Beardsley, ms.).
1964 6-7 Mar. 2 7 birds seen. No nests found (POBSP,
BSFW).
25. July ? 3 birds seen, 2 of which appeared to
be paired (BSFW).
23-24 Sept. 150 No evidence of nesting found (POBSP,
BSFW).
1965 13-14 Mar. 100-150 No nests found (BSFW, POBSP).
1966 28 July- 150 Six old nests found 1 August (Berger,
1 Aug. UO 72is HELO)
1967 8-9 Mar. ? Ten to 15 birds seen. Some singing
males heard (POBSP, BSFW).
13-14 Sept. 625 Estimate based on transect censuses
(BSFW) .
1968 7-9 Mar. 615 Estimate based on transect censuses.
No nests found but a recently fledged
young observed (POBSP, BSFW).
24-27 Aug. 2 A nest with two eggs found 25 August
(Berger, 1972: 1101.
1969" 21 Marz. 41 Based on transect censuses. No evidence
(0-123)* of nesting was found (BSFW).
29 May- . 498 lst estimate based on transect censuses
10 June (211-785) *; 30 May; 2nd estimate based on transect
493 censuses 6 June (BSFW). A nest with
(285-701)* two small young found 30 May and one
with a near fledging young found 2 June
(Berger 91972: U0).
19707 915 Aug: 384 Based on transect censuses. No nests
(134-477)* found (BSFW).

113

Table 26. (Continued)

Population

Date of Survey Estimate Breeding Status, Remarks, and References

1971 18-19 Aug. 273 Based on transect censuses (BSFW).
(91-454)*

15 Sept. ? About 10 birds seen (BSFW).

1972 26 Sept. 592 Based on: transect censuses (BSFW).
(334-850)*

973° 31> July 198 Based on transect censuses (BSFW).
(46-350)*

* Figures in parentheses represent confidence limits for the population
estimate at a 95% confidence level.

Table 27. Nihoa Millerbird specimens from Nihoa Island

ae
Museum Museum and Museum Date
Museum Males Nos. Females Nos. imm. Nos. Collected Collector
USNM 5 287888 1 301127 10 301128- 12-15 June Wetmore
(type), 137 1923
301126,
301138-
140
PAS 4 146144- 5 146143, 16 Aug. Vanderbilt
145, 146146- 1940
146149, 148,
146152 146150
AMNH Ll 2325632. 16) Aec: Vanderbilt
1940
NIHOA FINCH Pstttarostra cantans ultima

Status

Abundant endemic breeder; maximum recent population estimate:
6,686. Present throughout the year. Commonly seen because of their
confiding nature. Prefers to nest in rocky outcroppings and nests
principally from late February through at least April.

114

Populations

Although recent estimates are variable, they suggest a population
on the order of 3,000 to 5,000 birds (Table 28). These estimates are
consistently higher than those estimates made by earlier observers but
we believe that this difference may be attributed to more reliable
recent estimation techniques than to any inherent change in the popu-
lation levels of this finch. Recent estimates do not suggest any
pronounced seasonal changes in populations.

Annual Cycle

Available evidence suggests a fairly pronounced spring breeding
season. A near fledging young seen by Richardson on 18 March 1954
indicates that egg-laying may begin in late February. This is cor-
roborated by the presenceof young early in March during 1967 and 1969.
No data is available for nesting activities during April and nests have
not been recorded later in the year. Observations by Berger (1972: 159-
161) on captive Nohia Finches may suggest the potential extent of the
breeding season in the wild. He recorded eggs during a period extending
from 17 December 1969 through 27 July 1970.

Breeding Habitat

Nihoa Finches are widespread over the island but often occur
commonly near the rocky outcroppings (Fig. 32) which serve as their
principal nesting sites. Several observers have noted that these birds
often congregate near seeps or pools of water with Wetmore noting that
as many as 50 could be seen in such situation at one time.

Figure 32. Nihoa Finch on rocky outcropping, 16 September 1972. BSFW
photograph by Eugene Kridler.

MES

Food Habits

Like the related subspecies on Laysan Island, Nihoa Finches are avid
egg eaters, a behaviorism that has been recorded by many observers. In
1923 Wetmore noted that literally thousands of tern, shearwater, and
petrel eggs had had their sides cut open by the finches. In July and
August 1966 both Berger (1972: 157) and Heiden (POBSP) noted this species
feeding on eggs of the Brown Noddy. In one instance Berger noted 20
finches within 30 feet of a single broken egg. Heiden also observed
this species feeding on the eggs of the Gray-—backed Tern, Red-footed
Booby, and Wedge-tailed Shearwater.

The only other information available on the food habits of this
species comes from Vanderbilt and de Schauensee (1941: 13) and Rich-
ardson (1954: 224). The former noted that several stomachs contained
",,.a gelatinous yellow substance (probably yolk of egg), minute black
seeds and microscopic pebbles.'"' Richardson repeatedly observed these
finches "eating the small, green flower heads of Chenopodtwn sand-
wicheum...[and noted]...one bird picking out the still partly green
seed of Portulaca cawmit".

Banding

A total of 336 Nihoa Finches have been banded by the BSFW. Four
adults were banded in September 1964; 8 nestlings were banded in March
1965; 12 adults were banded in July 1966; 46, all adults except for one
immature, were banded in March 1967; 22 nestlings were banded in March
1968; 37, including 7 adults and 30 immatures were banded in August
1968, and another 207 were banded in June 1969. Forty-five of the 46
birds banded in March 1967 were shipped by BSFW to French Frigate Shoals
with 32 later being introduced to Tern Island and with 10 being intro-
duced to East Island. The East Island population did not survive. As
Many as 9 were still present on Tern Island in September 1971 (BSFW).

Specimens

Fifty-nine specimens of Nihoa Finches are listed by location in
Table 29. There are in addition a skeleton (USNM 289277) and an
alcoholic (USNM 289329) both of which were collected by Wetmore in
June 1923.

116

Table 28. Observations of Nihoa Finches on Nihoa Island

Population

Date of Survey Estimate Breeding Status, Remarks and References

1885 22. July ? Species seen by Sanford B. Dole
(Munro, 1960: 130).

1915 « .18..Mar.. 7000. ox (Munter, 1915: 132). The presence of

more" this bird was later noted by Bryan

(1916), s

1916 12 Feb. ? 5 specimens collected by Munter (ms.)
were later described as a new species
Telesptza ultima by Bryan (1917).

1923 11-16 June 800 Birds had finished breeding and adults

were in molt (Wetmore, ms.).

1940 7-15 Aug. 500-1, 000 Birds not nesting but 2 old nests found
(Vanderbilt and de Schauensee, 1941: 13).

1953 21-22 Dec. "reasonably 2 old nests found (Richardson, 1954:
numerous" 224)".
19547 928 Mar’. = 1 near-fledging young found in 1 of the

nests seen the previous December
(Richardson, 1954: 224).

1961 9-16 Dec. 800-1,200 No nests found (Kramer, ms.).

1962 10 June "as abundant as No nests found (Kramer and Beardsley, ms.).
in December 1961"

1964 6-7 Mar. 2,500- 1 bird seen nest building (POBSP).
5,000
23-24 Sept. 4,500- No evidence of nesting found (POBSP,
5,000 BSFW).
1965 13-14 Mar. 4,000 6 nests with young and 1 containing 3

eggs found. Of 4 nests examined, 2
contained 2 young and 2 contained 3
young. 8 nestlings were banded (POBSP,

BSFW) .

1966 28 July- 5,000 No evidence of breeding found (POBSP,
1 Aug. BSFW).

1967 8-9 Mar. 5, 000 42 birds captured for introduction to

Tern and East Islands, French Frigate
Shoals (POBSP, BSFW). Eggs and young
found (Berger, 1972: 158).

Table 28.

Date of Survey

1968

1969

1970

LO71

1972

1973

(Continued)
Population
Estimate Breeding Status, Remarks, and References
13-14 Sept. 5,000 Estimate based on transect censuses
(BSFW).
7-9 Mar. 6,600 Estimate based on transect censuses.
Birds apparently at a peak of nesting.
Birds seen building. 13 nests found
with a total of 28 nestlings and 10
eggs. The maximum number of young in a
nest was 4. 22 nestlings were banded
(POBSP, BSFW).
24-27 Aug. 6,686 Estimate based on transect census
(4,881-8,491)* (BSFW).
21 Mar. 2,993 Estimate based on small number of trans-
(1,913-4,073)* ect censuses and possibly subject to
error. 2 nests found, 1 with 1 egg and
1 with 1 young (BSFW).
29 May 2,987 lstiestimate based on transect censuses
(2,206-3,948)* 30 May; 2nd estimate based on transect
Pis28 censuses 6 June (BSFW).
(924-2 ,132)%*
15 Aug. 2,34) Estimate based on transect censuses.
(1,637-3,045)* Several old nests found; no evidence of
breeding noted (BSFW).
18-19 Aug. 3,759 Estimate based on transect censuses.
(2,707-4,811)*No active nests found (BSFW).
15 Sept. common (BSFW).
everywhere
16 Sept. 3,799 Estimate based on transect censuses
(3,009-4 ,589)*(BSFW).
a1 uly 1,318 Estimate based on transect censuses

(892-1,744)* (BSFW).

* Figures in parentheses represent confidence limits for the population
estimate at a 95% confidence level.

LU,

118

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Mammals

Only one species of mammal is known to occur at Nihoa and its

appearance there is uncommon.

HAWAIIAN MONK SEAL

Status

Uncommon visitor.

Observations

Monachus schautnslandt

Not known to breed at Nihoa.

All observations of this species at Nihoa are summarized in Table

30).
Table 30. Observations of Hawaiian Monk Seals at Nihoa Island
Number
Date of Survey Seen Remarks and References
1657. 23 Apr. about a On beach. Several shot by King
dozen Kamehameha IV (Paty tm Emory, 1928: 9).
1940 7-15 Aug. 0 (Vanderbilt and de Schauensee, 1941: 8).
1953 21-22 Dec. 0 (Kenyon and Rice, 1959: 217).
1954 18 Mar. 0 (Kenyon and Rice, 1959: 217).
1957 28 Dec. 0 (Ritee, 1960". 377).
1964 6-7 Mar. i 1 lying on small beach on west side
of Adam's Bay 6 March. Adult seem
swimming offshore later in day (POBSP).
23-24 Sept. 1 Adult swimming near landing site (BSFW,
POBSP).
1965 13-14 Mar. 6 3 seen each day: 2 adults, male and
female; 4 subadults, 1 female (BSFW,
POBSP).
1966 28 July- 0 (BSFW, POBSP).
1 Aug.
1967 8-9 Mar. 0 (BSFW, POBSP).
13-14 Sept. 0 (BSFW).

ig

120

Table 30. (Continued)

Number

Date of Survey Seen Remarks and References
1968 7-9 Mar. 0 (BSFW, POBSP).

24-27 Aug. 0 (BSFW) .
1969 21 Mar. 0 (BSFW) .

29 May 1 (BSEW) .
1970 15 Aug. 0 (BSFW) .
1971 18-19 Aug. 0 (BSFW) .

15 Sept. 1 Found basking on sandy beach (BSFW).
1972 16 Sept. 3 (BSFW).
19732 oe July, 4 Basking on sandy beach (BSFW).

Reptiles

Two species of reptiles, a lizard and a turtle, are known to
occur at Nihoa. Both are uncommon and there is no adequate evidence
that either species has ever bred there.

MOURNING GECKO Leptdodactylus lugubris

Status

Uncommon resident; probably a recent introduction that may not
be established.

Observations

This species has only been twice seen on Nihoa. A single specimen
was collected in HZragrostis clumps in September 1964 (Beardsley, 1966:
160) and another specimen was collected in March 1965 by Walker. The
absence of observations from earlier visits, particularly those in 1923
and 1940, suggests that the species is probably a recent introduction,
most likely stemming from 1961 and 1962 when oil barrels, boards, and
much other miscellaneous material was ferried to the island by the
helicopters of the HIRAN operation. The absence of more recent ob-
servations suggests that the gecko may not have become established on
the island.

GREEN TURTLE Chetonta mydas
Status
Uncommon visitor.

Observations

Emory (1928: 8) remarked that "Turtles are fairly common [at
Nihoa]"" but we have been unable to find any historical basis for his
statement. Perhaps some were seen during his visit in 1924. Recent
observations of turtles at Nihoa are few (Table 31) and indicate the
species is only an occasional visitor to the island. They are most
likely visitors from the nearby large breeding colony at French Frigate
Shoals (see Amerson, 1971: 79-92).

Table 31. Observations of Green Turtles at Nihoa Island

Number

Date of Survey Seen Remarks and References
1964 6-7 Mar. 0 (BSFW, POBSP).

23-24 Sept. 2 Seen just offshore (BSFW, POBSP).
1965 13-14 Mar. if About 2 1/2 feet long (BSFW, POBSP).
1966 28 July- 0 (BSFW).

1 Aug.
1967 8-9 Mar. il 1 large turtle seen offshore on 9

March (BSFW, POBSP).

12a:

Wa

Table 31. (Continued)

Number
Date of Survey Seen Remarks and References
1967 13-14 Sept. 0 (BSFW).
1968 7-9 Mar. af Seen swimming offshore on 8 March
(BSFW, POBSP).

24-27 Aug.. 0 (BSFW).
IGS) 2S Mars 0 (BSFW) .
LOO eS AU, 0 (BSFW).
1971 18-19 Aug. 0 (BSFW).

15 Ssept.. S) 2 were on a low rock at the base of
the northwest cliff; the other was
seen swimming around the ship (BSFW).

19/2 M6ASept .« Hl (BSFW).
1973 231 July 0 (BSFW).
ACKNOWLEDGMENTS

Many persons contributed much time and effort towards the com-
pletion of this report. Michio Takata, Director, Hawaii Division of
Fish and Game, kindly allowed us to use unpublished reports in the
Division's files. Dr. Alexander Wetmore, Smithsonian Institution,
Washington, D.C., graciously gave us full use of the information: ob-
tained by him during the 1923 Tanager Expedition, and Dr. Frank
Richardson, University of Washington, Seattle, Washington, communicated
population estimates made during his visits to the island in December
1963 and March 1964. Mr. Edwin H. Bryan, Jr., Manager of the: Pacific
Science Information Center of the Bernice P. Bishop Museum, Honolulu,
aided by allowing us full access to his files on many different
occasions. Dr. Philip S. Humphrey, University of Kansas, Kansas,
principal investigator of the POBSP, is to be thanked for his constant
support during the many frustrations encountered during the preparation
of this account.

Many others, particularly personnel of the Pacific Ocean Biological
Survey Program, Bureau of Sport Fisheries and Wildlife, and Hawaii
Division of Fish and Game, contributed field data which constitute a
large proportion of the information presented herein. These persons are
listed in Appendix Table 1.

We also thank A. Binion Amerson, Jr., and Dr. Philip C. Shelton
who made valuable comments during the preparation of the manuscript.
Dr. F. Raymond Fosberg kindly read and commented on the vegetation
section of this report. A. Binion Amerson critically read the entire
final draft.

Field work on Nihoa was made possible by a cooperative agreement
between the Department of the Interior, Bureau of Sport Fisheries and
Wildlife, and the Smithsonian Institution. We are also most grateful
to the U.S. Coast Guard whose logistic support made possible most of
the visits to Nihoa.

The camera copy was typed by Barbara B. Anderson with funding
through a contract with the Bureau of Sport Fisheries and Wildlife,
Department of the Interior (contract number 14-16-008-596, February
Bi nO).

28

124

LITERATURE CITED

Agassiz, A., and H.L. Clark. 1907-1912. Hawaiian and other Pacific
Echini. Mus. Comp. Zool. Harvard, Mem. 34: vii + 383 pp.

Alexander, W.B., et al. 1965. The families and genera of petrels and
their names. Ibis 107: 401-405.

Amerson, A.B., Jr. 1968, Tick distribution in the central Pacific as
influenced by seabird movement. J. Med. Ent. 53(3): 332-339.

----- 1971. The natural history of French Frigate Shoals, northwestern
Hawaiian Islands. Atoll Res. Bull. 150: xv + 383 pp.

Amerson, A.B., Jr., R.B. Clapp, and W.0. Wirtz II. 1974. The natural
history of Pearl and Hermes Reef, northwestern Hawaiian Islands.
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Amerson, A.B., Jr, and P.C. Shelton. In press. The natural history
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----- Committee on Classification and Nomenclature. 1973. Thirty-
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125

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<—<——— 1917. Description of Telesptza ultima from Nihoa Island. Auk
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aoe [1965]. Hawaiian Islands National Wildlife Refuge...[survey]
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----- [1966]. Hawaiian Islands National Wildlife Refuge trip
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=—=——— [1966]. Hawaiian Islands National Wildlife Refuge trip
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----- [1967]. Refuge log book for 6 March - 1 April 1967. 39 pp.

----- [1967]. [Partial report of] Hawaiian Islands National Wildlife
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14 pp.

----- [1968]. Field notes kept during the March 1968 survey of
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----- [1968]. Field notes kept during the August 1968 survey
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----- [1969]. Hawaiian Islands National Wildlife Refuge spring
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----- [1972]. Expedition Report. Hawaiian Islands National
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---== 1949. Ophiuroidea of the Hawaiian Islands. B.P. Bishop Mus.
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es L906. ~Bhe Starfishes of the Hawaiian Isilands. U.S. Fish Comm.
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128

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----- 1968. Records of Hippoboscidae (Diptera) from the central
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rar

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----- 1924. <A warbler from Nihoa. Condor 26: 177-178.

----- 1925. Bird life among lava rock and coral sand. Nat. Geog.
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133

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Thysanoptera. Univ. of Hawaii Press, Honolulu. viii + 475 pp.

----- 1948b. Insects of Hawaii, Vol. 3. Heteroptera. Univ. of
Hawaii Press. sv o+: 255 pp.

em 1958a. Insects of Hawaii, Vol. 7. Macrolepidoptera. Univ.
of Hawaii Press, Honolulu. ix + 542 pp.

----- 1958b. Insects of Hawaii, Vol. 8. Lepidoptera: Pryaloidea.
Univ. of Hawaii Press, Honolulu. ix + 456 pp.

134

Appendix Table l.

Date
1858?

1885* 22 July

1891* 26-27 May

1902* 1-3 June
5-9 Aug.
LOUD A Dec.

1914 7, Sept’.

TOMS eS) Matar

1916°>-12 Reb.

1923 24-25 May

11-20 June

Scientific visits to Nihoa Island, 1885-1973

Personnel
Dr. Rooke

Princess Liliukolani
E.M. Beckley

Sereno E. Bishop
W.E.H. Deverill
Sanford B. Dole

Mr).. Hall

Mr. Jaeger

Mr. Williams

Some 200 others

Rothschild Expedition

Henry ©. Palmer
George C. Munro

Albatross Expedition

Charles H. Gilbert (SU)***
Walker K. Fisher (SU)
Charles C. Nutting (SUI)
John 0. Snyder (SU)

George Willett (BBS)

Capt. James H. Brown (U”. 3)
Carl Elschner

Capt. James H. Brown (USCG)
Lt. William H. Munter (USCG)
4 members of crew

Lt. William H. Munter (USCG)
Crew of THETIS

Tanager Expedition

David L. Thaanum (BPBM) (conchologist)
Theodore T. Dranga (conchologist)
Chapman Grant (naturalist)

Tanager Expedition

Alexander Wetmore (BBS) (ornithologist)
William G. Anderson (collector)
Avi. Ge Atkinson (BAR)

Vessel

?

IWALANT

KAALOKAT

ALBATROSS

THETIS

THETIS

THETIS

THETIS

TANAGER

TANAGER

135

Appendix Table 1. (Continued)
Date Personnel Vessel

Edwin H. Bryan, Jr. (BPBM) (entomologist)

Bruce Cartwright (BPBM) (assistant in
hydrographic work)

Chapman Grant (BBS) (naturalist)

Charles J. Judd (forester)

Edward L. Caum (BPBM) (botanist)

Harold S. Palmer (BPBM) (geologist)

Eric L. Schlemmer (assistant to Wetmore)

David L. Thaanum (BPBM) (conchologist)

George Higgs (cook)

C. Montague Cooke, Jr. (BPBM) (malacologist)

1924 9-13 July Tanager Expedition TANAGER

Harold S. Palmer (BPBM) (geologist)
William G. Anderson (collector)

William Bush (collector)

Erling Christophersen (BPBM) (botanist)
Theodore T. Dranga (BPBM) (conchologist)
Kenneth P. Emory (BPBM) (archaeologist)
Kenneth I. Hobson (collector)

A. Landgraf (topographer)

OIG. 3) Mar . A.D. Trempe RELIANCE
B.L. Bassham (USCG)
Other members of crew

1940 7-16 Aug. Mr. and Mrs. George Vanderbilt NAVIGATOR
Clifton Weaver

Bosal y George Vanderbilt Pacific Equatorial PIONEER
Expedition

George Vanderbilt
Vernon E. Brock (HDFG)
B. Green

Robert R. Harry (SU)
Anita Vanderbilt
Lucille Vanderbilt

T. Ivar Vatland

1953 21-22 Dec. Frank Richardson (UW) BUTTONWOOD
O54" 13) Mar. Frank Richardson (UW) BUTTONWOOD
1955 21-24 Aug. Ivan T. Rainwater (USDA) AUKAKA

George Carter
David G. Nottage
Peter Nottage

Ed Sheehan

136

Appendix Table 1.
Date
1957 28 Dec.
1961* 2 Mar.
10-15 Dec.
(1600-0830) **
1962 10 June
(0615-1345)
1963* 5-6 June
1964 6-7 Mar.
(1000-0800)
29 Judy
(ea. 1000-
1500)
23-24 Sept.
(0930-1700)
1965 13-14 Mar.
(1045-1545)
1966* 18-20 Mar.
28 July-
1 Aug.

(Continued)
Personnel

Karl W. Kenyon (BSFW)
Dale W. Rice (BSFW)

David H. Woodside (HDFG)
Raymond J. Kramer (HDFG)

Raymond J. Kramer (HDFG)
Gerald Swedberg (HDFG)
HIRAN II personnel

David B. Marshall (BSFW)
John W. Beardsley (HSPA)
Raymond J. Kramer .(HDFG)
David H. Woodside (HDFG)

A. Binion Amerson, Jr. (POBSP)
Fred C. Sibley (POBSP)

Eugene Kridler (BSFW)

A. Binion Amerson, Jr. (POBSP)
Loren Kroenke (UH)

Edward O'Neill (BSFW)

Ronald L. Walker (HDFG)

George S. Wislocki (POBSP)

Eugene Kridler (BSFW)

Eugene Kridler (BSFW)
John Beardsley (UH)
Robert R. Fleet (POBSP)
Charles R. Long (POBSP)
Ronald L. Walker (HDFG)

Eugene Kridler (BSFW)
Winston Banko (POBSP)
Chandler S. Robbins (BSFW)
Ronald L. Walker (HDFG)

Eugene Kridler (BSFW)
Andrew Berger (UH)
Nelson Rice (HDFG)
Ronald Walker (HDFG)

Eugene Kridler (BSFW)
Andrew J. Berger (UH)
Richard S. Heiden (POBSP)
Ernest Kosaka (HDFG)

Vessel

Aerial Survey

PLANETREE

FLOYD COUNTY

STONE COUNTY

Helicopter

TAWAKONI

PLANETREE

CHARLES H.
GILBERT

BASSWOOD

BLACKHAW

BUTTONWOOD

CHARLES H.
GILBERT

137

Appendix Table 1. (Continued)
Date Personnel Vessel

1967 8-9 Mar. Eugene Kridler (BSFW) BASSWOOD
(0945-1040) C. Douglas Hackman (POBSP)
Ernest Kosaka (HDFG)
John Maciolek (BSFW)
Richard Wass (UH)

13-14 Sept. Eugene Kridler (BSFW) BUTTONWOOD
(0855-1420) Robert Ballou (BSFW)

John L. Sincock (BSFW)

Ronald L. Walker (HDFG)

1968 7-9 Mar. Eugene Kridler (BSFW) IRONWOOD
(1130-1030) Roger B. Clapp (POBSP)
Karl W. Kenyon (BSFW)
Ernest Kosaka (HDFG)
John L. Sincock (BSFW)

24-27 Aug. Eugene Kridler (BSFW) BUTTONWOOD
(0900-1400) G. Brent Dalyrymple (USCG)

Richard R. Doell (UDCG)

C. Robert Eddinger (UH)

Derral Herbst (UH)

John L. Sincock (BSFW)

1969 21 Mar. Eugene Kridler (BSFW) BUTTONWOOD
(0900-1730) Karl W. Kenyon (BSFW)
George Laycock (NAS)
David L. Olsen (BSFW)
John L. Sincock (BSFW)

29 May- David L. Olsen (BSFW) MAHL
10 June Ernest Kosaka (HDFG)

James McVay (UH)

William Patzert (UH)

John L. Sincock (BSFW)

Douglas Yen (BPBM)

1970 15 Aug. Eugene Kridler (BSFW) BUTTONWOOD
(0830-1600) Joseph Mazzoni (BSFW)
David L. Olsen (BSFW)
John L. Sincock (BSFW)
David H. Woodside (HDFG)

1971 18-19 Aug. David L. Olsen (BSFW) TERITU
(0800-1230) David Childs (SI)
Richard Grigg (HIMB)
Robert J. Shallenberger (OI)
James Vansant (UH)
William Worcester (UH)

138

Appendix Table 1. (Continued)

Date

UOT

Personnel Vessel

15> Sept. Eugene Kridler (BSFW) BUTTONWOOD

(1300-1800) Erwin A. Bauer
Kenneth S. Norris (OI)
John L. Sincock (BSFW)
Eric L. Schlemmer

16 Sept. Eugene Kridler (BSFW) BUTTONWOOD

(0700-1500) Russel Apple (USNPS)
Bruce Benson (HA)
Ernest Kosaka (HDFG)
David L. Olsen (BSFW)
John L. Sincock (BSFW)

1973 31 July David L. Olsen (BSFW) BUTTONWOOD

*

kX

KKK

(1030-1830) John L. Sincock (BSFW)
Leighton Taylor (BSFW)
Thomas Telfer (HDFG)

No landing made on island.

Time of arrival and departure, where known, is listed under the
date of visit for surveys made during the 1960's.

Glossary of Abbreviations: BBS, Bureau of Biological Survey;
BPBM, Bernice P. Bishop Museum; BSFW, Bureau of Sport Fisheries
and Wildlife; HA, Honolulu Advertiser; HBAF, Hawaiian Board of
Agriculture and Forestry; HDFG, Hawaii Division of Fish and Game;
HIMB, Hawaii Institute of Marine Biology; HSPA, Hawaiian Sugar
Planters Association; NAS, National Audubon Society; OI, Oceanic
Institute, Waimanalo, Hawaii; POBSP, Pacific Ocean Biological
Survey Program; SI, Smithsonian Institution; SU, Stanford
University; SUI, State University of Iowa; UH, University of
Hawaii; USDA, United States Department of Agriculture; USCG,
United States Coast Guard; USCGS, United States Coast and Geodetic
Survey; USNPS, United States National Park Service; UW, University
of Washington.

1359

Appendix Table 2. Results of scientific visits to Nihoa Island,

1885-1973
Date Results
1858 ? According to Hillebrand (1888: 451), seeds of the

endemic palm were brought to Honolulu by a Dr. Rooke.
We know nothing further of this visit.

1885 22 July Topographic and geologic observations. Observations
and collections of birds, none of which was sub-
sequently reported. Archaeological material collected
included a stone bowl and dish, a coral rubbing stone,
and a coral file.

1891 26-27 May Birds observed from offshore. 3 Red-footed Boobies
collected and other birds observed (Munro, 194la:
1941b).

1902 1-3 June Observations of birds from offshore (Fisher, 1903).
5-10 Aug. Collected offshore: corals, molluscs, hydroid,

schizopod, crabs, fish, echinoderms, and bird
specimens (at least 6 birds skins representing 3

species).

LOZ 7 aDecs Observations of birds from offshore.

1914 D7 Sept. Island and its geology described; seeds and por-
tions of Pritehardta taken to Honolulu. Plants
collected.

OMS ely Mag) Observations of birds and estimates of the numbers

present; first published mention of the Nihoa Finch,
palm seeds collected (Munter, 1915).

1916 12 Feb. Observations of birds; photographs of flora and
fauna; collected: plants; 5 specimens of the Nihoa
Finch, reported and later described by W.A. Bryan
GHOPGrR aL Sir7)y

1923 24-25 May Observations of birds and description of the

11-16 June Nihoa Millerbird (Wetmore, 1924, 1925). A new bird
distributional record from the June 1923 visit was
later reported by Clapp and Woodward (1968). 109
bird specimens (skins) of 17 species collected.
Collections of: crustacea, echinoderms, foraminifera;
fish, mollusca; marine algae, insects, vascular
plants, rocks. Geology, topography, and archaeology
described.

231-338 O- 77-17

140

Appendix Table 2. (Continued)

Date Results
1936 3 Mar. Observations of birds and their breeding status. ‘

1940 7-10 Aug. Observations and annotated bird list (Vanderbilt
and de Schauensee, 1941); photographs taken and
color movie of birds made. 48 birds of 12 species
collected.

POS 1 ax Juiky, Fish collected; 12 Nihoa Finches captured for trans-
port to Honolulu Zoo of which 6 evidently reached
the Zoo (Herald, -1952:.71'5)s

1953. 21-22 Dec. Observations of seabirds and their breeding status
(Richardson, 1957); seal census taken but none found.

1954 18 Mar. Observations of seabirds and their breeding status
(Richardson, 1957); seal census taken but none found.
Notes on status of Nihoa Finch and Millerbird (Rich-
ardson, 1954).

1955 21-24 Aug. Charcoal collected for radioactive carbon dating;
3 or 4 Great Frigatebirds captured for transport to
Honolulu Zoo; plants collected; an unfinished adz
and a stone bowl collected; movies, photographs
and tape recordings made.

1957- 28.Dee; Seal census taken but none found; aerial census of
albatrosses (Rice and Kenyon, 1962).

1961 2 Mar. Bird observations from offshore.

10-15 Dec. Observations of birds with particular emphasis on
Nihoa Finches and Millerbirds; survey of vegeta-
tion with particular emphasis on status of Nihoa
Palm; vegetation photostations established; effect
of military activities investigated; plants col-
lected; seeds of Pritehardia and Chenopodium
collected for artificial propagation.

19625200 June Observations of birds with particular emphasis on
the Nihoa Millerbird; brief notes on vegetation;
collection of 15 species of insects and plant
associates; effect of military activities investi-
gated; photographs taken.

1963 5-6 June Birds observed offshore; 2 Bulwer's Petrels col-
lected.

Appendix Table 2.

Date

1964

1965

2a uly

23-24 Sept.

13-14 Mar.

1966 28 July-

1967

1968

1 Aug.

8-9 Mar.

13-14 Sept.

24-27 Aug.

(Continued)
Results

Observations of birds; seals and turtles censused;
refuge signs erected; vegetation photographed.
Collected: plants; limpets, and algae by Walker;

2 petrel chicks; arachnids. 99 birds of 11 species
banded.

Observations of birds and their breeding status.

Observations of birds with particular emphasis on
the millerbird; census of turtles and seals; 57
birds of 5 species banded. Collected: plants,
isopods, arachnids, insects, 1 lizard.

Observations of birds; turtles and seals censused;
312 birds of 9 species banded. Collected: 1
Sooty Storm Petrel, 1 lizard.

Observations of birds with particular emphasis

on surveys of Nihoa Finch and Millerbird; turtles
and seals censused; photographs taken of terrain;
refuge sign erected. Collected: bird specimens;
limpets by Berger. 1,544 birds of 8 species banded.

Observations of birds; turtles and seals censused;
45 Nihoa Finches captured by BSFW for introduction
to French Frigate Shoals; 1 Millerbird and 46 Nihoa
Finches banded. Collected: marine inshore or-
ganisms, hippoboscid flies, 1 finch.

Observations of birds with particular emphasis on
Nihoa Finches and Millerbirds; transect censuses
made of finch and Millerbird populations; 1 gecko
and 1 finch collected.

Observations of birds with particular emphasis on
Nihoa Finches and Millerbirds; transect censuses
made of finch and Millerbird populations; turtles
censused; plants collected; 105 birds of 5 species
banded.

Observations of birds with particular emphasis on
Nihoa Finches and Millerbirds; transect censuses
made of finch and Millerbird populations; turtles
and seals censused; collection of rock samples for
analysis of magnetic properties; observations of
vegetation; ectoparasites collected from finches;
42 birds of 3 species banded.

141

142

Appendix Table 2.

Date

1969

1970

LOZ

OT2

O73

21> Mark

29 May-
10 June

15 Aug.

18-19 Aug.

14-15 Sept.

16 Sept.

SE July

(Continued)
Results

Observations of birds, censuses of turtles and
seals; island vegetation cover mapped; partial
censuses made of finch and Millerbird populations.
2 Sooty Storm Petrels banded.

Observations of birds with particular emphasis on
Nihoa Finches and Millerbirds; transect censuses

of finch and Millerbird populations; ethnobotanical
surveys, marine survey of surrounding waters,
retrieval of current meters, vegetation photosta-
tion photographs obtained. 224 birds of 2 species
banded.

Observations of birds, transect censuses of finch
population.

Observations of birds; transect censuses made of
finch and Millerbird populations. Offshore
marine observations.

Cursory observations made of birds, seals and
turtles. Several archaeological samples obtained
for Carbon-14 dating.

Observations of birds, seals and turtles; transect
censuses made of finch and Millerbird populations.

Cursory observations of birds. Seals noted;
transect censuses made of finch and Millerbird
populations.

icaeseenaoaRaein

143

Appendix Table 3. Publications on collections and studies (with
the exception of birds) made on Nihoa Island,.

1885-1973%
Protozoa
Cushman in Edmondson Reports 15 species of foraminifera collected
et al. offshore by the Tanager Expedition.
Coelenterata

Nutting, 1905. Reports 1 species of hydroid collected
south of Nihoa by the Albatross Expedition.

Vaughan, 1907. Reports 6 species of corals (Madreporia)
collected offshore by the Albatross Expedi-
tion.

Nutting, 1908. Reports 11 species of coral (Alcyonaria)
collected by the Albatross Expedition; most
are described as new species.

Mollusca

Pelsbry, 1927. Lists a barnacle collected by the Tanager
Expedition.

Dati. et al., 1938. Lists two species of pelecypods collected
offshore by the Albatross Expedition.

Annelida
Hartman, 1966. Summarizes published records of polychaetes

{1 species); gives current taxonomy.

Arthropoda
Arachnomorpha (Arachnida)
Bryan, €¢ al., 1926. States that bird ticks were found abundantly.

Jacot, 1929. Reports an orabatid mite (Acarina) from
collections made by the Tanager Expedition.

Beardsley, 1966. Reports 5 Araneida and an undetermined
pseudoscorpion from collections made in
September 1964. First record of the oc-
currence of pseudoscorpions.

Amerson, 1968. Reports the distribution and hosts of ticks
from collections made by the POBSP.

1d

Appendix Table 3. (Continued)

——

Crustacea
Ortmann), 1905. Reports a single species of schizopod
collected in the vicinity of Nihoa by the
Albatross Expedition.
Rathbun, 1906. Reports brachyuran and macruran crabs col-

lected offshore by the Albatross Expedition.

Edmondson in

Edmondson et al., Reports 3 species of decapods collected by
Lg25% the Tanager Expedition.
Bryan et av*.,- 1926. Indicates that isopods were collected by

the Tanager Expedition.

Beardsley, 1966. Lists 2 species of isopods from collections
made in September 1964.

Labiata (Hexapoda - Insects)

Timberlake, 1924. Records a chalcid fly collected by the
Tanager Expedition.

Bryan et al., 1926. Reports ca. 67 species of insects collected
by the Tanager Expedition.

Wheeler, 1934. Lists 4 species of ants on the basis of
earlier publications.

Lopes, 1938. Describes a new species of sarcophagid
fly from collections of the Tanager Expe-
dition.

Usinger, 1942. Reports 3 species of Nystus (Hemiptera:

Lygaeidae), 2 described as new, from col-
lections by the Tanager Expedition.

Zimmerman, 1948a. Lists 8 species of insects (1 thysanuran,
4 cockroaches; 1 embiopteran, and 1 earwig.
Zimmerman's various distributional records
in the Insects of Hawaii series derive from
the Tanager collections, but extensively
revise taxonomy, reidentify specimens, and
identify to species hitherto unidentified
specimens; several new distributional records
are listed in the series.

Zimmerman, 1948b. Lists 6 species of Hemiptera (4 lygaeids,
1 nabid, and 1 anthocorid).

Appendix Table 3.

Ross, 1951.

Zimmerman, 1958a.

Zimmerman, 1958b.

Maa, 1962.

Hardy, 1964.

Yashimoto, 1965.

Hardwick, 1965.

Beardsley, 1966.

Maa, 1968.

Echinodermata

Fisher, 1906.

Fisher, 1907.

Agassiz and Clark,
1907-1912.
Clark, 1908.

Clark tm Edmondson
et al., 1925

(Continued)

States that the embiopterid collected by
the Tanager Expedition and reported as
Oltgotoma tnsularis (Bryan et al., 1926)
is actually Oligotoma (Aposthonta) oceanta
Ross sp. nov.

Lists 2 species of noctuid moths.

Lists 1 pyralid moth and 1 pterophorid
moth.

Reports specimens of Hippoboscidae col-
lected by the Tanager Expedition.

Identifies a dolicopodid fly not specifically
identified in Bryan et al., 1926.

Lists 2 species of eulophids (Hymenoptera:
Chalcoidea).

Describes a noctuid moth from specimens
collected by the Tanager Expedition.

Reports 110 species of insects collected in
June 1962 and September 1964, 41 of them
new distribution records. Lists earlier
records of insects but does not include
Mallophaga.

Reports POBSP collections of hippoboscid
flies.

Reports 9 species of starfishes (Asteroidea)
from collections made by the Albatross
Expedition offshore.

Records 3 sea cucumbers (Holothuroidea)
collected by the Albatross Expedition.

Records 9 species of Echinoidea collected
by the Albatross Expedition.

Reports a crinod collected from offshore
by the Albatross Expedition.

Reports 3 species of Echinoidea collected
by the Tanager Expedition

145

146

Appendix Table 3.

Clark, 1949);

Chordata
Vertebrata
Pisces

Snyder, 1904.

Gilbert, 1905.

Fowler and Ball,
1925

Strasberg, 1956.

Reptilia

Beardsley, 1966.

Mammalia
Batiyeelooine
Kenyon and Rice,

1195:9'-
Rice, 1960b.

Tomich, 1969.

Flora

Beccari, 1889.

Bryan, 1916.

(Continued)

Reports 10 brittle stars (Ophiuroidea)
collected by the Albatross Expedition.
Summarizes previous records for echinoderms.

Reports 1 species collected by the Albatross
Expedition.

Records 22 species of deep sea fishes col-
lected in the vicinity of Nihoa by the
Albatross Expedition.

Reports 7 species of fish collected by the
Tanager Expedition.

Revises taxonomy of Hawaiian blennioid
fishes and records 1 species from Nihoa.

Gives first record of a lizard, Leptdodactylus -
lugubrts, from a collection made in September
1964.

First report of seals at Nihoa.

Reports no seals seen during visits by
Richardson in 1953, and 1954.

Reports no seals seen on aerial survey
December 1957.

Reports occurrence of monk seal in March
E965.

Describes endemic palm (Pritehardia remota)
from cultivated specimen from Honolulu.

Indicates palm seeds were collected during
the April 1915 cruise of the THETIS. The

Appendix Table 3. (Continued)

Beccari and Rock,
1921.

Christophersen and
Caum, 1931

Magnusson, 1942

Lamoureux, 1964.

Tsuda, 1966.

Geophysical
Bishop, 1885a and b.

Elschner, 1915.

Rowers, 1920.

Washington and Keyes,
O26;

Balmer, 1927. ¢

Kroenke and Woollard,
1965).

Archaeology

Emory, 1928.

THETIS visited Nihoa in March not April
1915 and two reports of the visit give no
indication that plants were collected then.
In actuality the seeds were collected in
September 1914 (see next entry).

Remarks on Pritchardta specimens collected
in September 1914.

Reports 20 species of vascular plants col-
lected by the Tanager Expedition.

Lists 19 species of lichens collected by
the Tanager Expedition.

Reports that the 11 species of vascular
plants collected by the HDFG in 1962 are
represented in the 20 species collected by
the Tanager Expedition in 1923.

Reports 2 species of marine benthic algae
from collections made in July 1924 and
March 1964.

Gives geological and topographical observa-
tions made in 1885.

Gives descriptions and geological comments
from observations made offshore in September
1914.

Records observations on a rock specimen
stated (erroneously) to have been collected
by W.A. Bryan.

Reports results of studies of rocks collected
in) OA Sand 923).

Gives geological and topographical observa-
tions made by the Tanager Expedition.

Reports gravity observations made in March
1964.

Reports on archaeological work conducted
by the Tanager Expedition and summarizes
all available information.

U. S. GOVERNMENT PRINTING OFFICE: 1977 O = 231-338

147

Nos. 208-219

ATOLL RESEARCH
BULLETIN

QE
= os
AEF)
= 7

208.

209.

210.

211.

212.

213.

Notes on Plants of the Genus
Cauler pa in the Herbarium

of Maxwell S. Doty at the
University of Hawaii

by Wm. Randolph Taylor
Marine Algae of the Te Vega
1965 Expedition in the
Western Pacific Ocean

by Wm. Randolph Taylor

Marine Algae Known from the
Maldive Islands
by H. E. Hackett

The Benthic Algal Composi-
tion, Standing Crop, and
Productivity of a Caribbean
Algal Ridge

by Judith L. Connor and

Walter H. Adey

Preliminary Observations on
the Algae, Corals, and Fishes
Inhabiting the Sunken Ferry
“Fujikawa Maru’ in Truk
Lagoon

by Roy T. Tsuda, Steven S.
Amesbury, and Steven C. Moras
Chemistry of Freshwater
Pools on Aldabra

by A. Donaldson and B. A.
Whitton

Issued by

214.

215.

216.

217.

218.

219.

May 1977

Observation on Redox Poten-
tial in Freshwater Pools on
Aldabra

by B. A. Whitton and M. Potts

Algal Flora of Freshwater
Habitats on Aldabra

by A. Donaldson and B. A.

W hitton

Terrestrial and Freshwater
Algae of Three Western
Indian Ocean Islands (Astove,
Farquhar and St. Pierre )

by B. A. Whitton and A.

Donaldson

Terrestrial and Swamp Algae
from Three Islands in the
Chagos Archipelago, Indian
Ocean

by B. A. Whitton, A. Donaldson,
D. J. Bellamy and C. Sheppard

The Holocene Reef Systems
of Eastern Martinique, French
West Indies

by Walter H. Adey, Patricia J.
Adey, Randolph Burke and

Leslie Kaufman

Island News and Comment

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

ATOLL RESEARCH BULLETIN
NO. 208

NOTES ON PLANTS OF THE GENUS CAULERPA
IN THE HERBARIUM OF MAXWELL S. DOTY
AT THE UNIVERSITY OF HAWAII

by Wm. Randolph Taylor

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

May 1977

NOTES ON PLANTS OF THE GENUS CAULERPA
IN THE HERBARIUM OF MAXWELL S. DOTY
AT THE UNIVERSITY OF HAWAII

by Wm. Randolph Taylor !

In 1966 on the invitation of Professor Maxwell S. Doty I spent
several weeks at the University of Hawaii to review his holdings of
certain algal genera, particularly those of Cauler pa (Chlorophyceae-
Siphonales), and especially the Pacific specimens. 13 In view was
the rectification of the determinations, where necessary, and the
organization of the genus as represented in the collection from a
single viewpoint. This I did at that time, but because of other work
in hand was unable to consider preparing a report for publication.

More recently, pressed to make such a report, I sent for and
received the more troublesome part of the material on loan, and have

1 Department of Botany, University of Michigan.

2 This study while in Honolulu was conducted under an Atomic Energy
Commission Contract no. AT(04-3)-235 to Prof. Maxwell S. Doty, and
later continued at the University of Michigan under Nat. Sci. Found.
grant GB-3186 in part, for which help the writer is most thankful.

3 The last names of the principal collectors of the materials reported
in this paper are associated with the collection number in each case,
but in order that their identity may be more accurately defined their
initials where available are given in the following list:

Alcala, A.C.; Almazan, M.; Alvarez, V.; Buggeln, R.; Cooper, J.;
DOLLY Moen PM GaAn; Cow es HONG, uu hiet a Gislmarstam.) sMure Hae lg. Bis
Haltisiteadi. Sear Hastings, CG. Ha HORWaEe zy ines KANG). WJ). Bier Kae, Grek,
Littler, M.M.; Long, C.R.; Matsui, T.; Menez, E.G.; Newhouse, G.;
Pages, P.; Randall, J.; Rogers, D.P.; Sachet, M.-H.; Santos, G.A.;
SeLDUnBENy weir IOlCGtdLEO, Aa. SOrokin, Yo7.otone, B.C. 3 Strasburg),
DeWey LEON) Who GeCw USudal, Ra welasquez, Gat. 7; Villaroya, Mt. 5
Wainwright, S.A.; Wreede, R.E. de.

(Manuscript received January 1974--Eds.)

gone over it again. On the basis of my 1966 notes, review of
material accessioned since then, and reconsideration of the specimens
loaned in 1972 I present here a list of representative western and
central Pacific Caulerpa material in that herbarium, excluding Japan,
Australia and New Zealand. This is probably the largest representa-
tion of that genus from the area in any herbarium. Because of the
very great duplication of specimens from certain localities I only
cite a few examples from each, but attempt to include all significant
places which could be located geographically.

When I agreed to go over the great bulk of material again and to
prepare this report I was unaware that G.C. Trono had (1968) published
on the Caroline Islands material, or was later to publish (Trono 1972a,
b; 1973) on Philippine material , leading to great duplication of
effort without commensurate yield of new data. However, Since this
present paper covers the central and western Pacific (south of Japan,
north or east of Australia) in general, and since the citation again
of Caroline Island and Philippine specimens could be kept to a
minimum, it seems wise to include them where necessary to complete the
tale of Honolulu holdings. ;

It does not seem needful to review here the literature on Pacific
green algae, as this is not an historical paper. Nor is ita
summation of all accounts of Caulerpa in the western Pacific.
Necessarily the basis for any account of the genus is the monograph by
Mme Anna A. Weber-van Bosse (1898). Mention of the genus occurs in
many of the more general papers. A good deal of this literature has
already been brought together, in Okamura (1916), Taylor (1950, 1966),
May (1953, 1966a, b), and Dawson (1956, 1957), while Gilbert has also
listed it (1959) and Abbott (1961), Trono (1968) and Womersley and
Bailey (1970) have extended it still further.

Nor can too detailed conclusions be drawn respecting the
geographical ranges of the species listed. For this all other major
herbaria concerned with the area should be consulted. Nevertheless
very many extensions of range are involved, and we do get some idea of
what species are commonest throughout the central and western Pacific.
It is clear that Caulerpa lentillifera, C. mexicana, C. peltata, C.
racemosa in particular, C. serrulata and C. urbilliana are by far the
commonest. The substantial representation of the very small C.
ambigua about Hawaii probably reflects the closer observation possible
from a well-equipped scientific center. Other small species seldom
mentioned, like C. elongata, C. webbiana and smaller forms of C.
fastigiata probably have been missed by reason of a lack of close
search.

Some species are nearly ubiquitous in the subtropics and tropics
around the world, and in this we could include, with others, C.
cupressoides, C. racemosa and C. sertularioides. On the other hand
C. lentillifera does not come into the Atlantic flora, though found
from the Marshall Islands to the Red Sea, parallelling C. urvilliana

ranging from the Line Islands west to Mauritius, absent from the
Atlantic. Disproportionate abundance is suggested in some cases, for
C. serrulata seems to be far more abundant in Pacific collections than
West Indian, though found in both oceans, while the reverse seems to
be the case with C. cupressoides.

Less familiar species are harder to evaluate as to range. They
may have a scattering, sparse distribution. First found in the
Marshall Islands, C. bikinensis is now known to reach the Tuamotw group.
From the Carolines to the Philippines is about the range of C.
lessonii. We must not discount the possibility of migration to new
areas after a long period of limitation to an old range. For instance,
C. scapelliformis, long known from the Red Sea to south Australia and
Japan has of recent years turned up in Brazil and Guadeloupe, and by
1966 abundantly in Barbados and Antigua, areas where such a relatively
large and distinctive alga could hardly have grown unrecorded through
the previous decades. If one is reluctant to deny its earlier
presence there one must remember the case of Codium fragile (Sur.)
Hariot, much more closely documented in its migration from the Pacific
to the eastern coast of the United States, eventually reaching
northern New England. There it was never seen until recent years, in
an area of intensive observation of the marine algae, but it is now
abundant enough to constitute a nuisance to the shellfish industry.

List of specimens

Caulerpa ambigua Okam.
HAWAII
LAYSAN I., from the s.w. side, Tsuda no. T-548, 6 xii 63.
KAUAI I., Anahola, Doty no. 10224, 7 ii 52.
OAHU I., Waikiki Beach, Choy, 16 iii 54, Doty no. 13063a, 4 xii
55, Ewa Beach, Littler no. 26500, 2 vi 70.

Caulerpa antoensis Yamada!
MARSHALL ISLANDS
ARNO ATOLL, Ine Village, Horwitz no. 9040, 10 vii 51.
CAROLINE ISLANDS
PTINGELAP I., Menez no. 21926, 1 vii 60.
MOKIL IDS., Urak I., Menez no. 21344, 29 vi 60.
PULUWAT I., Menez no. 23236, 7 viii 60.
EFALUK IDS., Ifaluk I., Menez no. 23055, 10 viii 60; Ella I.,
Menez no. 23232, 10 viii 60; between Ifaluk I. and Falalap I.,
Menez no. 23102, 10 viii 60.

A fragment of Yamada's original material (Yamada 1941, 1944a, b)
from Ant Atoll near Ponape Island in the eastern Carolines, which he
kindly sent me after my Bikini book appeared, showed very short axes

1 ¢. arenicola Taylor 1950.

and irregularly disposed ramelli, but his 1941 figure, far better than
that of 1944a which accompanied his formal description, shows them
bilaterally disposed in large part. The ramelli on his plant are a
little thicker than on the Bikini plants, reaching at least 200 um.

On some of the Caroline Island specimens they were only 135 um diam,
but in others: reached 264 -um.

Caulerpa bikinensis Taylor
TUAMOTU ARCHIPELAGO
RAROIA I., Tomokgolu (? Tomogagie), Doty & Newhouse no. 11426, 29
Vil (5.20

Caulerpa brachypus Harv.
LINE ISLANDS
PALMYRA I., Long no. 2833, 27 xi 64.
MARSHALL ISLANDS
MAJURO ATOLL, Seiburth & Sorokin, no number, ii 70.
CAROLINE ISLANDS
PULUWAT I., Menez nos. 23064, 23091, 7 viii 60; these included
strongly dentate specimens.
YAP I., Menez no. 21489, 19 viii 60.
PHILIPPINE ISLANDS
SAMAR I., Samar Prov., Guiuan, Santos no. 20466, 29 iv 48 (not
dentate), Guiuan, Hinatunglan, Santos no. 26462, 17 vi 69 (very
attenuate).
NEGROS I., Negros Oriental Prov., Siquijor I., Carrio Solong-on,
Alcala no. 20522, n.d., Mefiez nos T-1132, T-1137, 28 ix 67,
Santos nos. 26042, 26047, 22 v 69, 26197, 26221, 26 v 69.
PALAWAN I., Palawan Prov., Cujo I., Menez 16537, 30 iv 58.
MINDANAO I., Surigao del Norte Prov., Surigao, Menez no. 18363,
n. d.; Zamboanga City Prov., Little Santa Cruz I., Santos 26544,
Poy, Nyalalal, Zils
SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., Doty &
Hair no. 25418, vi 66; Siasi Group, :Siasi I., Doty & Hair no.
25537; Tawitawi Group, Bilitan I., Doty & Hair noi/25499\. Siiway
66; Sibitu Group. Tumindao I., Sitangkai, Doty & Hair no..
25451], ivi. 166".

Caulerpa cupressoides (West) C. Ag.
LINE ISLANDS
FANNING I., Tenupa, Wainwright no. 20101, 2 vii 63
MARSHALL ISLANDS
ARNO ATOLL, Ine Village, Doty no. 9001, 26 v 51.
CAROLINE ISLANDS
KUSAIEI., Lele Harbor, Menez no. 15852, 16 vii 60, Tafsanak
Village, Mefiez no. 23407, 16 vii 70; Utwa I., Menez no. 23248,
7 “Wall 560%
PINGELAP I., Menez nos. 21312, 21935, 7 viii 60.
TRUK IDS., Quoi I., Menez no. 23191, 2 viii 60; Moen I., Menez
15901, 31 vii 60; Fefan I., Menez no. 23081, 8 viii 60; Falas I.,
Menez no. 23513, 30 vii 60.

ULUL I., Mefiez no. 15883, 6 viii 60.

PULUWAT eT oneMenezenos. .23131, 23235, 7, viii 60.

IFALUK I., Menez no. 23126, 10 viii 60; between Ifaluk I. and
Falalap I., Menez no. 23063, 10 viii 60.

SOROL I., Menez nos. 23049, 23162, 13 viii 60.

YAP I., north of harbor channel, Menez nos. 21572, 23092, 18
viii 60.

HELEN I., reef, Mefiez no. 15415 p.p., 28 viii 60.

PHILIPPINE ISLANDS

LUZON I., Quezon Prov., Baler, Diksalarin, Velasquez no. 16469B,
2351766).

CATANDUANES I., Catanduances Prov., Benticayan s. of Vinticayan
Point, Doty & Velasquez no. 16655, 21 v 58.

SAMAR I., Samar Prov., Guiuan, Pagnamiton, Doty & Alvarez no.
WAS24), (Aaa oO 5).

MINDANAO I., Zamboanga City Prov., Sacol I., Doty & Hair no.
25308 a2 2a Oo

SULU ARCHIPELAGO, Jolo Group, Pangasinan I., Doty 25421, vi 66;
Tap Group, —Siasi.,.. ,~voty,& Hair no. 25525, tO yiii 66;
Tawitawi Group, Tawitawi I., Balimbing Pt., Menez no. 1101,
235x Gi. Bilavtan ©, Dotyes Hair no. 25484, 5 vill 66.

Caulerpa elongata Weber-van Bosse
MARSHALL ISLANDS
ARNO ATOLL, Horwitz no. 9046, 10 vii 51.
CAROLINE ISLANDS
VAP i. .Menezeno. 23780, 19) vidi 6O.
PHILIPPINE ISLANDS
LUZON I., Sorsogon Prov., Bulusan, Kraft no. 349, 17 Mar. 68.

Caulerpa fastigiata Mont.
CAROLINE ISLANDS
KUSAIEI., Tafansak, Menez no. 21955, 16 vii 60; Utwa I., Menez
MOw ZOLA ley vals (GO!
PHILIPPINE ISLANDS
SAMAR I., Samar Prov., Guiuan, Pagnamiton, Doty & Alvarez no.
WAS s> 4) atal (OS).
NEGROS I., Negros Oriental Prov., Siquijor I., Carrio Solong-on,
Santos nos. 26052, 26160, 22 v 69.

Caulerpa fergusonii Murray
PHILIPPINES

LUZON I., Sorsogon Prov., Bulusan, Santos no. 26522, 9 viii 71.

Caulerpa lentillifera J. Agardh
CAROLINE ISLANDS
TRUK IDS., Fefean I., Menez without number, 28 vii 60.
PHILIPPINE ISLANDS
LUZON I., Batangas Prov., Matabungkay, Santos no. 20450 p.p.,
3 iv 68. Pangasinan Prov., Hundred Islands, Virgin Cave I.,
Doty & Menez 16342, 15 ii 58.

CATADUANES I., Cataduanes Prov., Virac Point, Doty & Velasquez
NO VO7AG6 ie 2 O) pVae Doe
SAMAR I., Samar Prov., Guiuan, Hinatunglan, Santos 26461, 17 vi
69.
CEBU I., Cebu Prov., Cebu City market, n. coll., no. 18318,
14 vii 58; Mactan I., Cordova, at Caulerpa-farm, Doty no.
16075, 17 i 58, Calawisan, Menez no. T-1027, 8 ix 67.
NEGROS I., Negros Oriental, Siquijor I., Solong-on, Alcala
nos. 20520, 20523, Santos nos. 26198, 26 v 69.
PALAWAN I., Palawan Prov., Pto. Princesa, Inagawan, Velasquez
no: (3079, ‘20 vi 51> Cujo. 1. Cujo, Menez: /6535), 30 mivaSor
MINDANAO I., Davao Prov., Davao, Sasa Wharf, Doty no. 18056,
26 vi 68; Zamboanga Prov., Little Santa Cruz I., Menez no.
T-1057, 10 ix 67; Sacol I., dwarf, Menez no. T-1096, 19 ix 67.
SULU ARCHIPELAGO, Sulu Prov., Tapul Group, Siasi I., Doty & Hair
no. 25530, 10 viii 66, Sibaud I., Santos nos. 26545, 26559,
14 ix- 71; Manubal; i: , Doty.& Hair no.,:25529)77 lO. viii76-) Tapaan
L., Doty & Hair no. 25574, Trono T4064, T4087, 29° xii 7O:
Tawitawi Group, .Bilitan.1.,, Doty & Hair nos: 255017" 255107;
5 viii 66; Tawitawi I., Balimbing Pt., Menez no. T-1102,
2 SON e

The size of the ramelli on plants assignable to this species
seems to vary greatly. In most specimens they are small, as we find
in Cebu I. no. 18318, where the diameter is about 1.5 mm. In contrast,
one places Bilitan I. no. 25511, where they are over 4 mm in the dry
state. Similar coarse plants were found at Rongelap lagoon (Taylor
1950, p. 67) and it is possible that we are dealing with two species,
rather than one very variable one.

Caulerpa lessonii Bory
CAROLINE ISLANDS
TRUKGIDS..,..nefocan lL. , Menez no. 23029) 28 vii, 6O-
YAP I., Yap Harbor, Menez no. 23144, 18 viii 60.
PHILIPPINE ISLANDS
LUZON I., Pangasinan Prov., Lingayan Gulf, Cabarruyan I., Anda,
Santos no. 20406a, 22 ii 68. La Union Prov., San Fernando,
Almazan no. 1300, 3 iii 68. 5

This group of Caulerpas from the islands gave me much concern
because of a marked similarity to C. cupressoides var. lycopodium f.

elegans (Crouan) Weber-van Bosse of the West Indies. They also
resemble the figures and descriptions of C. lessonii Bory and
C. distichophylla Sond. However, Prof. H.B.S. Womersley informs me

that the ramelli of C. distichophylla are compressed, not terete as
described by Mme Weber-van Bosse (1898, p. 341), and as are those of
our plants, and as the type locality for the species lessonii is in
the Caroline Islands the probability is strong that plants like these
were what Bory had in hand, though I have not actually had the type to
examine. The relation of the v. lycopodium f. elegans to C.
cupressoides seems to me rather insecure, and these lessonii plants
would have even less claim to relationship.

Caulerpa macrodisca Decne.
PHILIPPINE ISLANDS
CEBU I., Mandaue (Mandawi), north of Cebu City, Quijano no. 429,
6 vii 64.

Caulerpa mexicana (Sond.) J. Agardh
HAWATIAN ISLANDS
KAUAI I., Haena Point, Randall no num., dwarf, 11 xi 51.
OAHU I., Hanauma Bay, Doty no. 3936, 13 xi 51; Lanikai,
Strasburg no num., 26 x 52; Maili, Rogers no nun.,
11 v 56; Waimanalo Beach, C.F. H7Tastings no. 282, 15 ix 68.
LANAI I., Huawai Bay, Degener no. 28691, 9 i 64; Kapihaa Bay,
Doty & Lee no. 22071, 27 xi 60; Kaumalapau, Doty & Lee no. 22111
AAS} 5% (EXO)e
MAUI I., Lahaina, Matsui no. 12862, ix 55.
PHILIPPINE ISLANDS
LUZON I., Sorsogon Prov., Bulusan, Santos no. 26253, 4 vi 69;
La Union Prov., San Fernando, Almazan no. 1306-2, 3 iii 65.

In western Atlantic waters C. mexicana and C. taxifolia are
readily distinguished. Only in dwarfed plants about 1 cm tall are
the characters so ill-developed as to lead to indecision. hast as
not so in the Pacific. Intermediates are not infrequent, and numbers
282, 12862, 22071, 22111, 26253 may be considered examples tending
that way. It seems to me possible that the basic stock simply
evolved two types clearly in one ocean but did not proceed so far in
another. To blindly disregard that achieved potential shown in the
Atlantic and reduce C. mexicana to synonymy seems to me quite the
wrong attitude. Better to recognize that some plants have, in the
Pacific, the full characters of C. mexicana and call them by that
name, others of C. taxifolia, and that intermediates exist either by
lack of complete evolutionary divergence or, as an alternative
hypothesis, by hybridization.

To point out the characters of these two species in their distinct
forms, I would suggest the following. For C. taxifolia I expect
relatively narrow blades, usually long, with a narrow axis, and with
ramelli of the order of O.75-1.20 mm in width, about 5-12 times as

long. There is little tapering toward the base, and tapering to the
tip is gradual. They are strongly compressed, but are only about 3
times as broad as thick. Bérgesen's illustrations (1913, figs. 104,

105) are good.

For C. mexicana I expect relatively broader blades, usually
short, with broader axes and broader ramelli up to 1.0-2.5 mm wide,
2.5-5.0 times as long. They show substantial narrowing at the base,
and at the tips they are more abruptly rounded-tapered as a rule.
They are flat, and may be 10-30 times as broad as thick. They are
often so close that they overlap distally, though in the var. laxa
(Weber-van Bosse) Taylor they are widely spaced. Mme Weber-van
Bosse's figure (1898) pl. XXIV no. 2 (as C. pinnata fa. mexicana),

although of a rather small individual, represents this species fairly
well. Perhaps mine (1960, pl. 12, fig. 5) is a little better.

Caulerpa peltata Lamx.
HAWAIIAN ISLANDS

LAYSAN I., west side, Tsuda no. T-523, 6 xii 63.

NECKER I., Long no. T-943, 25 ix 64.

KAUAI I., Anahola Bay, Doty no. 10022, 2 ii 52.

OAHU I., Oli Bridge near Koko Head, Doty no. 8382; Kaneohe Bay,
Moku-O-Loe (Coconut) Island, Littler no. 20538, 10 x 68.

LINE ISLANDS
FANNING I., Danger Point Pool, Wainwright no num., 9 vii 63.
MARSHALL ISLANDS

ARNO ATOLL, Ine Village, Horwitz no. 9080 with disks to 9 mm diam.

dry. 26 7 Si.
CAROLINE ISLANDS

PONAPE I., Epwelkapw, Menez no. 15675, 20 vi 60.

TRUK IDS., Truk I., Menez no. 21059, 31 vii 60; Moen I., Menez
no. 23693, 29 vii 60; between Moen and Falo Ids., Menez no.
23383, 29 vii 60: d

PALAU I., Iwayama Bay, Menez no. 15102a, 22 viii 60.

PHILIPPINE ISLANDS

LUZON I., Quezon Prov., Baler, Digisit, Velasquez no. 16989,

24 iv.58; Sorsogon Prov. , Bulusan, Santos no. ,26513,.6.viai 7.

MINDORO I., Mindoro Oriental Prov., Puerto Galera, Sabang Cove,
Velasquez no. 1632, 8 v 48.

NEGROS I., Negros Oriental Prov., Dumaguete City, Santos no.
26012, 20 v 69, Matio Beach, Santos no. 26080, 24 v 69,
Uy-Pitching Beach, Santos no. 26149 (very good), 25 v 69,
Mataio Beach, Santos no. 26024 (very good), Lo-Ok Beach,
Santos no. 26244 (superb), 28 v 69.

MINDANAO I., Zamboanga Prov., Zamboanga City, Arena Blanco,
Santos no. 26535, 24) vada le.

SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., Doty &
Hair no. 25245, 1966.

Caulerpa pickeringii Harv. & Bail.
TUAMO TU ARCHIPELAGO
RAROIA ATOLL, outer reef, Doty & Newhouse no. 11162, 9 vii 52;
Okarekare, Doty & Newhouse no. 11228, 1 viii 52.

Caulerpa racemosa (Forssk.) J. Agardh
HAWAIIAN ISLANDS

LAYSAN I., Tsuda no. T-541, T-594, 6 xii 63.

NECKER I., Long no. T-944, 25 ix 64.

KAUAI I.,.Anahola Bay, Doty.nos...9999,; 10231,,.2-7.ii1 52.

OAHU I., Hanauma Bay, Doty no. 10672, 3 v 53, no. 10616, 5 iv 53;
Manana (Rabbit) I., Doty no. 8833, 22 iv 51; Kaneohe Bay,
Doty no. 8459, 24 xi 50; Ulupau Head, Kii Point, Strasburg no.
8596, 8 ii 51; Maili, Rogers no num., 11 v 46.

MAUI I., LaPerouse Bay, Doty no. 13187, 17 xii 55; McGregor Point,
Matsui & Gilmartin no num., 24 vi 65.

HAWAII I., Kona, Kealakekua, Randall no. 10041, 10 ii 52.
GILBERT ISLANDS
TARAWA ATOLL, Betia, Cooper no. 18823, vii 62 (with several
varieties).
MARSHALL ISLANDS
ARNO ATOLL, Ine, Horwitz no. 9047, 10 viii 51.
CAROLINE ISLANDS
TRUK IDS., Dublon I., Menez no. 273, Nb. 59, 31 vii 60; Fefan I.,
Menez no num., 28 vii 60.
YAP I., Yap harbor, Menez nos. 21985, 23538, 19 viii 60.
PALAU IDS., Iwayama Bay, Menez no. 361, Nb. 59, 22 viii 60;
Peleliu, Doty no. 20530, 23 ix 68.
PHILIPPINE ISLANDS
LUZON I., Albay Prov., Calayucay Bay, Doty & Velasquez no. 10891,
19 v 58; Sorsogon Prov., Bulusan, Santos no.s 20490, 15 v 58,
26771, 3 vi 69; Batangas Prov., Matabungkay, Santos nos. 1306,
20 iii 65, 20449, 3 iv 68; Pangasinan Prov., Lingayan Gulf,
Hundred Islands, Doty & Menez no. 16267, 16 ii 58, Alaminos,
Cangaluyan Reef, Menez no. T-1015, 4 ix 67, Cabarruyan I.,
Anda, Santos no. 20401, 22 ii 68.
CATANDUANES I., Catanduanes Prov., Benticayan Bay, Doty &
Velasquez no. 16654, n. d.; Virac Point, Doty & Velasquez
nos. 16743, 6744, 20° v 58:
MINDORO I., Mindoro Oriental Prov., Puerto Galera, Doty no.
10909, 2-4 xii 53, Baléte Cove, Velasquez no. 5306, 23 iv 62.
SAMAR I., Samar Prov., Guiuan, Santos no. 20465, 26 iv 68,
Hinatunglan, Santos nos. 26368, 26421, 15 vi 69.
CEBU =L4,--Cebu’ Prov. ; Danao, Pages ‘no. 235, -3' xi 59; Liloan,
Pages no. 78, 10 vi 60; Mactan I., Calawisan, Santos no. 26507,
21 vii 71, Beramis Dam, Santos no. 26507, 21 vii 71, Caulerpa-
farm, Doty & Alvarez no. 14625, n. d.; Olango I., Sta. Rosa,
DONE INS)S” IUGOW/ 7/5, TalS le
PALAWAN ARCHITELAGO, Palawan Prov., Inlulutoc Bay, no coll.,
no. 5/50, 29. 1v 6a. "Cuyo! tds. Cuyo, no. coll. ;"6000a, I'2°v 64.
Menez no. 16536, 30 iv 58, Sandoval-Paras, no. 13066, vi 65.
MINDANAO I., Surigao del Norte Prov., Hinatuan Mawes I., Mefiez
no. 18005, 7 vii 58; Zamboanga City Prov., Sacol I., Doty &
Hair no. 25394, 22 vi 60, Pitogo, Doty & Hair no. 25353,
PAO) aval; (XS)e
SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., Doty &
Hair no. 25419, 25424, 1966; Tapul Group, Siasi, Sibaud I.,
Santos’ nos. 26556, 26558, 14 ix 71, Manubal I., Doty & Hair
nos. 25524, Tapdan i., Doty & Harr no. 25568, TO viii ‘6;
Tawitawi Group, Bilitan I., Doty & Hair nos. 25498, 25500,
25504, 5 viii 66; Sibutu Group, Tumindao I., Sitangkai, Doty
& Haay nos. 25450, 25452), 1966.
INDONESTA
JAVA, Bay of Banten, Pulau Dua, Soegiarto no. 8, n.d.
Since this species often grows tightly attached to the exceedingly
rough, eroded intertidal rocks characteristic of tropical coral
islands, it is commonly difficult to detach, spread out and display

10

the plants well. In such exposed places they form compact interlaced
masses, and do not clearly develop distinctive varietal characters.

In fact, plants which grow in relatively quiet water are those which
show them. Pacific Caulerpas show features similar to those shown

by western Atlantic individuals, although not always the same in range.
For the purposes of this list it did not seem wise to attempt to list
the varietal type of each collection, although the specimens as they
were studied were annotated when appropriate. It may be of interest
to point out a few distinctive forms. The last specimen listed, for
instance, Soegiarto no. 8, appears to be the var. corynephora (Mont.)
Weber-van Bosse. The commonest variety of most oceans, var.
clavifera (Turn.) Weber-van Bosse, seems to be approximated by Bilitan
I. no. 25504 and Bulusan no. 20490. Mactan I. no. 26507 may be
associated with var. lamourouxii (Turn.) Weber-van Bosse. For var.
laetevirens (Mont.) Weber-van Bosse we may pick Olango I. no. 16077
and Bilitan I. no. 25498 as reasonable examples. Under var.
occidentalis (C. Agardh) Bé¢rg. may come Andanno. 20401 and Siasi I. no.
26556. Var. turbinata (J. Ag.) Eubank (which I have previously
called var. chemnitzia) shows a wide range of size. Manubal I. no.
25524a and several others seem not too far from the ordinary concept
of the variety. However, Guiuan no. 26421 is very much taller than
usual, and Bulusan no. 26771 is an extremely lush variant which it
would be hard to distinguish from C. matsueana Yamada (1944a p. 28),
which may not be worthy of specific rank.

Caulerpa selago (Turn.) C. Agardh
PHILIPPINE ISLANDS
SULU ARCHIPELAGO, Sulu Prov., Sibutu Group, Tumindao I., Sitangkai,
Doty & Hair no. 25449, 1966.

Caulerpa serrulata (Forssk J. Agardh, var. serrulata
HAWAIIAN ISLANDS
OAHU I., Hanouma Bay, Doty no. 10670, 3 v 53; Hanula Park, Doty
NO.) S66L.,.:°22 iv 51; Haleiwa, Fong no. -num.s, +17 )-55:.
MAUI I., Kihei, Matsui no. 12874, n..d.
LINE ISLANDS
PALMYRA I., Halstead no. 12190, 27 iv 53; Barren I., Doty no.
18586, .280x11 .59:.
GILBERT ISLANDS
TARAWA ATOLL, Betio, Cooper no. 18809A, 18810, vii 62.
CAROLINE ISLANDS
PONAPE IDS., Mantapeitak I., Menez no. 21543, 20 vi 60.
TRUK, /EDS..,. Quoi oL.,,. Menez no. 72101472 viii) 60.
PHILIPPINE ISLANDS
LUZON I., Sorsogon Prov., Gubut, Villaroya no num., iv 65; Albay
Prov., Albay Gulf, Lubas Point, Doty & Velasquez 16846B, 18 v
58; Calayucay Bay, Doty & Velasquez no. 16892, 19 v 58; Quezon
Prov., Baler, Digisit, Velasquez 16986, 24 iv 58; Batangas Prov.,
Matabungkay, Santos no. 1306, 20 iii 65; Pangasinan Prov.,
Caburruyan (Anda) I., Santos no. 20406b, 22 ii 58.

ial

MINDORO I., Mindoro Oriental Prov., Puerto Galera, Medio Ids.,
Velasquez no. 3495, 12 v 53.

CEBU I., Cebu Prov., Danao, Sabang, Pages no. 99, 23 xi 60.

MINDANAO I., Zamboanga Prov., Sacol I., Doty & Hair no. 25392,
22 vi 66.

SULU ARCHIPELAGO, Sulu Prov., Tapul Group, Mandubal I., Doty &
Hair no num., 10 viii 66; Tawitawi Group, Bilitan I., Doty &
Hair no. 25502, 5 viii 66; Sibutu Group, Tumindao I.,
Sitangkai, Doty & Hair no. 25448, 1966.

Caulerpa serrulata var. boryana (J. Agardh) Weber-van Bosse
CAROLINE ISLANDS
PONAPE IDS., Nanmatol I., Menez no. 23311, 23 vii 60
VAPIOUDSi0,, Yap, Menez no. 215537 18 vari 60.
MARIANAS ISLANDS
SAIPAN I., Maniagassa I., Doty no. 20326, 24 xi 67.
PHILIPPINE ISLANDS
LUZON I., Pangasinan Prov., Hundred Ids., Cuenco I., Cuenco Cave,
Doty & Menez no. 16204, 15 ii 58.
MINDANAO I., Davao Prov., Davao s. of Sasa Wharf, Doty no. 18055,
A,» ip Bike

Caulerpa serrulata var. spiralis Weber-van Bosse
LINE ISLANDS
PAEMYRA I., King no. 0835, 8 1 53; Barren E:, Doty no. 1860,
Sandie Doty NOneteoalii) 29i xa 59).
TUAMOTU ISLANDS

RAROIA ATOLL, Doty & Newhouse no. 11010, 30 vi 52.
SOCIETY ISLANDS

MOPTHAWA be sachet no. 9927 18 vai (63).

GILBERT ISLANDS

TARAWA ATOLL, Betio, Cooper no. 18803, vii 62.
MARSHALL ISLANDS

ARNO ATOLL, Arno I., Doty no. 9225, 30 vii 51; reef off Motol-En

in, HoLwitz) now 94947) 2) yas: Sil Kabunlok £., Horwitz no. Gil64',
imaGle
CAROLINE ISLANDS

PONAPE I., Mantapeitak I., Menez no. 21678, 20 vi 60.

TRUK IDS., Moen I., Menez nos. 15885, 29 vii 60, 15926, 1 viii 60.
PHILIPPINE ISLANDS

LUZON I., Sorsogon Prov., Gubat, Villaroya no. 13067, 6 v 65.

Batangas Prov., Matabungka, Santos no. 13065, 20 iii 65.
CATANDUANES I., Catanduanes Prov., Benticayan Bay, Doty &
Velasquez no. 16653, 21 v 68.

MINDORO I., Mindoro Oriental Prov., Puerto Galera, Uyenco no.
ESOGS) as de

SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., Doty &
Hatten, 254s nn vas 166%

When the Caulerpa materials were studied in Honolulu in 1966 and
reviewed in 1967 the serrulata specimens were recorded as typical or
not distinguishably different, or, as nearer one of two particular
varieties. They were not studied again in 1973, and are here reported
as annotated in 1966-67. Trono's attempt to separate off a non-

12

spiralled form (Trono 1973, p. 9, fig. 19) as C. hummii Diaz-Pifferer
cannot be supported, for various degrees of twisting or flatness are
to be found in C. serrulata. This is as true in the West Indies as
in the Pacific. From the Diaz-Pifferer and Trono illustrations I
would certainly place their plants in C. serrulata, and, if one
wanted a varietal designation, near var. pectinata as understood by
Mme Weber-van Bosse (1898), near her figures 4-6 on Pl. XXVI.
Actually, the illustrations labeled C. hummii by these authors are
very much closer to the type of C. serrulata (as Fucus serrulatus
Forsskal), a photograph of which is before me, than are the
elaborately spiralled forms, such as that illustrated by Trono (1973,
Salto fe nA)) The name C. hummii Diaz-Pifferer (1969, pp. 12-17) should
be placed in the synonymy of C. serrulata (Forssk.) J. Ag.

It has been put to me that the difficulty of distinguishing
between C. cupressoides, C. serrulata and C. urvilliana is very great
in Pacific material. This may at times be so, and when only dwarfed
or abnormally attenuate specimens are available will always be so.

To start with, C. cupressoides.typically has terete erect axes,
and the ramelli are terete, short in some forms, longer in others.
When the ramelli are reduced to 3 rows the axis may be less clearly
terete than when the ramelli are pluriseriate or irregularly placed.
When they are biseriate the ramelli may be quite long, as inC.
cupressoides var. lycopodium fa. elegans, or reduced nearly to teeth,
as in var. flabellata, and then the axis is compressed.

In C. urvilliana the erect axes are terete in the stalk part, but
become compressed upwardly, and in full development above triangular,
where the teeth are triseriate. Ramelli are, then, represented by
teeth, not by cylindrical structures. While often somewhat flexuous
the axes do not become spiralled.

Also showing the lower axis at least briefly terete, C. serrulata
axes are flattened above, sometimes broadly, and the margins are
dentate. The type specimen is very small, with the flat branch axes
not spiralled. The branch axes in this species do range from straight
to elaborately spiralled, and in these the teeth may become somewhat
dislocated and those on the inner border of the spiral may be reduced
or suppressed. The illustrations purporting to be this species in my
Bikini book: (1950), pl.°30, f£igs...1,. 2) .were incorrectly identasied:
they are forms of C. urvilliana. That on pl... 29, fig. shouldgbe
referred to C. serrulata var. boryana: the name used at that time
would not satisfy the present rules of nomenclature.

Caulerpa sertularioides (Gmel.) Howe
HAWAIIAN ISLANDS

KAUAI I., Anahola Bay, Doty no. 9998, 2 ii 52; Aliomanu Beach,
DOEV NOs 79925) paolo 2)

OAHU I., Mokuleia, Doty no. 12234, 9 xi 52; Kaneohe Bay, Moku-O-
Loe. I..,. Doty,no. 8115, .3 x 50;° Haliona, Rogers no num. ,.. 3) Vv 46;
Kuapa Pond, Engard no num., 21 iv 43; Mali near Lua-Lua-Lei,
Doty nOse S7SAT sl iisis. 5 Jus

3}

GILBERT ISLANDS
TARAWA ATOLL, Betio, Cooper no. 18802, vii 62.
CAROLINE ISLANDS
TRUK IDS., Moen I., Baker Docks, Stone no. 2181, vii 57.
PHILIPPINE ISLANDS
LUZON I., Quezon Prov., Baler, Diksalarin, Velasquez no. 16469,
2351s So, Digaistt, no. 6985, 24 ix 58); Rizal) Prov) sManila
Bay, Velasquez no. 5577, 4 ii 64 (excellent), South Harbor,
Doty & Menez no. 16297, n. d.; La Union Prov., San Fernando,
Almazar no. 1306, 3 iii 65.
CATANDUANES I., Catanduanes Prov., Benticayan, Doty & Velasquez
no. 16685b, Virac Point, Doty & Velasquez no. 16471, n. d.
CEBU I., Cebu Prov., Mactan I., Lapu Lapu, Calawisan Caulerpa-
farm, Doty & Alvarez no. 14626, 4 iv 65.
PALAWAN ARCHIPELAGO, Palawan Prov., Cujo Ids., Cujo I., Menez
NOPE LO SSO eniewacls
SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., n. coll.,
no. 25416, n. d.; Tapul Group, Tapaan I., Doty & Hair no.
25573, 10 viii 66, Tapaan Lagoon, Santos no. 26547, 10 ix 71,
Sibaud I., Doty & Hair no. 25564, 10 viii 66.

Both fa. brevipes (J. Agardh) Sved. and fa. longiseta (Bory)
Sved. are represented in this material, though perhaps less distinct-
ively than in Caribbean material. To the former we may assign Kauai
no. 9998 and Cujo no. 16539, to the latter Kaneohe Bay no. 8115 and
Mochi no. 21/S8il: The plants from South Harbor no. 6297 showed
exceptionally long and slender ramelli, which reach a length of 18 mm.

Caulerpa seuratii Weber-van Bosse
TUAMOTU ARCHIPELAGO
RAROIA ATOLL, Raroia, Doty & Newhouse 11639, 9 viii 52; Oneroa,
Boty & Newhouse no. 11499, 29 vii 52 (excellent); Tomongoru
(? Tomogagie), Doty & Newhouse no. 11427, 29 vii 52;
Teuriamote, Doty & Newhouse no. 11476, 26 vii 52.

Caulerpa taxifolia (Vahl) C. Agardh
CAROLINE ISLANDS
TRUK SED Sy, Ouoly lh. Menezeno. 253077) 2 svasial (60.
YAP ,I., Menez no. 21897, 18 viii 60.
PHILIPPINE ISLANDS
NEGROS I., Negros Oriental, Siquijor I., Menez T-1138, 28 ix 67,
Carrio Solong-on, Santos no. 26187, 26 v 69.
SULU ARCHIPELAGO, Sulu Prov., Mandubal I., Doty & Hair no. 25529a,
OP VaAnhh (66

Dawson's figure 17 (1956, p. 36) I would unhesitatingly ascribe
to C. mexicana (Sond.) J. Ag. The broad axes, with the ramelli
clearly contracted toward the base, are characteristic.

14

Caulerpa urvilliana Mont., var. urvilliana
LINE ISLANDS
PALMYRA I., Sand I., Doty no. 186211, 29 xii 59.
FANNING I., North Pass, Wainwright no num., n. d.; Tempua,
Wainwright no. 20101, n. d., Greig Point, DeWreede no. 1222
IDO) Da eek Oe
TUAMOTU ARCHIPELAGO
RAROIA ATOLL, Kukina, Doty & Newhouse no. 11454, 3 vii 52;
between Temari and Kumekume, Doty & Newhouse no. 11021,
I Aa Sys
MARSHALL ISLANDS
ARNO ATOLL, Ine Village, Horwitz no. 9044, 10 vii 51.
CAROLINE ISLANDS
PINGELAP I., Menez nos. 21454, 21925, 1 vii 60.
MOKIL IDS., Urak I., Menez nos. 21227, 21285, 21360, 29 vi 60.
TRUK “EDS!.!,, Quol, P-Menez, no.) 26027. 2) viii. 6O).
IFALUK IDS., Ifaluk I., Menez nos. 23111, 23126, 10 viii 60;
Ella I., Menez no. 23230, 10 viii 60.
YAP I., south of the channel,.Menez nos. 21567, 15817 (very
good), 19 viii 60, north of the channel, no. 21896, 18 viii 60.
PAGAU IDS...,..Peleliu lL.) Doty no. \20529),).23.xi 68’.
HELEN I., west side, Menez nos. 15480, 15519, 15537, 30 vii 60,
Helen Island Reef, Menez no. 15390, 28-30 viii 60.
PHILIPPINE ISLANDS
LUZON I., Sorsogon Prov., Bulusan, Santos no. 20513 p.p.,
L7hini 6S 2
NEGROS I., Negros Oriental Prov., Siquijor I., Carrio Solong-on,
Menez no. 1135, 28 ix 67, Santos nos. 26208, 26055 (very good).
Tending toward v. vitiensis, Alcala no. 20518, 28 v - 29 vi 68,
Santos nos. 26041, 26182, 22 v 69.
PALAWAN ARCHIPELAGO, Palawan Prov., Cujo Ids., Cuyo, no coll.,
NOW, 6540), 3Oriv 58).
SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., Doty
& Hair, no. 25447, vi 66; Tapul Group, s. of Siasi I., Doty
& Hair no. 25529b, 10 vila 66; Tapaan 1. ; ‘Trono; ino. T4077,
29 xii 70; Tawitawi Group, Tawitawi I., Balimbing Point,
Menez no. T-1101, 23 ix 67.

Caulerpa urvilliana var. vitiensis (Sond.) Weber-van Bosse
TUAMOTU ARCHIPELAGO
RAROIA ATOLL, Ohehoa, Doty & Newhouse no. 11503, 2 viii 57
(excellent).
PHILIPPINE ISLANDS
SULU ARCHIPELAGO, Sulu Prov., Jolo Group, Pangasinan I., Doty no.
25422, vi 66.

Caulerpa verticillata J. Agardh
HAWAITAN ISLANDS
OAHU I., Kaneohe Bay, Moku-O-Loe (Coconut) Island, Doty no. 8117,
3 x 50 (excellent), Jones no. 10433, 7 i 52, Wainwright no num.,
17 vii 59, Buggeln & Tsuda no. 14186, 15 vi 64, Littler no.
20539), 10) x 168 -

15

CAROLINE ISLANDS

PONAPE.,
MANTAPEITAK I., Menez no. 23409, 20 vi 60.

KOLONIA I., mud flats toward Sokehs I., Menez no. 21108, 25 vi 60
PHILIPPINE ISLANDS
CEBU I., Cebu Prov., Davao Bay, Boty & Alvarez no. 14350, 3 ii
65; Mactan I., Calawisan, Menez no. T-1029, 6 ix 67.
NEGROS I., Negros Oriental Prov., Siquijor I., Carrio Solong-on,
Santos no. 26161, 26 v 69.

Caulerpa webbiana Mont.
HAWAIIAN ISLANDS
LAYSAN I., Tsuda no num., 6 xii 63.
KAUAI I., Kuaehu Point, Abbott no num., 21 ii 46.
GCAHUN LE. 7, Mokuleta, Doty now 222335, 9 xi 525 Laie, Doty no. 19287,
30 i 60; Waikiki, Doty no. 8260, 3 xi 50; Lua-Lua-Lei, Maili,
Doty no. 8733.
EASTER ISLAND, Hotuiti, Matsui no. 20671, 3 iv 69.
MARSHALL ISLANDS
ARNO} ATOLL, Enen Edrik ~£., Horwitz no. 9032, 7 vii 5.

References

Abbott, 1.4. 1961. A check list of marine algae from Ifaluk Atoll,
Caroline Islands. Atoll kes.) Bulls 71: l—-5.

Bérgesen, F. 1913. The marine algae of the Danish West Indies.
I. Chlorophyceae. Dansk, Bot. Arkiv, 1(4)2 1-158 + 2,/-§26 figs),
map.

Dawson, E.Y. 1956. Some marine algae of the Southern Marshall

Islands. Pactvfiic Ser. LO()): 25-66, 66 text—figs.

Dawson, E.Y. 1957. An annotated list of marine algae from Eniwetok
Atoll, Marshall Islands. Paciimicey sei. Wily 92-132) 30 text—frigs .

Diaz—-Pitferer, M. 1969. Caulerpa hummii, a new species of marine
algae (Chlorophyta, Caulerpales) from Venezuela. Phycologia
TG) een ae

Gilbert, W.J. 1942. Notes on Caulerpa from Java and the Philippines.
Pap. Michigan Acad. Sci., Arts & Lett. 27: 7-26, 5 text-figs.

Gilbert, W.J. 1946. Philippine Chlorophyceae II. Bulli. LOTT «BOt.
Clubins Ch) 75-99)

Gilbert, Wad. | 196L. An annotated checklist of Philippine marine

Chlorophyta. Philippine Journ. Sci. 88(4): 413-451, 1 text-fig.,
pl GLO59)))<

231-339 O- '717 - 2

16

Gilbert, W.J. 1962. Contribution to the marine Chlorophyta of
Hawaii, I... Pacific Sci. .135-144, 8 text-figs.

May. Ve, 953. Some marine algae from New Caledonia collected by
Mrs. R. Catala. Contr. New South Wales Nat. Herb. 2(1):
36-66.

May, V. 1966a. Algae of the Gilbert Islands. Contr. New South
Wales Nat. Herb. 4(1): 13-16.

May, V. 1966. Further records of algae from New Caledonia
collected by Mrs. R. Catala. Contr. New South Wales Nat. Herb.
An) Lie.

Okamura, K. 1916. List of marine algae collected in the Caroline
and Mariana Islands, 1915. Bot. Mag. Tokyo 30: 1-14, 9 text
EC iSsei, 1b yolks

Taylor, Wm. Randolph. 1950. Plants of Bikini. XV. +22 7-- te leap pee,

frontisp., 79 pl. Univ. Mich. Press, Ann Arbor.

Taylor, Wm. Randolph. 1966. Records of Asian and Western Pacific
marine algae, particularly from Indonesia and the Philippines.
Pacific Sci. ~20(3): 3422359. - 2 text—rags <

Taylor, Wm. Randolph. 1973. Marine algae of the Smithsonian-Bredin
Expedition to the Society and Tuamoto Islands. PaCLEVe) Sei.
27(1)-: 37-43, 11 text—-figs:

TrOnO), \GaGsy, L96G. The marine benthic algae of the Caroline Islands,
I. Introduction, Chlorophyta, and Cyanophyta. Micronesica
4(2) > 137-206, 19 pls.

BROnO,.~GsGe, . OnZas Notes on some marine benthic algae in the
Philippines. Kalikasan Philippine Journ. Biol. 1: 126-147,
21 figs.

MrOnG, 1G.C.. L972b:: The marine benthic algae of Siasi Island and
Vicinity. I. Introduction and Chlorophyta. Kalikasan

Philippine Journ. Biol. 1: 207-228.

TEONO, Gace. L973): Studies on the marine benthic Chlorophyta of
Puerto Galera, Oriental Mindoro, Philippines. Univ. Philip.
Nat. Sci. Res. Center, Techn. Rep. 1: 1-25, 24 text-figs.

Tsuda, R.T. & Belk, M.S. 1972. Additional records of benthic algae
from Yap, Western Caroline Islands. Atoll.Res. Bull., 156: 1-5;

Yamada, Y. 1941. Species of Caulerpa in the South Sea. Kagaku
Nanyo 4: 95-105, 16 text-figs.

sE7/

Yamada, Y. 1944a. New Caulerpas and Halimedas from Micronesia.
SC? fap~ -EnSt. ALgol. Res. Hokkaido Univ. 3(U): 27-29, pills). 1-5.

Yamada, Y. 1944b. A list of the marine algae from the Atoll of Ant.
Sci. srap. (nse. Algol. Res. Hokkaido Univ. 3)(1.)> 31-45, pls. 6,
fis

Weber-van Bosse, A.A. 1898. Monographie des Caulerpes. Ann. Jard.
Bot. Buitenzorg, 15: 243-401, pls. 20-34.

Weber-van Bosse, A.A. 1910. Note sur les Caulerpa de l1'fle Taiti
et sur un nouveau Caulerpa de la Nouvelle-Hollande. Ann. de
i fast. Oceanogr. 2: 1-8, pl. 1, 2.

Womersley, H.B.S. and Bailey, A. 1970. Marine algae of the Solomon
Islands. Philos. Trans. Roy. Soc., B, Biol. Set, 259: 257-352,
pls. 24-27, 10 text-figs.

ATOLL RESEARCH BULLETIN
NO. 209

MARINE ALGAE OF THE TE VEGA 1965 EXPEDITION
IN THE WESTERN PACIFIC OCEAN

by Wm. Randolph Taylor

Issued by
THE SMITHSONIAN INSTITUTION

Washington, D. C., U.S.A.
May 1977

MARINE ALGAE OF THE TE VEGA 1965 EXPEDITION
IN THE WESTERN PACIFIC OCEAN

by Wm. Randolph Taylor !

The widespread interest in oceanography currently developing has
revived the practice of research cruises, but these have generally
been devoted to problems of the open ocean rather than of the shore-
line, the littoral and sublittoral. The marine institutes of both
Atlantic and Pacific coasts of North America have been most active in
this open sea work. It is particularly noteworthy that one series
promoted by Leland Stanford Jr. University with the M/V Te Vega did
afford opportunity for work in shallow water along shore.“ Professor
Lawrence R. Blinks, Director of the Hopkins Marine Station of that
University was Chief Scientist for the 1965 cruise with which this
report is concerned, and the algal material was collected by
Dr. Charles F. Cleland, then at Stanford University. ° The writer is
greatly indebted to him for the opportunity to study the material,
which was accompanied by extensive field notes and partial identifi-
cation. For the most part these data could only have been fully
utilized by the person who had done the work in the field, but the

Department of Botany, University of Michigan.

2 The field work of this M/V Te Vega cruise of the winter of 1965 was
conducted under National Science Foundation Grant no. G-17465 to
Stanford University. Part of the laboratory research on the algae
was done under National Science Foundation Grant no. GB-3186 to the
University of Michigan. For these supporting grants the collector
and the writer are most grateful. A selection of voucher specimens
has been placed in the Herbarium of the University of Michigan and
the balance returned to the collector.

3 Present address: Smithsonian Radiation Biology Laboratory,
12441 Parklawn Drive, Rockville, Maryland 20852.

(Manuscript received April 1974 -- Eds.)

general characters of the stations are outlined on following pages,
based on the notes provided. Unfortunately much time has elapsed
Since the material was collected and the large proportion which was in
formaldehyde has suffered accordingly; nevertheless much of value
remained and it is possible now to give a report in which a large
proportion of records are new for the areas concerned. So far as
these collections are concerned, the reports start with observations
near Singapore, Swing up past northern Borneo and through the southern
Philippines, then down through the Bismark Archipelago to New Britain
Island, over to Bougainville Island in the Solomon Islands, with the
last sampling from Guadalcanal Island.

The stations visited by the Te Vega Expedition in question are so
distant from each other that a thorough review of the pertinent phyto-
geographical literature is impracticable. Most curiously, Singapore,
which has been a famous botanical center for many decades has never
been favored with a major published algal flora, although the former
Director of the Botanical Garden, Dr. H.M. Burkill, collected there
extensively, and so the first station, the nearby Hantu Island, has no
marine phycological history. Nevertheless, the Siboga Expedition
reports of Mme A. Weber-van Bosse (1913-1923) apply very well, even
though ranging far to the east of Indonesia.

There have been several papers on Philippine marine algae,
summarized by me (1966) and supplemented by others, as Trono (1972).
The Caroline Islands have been visited by E.G. Menez and reported upon
by Trono (1968, 1969), and Tsuda both collected there and reported his
Resubtes wh 972). The Bismark Archipelago has less algal history.

The Zaca Expedition collected in the Solomon and Santa Cruz Islands in
1933 and Setchell reported on the plants (1935), while the Solomons
have had the benefit of a recent study by Womersley (1969, 1970).
Chapman has recently published on the algae of Fiji (1971). Setchell
(1924) dealt with the algae of Samoa. The Society and Tuamotu island
groups far to the east have been visited several times: Setchell
reported on Tahitian algae (1926) and those collected by a Smithsonian
Institution group in 1957 were studied by me (1973). While very

much to the north of the Te Vega Expedition route, the algal flora of
the Marshall Islands shows much similarity, and my account (1950) and
those of Dawson (1956, '1957) are’ useful for consultation.

From the literature cited in these papers a fairly complete idea
of the Indonesian-Polynesian marine flora can be obtained, and it leads

one to a wealth of minor scattered records. In addition, the
bibliographic compilations of Walker (1947) and of Trono (1966) must
be considered. When all this is taken into account it seems clear

that the information on the algal flora of this vast area is still too
fragmentary to justify any firm conclusions as to floristic trends,
although attempts were early made to do this (Weber-van Bosse 1913-
1923; Yamada 1926).

List of Stations
Station I. Gums O65

MALAYSIA, Pulau Hantu. An islet 11-16 km west of Singapore, off the
Malay Peninsula.

The area studied was in general a firm but silty shoal, with

scattered rocks exposed, and a few mangroves. The dominant alga
reported was Sargassum, but none was collected. A bit higher
Turbinaria conoides was prominent, and Bryopsis pennata seems to have
been frequent on these coarse algae. Various small species were

collected, but none seemed to occur in conspicuous communities.
Interesting genera include Microdictyon, Bornetella, Martensia and
Tolypiocladia. Notable near high tide line were communities of
Halophila-ovalis (R. Br.) J.D. Hooker, a widespread marine phanerogam.

Statron 2.0823 i 1965
SOUTH CHINA SEA at about 1° 47' N.L., 107° 47' E.L.

A few floating algae were collected, mostly Sargassum and its
epiphytes, none in conspicuous quantity.

Seaeion os. 24 2 W965
SOUTH CHINA SEA at about 2° 31' N.L., 110° 42' E.L.

This was a small collection from floating material, basically
portions of Sargassum plants with some epiphytes.

Station 4. 26 I 1965
SOUTH CHINA SEA at about 6° 33' N.L., 115° 51" E.L.

This sample was reported to have had a few more items than had
Sta. 3, but Leveillea jungermannioides and Sphacelaria furcigera were
the most common epiphytes on the Sargassum, and only these species
yielded voucher specimens.

SHEGIENCINS, (pe 7.5 Sk ap ALS

MALAYSIA, Pulau Gaya area. These stations were in a complex of
islands off the east coast of North Borneo.

Station 6 is on the west side of Pulau Mantabuan. The aspect is
reported to be one of sparse vegetation over a sandy bottom. The
chief alga seems to have been Enteromorpha lingulata, but various other
things were here or at Station 7, mostly Champia parvula and Laurencia
papillosa. While the Champia might have been an epiphyte, the other

things probably grew on rocks and shells exposed above the sand.
Station 7, off the south side of Pulau Gaya nearby, is reported to
have involved a reef of live corals nearly free of algal colonies,
though a little Halimeda opuntia was noted, but not collected.

Stations? 8-10.54 Gir ie tos 6) tay; L965
MALAYSIA, Pulau Gaya area.

Station 8 consisted of a shoal sand-spit between two islands, part
of the crest being barely uncovered at low water, and part of the shore
of Pulau Tatagan nearby. Pieces of shell and dead coral lay about and
live corals became more abundant as one moved into deeper water. The
vegetation involved a conspicuous mat of Valonia aegagropila over
which the water, 1.8-2.4 meters deep, was at times found to reach a
temperature of 38°C. Dictyosphaeria calvernosa was common and grew
especially well in the shelter of the Valonia and other larger algae.
Valonia ventricosa was also notable here, and one large individual
displaced 90 ml. Spongocladia dichotoma, a most curious plant, was
also notable here. A little deeper, scattered rocks bore a good
vegetation of the very common Halimeda opuntia, H. macroloba, several
Caulerpas and Microdictyon montagnéi. Patches of turtle-grass,
Enhalus acoroides (L., f.) Royle, showed colonies of Padina associated
Wiehe iit.

Station 9k. Sltdasco: Gud WO65
MALAYSIA, Pulau Gaya area, Pulau Tatagan.

This material is chiefly from one of the islets adjoining
Station 8, with which collection the material was bottled, and it
involved part of the north shore of the sand spit. The islet was
fairly rich in algae along the northwest side. Here Caulerpa
racemosa and Dictyota dichotoma were common on the scattered rocks,
where also there was quite a bit of Chlorodesmis comosa. Mangrove
roots near shore carried Lobophora variegata and smaller epiphytes.
Halophila ovalis was quite common on the sandy areas, and a variety of
Caulerpa mexicana was found with it.

Station lO. 4) 5 i, L965
MALAYSIA, Pulau Gaya.

Collections were made progressively along the shoal and beyond the
collection Stations 8, 9, and thus beyond the Valonia aegagropila
community. There were occasional rocks and broken coral masses here.
On the lower sides of them and not found elsewhere, appeared Caulerpa
verticillata and Udotea javensis. Valonia ventricosa plants grew in
some numbers on dead coral: Spongiocladia dichotoma grew rather
abundantly in a few patches on old coral near shore, and
Dictyosphaeria cavernosa was common where somewhat shaded, while in

5
deepest shade D. versluysii also occurred. In the sand Avrainvillea
erecta was common. Caloglossa is reported on mangrove roots near the

end of the route, but did not appear among the specimens preserved.
This was a station with a relatively richly varied flora.

Steayeioin thik, Zh ey acig USVeS)
MALAYSIA, Pulau Gaya area, Pulau Mantabuan.

This collection was made off a sandy beach south of the pearling
station. Avrainvillea erecta was again common. One large clump of
Hydroclathrus clathratus was seen.

Sue giles als IAB Ge ALES LCS):
MALAYSIA, Pulau Gaya area, Pulau Mantabuan.

On the south side there was a community of Valonia aegagropila
and associates similar to that of Station 9. On the north and west
a broad sandy flat with patches of turtle-grass was bounded outside
by patches of coral, beyond which there is a deeper lagoon-like zone
about a. meter deep at low tide, where there was a rather different

flora from that elsewhere. It especially consisted of genera favoring
sheltered water, such as Hypnea, Ceramium, Champia, Spridia,
Centroceras, and Tolypiocladia. The broken coral reef bordering this

on the seaward side had a heavy growth of Enteromorpha clathrata,
Trichosolen and Polysiphonia.

Statrony ls. “13° ir 1965
PHILIPPINES, Mindanao Island, Zamboanga, off Zamboanga City.

This station involved a small offshore islet and lagoon. Ulva
reticulata was very abundant on the sandy bottom, and Cladophoropsis
Philippinensis with Valonia aegagropila formed large masses with it.
Boergesenia was reported aS common on the bottom, but no voucher
specimens were received.

Statvon) 1425.26 11-4 rrr 1965
BISMARK ARCHIPELAGO, New Britain Island, Rabaul area.

This station was on the north side of Crater Peninsula opposite
the town of Rabaul. Sargassum cristaefolium in quantity, and
Galaxaura oblongata were found here, washed ashore.

Stattonelo.) 26m 0-4. ii 965

BISMARK ARCHIPELAGO, New Britain Island, Rabaul.

This material was collected from a small, rocky island in Rabaul

Harbor. On the rubble of dead corals and rocks there were a few
scraps of algae, but only Actinotrichia rigida was in fair condition.
By diving Galaxaura veprecula was obtained from a depth of about 20
meters.

Station 16. 26bl—4 ies 96S

BISMARK ARCHIPELAGO, New Britain Island, Rabaul area, Duke of York
Islands.

A general and rather varied collection, some 17 species being
represented in the material recovered, reflecting the variety of areas
concerned. The species of Halimeda were the most noteworthy elements,
but large plants of Valonia ventricosa with a displacement up to 75 ml.
seem to have been prominent. There were 3 Caulerpas, and the
occurrence of large plants of Udotea argentea was notable.

Station 17. ©26 11-4 IIr 1965
BISMARK ARCHIPELAGO, New Britain Island, Rabaul harbor.

On an old ship null there was a growth of Caulerpa serrulata, and
some Bryopsis which was not preserved.

Station! 183.4 26, Pi—4° Pile 1965
BISMARK ARCHIPELAGO, New Britain Island, Rabaul, Matupi I.

This station involved a turtle-grass bed of Cymodocea serrulata
(R. Br.) Asch. & Magn. and Halodule univervis (Forssk.) Asch., with
associated Halophila ovalis, in 1.2-2.4 meters' depth of water. It
yielded more Udotea argentea, Dictyopteris repens and several little
things, such as Neomeris annulata, Acetabularia major and Sphacelaria
tribuloides.

Station 19. 26 II-4 III 1965
BISMARK ARCHIPELAGO, New Britain Island, Rabaul.

On a coral patch at this station there was some Caulerpa serrulata,
and on exposed rocks Turbinaria ornata was reported but without
voucher specimens.

Station 20) 26, i1-4 Trt. 1965
BISMARK ARCHIPELAGO, New Britain Island, Rabaul area.

On the shore of Point Gazelle 16-32 km south of Rabaul there was
a collecting area with considerable surf, where at depths of 0.3-1.0
meter Chlorodesmis comosa, Hydroclathrus clathratus and Ceratodictyon
spongiosum were found. Boergesenia forbesii was reported very common
in a narrow zone at a depth of about O.3 m.

Stesnedom ily S}cwiciSwS aceite I's)
SOLOMON ISLANDS, Bougainville Island, Kieta area, Puk Puk Islet.

This collection was made on a protected southern point with
Sargassum and Padina australis growing on hard objects over the sand
or on patches of coral reef at depths of O.6-1.8 meters. There were
many smaller things in addition. Most notable for this station were
Chlorodesmis comosa, Neomeris dumetosa, and the spectacular Halymenia
durvillaei.

Seaton i260 Skil 5) Lit L965

SOLOMON ISLANDS, Bougainville Island, Kieta area, Puk Puk Islet.
While made from the east side of the islet the vegetation was

similar, but because of the lack of segregation of the Kieta area

materials little beyond Galaxaura apiculata could be directly

aberlbuced= to) ate

Sitaivone25. 16) Loli So teri e965

SOLOMON ISLANDS, Bougainville Island, Kieta area.

The material of this collection came from the fringing reef several
miles from shore, separated from it by deep water and exposed to

moderate-to-heavy surf. ‘Inside this was a lagoon-like area with much
dead coral and a sandy shoal, free from surf but with substantial surge
currents. For this area most notable was Tydemannia expeditionis
which was present in quantity, a splendid acquisition. Caulerpa
pickeringii among the 4 Caulerpas found was less showy, but almost as
important. Halimeda incrassata and H. micronesica occurred here.

Stations 21 and 23 each yielded about 20 species of algae.
SeaeicimnSs 25, 2x fe) apaeaesau's) aeigac alSKey5}
SOLOMON ISLANDS, Bougainville Island, Kieta area, near Marowa Point.

This material came from the first point east of Marowa Point.
This was exposed to moderate surf. Many of the algae were growing in
shade, due to the neighboring cliff with overhanging vegetation. The
substrate was rock, more or less sand-covered, and corals were absent.
Halimedas were reported in evidence, but not collected; Cladophoropsis
? zollingeri formed extensive mats. Chnoospora minima in a small form
was the only conspicuous brown alga. Actinotrichia rigida was
frequent on the rocks, as were Galaxaura squalida and G. apiculata.

Sieaealioi., 275 Aah acacigs ALS
SOLOMON ISLANDS, Guadalcanal Island, Kamembo Reef.

This station was 16-32 km north of Honiara. A companion on the

expedition contributed to the Cleland collections a small plant of
Halymenia durvtllaei from this station, found on a coral head at a
depth, of 330-425. m.

List o£ species
CHLOROPHYCEAE
ULVALES
Ulvaceae

Enteromorpha lingulata J. Agardh MALAYSIA, Pulau Gaya area,
Stask 6a h27
Enteromorpha clathrata (Roth) J. Agardh Pulau Gaya area,
Sta. 12.
Ulva reticulata Forssk. PHILIPPINES, Mindanao I., Sta. 13.
CLADOPHORALES

Cladophoraceae
Chaetomorpha crassa (C. Agardh) Kutz. Mindanao I>, Sta. o5-
Chaetomorpha linum (MUll.) Kutz. SOLOMON ISLANDS, Bougainville
MEA pe seeks alge s
Rhizoclonium hookeri Kutz. Pulau Gaya area, Sta. 9;

Bougainvidle) i.) Sta. 225%

Filaments in this material range from 35 um to 100 um, and so

cover the widest range I have ascribed to the species.
Cladophora sp. Material of this genus was not a feature of

these collections, though a small specimen which I could not

assign to any species, did occur in the Pulau Gaya area

material.
SIPHONOCLADIALES
Dasycladaceae
Neomeris annulata Dickie BISMARK ARCHIPELAGO, New Britain I.,
Sical.culerns
Neomeris dumetosa Lamx. MALAYSIA, Pulau Hantu, Sta. 1;
Bougainville £., -Sta.- 21.
Bornetella oligospora Solms-Laub. Pulau vHantuy Staci
Acetabularia major Solms-Laub. New, Britain L., Sta. 267
Acetabularia parvula Solms-Laub. Pulau Gaya area, Sta. 10;
Bougatnviller dD. , Sta. 2lc
Valoniaceae

Valonia aegagropila C. Agardh Pulau Gaya area, Stas. 8-10;
Mindanao i. Sta. 3; Bougainville, ia 1Stase, 20021.

Valonia ventricosa J. Agardh Pulau Gaya area, Stas. 8, 10;
New Britain I. area, Duke of York Islands, Sta. 16;
Bougainvall Ley ten, sotase. 21a 22.

+ In listing the localities where the various species were found the
name of the general area: PHILIPPINES, SOLOMON ISLANDS, etc., will
be given only at the first appropriate place, since, like the
detailed site characterization, it can be ascertained through the
station number in the preceding descriptions of localities.

Boergesenia forbesii (Harv.) Feldm. Pulau Gaya area, Stas. 8,
oS NeweBreitainwl., Sta. .20;) Bougainville ia, eSta. 25.
Valoniopsis pachynema (Mart.) Bgrg. Bougainville I., Sta. 25.

Dictyosphaeria cavernosa (Forssk.) Bérg. PudltaweHaneu, sical. 1!
Pulau Gaya area, Stas. 8, 10.
Dictyosphaeria versluysii Weber-van Bosse. Pulau Gaya area,

Sittas LO} New Bisittaim ee, Sittase. 2h 22)
Cladophoropsis philippinensis W.R. Taylor. Mindanao I., Sta. 13.
Cladophoropsis zollingeri (Kutz.) Borg.? Bougainville I.,

Sittares2o

Spongocladia dichotoma (Zan.) Murr. & Bood. Pulau Gaya area,
SivaleaeOF

Microdictyon montagnei Harv. Pulau Hantu, Sta. 1; adrift,
South China Sea, Sta. 2; Pulau Gaya area, Sta. 8.

Anadyomene plicata Zan. Dukenor, York sids,. ; “Staqe6; sBougain—
Tilley Wop Season Zip ASa 256

Anadyomene wrightii Gray. Bougainville I., Sta. 23.

SIPHONALES
Bryopsidaceae

Bryopsis pennata Lamx. Pulau Hantu, Sta. 1; New Britain I.,
Stave l/s ssougdi ny nL her ts Sttasteeaihe 1235

Trichosolen parva (Dawson) Taylor PulaueGayalstden kee These

plants were 1-2 cm tall, taller than those reported by Dawson
from Viét Nam (1954, p. 393) and more densely clothed with
ramelli, but the dimensions of axes and ramelli, the shape
and range of measurements of the gametangia are close.
Certainly in view of the little we know of the range of
variation in the genus one had best regard these plants as
a relatively luxuriant form of T. parva.

Caulerpaceae

Caulerpa ambigua Okam. Bougainville T., Sta. 2i'.

Caulerpa lentillifera J. Agardh. Pulau Gaya area, Sta. 9;
DUKCHOE YOLK mdse, Sitas W6).

Caulerpa lessonii Bory. Pulau Gaya area, Sta. 8.

Caulerpa peltata (Lamx.) Weber-van Bosse. Pulau Gaya area,
Stas, 9), si2ei New Britain i. ~anrea,’Sta.'16;" Bougainville I.,
Staean25\

Caulerpa pickeringii Harv. & Bail. Bougainvitte: i. “Sta. 23:
Caulerpa racemosa (Forssk.) Weber-van Bosse Pulau Gaya area,
Stas. §S, lO} Duke of York ods.; Sta. 16; Bougainville” E.,

Sta. 23

Caulerpa serrulata (Forssk.) J. Agardh. Pulau Hantu, Sta. 1;
PulausGayavarcay,, sStas.) S)LO; New! Britain f.,- Stas: 17), 19);
Bougainville” is, Sta. 2k:

Caulerpa sertularioides (Gmel.) Howe Pulau Gaya area, Stas. 8,

Dis
Caulerpa taxifolia (Vahl) C. Agardh ? Pulau Gaya area, Sta. 9.
Caulerpa urvilliana Mont. DUKey OE YORK dS. 7 StaeLo.
Caulerpa verticillata J. Agardh. Pulau Gaya area, Sta. 10.
Codiaceae
Chlorodesmis comosa Bail. & Harv. Pulau Gaya area, Sta. 9; New

lsealeelalin Ilo p Steels (AOR Ilowblopelaion Alle) ah4 5 Sheeve - Zils

10
Avrainvillea erecta (Berk.) A. & E. S. Gepp. Pulau Gaya area,
Sitas soi
Avrainvillea lacerata J. Agardh. Bougainvillegis,, Stas i23%
Tydemannia expeditionis Weber-van Bosse. Bougainville I.,
Stam ar
Udotea argentea Zan. Duke of York Islands, Sta. 16; New
Britainedcke ota. le
Udotea javensis (Mont.) A. & E. S. Gepp Pulau Gaya area,
Sta. 10; Bougainville I., Sta. 21.
Codium tenue Kutz. Rabaul, Sta. 15; Duke of York Ids., Sta.
16; Kieta, Sta. 23.
Halimeda cylindrica Decne. Duke of York Ids., Sta. 16; New
Britain I., Stas. 18, 19.
Halimeda discoidea Decne. Pulau Gaya area, Sta. 9; Bougainville
iy ae Stanw2.
Halimeda incrassata (£11.) Lamx. Bougainville i) Stamecse
Halimeda macroloba Decne. Pulau Hantu, Sta. 1; Pulau Gaya
area, Sta. 8; Duke of York Ids., Sta. 16.
Halimeda micronesica Yam. Bougainville I., Sta. 23.
Halimeda opuntia (L.) Lamx. ~ Pulau Hantu Sta. 1; Pulau Gaya
area, Stas. 8, 9; Mindanao I., Sta. 13; New Britain I., Sta.
15; Duke of; York Ids. ,; Sta.6; Bougainville tl. .Stas also
PHAEOPHYCEAE
SPHACELARTALES
Sphacelariaceae
Sphacelaria furcigera Kutz. South China Sea, Sta. 4.
Sphacelaria novae-hollandiae Sond. Bougainville; i.;, Sta.) 23.
Sphacelaria tribuloides Menegh. New Britain I., Sta. 18;
Bougainvilie.sI.,Sta. 21.
DICTYOTALES
Dictyotaceae

Dictyota apiculata J. Agardh, var. jedanensis Weber-van Bosse?
Pulau Gaya area, Stas. 8, 12.

Dictyota ceylonica Kutz., var. rotundata Weber-van Bosse?

Pulau Gaya area, Sta. 12.

Dictyota dichotoma (Huds.) Lamx. Pulau Hantu, Sta. 1; Pulau
Gaya area, Sta. 9; Duke of York Ids., Sta. 16; Bougainville
IEGia Sele Asie

Dictyota divaricata Lamx.? Pulau Gaya area, Sta. 12; Duke of
York Widsii, iStacilGi.

Dictyopteris repens (Okam.) Bé¢rg. New Britain 1. :Staz e.s)
Bougainville L.,)Sta. 21.

Padina australis Hauck Pulau Hantu I., Sta. 1; New Britain I.,
Sta. v8, Bougainva lle i. ):Sta.-21,.

"Dictyerpa" or "Vaughniella" phase of Padina. Pulau Gaya area,
Stas. 9,10; Mindanao ‘1..,./Sta. 133 ‘Bougainville 2. ,)/>Sta.237
Some of the P. australis material is transitional from this
phase.

Lobophora variegata (Lamx.) Womersl. Pulau Gaya area, Stas. 9,
lO; -Bougainvel le Viti, (Stacwe232

ita

PUNCTARIALES
Asperococcaceae
Chnoospora minima (Her.) Papenf. Bougainville —., Sta. 26.
Hydroclathrus clathratus (Bory) Howe Pulau Gaya area, Stas.
8, 10; New Britain I., Sta. 20.
FUCALES
Sargassaceae
Sargassum. Several specimens of Sargassum in the Cleland
material defied identification with the reference materials
available, S. cristaefolium and S. swartzii being the ones

clearly recognizable. The best that I can do at present is
to suggest the possible relationships of the more distinctive
of them. In addition to the 4 doubtful ones, other pieces

in the collection may be variants of these, or may be other
less distinctive forms.

Sargassum capillare Kutz.? Pulau Gaya area, Pulau Mantabuan,
Steal. Zale Tall, exceeding 6.5 dm, much branched, the axes
terete, nearly smooth. The leaves crowded on the lateral
branches, about 2 mm broad, 10 mm long, linear-lanceolate,
the margins sharply serrate, elaborately proliferous, as to
lesser degrees are the leaf faces, the stems and the vesicles.
Cryptostomata few, on each side of the subpercurrent costas.
Vesicles small, short-stalked. Receptacles sparingly forked,
exceeding the leaves.

Sargassum cristaefolium C. Agardh New Britain Island, Sta. 14.

Sargassum myriocystum oe Agardh ? Pulau Gaya, Sta. 9. To) 2
dm tall, the axes terete, densely spinulose. The leaves to
about 7 mm broad, 30 mm long, broadly lanceolate, the costae
percurrent, the leaf margins erose-dentate, the cryptostomata
small, numerous, scattered.

Sargassum ornatum Grev.? Rawk, “South, China, SearSta. Ss. TO
at least 3 dm tall, the axes terete, smooth. The leaves
crowded on the lateral branches, to 5 mm broad, 35 mm long,
oblong-ovate, broad to the rounded apex, sharply serrate.
Costa not reaching to the leaf tip; cryptostomata small,

numerous, scattered. Vesicles short-stalked, short oval,
apiculate.

Sargassum polycystum J. Agardh ? Adrift, South China Sea, Sta.
Ne Plants very delicate, the terete axes spinulose. The

leaves chiefly on the upper axes and the outer parts of the
lateral branches, small, about 2.0-2.5 mm broad, 1 cm long,
lanceolate, strongly serrate. Costa subpercurrent;
cryptostomata few, somewhat irregularly dispersed.

Sargassum swartzii (Turn.) J. Agardh Adrift, South China Sea,

Stan a2. Plants to at least 3 dm tall, the stems flat, smooth.
The leaves to 3.0-3.5 cm long, narrowly lanceolate, long-
attenuate, shallowly dentate. Costa percurrent, with the
cryptostomata in an irregular row on each side. Vesicles

large, spherical.

231-339 O-'77 - 3

12
Sargassum sp. Bougainville I., Kieta area, Sta. 22. About
3 dm tall, much branched, the stems terete, smooth. The
leaves rather crowded, small, to 2.5 mm broad, 12 mm long,
Narrow, acute, sharply dentate. Costa inconspicuous, about
0.6 the length of the leaf; cryptostomata rather numerous,
scattered on each side of the costa. Vesicles numerous,
very small, round to oval, frequently apiculate.
Turbinaria condensata Sond., prox. Pulau Gaya area, Sta. 10.
Turbinaria conoides J. Agardh Pulau Hantu, Sta. 1; Pulau Gaya
area (Stan. lO.
Turbinaria marrayana Bart. New Britain I., Sta. 20.
Turbinaria ornata (Turn.) J. Agardh Bougainville. f., Sta. 21.
RHODOPHYCEAE
BANGTIALES
Bangiaceae
Erythrotrichia carnea (Dillw.) J. Agardh Pulau Gaya area, with
Tricnosolen, ota. a2.
NEMALITONALES
Chaetangiaceae
Actinotrichia rigida (Lamx.) Decne. New Britain I., Sta. 15;
Bougainville. 7Stas. 2-225
Galaxaura apiculata Kjellm. Bougainville L., Stas. 22) a 256
Galaxaura fasciculata’ Kjecllim., prox. Bougainville Et. ,. sStacsw2ay
Galaxaura oblongata (Sol.) Lamx. New Britain I., Sta. 14.
Galaxaura squalida Kjellm. Bougainville <i. 7 Stax. 25:
Galaxaura ventricosa Kjellm. New Britain £2; Sta. ais. This
material agrees well with the description of G. ventricosa,
except that the marginal spinulose cells were minutely
apiculate.
CRYPTONEMIALES
Rhizophyllidaceae
Desmia hornemanni Lyngb. Bougainville’ i., Sta. 23:
Corallinaceae

Fosliella farinosa (Lamx.) Howe Pulau Gaya area, Sta. 12;
Duke o£ York rds,;.Sta. wiG.
Lithophyllum moluccense Fosl. Bougainville I., Sta. 23.

Amphiroa anceps (Lamk.) Decne. Bougainville io; Stage 2a

Amphiroa foliacea Lamx. New Britain I., Sta. 20; Bougainville
eG iteals yes

Amphiroa fragilissima (L.) Lamx. Pulau Gaya area, Sta. 8;

Duke of ‘York Ids, Sta.) 16.
Amphiroa tribulus (Sol.) Lamx.? Pulau Gaya area, Stas. 8, 9.

Jania tenella Kutz.? Bougainville £. Sta. “26. This small
sample at first seemed to resemble J. rubens, but the upper
branching was more spreading. The diameter of the lower

axes ranged from 120-260 um, which is not too different

from J. rubens, but the segment length below was only 1.5-
1.75 times the diameter and above to 2.0-2.5 times, which

LS» Coo; Short. ‘for thatkvspeciies: In face, is rather shore
for J. tenella’ as figured by Kititzing, -(l858,-8, pl. “Ss Ene

iS}

Grateloupiaceae
Halymenia durvillaei Bory. New Britain a. , ctase 2,122;
Guadacanal I., Sta. 27.
GELIDIALES
Gelidiaceae

Gelidiella acerosa (Forssk.) Hamel Pulau Gaya area, Stas. 8,
9;""Bougainville 2I., Stas. 21; 25.

Gelidiella myrioclada (Bgrg.) Feldm. & Hamel Bougainville I.
Stat 25%

Gelidiopsis intricata (C. Agardh) Vick. Pulau Gaya area, Stas.
3}, 1KOr IDK Cie Yaoidk< NCIS, Siteeion SGA Ney whesesigha\ ito, Siesin alse
Bougaimvael Vem Sy Stcals 251.

Gelidiopsis repens (Ktitz.) Schm. Bougainville I., Stas. 21,
DS\¢ The upper branching is more regularly digitate than
figured by Okamura (1936, p. 636) but the resemblance to
his figure is otherwise close.

Gelidium latifolium (Grev.) Thuret & Bornet, prox. Pulau Gaya
aGbeal me sizaei >; sBOUGaIN VA lale! seca 125)
Pterocladia caloglossoides (Howe) Dawson, prox. Bougainville

Mop Sess 23, 255 26 The fragments submitted are so small
that the full habit cannot be confirmed, but in detail there
is agreement.
GIGARTINALES
Hypneaceae

Hypnea cervicornis J. Agardh Pulau Gaya area, Stas. 10, 12;
DUKeROR York sds, Sta. Ge

Hypnea nidulans Setch.? Bougainvallerhey Stas 25

Hypnea spinella (C. Agardh) Kutz. Bougainva lle riz Sita. 23)
Sphaerococcaceae
Ceratodictyon spongiosum Zanard. Dukewor York dsr, Sta. UG
NGI Isheslicelalig: Ibo, Sieelg ZOc
Gracilariaceae
Gracilaria debilis (Forssk.) Bégrg. Mindanao i), Sta. L3r
Solieriaceae
Eucheuma crassum Zanard. PullauNhantulyy) Sta ak.
RHODYMENTALES
Champiaceae

Lomentaria hakodatensis Yendo ? Kieta area, Sta. 21. These
plants are smaller and less branched than L. sinensis Howe
@ig247 “p.. 39) pl tig. 1), whitch has been” reduced to
synonymy under L. hakodatensis, and the tetrasporangial
branchlets seem to stand erect and to be sagittate to

lanceolate in shape. The lower branches often curve
down and attach at the tips.
Champia parvula (C. Agardh) Harv. Pulau Gaya area, Stas. 6,

ie? -eMancdanao) sly ppactan Lo DUKE Ot York lds. , Sita. iG)
Bougainville £., Sta. 23.
Champia viellardii Ktitz.? Bougarmvyanalen ley Stas 21e 2S.
Coelarthrum boergesenii Weber-van Bosse. Bougainville I.,
Sesio, Alks

14

CERAMTALES
Ceramiaceae
Antithamnion sp. Pulau Gaya area, Stas. 12, 12a. There was
abundant material of this plant, but unfortunately it was all
sterile. The plants were about 5 mm tall, growing on Enhalus
leaves. The main filaments reached a diameter of 15-47 um.
The branchlets were opposite or verticillate, of equal length,
sub-simple low on the axes, but above closely alternately
branched, not secund. The cells near the bases of the
branchlets were 12-14 um diam., near the tips 7.7-9.3 um.
No gland cells were found.
Wrangelia bicuspidata Bérg.? New Britain I., Sta. 15;
Bougainville, 1.\,, Sta 21%
Griffithsia sp. Pulau Gaya area, Sta. 12. Not adequate for
identification to species.
Ceramium sp. Duke of York Ids., Sta. 16. Same observation.
Centroceras clavulatum (C. Agardh) Mont. Pulau Gaya area,
Sta. 12; Bougainville I., Sta. 26.
Spyridia filamentosa (Wulf.) Harv. Pulau Gaya area, Stas. ll,
12
Delesseriaceae
Martensia flabelliformis Harv.? Pulau Hantu, Sta. l.
Rhodomelaceae
Polysiphonia flaccidissima Hollenb., var. New Britain I.,
Star U9: Determined by the kindness of Dr. George F.
Hollenberg.
Polysiphonia savatieri Hariot. Pulau Gaya area, Sta. 12.
Determined by the kindness of Dr. George F. Hollenberg.
Tolypiocladia glomerulata (C. Agardh) Schmitz? Pulau Hantu,
Sta. 1;- Pulau ‘Gaya; area, :Sta..12;,.New Britain, L..j7Stapel5);
Bougainville, 2.) Sta. ¥23%
Bostrychia binderi Harv. Bougainville I., Sta... 25;
Bostrychia calliptera Mont.? Pulau Gaya area, Sta. 10.
Herposiphonia secunda (C. Agardh) Ambronn, prox. Bougainville I.,
StajeZiis
Herposiphonia tenella (C. Agardh) Ambronn prox. Pulau Gaya
area, (Sta. 107) (Bougainville I 2)" Stay 21.
Leveillea jungermannioides (Mart. & Her.) Harv. South China
Sea, Sta... 4; “Bougainviller...,. Stac24..
Amansia glomerata C. Agardh ? Bougainville I., Stas. 21, 23.
Acanthophora spicifera (Vahl) Bérg. Pulau Hantu, Sta. 1;
Pulau, Gaya, Sta. 11 7) Bougainvid lev l..,. Stas. 21,122.
Laurencia obtusa (Huds.) Lamx. Pulau Gaya area, Sta. 12.
Laurencia papillosa (Forssk.) Grev. Pulau Gaya area, Stas. 6, 8;
Duke of York Jds., Sta. 16;)/Bougainville: 2: ,.Stas..22,.023%

References

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a5)

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16

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ATOLL RESEARCH BULLETIN
NO. 210

MARINE ALGAE KNOWN FROM THE MALDIVE ISLANDS
by H. E. Hackett

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

May 1977

CONTENTS

Introduction

Maldive reef features
Environmental features
Habitat descriptions

Algal ridge

Leeward atoll margin

Seaward reef flat

Calcareous debris, beach rock and pavements
Mangrove

Cymodocea-Thalassia flats

Lagoon reef flat

Velu

Lagoon floor

Knoll reef flat and slope

The rubble pool at Filadu, Tiladummati Atoll

Stations of cruise B of the R/V Te Vega
Stations of other expeditions
Systematic list

Cyanophyceae

Rhodophyceae

Chlorophyceae

Phaeophyceae

Acknowledgement

References

MARINE ALGAE KNOWN FROM THE MALDIVE ISLANDS
by H. E. Hackett !

Introduction

At the opening of the International Indian Ocean Expedition in
1964, 17 red algae, 5 green algae, and 2 brown algae were known from the

Maldive Islands. These had been collected during the J. Stanley
Gardiner Expedition (Barton.1903; Foslie 1903, 1907; Weber van Bosse
and Foslie 1904) and the John Murray Expedition (Newton 1953). The

coralline algae are in the Foslie Herbarium, others are in the British
Museum.

During the Cambridge Expedition to Addu Atoll in the summer of
1964, Sigee (1966) collected algae from the Southern part of the atoll
and published ecological notes. Specimens were identified by Tsuda
and Newhouse (1966). They added 7 bluegreens, 20 reds, 25 greens, and
7 browns to the list of known algae. Specimens are at the University
of Hawaii. The Expedition reports (Stoddart 1966) provide extensive
references on the natural features of the Maldives.

During the U. S. Navy Biological Expedition to the Chagos in
August 1967, C. Rhyne collected at Addu. The species are reported in

this paper, and vouchers are at the U. S. National Museum. As part of
the International Indian Ocean Expedition, Cruise B of the R/V TE VEGA
went to the Maldive Islands in 1964. Extensive algal collections were

made at nine atolls.2

An area was usually sampled by removing algae from a square meter
plot or by line transect. On occasion SCUBA, a two foot wire dredge
or a six foot beam trawl were used. Some specimens were collected by
other members of the scientific party.

The collections bring the total of known Maldive algae to 21
bluegreens, 163 reds, (about 30 additional entities from dredge hauls
are too fragmentary for analysis), 83 greens, and 18 browns.

Hackett (1969b) compared the Maldive algae to those of the Marshall
Islands, contrasted floristic differences between northern and southern
atolls, and gave indicator genera for the Maldive ecosystem. Aregood

Present address: Little Diamond Island, Portland, Maine 04109
(Manuscript received November 1974 -- Eds.)

The field portion of this work was supported by the National Science
Foundation U. S. Program in Biology.

2

and Hackett (1971) reported a new species of Dictyurus from the
Maldives.

This paper reports habitat descriptions from Cruise B and a list of
known Maldive algae. The list is based in part upon an unpublished
PhD dissertation at Duke University (Hackett, 1969a). Specimens are
at Duke University, Durham, N. C. and at the U. S. National Museum.

Maldive reef features

The Maldives form a double row of atolls on the Chagos-Laccadive
Bank. They extend northward from the equator along the 73°E meridian
for more than 600 miles, and include more than 2,000 islands. The
islands are for the most part transient, building and eroding rapidly.
Large populated islands have completely disappeared within the memory
of living natives {(Hasen Didi, 1964 personal communication). Probably
because of this transient quality, the Maldivian language, Divehi,
emphasizes reef types rather than islands. These terms are pertinent
and should be accepted as part of contemporary atoll terminology.

Tracey, Cloud and Emery (1955) have standardized terms for atoll
reef features. Their classification, while applicable to most Pacific
atolls, is not adequate for many Maldive atolls. The Maldives have a
more complex atoll structure than is typical in the Pacific. Most of
the larger atolls are best described as atolls within atolls or
composite atolls. In addition, because of the alternating monsoon wind
pattern, the Maldives do not have typical windward and leeward sides.
The terminology of Tracey, Cloud and Emery is followed where appropriate,
but some new terms apply only to Maldive reef features.

Faru means a reef, part of which is above water at low tide. The
faru may have an island, and it may be a part of the atoll rim or a
formation in the lagoon.

Farus in the lagoon probably developed on cora-algal knolls.
Such farus are frequently so well developed that each has a central
lagoon and, thus, becomes a microatoll. A shallow lagoon of this type
is called a falu. A deeper lagoon is called a velu (vilu), although
the distinction between them is not always clear. }

Farus on the atoll rim may enclose a falu or velu. These are
usually present when massive reef corals grow on the edge of the lagoon
reef flat.

Atoll or atolu is the only Maldivian word placed into the English
language. Yet, atolu in Divehi does not mean a circle of reef and
islands enclosing a lagoon. Atolu refers only to a political division.

! Spelling is based on recent interpretations from Divehi. These

were prepared by N. T. Hasen Didi, during Cruise B. The Admiralty
Chart name appears in parentheses after the corrected name.

The political unit usually corresponds to the English language concept
of an atoll. However, the largest emergent atoll in the world is in
the Maldives and is made up of two political units. The northern half
is called Tiladummati Atoll and the southern half Miladummadulu Atoll.

Both Agassiz (1903) and Gardiner (1903) made extensive comments on
the structure of Maldivian atolls, and Hass in 1962 proposed a new
theory of atoll formation based on his expedition to the Maldives.
Plates I and II show in cross section the conspicuous features of a
typical Maldive atoll.

Environmental features

The only extensive records of air temperature are kept by the
R. A. F. Meteorological Office at Gan, Addu Atoll. But some air and
sea temperature records are available from expedition reports and
Maldivian sources.

From the Addu records of 1959 to 1964 the lowest air temperature
was a Jan. reading of 20.6°C, and the highest in April 32.8°C (Scorer,
Mar. 1964 personal communication).

It is likely that surface water temperatures are similar throughout
the year. Water temperatures of above 31°C were recorded in March
and October from Cymodocea beds and lagoon reef flats. Lowest water
temperatures are 27.2°C for lagoon surface readings in January.
Agassiz (1903) showed that there is probably no appreciable temperature
change from the top to the bottom of the lagoon.

Rainfall data from the northern islands are scant. Figures from
the Maldive Ministry indicate that the northern islands have a rainfall
that is nearly equal to that of Addu, whicH# is the southern most atoll.
But the rain of the northern monsoon atolls is seasonal with a peak
during May at the beginning of the Southwest Monsoon. In contrast the
continuously unsettled weather of Addu results in more even monthly
rainfall.

The prevailing monsoon wind regulates the seasonally reversing
currents. At least in the northern Maldives the cross bank currents
move westward from December to April, during the Northeast Monsoon;
and they move more or less eastward from May through August, during the
Southwest Monsoon.

Tidal data are scant, but the U. S. Naval Oceanographic Office
(1951) states that the mean high water interval for Malcolm or Mukundu
Atoll in the northern Maldives is 10 hours, 20 minutes. Spring tide
range is 2.7 feet. Mean tide range is 2.1 feet. Tidal streams are
irregular and much variation occurs.

The best salinity records are those for Addu Atoll kept by Stoddart
(1966).

Habitat descriptions
Algal ridge

The algal ridge, or windward atoll margin, is a sporadic feature
in the Maldives. A supposed prerequisite for its development is a
continuous onshore wind that provides the water agitation needed for
massive coral growth. Where corals reach the surface a pavement of
calcareous red algae develops in the surf.

Algal ridges are seen in the atolls of the eastern side of the
archipelago on their eastern and northeastern rims and in the western
atolls on their western and southwestern rims. These are the sites
that face the open sea and the alternating monsoon winds.

The habitat was studied on Himmafuri Faru in the northeastern
sector of Male Atoll. There were no surge channels and the pavement
was of Porolithon and Lithophyllum. Crevices continuously flushed
with water contained a richly developed algal turf, dominated by small
creeping species of Chondria, Laurencia, Hypnea, and Polysiphonia.
Occasional are Amphiroa, Anadyomene, Caulerpa peltata, Dasya,
Dictyosphaeria, Halimeda, Jania, Oscillatoria, Peyssonelia, Pocockiella,
Rhipidiphyllon, a tiny Sargassum, Schizothrix, Valonia, and Valoniopsis.
Heteroderma is a common epiphyte.

Leeward atoll margin

Although there are no true leeward reef margins in the Maldives,
comparable situations are common where the atoll rim does not face
the open sea. The habitat was studied on the eastern side of North
Malosmadulu, facing the interatoll flat. Surf is negligible and the
reef corals do not reach the surface, but slope gradually to deep water.
Lithophyllum dominates. Confined to crevices are turfs of Chondria,
Herposiphonia, Hypnea and Tolypiocladia. Occasional are Codium,
Boodlea, Gelidium, Halimeda, Jania, Turbinaria, and Vidalia. Epiphytes
are Acrochaetium, Enteromorpha, and Jania.

Seaward reef flat

The seaward flat is usually behind an algal ridge. If there is
no ridge the seaward reef flat slopes gradually into deep water and is
probably not distinguishable from the leeward atoll margin in the
Maldives. The community at Himmafuri Faru, behind the algal ridge, is
in less than a meter of water, and in places very broad and flat with
high water temperatures. The sponge symbioses, stinging corals, blue
corals, and true corals are abundant. Tridacna is occasional.

Most of the algae are small and confined to crevices. Occasional
are Actinotrichia, Amphiroa, Caulerpa serrulata, Caulerpa racemosa,
Dictyota, Goniolithon, Halimeda, Halymenia (rare), Jania, Padina,
Peyssonelia, Pocockiella and Turbinaria. Turf formers are Boodlea,
Chondria, Champia, Cladophora, Herposiphonia, Hypnea, Microdictyon,
Polysiphonia, and Tolypiocladia. Acrochaetium, Erythrocladia,

Fosliella, Heteroderma, Schizothrix, and Sphacelaria are epiphytic.

The turf algae are mostly infertile, partially developed, and of
divergent growth forms. This habit is apparently due in great part to
the feeding of various reef herbivores. In areas of greater protection,
as interisland flats, occasional large clumps of Ceramium may develop.
In areas of considerable water flow Calothrix turfs may be extensive.

Calcareous debris, beach rock, and pavements

Distinct zonation was observed only once. The supralittoral black
zone of Entophysalis and some Calothrix develops where the calcareous
debris is stable. Below this is a supralittoral lighter band of the
same genera. Its origin is unknown. The upper limits of the littoral
are marked by a heavy growth of Entophysalis and Calothrix. The color
given to the substrate by these algae gradually decreases down to low
tide. This would appear to be related to the thickness of the brown
sheath of Calothrix. The sheath is not so thick on those trichomes
from the lower littoral. In the sublittoral, most debris is covered
with Entophysalis and penetrated by Gomontia and Ostreobium filaments.

Beach rock is usually covered with Entophysalis, which has
considerable endolithic development. There are several other bluegreen
and green endolithic filaments.

In the northern atolls shallow depressions in beach rock and
littoral pavements near iow tide may have minute algal turfs of Boodlea,
Calothrix, Ceramium, Cladophora, Dictyosphaeria, Erythrotrichia,
Microcoleus, Oscillatoria, Polysiphonia, Schizothrix, Spirulina and
hydroids.

At Addu, beach rock has well developed turfs. Crevices contain
Anacystis, Caulerpa racemosa, C. serrulata, C. taxifolia, Ceramium,
Centroceras, Chondria, Cladophora, Dictyurus, Ectocarpus, Enteromorpha,
Hormothamnion, Jania, Microcoleus, Oscillatoria, Padina, Polysiphonia,
Schizothrix, Sphacelaria, and Wrangelia. Thick turfs entirely of
Calothrix were collected on debris throughout the littoral and sub-
iIslieeergel <

Mangrove

Rhizophora mucronata is occasional in protected sites along lagoon
beaches. Where the prop roots of this mangrove extend into the
sublittoral, an algal turf of Gelidium, Jania, and Caulerpa racemosa
develops.

Cymodocea-Thalassia flat

Flats of Cymodocea and Thalassia are rare and turf development is

usually sparse. They occur mostly on seaward reefs or adjacent to
channels crossing the atoll rim, although small tufts of Thalassia may
be in most any protected area. In the northern atolls, flats are

composed of Cymodocea rotundata with some Thalassia hemprichii. The

6

latter is usually poorly developed and heavily epiphytized. The flat
at Addu has Thalassedondron ciliatum in addition to Cymodocea and
Thalassia.

Flats contain many algae that are not seen elsewhere. Near
Himmafuri Village, Male Atoll, poorly developed specimens of Acanthophora,
Chondria, Gracilaria, Hypnea, and Leveillea were collected along with a
Single Acetabularia. Dominant are large clumps of Jania and
Goniolithon, the latter often overgrown with Peyssonelia. Occasional
are Boergesenia, Bryopsis, Caulerpa, Chaetomorpha, Champia, Cladophora,
Dictyosphaeria, Halimeda, Padina, Sphacelaria, and Tolypiocladia.
Acrochaetium, Entocladia, Entophysalis, Erythrotrichia, Fremyella,
Hormothamnion, Microcoleus, Oscillatoria, and Schizothrix are epiphytic
on larger algae. Thalassia is commonly epiphytized by Ceramium,
Cladophora, Enteromorpha, Fosliella, Jania, Ulvella, and many bluegreens.

The algal composition of the flat at Addu Atoll varied. In May
the site had considerable silting with poor water visibility.
Occasional are Brachytrichia, Caulerpa, Centroceras, Ceramium, Champia,
Chondria, Cladophora, Dictyosphaeria, Dictyota, Ectocarpus, Enteromorpha,
Fosliella, Herposiphonia, Jania, Mastigocoleus, Microcoleus,
Pocockiella, Turbinaria, and Udotea javensis. Subsequent summer
collections found Boergesenia, Boodlea, Hydroclathrus, Rosenvingea,
Spyridia, Tolypiocladia, and Udotea orientalis.

Lagoon reef flat

In the northern atolls, if reef building corals occur on the
lagoon periphery of the flat, masses of Porolithon craspedium and P.
gardineri occur among them. In either case Actinotrichia, Bryopsis,
Cladophora, Dictyopteris, Gelidium, Goniolithon, Halimeda, Hypnea, Jania,
Microdictyon, Peyssonelia, Pocockiella, Tydemania (rare), Valonia, and
Vidalia occur in crevices. Acanthophora, Amphiroa, Boodlea, Caulerpa
racemosa, C. serrulata, C. taxifolia, Centroceras, Champia, Cladophorop-
sis, Derbesia, Ectocarpus, Enteromorpha, Hypnea, and Sphacelaria grow on
the lower parts of corals. Epiphytes are Acrochaetium, Asterocystis,
Microcoleus, Schizothrix and Spirulina.

At Addu Atoll both algal diversity and development is much greater.
In crevices are Acrochaetium, Amphiroa, Asterocytis, Boodlea,
Botryocladia, Bryopsis, Caulerpa, Ceramium, Chaetomorpha, Champia,
Chondria, Cladophora, Cladophoropsis, Codium, Derbesia, Dictyopteris,
Dictyota, Dictyurus, Ectocarpus, Enteromorpha, Erythrotrichia,
Galaxaura, Gelidium, Halimeda, Herposiphonia, Hypnea, Jania, Laurencia,
Martensia, Microcoleus, Oscillatoria, Pocockiella, Polysiphonia,
Schizothrix, germinating Sphacelaria propagulae, Spongomorpha, Struvea,
Taenioma, Tolypiocladia, and Turbinaria.

Velu
The only small lagoon or velu studied was on Himmafuri Faru. Tet

is on the lagoon reef flat and separates the massive lagoon reef
structure from the island. The velu has little water circulation and

the bottom at three to four meters is of bare calcareous sand with
occasional clumps of Acropora formosa. Algae occurring as a turf on
the lower parts of the coral are Caulerpa, Dasya, Gelidiella,
Herposiphonia, Jania, and Spyridia.

Lagoon floor

The lagoon floor in northern atolls is rich in algae. Over 20%
of all Maldive algal species were collected there only. Undoubtedly
many were overlooked and several are too fragmentary for analysis.
Dredge hauls were dominated by Microdictyon and Pocockiella. Other
genera are Anadyomene, Antithamnion, Botryocladia, Bryopsis, Callitham-
nion, Caulerpa, Ceramium, Chaetomorpha, Champia, Chondria, Chrysymenia,
Dasya, Dictyopteris, Dictyota, Dictyurus, Enteromorpha, Entocladia,
Entophysalis, Erythrotrichia, Fauchea, Gracilaria, Halimeda, Haloplegma,
Heteroderma, Herposiphonia, Hypoglossum, Laurencia, Lophocladia,
Martensia, Neomeris, Oscillatoria, Padina, Polysiphonia, Schizothrix,
Sphacelaria, Spongomorpha, Struvea, the flabellate form of Tydemania,
Udotea, and Valonia.

Melobesioid algae were common only in a dredge haul in a channel
crossing the atoll rim.

In contrast to northern atolls the lagoon floor at Addu Atoll seems
to have a sparse algal community. Caulerpa verticillata, Chaetomorpha,
Cladophora, Heteroderma, Mastigocoleus, Schizothrix, and Udotea occur.

Knoll reef flat and slope

Coral knolls are farus in the atoll lagoon. The three sites
observed are Dunidu in Male Lagoon, Walla in South Nilandu Lagoon, and
Madugali in Ari Lagoon. All have flats about 25 meters in width that
slope away from emergent islands to a depth of 4 meters before dropping
sharply to the lagoon floor. Dunidu is a disturbed site. The
island is the British Administrative residence. Walla and Madugali
are uninhabited.

Conspicuous at Dunidu is Tydemania. It is the only alga growing
profusely above the surface of the corals. Caulerpa, Chaetomorpha,
Champia, Enteromorpha, Halimeda, Herposiphonia, Hypnea, Struvea, and
Tolypiocladia are in crevices. Most dead corals were covered with
mixtures of Entophysalis, Hormothamnion, Microcoleus, Schizothrix, and
the arcuate tufts of Oscillatoria.

At Walla coral crevices contained Acrochaetium, Amphiroa, Caulerpa,
Ceramium, Chlorodesmis, Cladophora, Coelarthrum, Codium, Derbesia,
Dictyurus, Dictyota, Asparagopsis (tetraspotophytes), Griffithsia,
Halimeda, Herposiphonia, Heteroderma, Hormothamnion, Hypoglossum,
Pocockiella, Porphyrosiphon, Schizothrix, Sphacelaria, Spirulina,
Struvea, Turbinaria, Udotea, and Vidalia.

At Madugali Halimeda is dominant but confined to crevices along
with Acrochaetium, Caulerpa racemosa, Ceramium, Chondria, Hypoglossum,

231-339 OO - 77-4

8

Jania, and Wrangelia.
The rubble pool at Filadu, Tiladummati Atoll

Near a channel crossing the reef flat, a circular mass of
calcareous debris enclosed a pool about 20 meters in diameter and a
meter in depth. The wall, which served as a tern rookery, allowed
algae to grow free from wave activity and grazing by herbivorous fishes.

Although large clumps of Halimeda occurred, the pool was dominated
by entangled and partially free floating mats of the Cnidarian Zoanthus
mixed with Caulerpa racemosa var. microphysa and C. sertularioides.

The Caulerpa was the most profusely developed of any shallow water alga
in the Maldives.

Stations of Cruise B of the R.V. Te Vega 1964
Male Atoll

2 Maia. Dunidu Faru. (73230 U Sunes Ac SOUND es Littoral beach
rock and smooth calcareous pavement on the east side of the coral knoll
island.

oye As above. Reef flat, reef margin, reef front, 1-2 meters of
water on the north side.

Al. Mar. 19. Himmafuri Faru. (73234 UE ac 8 3075N) 7 Cymodocea
bed just below low tide on the north side of the island near the village.

Be Mar. 21". As above but on the south side of the island.
5a. Seaward reef flat, 1 meter or less of water.

5b. Algal ridge.

5c. Velu, 3-4 meters of water.

6. Mane 22% South of Kagi Faru. (73°34"E,. 4°32—351N)2 Lagoon
floor in 60-66 meters, beam trawl.

ke Mare: 92:2). Kagi Faru. Cig 290455 a Ge 4OeNiie Seaward reef flat
on the north side of the Faru, 2-6 meters of water.

Fadiffolu Atoll

Sk Marea 36 Between Mako (Maro) and Mafilafuri (Mafilefuri).
(32:52 AE ee Sic 701 atne OLIN) es Inter-island flat of calcareous debris, less
than 1 meter of water with current.

Qe Mar 24. As above but nearer the lagoon side, sheltered water
akp the, pount vot Vow tide

9a. Rhizophora roots.

9b. Coral debris especially Acropora.

9c. Sparse Thalassia bed.

Oe Mars 25) East of Lohi and Lasalafuri. @WB229 Eva 5 2 20.4N)) i=
Lagoon floor in 50-70 meters, beam trawl.

ie Mare 25). Near Maduvvari. C8229 UE a Silks UN) pe Channel across
the atoll reef in 50-70 meters, beam trawl.

Miladummadulu Atoll

iene 1 Mar 27 Kendikolu. (UBD Mae SSE 7/ Np) S Lagoon reef flat, 3
meters of water.

Se Marty 6 2i7>. Just west of Kendikolu. Lagoon floor in 50 meters,
wire dredge.

14. Mar. 28. Kendikolu. Patch reef adjacent to the lagoon beach,
less than 1 meter of water.

iS Mars. 129}. Near Bomasdu. G13: 19) GEy 2 5° SSN) x: Lagoon floor in
72 meters, beam trawl.

Tiladummati Atoll

eG. aeMar.? <3... Filadu. G/3-AO2, I62755¥ NN). Enclosed rubble pool on
the reef flat.

17. As above. Lagoon reef flat outside the pool, 5 meters of water.
Male Atoll
PGaen apes (19 . Dunidu Faru. See station 2. Littoral pools on

pavement and the pavement at the point of low tide.
IS), Nore AES )c As station 3.
North Malosmadulu Atoll

AO) Nowe YEE Ugu Faru (Ongu). (73 ORE «4 240" N))< Site similar to a
leeward reef flat, 1-3 meters of water.

2 ApY. 21. Near Ugu Faru. (73208 E, 4°41."N)): Lagoon floor in 44
meters, wire dredge.

Ari Atoll

DBs INSG5 PE Madugali. (C7224 5 UR 4 6 UN). Beach rock and the reef
flat of a coral knoll island, to 2 meters of water.

South Nilandu Atoll

236), sAprE. 23). Kuda Huvadu. (2255 UE 2 cao UN) ie Floating algal mat
from the lagoon beach.

10

24. Apr. 24. Walla Faru (Wala). G225 5. eae ADEN) Be Reef flat of
a coral knoll island.

Addu Atoll

Ps iNeed LAS}. Gan Island. (SiS) A OCALA) Littoral beach rock
from the lagoon beach near the inter-island flat.

26. HAD ssO% Wilingili. (732 TE O74 eS) Lagoon reef flat, less
than 1 meter of water.

27. ‘Mayr 4% As station 25. Coral rubble surrounding the pier on the
lagoon beach, 1 meter of water.

28.) . ‘Mayii5 North of the Gan-Fedu gap. (732 9NEVROSAMAUS)) i Lagoon
floor in 40-50 meters, wire dredge.

29. May 8. As station 25. Small rubble island on the inter-island
flat between Gan and Fedu. Cymodocea-Thalassia beds and littoral
calcareous debris.

30; May lO. BusbyIsland. C73 °NOUR RT OCAOMUS i Lagoon reef flat on

the western side of the island, coral rubble and shells in 1 meter of
water.

Stations of other expeditions

Barton. Algae collected during the J. Stanley Gardiner Expedition to
the Maldives were listed by Barton (1903). All the Maldive plants
were dredged in water 25-45 fathoms. The locations are given in the

systematic list.

Foslie. Coralline algae collected during the Gardiner Expedition were
listed by Foslie (1903, 1907) and Weber Van Bosse and Foslie (1904).
The locations are given in the systematic list.

DS. Algae collected by D. Sigee during the Cambridge to Addu Atoll
were listed by Tsuda and Newhouse (1966). For locations consult that
paper.

Rhyne. Algae were collected at Addu Atoll in August 1967 by C. Rhyne
during the U.S. Navy Biological Expedition to Diego Garcia. All were
found on the shallow inter-island flat between Gan and Fedu.

iLal
Systematic list
Cyanophyceae

Anacystis aeruginosa (Zanardini) Drouet and Daily
Stations: 23, 25; on Enteromorpha and Schizothrix or interstitial.

Brachytrichia maculans Gomont
Stations: 29; on calcareous debris with Calothrix in Cymodocea beds.

Calothrix crustacea Schousboe and Thuret
Stations: 4, 18, 22, 29; on coral debris, beach rock and on
Cymodocea.

Calothrix pilosa Harvey
Stations: . 167 6, 929) BDS 1202 Btonmingra tusk onscoral ~mubbie iin
Cymodocea beds or blackening supralittoral debris in undisturbed
areas.

Coccochloris elabens (Brebisson) Drouet and Daily
Stations: 15; Several irregular and often medianly constricted cells
of this tentatively identified alga (Drouet, 1967 personal communica-
tion) are within many of the blade cells of the monostramatic
specimens of Dictyopteris.

Entophysalis conferta Drouet and Daily
Seatlonsiin |S), Dalp Dbp Sc, 1S), LS on Microcoleus, SchiZocehnrix,
Polysiphonia, and Valoniopsis.

Entophysalis deusta Drouet and Daily
Stations: 3, 4, 8, 9, 13, 16, 18, 26; ‘onjand in calcareous,debrisis

Fremyella vitiensis (Askenasy) De Toni
Stations: 4, on various algae in Cymodocea beds.

Hormothamnion enteromorphoides Gritinow
Stations: 4, 25; on Bgergesenia, Thalassia, and Cymodocea or forming
a crust on sand in crevices of beach rock.

Hormothamnion solutum Bornet and Griinow
Stations: 9; Rhyne 814a; in the algal turfs on Rhizophora roots
and on Thalassia.

Mastigocoleus testarum Lagerheim
Stations: 9, 28, 29; penetrating beach rock and shell debris.

Microcoleus lyngbyaceus (Ktitzing) Crouan
Sueaclemss Sp 4, Sap Slo, Se, Ye NO, aly, Us, Aaa 2G, 295 SOF wohisaes
816b; on coral debris, beach rock, in sand or epiphytic, never
dredged.

q°2

Oscillatoria submembranacea Ardissone and Strafforello
Stattonsiz 93), 144 5a, SC, 2, Oa Oy ie 2S SOs Rhyme) 1Si2 5)
forming erect tufts on beach rock or coral debris, or forming spheres,
binding sand, or epiphytic.

Oscillatoria lutea Agardh
Stations: 4, 26; ‘on calcareous debris in Cymodocea beds.

Porphyrosiphon notarisii (Meneghini) Gomont
Stations: 115,24; on a hydvoid, on: Microdictyon.

Rivularia polyotis (J. Agardh) Bornet and Flahault
Stations: 9, on Thalassia, on shells.

Schizothrix arenaria (Berkeley) Gomont
Stations: -5a,"5e) lOa, 257 sRhyne 85; “in sand.

Schizothrix calcicola (Agardh) Goment
SPaesdons: 243i, 247577 On) Oy UO, eg, LS noel, e 26), Ore DSOor
a common algal epiphyte and often in the tissues of sponges and
Zoanthus.

Schizothrix mexicana Gomont
Stations: “5a, 5b, 5c, 9), 10a, lb, 15,.17, 2207242267) (26) eeRhyne
816a; with other bluegreen algae in turfs especially on the lower
parts of coral, or epiphytic on red algae.

Schizothrix tenerrima (Gomont) Drouet
Stations: 20, on Gelidium.

Spirulina subsalsa Oersted
Stations: 9, 16, 17, 18, 20; in turfs with other bluegreen algae
especially Microcoleus.

Rhodophyceae

Acanthophora spicifera (Vahl) Bérgesen
Stations: 4, 9b; in Cymodocea and Thalassia beds, rare. Plants
of small stature and heavily epiphytized.

Acrochaetium sargassicola Bérgesen
Stations:. 20, 30; on Dictyota, Turbinaria, and coralline algae.

Acrochaetium sp. 1
Stations: 17, 22, 26, 30; on various algae. Plants resemble
A. gracile Bérgesen, but with larger dimensions.

Acrochaetium sp. 2
Stations: 4, 24; on Champia and Hypnea from Cymodocea beds, on
Dictyota from the reef flat. Plants O.2mm - 0.4mm high, sparsely
branched; basal cell 18 uw diam., spherical, at least in young plants,
ultimately forming a filamentous creeping axis or a disk like

IE3)

structure; lower cells of the upright axis 7-8 uw diam., 10 u long,
ultimate unicellular branchlets usually arising from each of the
2-4 cells of a side branch; plastid band shaped to fragmented.

Actinotrichia fragilis (Forskal) Bgérgesen
Stations: 7, 17; occasional in crevices.

Amphiroa anastomosans Weber Van Bosse
Stations: 5b, 7, 24, 26; in crevices of the reef flat and in mixed
algal turfs of the algal ridge.

Amphiroa fragilissima Lamouroux
Stesneilomiss Slo, Slop — all enleeul ie bbeesi-

Amphisbetema indica (J. Agardh) Weber Van Bosse
Statrons: | DS, al2-

Antithamnion breviramosus Dawson
Stations: 7, 10a; on Valonia from the reef flat, on dredged
Halimeda. The axis diameter is smaller than variety simplex
Dawson (1957) from Eniwetok. Plants are poorly developed, and not
all have gland cells.

Antithamnion butleriae Collins
Stations: 10a. Species previously known only from the West Indies.

Antithamnion pseudocorticatum Dawson
Stations: 15, on Microdictyon. Plants are better developed than
the type. The lower side of the basal cell of the ramuli cuts off
a spherical but not densely staining gland-like cell.

Antithamnion sp. 1
Stations: 15, on Microdictyon and Dictyopteris. Plants are like
A. sublittorale Setchell and Gardiner, but have di-trizonate gland
cells.

Antithamnion sp. 2
Stations: 10a, on calcareous debris. Plants fragmentary, indeter-
Minate axes unbranched, cells 15 u diam., 50 u long; determinate
ramuli arising near the upper end of the axis cell, opposite,
unbranched, to 175 yw long, lower cells 10 u diam., 17 wu long,
terminal cell 5 uw diam., 12 uw long, blunt; flattened gland cells
10 uw long, laterally cut off from lower cells of the ramuli.

Antithamnion sp. 3
Stations: 15, on Microdictyon. Plants fragmentary, axis cells
20 uw diam., 70 uw long, each giving rise in a spiral arrangement to
two adjacent and thus not opposite, unbranched, determinate ramuli
to 180 uy long, one often deciduous in older parts; cells of lower
ramuli 8 uw diam., 20 uw long, terminal cell blunt; supposed gland
cells not as distinctively dark at maturity as most species, -
ovoid to spherical 10 u — 15 yw borne usually on the lower side at
a midpoint on the ramuli.

14

Archaeolithothamnion erythraeum (Rothpletz) Foslie
Stations: Foslie, South Nilandu.

Archaeolithothamnion schmidtii Foslie f. dissita Foslie
Stations: Foslie, South Nilandu and Male

Asparagopsis taxiformis (Delile) Collins and Hervey
Stations: 24, 26; tetrasporophytic plants only.

Asterocytis ornata (C. Agardh) Hamel
Stations: 12, 17, 24, 26, 30; on Sphacelaria, Entophysalis,
Polysiphonia and other turf algae in crevices.

Botryocladia skottsbergii (Bérgesen) Levring
Stations clon 215 26 DS w238e

Callithamnion sp.
Stations: 10a, 15: Plants erect to lcm, axes monosiphonous,
alternate to dichotomously branched, branches spirally arranged
and apparently indeterminate and not differing from the axes, usually
straight; lower cells 37-55-75 uw diam., 212-250 uw long, somewhat
expanded at the ends; upper cells 15 u diam., 100 u long; terminal
cells 8 u diam., 26 yw long, apex wall very thick, blunt; not
reproductive.

Centroceras clavulatum Montagne
Stations: 9b, fragments numerous in algal turfs on dead coral.

Centroceras minutum Yamada
Stations: 9b, 26, 29; in algal turfs on dead coral.

Ceramium fibriatum Setchell and Gardner
Stations: 8, DS 105; on sandy areas of the reef flat, rare.
These plants are the only conspicuous Ceramium species found.

Ceramium gracillimum (Griffiths) Harvey var. byssoideum (Harvey)
Mazoyer
Stations: 4, 7, 22, 24, 29; Rhyne 804, on Cymodocea, Thalassia,
Halimeda, Dictyota, and Amphiroa.

Ceramium mazatlanense Dawson
Stations: 10, 11, 21, 24; on Codium and Dictyota from the reef flat
and on Bryozoans, Microdictyon, and Pocockiella from the lagoon floor.
Plants are less developed than typical.

Ceramium sp. 1
Stations: 25, 30; on beach rock and debris. Plants have a greater
number of tetrasporangia at a node and a more regular arrangement of
cortical cells than C. huysmansii Weber Van Bosse.

Ceramium sp. 2
Stations: 9c, on Thalassia. Plants to 3mm high, sparingly and
irregularly branched, apices curved; mature axis cells 85 u diam.,

15

SON ie vong; mature cortical band 200 u diam., 100 yu high with some
secondary growth, cells 8-24 uy diam., irregular not organized into
zones; forming bisporangia, 27 wu diam., 46 u long, up to 10 ata
node, involucrate.

Ceramium sp. 3
Stations: 18, on beach rock. Plants to 2mm high, sparingly and
irregularly branched, apices straight, mature axis cells 125 u dian.,
187 uw long; mature cortical band 150 uw diam., 125 uw high with much
secondary growth, cells usually round 13 u, not organized into zones,
not reproductive.

Ceramium sp. 4
Stations: 30, on coral debris. Plants creeping to erect, minute,
rarely branched, apices straight; mature axis cells 33 u diam.,
90 u long; mature cortical bands 43 uw diam., 24 wu high; cells
irregular 8-13 uw, irregularly arranged into more or less three
transverse zones, rhizoids occasional arising from the bands, not
reproductive.

Ceramium sp. 5
Stations: 15, on Microdictyon. Plants to 3mm high, abundant
dichotomous to alternate branching, apices straight; mature axis
cells 163 wu wide 188 uw long, bulging; mature cortical bands 160 4
wide, 62 u high; cells small highly irregular with some gland
cells, not reproductive.

Ceratodictyon spongiosum Zanardini
Stations: DS 69.

Champia parvula (C. Agardh) Harvey
Sizmclonmss sip 4 Wp Soi Sten Aare AS wis} IEAZe ye islanisnye) te O)F Voyal
Thalassia, Gelidium, Dictyosphaeria, Padina and other algae.

Champia salicornoides Harvey
Statkvonsias 21) SDSel22b-

Champia sp.
Stations: 13; Plant fragmentary to 3cm high; .5-1mm dian.,
sparingly branched; not constricted at septa; wall of one layer,
cells 34-84 u long, 28 uw diam., small cells interspersed 10 yp diam.,
medullary filaments 5 wu diam.

Chondria dasyphylla (Woodward) C. Agardh
Stations: 4, rare. Plants are poorly developed.

Chondria simplicuscular Weber Van Bosse
SEAETONS 4), 5b), Cin, 20)) 257) .aecommon component of tunis.

Chondria spp.
Four additional species of Chondria from the reef flat and lagoon
floor are being studied.

16

Chrysymenia sp.
Stations: ll. Plants fragmentary but resemble C. enteromorpha
Harvey but have cells twice the size.

Coelarthrum boergesenii Weber Van Bosse
Stations: 24, rare, in crevices. Gland cells are not distinctive
or numerous.

Crouania minutissima Yamada
Stations: hs

Cruoriella sp.
Stations: 4, 20; on calcareous debris from the reef flat and on
Conus from a Cymodocea bed.

Cryptonemia crenulata J. Agardh
Stations: 15. Surface cells are slightly smaller than typical
West Indian plants.

Dasya spp. .
Three species of Dasya from the lagoon floor and algal ridge are
being studied.

Dasyopsis geppii Weber Van Bosse
Stations: DS 95,101.

Dictyurus maldivensis Hackett and Aregood
Stations: 13, 15; see Aregood and Hackett (1971).

Dictyurus purpurascens Bory in Belanger and Bory.
Stations 124777925), (26). DS 247

Erythrocladia subintegra Rosenvinge
Stations: 7, 26; on Valonia and Struvea from crevices.

Erythrotrichia carnea (Dillwyn) J. Agardh
Stations: 4, 10, 25, 26, 30; ‘on Champia, Hypnéea, Dictyota;and
Thalassia from the reef flat, on Pocockiella from the lagoon floor.

Fauchea peltata Taylor
Stations: (63° .DS: 764%

Fauchea repens (C. Agardh) Montagne
Stations: “15%

Fosliella farinosa (Lamouroux) Howe
Stations: 4, 7, 9c, 26, 29; on Cymodocea, Thalassia, and various
algae.

Fosliella sp. 1
Stations: 4, on Padina and Dictyosphaeria. Plants mostly three cell
Layers thick; ‘cells 15‘ diam::, "20 u“long;-cap*cel ls lacking; sneters
ocysts 25-32 yw diam., 32-37 u long, colorless, spherical to ovoid.

157

Fosliella sp. 2
Stations: 4, 12; on Cymodocea and Valonia. Plants mostly one cell
layer) thick.) cells variable LO diam, 15 jt llong,, cutting, off cap
cells that are prominent on the surfaces of immature conceptacles;
heterocysts 12 uw diam., 26 uw long, not particularly distinctive.

Galaxaura filamentosa Chou
Stations: 26, on calcareous debris.

Galaxaura marginata (Ellis and Solander) Lamouroux
Staclonsem se DsmlO4 =

Galaxaura rudis Kjellman
Stations.) DS e21

Gelidiopsis sp.
Stations: LO) Plants fragmentary.

Gelidium crinale (Turner) Lamouroux
Stations: 9a, on roots of Rhizophora mucronata.

Gelidium divaricatum Martens
Sitaitslonsi) 20); 267) DS 72; In aligal “eurEs.

Galidium pusillum (Stackhouse) Le Jolis var. miniscula Weber Van Bosse
Sitatvonsi: | 26.

Gelidiella sp.
Stations: 5c, 26, in algal turfs on Acropora.

Gracilaria corticata J. Agardh
Stations: 4, rare. Plants poorly developed.

Gracilaria sp.
Stations: 4, rare. Plants are poorly developed but resemble
G. verrucosa. Central cells are smaller than typical and branches
do not show the basal attenuation of Ceylon specimens.

Griffithsia rhizophora (Griinow) ex Weber Van Bosse
Stations: 24; DS 80; in algal turfs on Dictyota, Caulerpa, and
Hypnea.

Griffithsia tenuis C. Agardh
Stations: 24, on Halimeda.

Goniolithon fosliei (Heydrich) Foslie
Stations: Foslie, Male.

Goniolithon frutescens Foslie
Stations: 4, 5a, 9b; DS 98, 113; Rhyne 798c, 828; Fosilile, Male
and Suvadiva.

18

Goniolithon frutescens Foslie f£. congesta Foslie

Stations:

Goniolithon laccadivicum Foslie £.

Stations:

9b; Rhyne 820a; Foslie, Male and Suvadiva.
typica Foslie

Foslie, Male.

Goniolithon myriocarpon Foslie

Stations:

Foslie, Male.

Goniolithon reinboldi (Weber Van Bosse and Foslie) Foslie

Stations:

Foslie, Male.

Goniolithon sp.

Stations:

4, on Goniolithon frutescens.

Haloplegma duperrayi Montagne

Stations:

Ss

Halymenia durvillaei Bory

Stations:
rare.

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Stations:

Herposiphonia
Hollenberg
Stations:

Day OA, aD Speleail Plants poorly developed and very

crassa Hollenberg ?
9b, on Dictyota from coral rubble.

delicatula Hollenberg
Oi, 23

dendroidea Hollenberg var. minor Hollenberg
30.

filifera Hollenberg
Si ZGis

obscura Hollenberg
ibe

pacifica Hollenberg
5a, 5c,., 9b, 20,:-29;' won’ Caulerpa) and other, algae:
parca Setchell

5c,:7, 24, 29; “DS 78; in Cymodocea beds and on Acropora’.

tenella (C. Agardh) Schmitz
24, on Pocockiella.
tenella (C. Agardh) Schmitz f. secunda (C. Agardh)

24.

Heteroderma lejolisii (Rosanoff) Foslie

Stations:

5b, 7, 21, 24, 26, 28; on Valonia, Chaetomorpha, Struvea,

Valoniopsis, Chondria, Microcoleus and other algae.

IS)

Heteroderma sp. 1
Stations: 5b, 7; on Hypnea, Pocockiella and Valonia. Plants one
cell layer, initial branhcing dichotomous, cells rectangular;
ultimate branching irregular but parallel, cells sigmoid 10 u diam.,
DA Nh Werereye Adjacent branches form a reticulate pattern the lense
Shaped openings are formed by two sigmoid cells; cap cells but no
heterocysts.

Heteroderma sp. 2
Stations: 5b, on Chondria. Plants one cell layer, in crusts to
125 wu wide, cells 7-12 wu diam., 10-17 uw long; crusts connected by
irregularly branched filaments, cells 12 uw diam., 25-32 yu long;
cap cells but no heterocysts.

Heteroderma sp. 3
Stations: 5b, 12; on Valonia and Laurencia. Plants one cell
layer, mostly of fan-like reticulate crusts formed by dichotomous
branching some plants with a more filamentous tendency, cells
irregular 8-10 uw diam., 10-25 uw long; conceptacles 85 u high, 50 u
basal diam., conical tapering to a single pore; cap cells but no
heterocysts.

Heterosiphonia sp.
Sittatwonsen bs) ells. Plants with distinctive acute apices.

Hypnea esperi Bory
SEAELONSI (5), 4, .0b, 7, 9C, 16, 20, 24, 26; DS 112, 18, 93; Rhyne
826.

Hypnea valentiae (Turner) Montagne
Stations: 4, 9b; DS 91; Rhyne 807.

Hypoglossum minimum Yamada
Sessoms . IOp_) snow 2An yee

Hypoglossum sp.
Stations: 28, on a tunicate. A single small spermantangial plant.

Jania capillacea Harvey
SREBieslLomse: Ap, Hop B67 yp Gatp Slop MO, wh, iy POR 2272237 PS, 25n Zor
30; DS 11; Rhyne 809a. Plants are almost always a component of
algal turfs, but best developed in Cymodocea beds. Two dredged
fragments were probably washed from the reef.

Janua ungulata Yendo
Sitatvons:) §20), on: Lurbinaria’.

Laurencia parvipapillata Tseng
Srataonsi:y ob

Laurencia spp.
Six additional species of Laurencia from the lagoon floor are being
studied.

20

Leveillea jungermannioides (Martens and Hering) Harvey
Stations: 4, rare.

Lithophyllum kotschyanum Unger f. typica Foslie
Stations, Sb, 2: "pSi92);.. Hosiiie,, (Maile:

Lithophyllum kotschyanum Unger f. madagascarensis Foslie
Stations: Foslie, Male.

Lithophyllum kotschyanum Unger f. subplicata Foslie
Stations: Foslie, Male.

Lithophyllum kaiserii Heydrich f. subplicata Foslie
Stations: Foslie, Male.

Lithophyllum spp.
Three additional species of Lithophyllum are being studied.

Lithothamnion maldivicum Foslie
Stations: Foslie, Male.

Lithothamnion indicum Foslie f. typica Foslie
Stations: Foslie, Male, Addu and South Nilandu.

Lithothamnion fructiculosum (Ktitzing) Foslie f. clavulata Foslie
Stations: Foslie, Male and South Nilandu.

Lithothamnion spp.
Two additional species of Lithothamnion including DS 117 are being
studied.

Lithoporella melobesioides (Foslie) Foslie
Stations: 4; Foslie, South Nilandu.

Lophocladia lallemandii (Montagne) Schmitz
Sitcationsis, lS.

Martensia fragilis Harvey
Stations: yWiO;, 26.

Peyssonelia calcea Heydrich
Stations: 3, on Acropora.

Peyssonelia gunniana J. Agardh
Stations: “DS 53:

Peyssonelia rubra (Greville) J. Agardh var. orientalis Weber Van Bosse
Stations: 12; Barton, Suvadiva.

Peyssonelia spp.
Five additional species of Peyssonelia from the reef flat are being
studied.

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Polysiphonia
Stations:

Porolithon craspedium (Foslie)

Stations:

Porolithon craspedium (Foslie) Foslie f.
Foslie, Male.

Stations:

Porolithon craspedium (Foslie) Foslie f. abbreviata Foslie
5b; Foslie, Male.

Stations:

Porolithon gardineri (Foslie)

Stations:

Porolithon onkodes

Stations:

exilis Harvey

24, with Herposiphonia on corals.

ferulacea Suhringer in J. Agardh

DS 35a.

flaccidissima Hollenberg

Zi SOs

howei Hollenberg

18.

pentamera Hollenberg

TESTA

poko Hollenberg var. langii Hollenberg

DS) 35:

scopulorum Harvey var. villum J. Agardh

die S829 AOS

spherocarpa Bérgesen

26.

tepida Hollenberg

Ze

upolensis (Grunow)

BOe DS) 29e%

12; Foslie,

12.

Porolithon sp.

Stations:

5b, the prime constituent of the algal ridge.
like P. craspedium in part, but are much encrusted and perforated

Foslie £.

Foslie £.

(Heydrich)
Spy obs, MSaG:

Hollenberg

Foslie

Foslie, Male.

and may be P. gardineri or P. sequinoctale.

Spermothamnion?
ByGl7 Ge Plants not reproductive.

Stations:

typica Foslie

compressa Foslie

typica Foslie

aX

Plants are

22

Spyridia filamentosa (Wulfen) Harvey in Hooker
Stations: 5c; DS 97: Rhyne 808a.

Taenioma nanum (Klitzing) Papenfuss
Stations: 26 in an algal turf on Acropora.

Tolypiocladia glomerulata (C. Agardh) Schmitz
Staitiions:-.3,: 4), v4, 20),) 26,7, 508.1 DS e447) ocRhyne SOM nor

Wrangelia argus Montagne
Stations: 22, 25.

Vidalia serrata (Suhringer) J. Agardh
Stations: 26") DS 19).

Vidalia sp.
Stations: 16, E720, 24; am “crevices: Plants erect to 4cm,
bushy and compact, axes flat to lmm broad; cross section .3mm thick
near margin, .37mm thick in center; alternately branched and
ultimately spinulose; upper apices inrolled; faintly costate on
staining; axis cells 100-200 u long, 75-90 yu diam., pericentrals
5; two layers of cortical cells, outer 5 wu diam., 10 uw long usually
in pairs; inner cells 100-125 uy long ovoid; gland cells ovoid
12 uw diam., 22 wu long often reaching the surface; spermatangial
clusters near the apices.

Chlorophyceae

Acetabularia sp.
Stations: 4. A single immature specimen.

Anadyomene wrightii Gray
Stations: 5b, 10, on Valoniopsis in the algal ridge and lightly

attached to sand on the lagoon floor.

Bgergesenia forbesii (Harvey) Feldmann.
Statvonsi::) 4, 30; Doo; Rhyne.

Boodlea composita (Harvey) Brand

Stakvons 5Ci,0 1S, ¢26) DS mO4 ye meRhy nem 799), Plants from northern
atolls are smaller in all aspects than what is usually reported and
resemble B. struveoides Howe. Addu plants are typical.

Boodlea van-bosseae Reinbold
Stationsi:* 9b, 20; 30),, on rubbilevor jini jaligalj tuts.

Boodlea sp.
Stations: “DS 63% Plants minute, inner cells 22 uw diam., 58-72 u
long; ultimate cells 22 u diam., 31 uw long, often curved, some
smaller and rhizoidal; hapteral cells various but usually square
Der ite

728)

Bryopsis pennata Lamouroux
Statrons: 26); DS) V4), 54%

Bryopsis sp.
Stations: 21, on Struvea. Axis .25mm basal diam., .2mm diam. where
ramuli diverge; ramuli alternate, 62 u diam., 1.5mm long, basally
constricted to 25 u, obtuse, mature ramuli often septate at base or
with ingrowths of the wall; plastids oval to spindle shaped
connected by fine strands.

Caulerpa ambigua Okamura
Stations: | 40 19926) 50.

Caulerpa crassifolia (C. Agardh) J. Agardh
Stdteronsi 23), lO, Lily 245

Caulerpa lentillifera J. Agardh
Stations: DS 8b.

Caulerpa racemosa (Forsk&l) J. Agardh var. clavifera (Turner) Weber
Van Bosse f. simplicissima Bgérgesen
Stations: 9c; Rhyne 80O2a.

Caulerpa racemosa (Forsk&l) J. Agardh var. macrophysa (Kiitzing) Taylor
SitatlOns: 07), Opal, eee yt eon2Oss Dosa; | Rhynes 611.

Caulerpa racemosa (Forskal) J. Agardh var. microphysa (Weber Van Bosse)
Taylor
Stations: 16.

Caulerpa racemosa (Forskdal) J. Agardh var. peltata (Lamouroux) Eubank
Stations: (5b, 5c, 247, 26, 30; Rhyne 802)(2)):

Caulerpa serrulata(Forskal) J. Agardh var. typica (Weber Van Bosse)
Tseng
Stations: 8, 9b, 25, 30; DS) 47; Rhyne 802(1), 812.

Caulerpa sertularioides (Gmelin) Howe
» SEENON UA YA 6

Caulerpa taxifolia (Vahl) C. Agardh
Stations 9c, 9b, 257) DS AO) Rhyne) 803).

Caulerpa verticillata J. Agardh
Stations: 28.

Caulerpa sp.
DtatLtonsic: e2alse Plants as C. serrulata but with reduced serrations

and no twisting.

Chaetomorpha brachygona Harvey
Stations: DS 66.

231-339 O-'77-5

24

Chaetomorpha crassa (C. Agardh) Ktitzing
Stations: < 4, 11, 5; DS LOG; Rhyne (8242

Chaetomorpha gracilis Kiitzing
Stations :iyell@) «clients. DS HOS:

Chaetomorpha javanica Ktitzing
Stations: 3, on coral with Enteromorpha.

Chaetomorpha linum (Muller) Ktitzing
Stations 7 147 alles as Sr

Chaetomorpha minima Collins and Hervey
Sitaitalionsi) Si a5) 26),, SOn

Chlorodesmis hildebrandtii A. and E. S. Gepp
Stations: 24, a single fragment.

Cladophora patentiramea (Montagne) Ktitzing?
Stations.) 221.

Cladophora crystallina (Roth) Kutzing?
SivatevOnsS capmlale ilesr all S ye lee

Cladophora inserta Dickie f. ungulata (Brand) Ktitzing?
Stations: 4, 7, 15, 30; epiphytic on various algae and Cymodocea.

Cladophora sp. 1
Stations: 4; DS 55; Rhyne 8097832. Plants are similar to an
undetermined species from Rongelap (Taylor 1950).

Cladophora sp. 2
Stations: 13, 28, on Dictyota, a Pecten shell and calcareous debris.
Plants are microscopic, prostrate with branching only in the plane
of the substrate; rhizoids terminating in a bulbous cell that
deeply penetrates the substrate or adheres to the surface of other
algae.

Cladophora spp.
Three additional species of Cladophora are being studied.

Cladophoropsis sp. 1
Stations: 10), 1, 15,>21,..0n Struvea’ and Microdictyon: Plants
attached by the expanded to crenulate basal portion on the original
coenocyte; in one plant basal portion developing colorless rhizoidal
filaments; mature segments generally dichotomously branched, 90 U
diam., to 575 uw long; ultimate segments at least in younger parts
moniliform without reference to septa; plastids discoid or
associated.

Cladophoropsis sp. 2
stations: 26, in algal tune: Plants fragmentary but resemble
C. suddanensis Reinbold.

25)

Cladophoropsis sp. 3
Stations: 26, on Caulerpa. Plants fragmentary and poorly developed
but resemble C. gracillima Dawson.

Cladophoropsis sp. 4
Stattons? 295, "e2i- Plants creeping to erect, branching irregular to
pectinate, often ending in rhizoids; rhizoidal segments 37-45 u
diam., 250-380 u long, rarely to 2mm long; lower axis segments
100 uw diam., 250-270 uw long, obtuse.

Codium arabicum Kittzing
Seaisltonsis | 20h 26e) DS A235

Codium edule Silva
Seacionsi 24,0263, DS 22.

Derbesia attenuata Dawson
Stations; 9247, 926 jp ainsalgall tums.

Derbesia sp. 1
Stations: 26, in algal turfs. Plants have smaller dimensions than
D. marina (Lyngbye) Kjellman.

Derbesia sp. 2
Stations: .30;.°DS 73: Plants erect to 5mm not particularly lax
irregular creeping axes 25-37 uw diam., erect axes dichotomous to
alternate; reproductive axes 75 u diam. tapering to 28 yu obtuse,
rarely with septa; sporangia oval, 90 uw diam., 142 uw long at
maturity; pedicle, as limited by septa, 20 u diam., 12 yu long;
plastids discoid, 5 wu diam.

Dictyosphaeria cavernosa (Forsk&l) Bérgesen
Stations: 4, 5b, 18, 30; DS 40 as D. intermedia var. intermedia;
Rhyne 830.

Enteromorpha compressa (L.) Greville
SHeciiciwoingn - Sip Slop Ale ilsy= exshe

Enteromorpha prolifera (Mtiller) J. Agardh subsp. prolifera typus III
Stations: 23, floating in calm water.

Enteromorpha spp.
Five additional species of Enteromorpha from the reef flat are being
studied.

Entocladia viridis Reinke
Stations: 4, 5c, 15, 30; in the sheath of Polysiphonia, Champia,
Lurencia, Schizothrix and on hydroids.

Gomontia polyrhiza (Lagerheim) Bornet and Flahault
Statmons-6.4, 5a,.9bs ean conalisy isheldis,; and. calcareous debrils.

26

Gomontia sp.
Stations: 16, 28; in shells. Axes and branching highly irregular,
cells 12-18 wu diam., 20-55 uw long, some empty others with spores,
but not other wise modified as sporangia, spores round 5 u.

Specimens of Halimeda are being studied by Colinvaux but the follow-
ing have been reported.

Halimeda discoidea Decaisne
Statvonsics 7 DS! alc

Halimeda copiosa Goreau and Graham
Statdvonsiii. 15),.92 16. See Colinvaux (1968), as H. hederacea.

Halimeda incrassata (Ellis) Lamouroux
Stations: DS 26, 75, 108; Barton, Addu.

Halimeda opuntia (L.) Lamouroux
Stations: DS 2a, 3, 4; Barton, Addu.

Halimeda tuna (Ellis and Solander) Lamouroux
Stations: Barton, Suvadiva.

Microdictyon pseudohapteron Gepp and Gepp
Stations: 5a, 9b, 11, 13, 15; M. pseudohapteron f. luciparense
Setchell is the only Maldive alga reported by Newton (1953) for
the John Murray Expedition. It was dredged from the lagoon floor
of Mulaku (Mulakadu) Atoll.

Microdictyon sp. 1
Stations: 11, 13, 15; several unreported fragments are in the
British Museum collection from the lagoons of Addu and Suvadiva.
They were dredged during the J. Stanley Gardiner Expedition.
Plants resemble M. agardhianum.

Microdictyon sp. 2
Stations: selon. 2K Plants belong to the Eumicrodictyon section of
the genus.

Microdictyon sp. 3
Stations: 21, on Halimeda. Plants belong to the Calodictyon
section of the genus and resemble M. japonicum.

Neomeris annulata Dickie
Sitationsiaasc le

Neomeris mucosa Howe
Stations. DS. OQ:

Ostreobium brabantium Weber Van Bosse
Stations: 4, 17, 28; in Pecten and Tellina shells, dead corals and
in coralline algae.

2

Ostreobium reineckei Bornet
Stations: 4, 9b, 28, 16, 17, 19; in Pecten and other shells, dead
corals and coralline algae.

Pseudochlorodesmis furcellata (Zanardini) Bgrgesen
Stations: 7, on Halimeda.

Rhipidiphyllon?
Stations: 5b, on Valonia. Plants as R. reticulatum (Askenasy)
Heydrich but anastomosing with distal crenulations or fibulae
rather than annulations.

Spongomor pha?
Stations: 6, 30; Plants are boodleoid with terminal cells from
i to. 2...7mm: ong.

Struvea anastomosans (Harvey) Piccone et Grunow ex Piccone
Siceltcrlonss SBS, Gp MO, ise Ailp Belay Wo.

Tydemania expeditionis Weber Van Bosse
Stations: Glomeruliferous plants 3, 17, DS 32; Flabellate plants 6.

Udotea javensis (Montagne) A. and E. S. Gepp
Seacvons: 21), 24, 26, 28, 30.

Udotea orientalis A. and E. S. Gepp
Stations: DS 15; . Rhyne 800, 817. Probably a seasonal alga.

Ulvella lens Crouan
Stations: 4, 5b, on Cymodocea, Chaetomorpha and Dictyosphaeria.

Valonia aegagropila C. Agardh
Stations: 5a, 30.

Valonia utricularis (Roth) C. Agardh
Stations: DS 103, 111; Barton, Suvadiva.

Valonia ventricosa J. Agardh
SeacLvonss 116, Wa LO Late lo, 2h. DS. oe.

Valoniopsis pachynema (Martens) B¢rgesen
Stations: 5; DS 99, 118, as Cladophoropsis sp.

Phaeophyceae

Dictyopteris repens (Okamura) Bégrgesen
Stations: 11, 12, 13, 15, 26; Barton, Suvadiva, as Haliseris
delicatula. Some plants are monostromatic and may represent a
second entity.

Dictyota patens J. Agardh
Stations: 7, 24, 26, 30; ° DS 5b, as Dictyota sp.

28

Dictyota bartayresii Lamouroux
Stations:;:. Rhyne. 789,, 789b,. 820; DS 5a. as, D.. friabilis; Barton,
Addu.

Dictyota spp.
Two additional species of Dictyota including DS 70 [as D. friabilis
in Tsuda and Newhouse (1966) ] are being studied.

Ectocarpus indicus Sonder
Stations: s-Obi,. 25), 267 50 Plants well developed at Addu only.

Ectocarpus irregularis Kltzing
Stations: 12, on Porolithon.

Ectocarpus variabilis Vickers
Stations: 25, on Thalassia.

Hydroclathrus clathratus (Bory) Howe
Stations: DS 45; Rhyne 890b. Probably a seasonal alga.

Padina commersonii Bory
Sieeticnvomikien “hs GAs Gy Ay Dabs

Padina sp.
Sieeticnlormimg Aeyp / iy W/o Plants are thicker than the previous and
oogonia are non indusiate.

Pocockiella variegata (Lamouroux) Papenfuss
Stabionsi. 47 - obs. O4 ale oD Oy, lds, 2, Moy liao ee Oe

Rosenvingea intricata (J. Agardh) Bérgesen
Stations: Rhyne 798a on Thalassia. Probably a seasonal alga.

Sargassum sp.
Stations: 5b, on Lithophyllum. Plants’.to 1.cm high.

Spatoglossum sp.
Statvons:, 7152 Plants lack a differentiated epidermal layer but
otherwise resemble S. cornigerum J. Agardh.

Sphacelaria novae-hollandiae Sonder
Stations Say 2 225: eo

Sphacelaria tribuloides Meneghini
Stations:s) Misi 24)) 308 Germinating propagulae are common, but
mature plants are rare and poorly developed.

Turbinaria ornata (Turner) J. Agardh var. ornata f. evesiculosa
(Barton) Taylor
Stations: 4, 7, 20, 24, 26, 29, in well protected crevices and
Cymodocea beds.

29
Acknowledgment

The cruise of the Stanford University Research Vessel Te Vega
to the Maldives in 1964 was made possible by the National Science
Foundation and the Maldive Government. The scientific party, directed
by Dixie Lee Ray, is indebted to N. T. Hasen Didi, Assistant Minister
of Home Affairs, who acted as interpreter and guide. The author is
grateful to those who examined and identified specimens or sent
specimens for study. Among these are Francis Drouet, George Hollenberg,
James Price, Charles Rhyne, Roy Tsuda, and Michael Wynne.

References
Aregood, Carol C. and H. E. Hackett. LOWS A new Dictyurus
(Rhodophyceae-Dasyaceae) from the Maldive Islands, Indian Ocean.

J. Elisha Mitchell Sci. Soc. 87(3):91-96.

Agassiz, A. INSIO)B}- The coral reefs of the Maldives. Mem. Mus. Comp.
Zool. Harvard Univ. 29:i-xxv, 1-168.

Barton, Ethel S. (Mrs. A. Gepp). 1903. List of marine algae
collected at the Maldive and Laccadive Islands by J. S. Gardiner
Esq. J= Linn. Soc. (London), Bot. 35:475-482.

Colinvaux, Llewellyn H. 1968. New species of Halimeda: a taxonomic

reappraisal. J. Phycology A(T) = 30-342

Dawson, E. Y. IS ISYSie Some marine algae of the southern Marshall
Islands. PACT ELC SCI. Ok 25—66)

SSSSSSSess tO Sire An annotated list of marine algae from Eniwetok
Atoll, Marshall Islands. PaGrale! SCN. 11:92-132.

Foslie, M. LIOSE The Lithothamnia of the Maldives and Laccadives.
imnGardiner, ww. (Si. IOS The fauna and geography of the Maldive
and Laccadive Archipelagoes. 1:460-471.

SSS SSSS=55 1907. The Lithothamnia. In Gardiner, J. S. 1907-1936.
Percy Sladen Trust Expedition to the Indian Ocean in 1905.
1 (2) Trans. Linn. Soc. (London), Second Ser. Zool. Daf MO

Gardiner, J. S. IGJO)3}- The fauna and geography of the Maldive and
Laccadive Archipelagoes. Welles -ilS2)- Cambridge University
Press, Cambridge.

Hackett, H. B- 1969a. Marine algae in the atoll environment:
Maldive Islands. Duke Univ. Unpub. dissertation.

SSSSSSoas= UEKSSI9)- Marine algae in the atoll environment: Maldive
Islands. Proc. Int. Seaweed Symp. @galis}7/—US)ie.

30

Hass, H. 1962. Central subsidence. A new theory of atoll formation.
Atoll Res. Bull. 91: 1-4,

Newton, Linda M. 1953/2 Marine algae. John Murray Expedition
Reports, 9(5) :395-420.

Sigee: (Denies 1966. Preliminary account of the land and marine
vetegation of Addu Atoll. Atoll Res. Bull. 116:61-74.

Stoddart; D., Rapwed 1966. Reef studies at Addu Atoll, Maldive
Islands. Preliminary results of an expedition to Addu Atoll in
1964. Atoll Res. Bull. 1G) 22

tracey, dis Lb. UL, Pi. Ee (Cloud and7K. C.hmenxy,. 1955. Conspicuous
features of organic reefs. Atoll Res. Bull. 46:1-3.

Tsuda, R. T. and J. Newhouse. 1966. Marine benthic algae from Addu
Atoll, Maldive Islands. Atoll Res. Bull. 116:93-102.

United States Naval Oceanographic Office. 19513 Sailing directions
for the west coast of India. Hydrographic Office Publ. 63.

Weber Van Bosse, Anna and M. Foslie. 1904. The Corallinaceae of the
Siboga Expedition. Siboga Expeditie 6l. E. J. Brill, Leyden.

CHAGOS TRENCH
OUTER SLOPE

CAVE and TERRACE ZONE

REEF FRONT

SEAWARD REEF MARGIN -ALGAL RIDGE
BOULDER ZONE

SEAWARD REEF FLAT

ATOLL REEF ISLAND

CYMODOCEA-THALASSIA FLAT

LAGOON REEF FLAT

LAGOON REEF MARGIN

LAGOON SLOPE
\
LAGOON FLOOR

CORAL KNOLL

Fig. 1. A composite cross section of the conspicuous features of a
large Maldivian Atoll from the eastern side of the archipelago.
Eastern half of the atoll.

MmHofG. ZAG

LAGOON FLOOR
KNOLL REEF SLOPE

KNOLL REEF FLAT
FALU

& ate

CORAL KNCLL ISLAND *
Su

BEACH ROCK and PAVEMENT
KNOLL REEF FLAT
KNOLL REEF SLOPE et

TIDE

LOW
TIDE

LAGOON FLOOR

LAGOON SLOPE
LAGOON REEF MARGIN
LAGOON REEF FLAT

20
FATHOMS

MANGROVE

ATOLL REEF ISLAND

SEAWARD REEF FLAT
SEAWARD REEF MARGIN
REEF FRONT
INTER ATOLL FLAT

A composite cross section of the conspicuous features of a
large Maldivian Atoll from the eastern side of the
archipelago. Western half of the atoll.

ATOLL RESEARCH BULLETIN

NO. 211

THE BENTHIC ALGAL COMPOSITION, STANDING CROP,
AND PRODUCTIVITY
OF A CARIBBEAN ALGAL RIDGE

by Judith L. Connor and Walter H. Adey

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

May 1977

= o.oo

THE BENTHIC ALGAL COMPOSITION, STANDING CROP,
AND PRODUCTIVITY
OF A CARIBBEAN ALGAL RIDGE !

by Judith L. Connor and Walter H. Adey 2

Abstract

The distribution and standing crop of benthic algal species ona
Caribbean algal ridge (St. Croix) and its associated carbonate pave-
ments is discussed and contrasted with that of other eastern Caribbean
algal ridges and a Pacific algal ridge. Mean standing crops of
3 kg/m? (wet weight) and a species richness of about 40 species (for
ten 0.25 m@ samples) were encountered on St. Croix. Non-calcified or
fleshy algae are greatly reduced in standing crop with depth away from
the high wave energy ridge crests and species richness increases.

This is correlated with the greater grazing abilities of fish and
invertebrates under less turbulent conditions.

On reefs of typical lower eastern Caribbean islands, only where
the force of water movement across intertidal algal ridges prevents
intense grazing by fish and echinoids are general high levels of algal
standing crop and productivity developed. However, in the more
eutrophic waters of higher islands and where wave action is
dense standing crops of larger fleshy algae can also extend
sublittorally to depths of at least 10 meters. We suggest that the
extensive filling of grazing niches in tropical reefs and a general
retreat of plant tissues to protective carbonate structures has caused
a reduction of primary productivity in typical reef environments.

greater,

1 Contribution no. 25 from the West Indies Laboratory of Fairleigh

Dickinson University. This project was supported by the Smithsonian
Research Awards Program.

2 smithsonian Institution, Washington, D.C. 20560.
(Manuscript received March 1975 -- Eds.)

Introduction

The windward reef margins of large numbers of Pacific atolls and
some high islands are rimmed by an algal ridge, a partly intertidal
and supratidal calcareous reef framework built primarily by crustose

coralline red algae. Less well known are the scattered but locally
abundant occurrences of algal ridges in the Caribbean and tropical
Atlantic. Adey and Burke (1976) describe the distribution of algal

ridges in the eastern Caribbean, and Adey (1975) treats in detail the
morphology, structure and geological history of the well-developed set
of algal midges, on? St. ;Crowx: The latter author also summarizes the
more important literature on algal ridges. In this paper, we describe
the fleshy algal flora of the St. Croix algal ridges and its richness
and ecology in terms of standing crop.

Few studies have been done on the standing crops of benthic
marine algae on algal ridges or coralline-rich reef areas (coralline
pavements). Taylor (1950) describes the difficulty of collecting on
the Bikini ridge (Marshall Is.). He gives an account of removing
algae from a particularly thick zone (a few millimeters to a
centimeter in thickness) on the ridge. Dahl (1971) describes an
algal turf association on the fringing reef flat of American Samoa.
His algal associations of low standing crop and high diversity decrease
in coverage across the reef flat to the seaward side where first
crustose algae and then, further seaward, living corals predominate.
Quantitative biomass ,measurements were not given by Dahl. Another
Pacific account, Soegiarto's work on Kaneohe Bay, Hawaii (in Smith
et al., 1973) indicates that standing crop and algal diversity are
greatest in the reef flat surf zone, where water movement is greatest
(250 g/m dry weight) and decrease in all directions from there.

Doty's (1969, 1971) studies on the standing crop on an algal ridge
and associated coralline pavements off Waikiki Beach, Hawaii are among
the few detailed quantitative studies treating the attached flora of a
coralline reef. Using Doty's data for November, 1967 as being about
average for the entire period covered, the mean standing crop reaches
a maximum of about 3400 g/m wet weight on the algal ridge and a
minimum of about 1300 g/m in the deeper zone immediately behind the
ridge (approximately 1 meter deep).

The upstream-downstream method of determining the organic
productivity of an ecosystem has become a standard method of marine
biology. Johannes et al. (1972), Gordon and Kelly (1962) and Odum
(1956) discuss the method and cite its use on several reef structures
in the Pacific, We have applied upstream-downstream analysis of
dissolved oxygen to determine the gross and net organic productivity
of a single lobe of an algal ridge on St. Croix.

Site Descriptions and Methods

Our study of fleshy benthic algae distribution was conducted on
several of the algal ridges along the eastern shore of St. Croix.
The collections were especially concentrated on the Boiler Bay algal
ridge in the northeast (Fig. 1).

The south shore algal ridges are more or less directly exposed to
a prevailing ESE trade wind of 10-20 knots and to seas with wave

heights of 1-2 m. The maximum elevations of these mostly actively
growing ridges is about 20-50 cm above mean low water spring tide
levels (m.l.w.sp.). Benthic algae were removed from quadrats

located on Robin, Beach and Isaacs Ridges (Fig. 1).

The Boiler Bay algal ridge, in an intermediate stage of
development, consists of a series of 30-35, more or less horseshoe-
shaped coralline frameworks ranging in diameter from about 50 m to as

Tittle as 2-3 m-. For convenience of reference, the larger of these
"boilers", "microatolls" or algal ridge "lobes" are named as reefs in
figure 2. The Boiler Bay algal ridge, essentially in its present

plan, was actively growing from about 2000 to 500 years before present.
However, wave action became blocked by growth of a coral reef at the
mouth of the bay, and as surface degradation by borers exceeds
carbonate accretion by corallines, the ridge is slowly being eroded.
The trade wind swell is refracted about 90° over the coral reef and
into Boiler Bay, and at the present time wave heights seldom exceed

30 em: Maximum elevations of the Boiler Bay ridge now range from
5-10 cm above m.l.w.sp. on West End Reef to 17 cm above m.l.w.sp. on
Shark Reef, though they were probably somewhat higher in the past.

Distinctive zones of crustose coralline algae, coral and
coralline-coral pavement are visible on and around the ridges and were
delineated by color patterns on aerial photographs. Color patterns
on the ridges and the surrounding pavements are often due to the
fleshy algae populations peculiar to each zone, which in turn are
largely dependent on depth, wave action and the grazing of animals.
Eight distinct algal zones were determined and are described below.
Zones 1 to 4 lie on the coralline-constructed algal ridge; the
remaining zones lie on the associated carbonate pavements or other
rock as described. The zone locations in Boiler Bay are indicated
in figures 3 and 4. Detailed maps of the area, along with the
methods used in mapping, are given in Adey (1975).

Zone 1 is the horizontal strip of algal ridge which lies above
mean low water spring tide levels. Between wave crests, it is
potentially exposed to desiccation and intense sunlight, though the
wave wash is rather constant and severe drying did not occur during
our two year stay in St. Croix. Zone 1 is characterized by a turf of
Rhodophyta species up to 10 cm thick dominated by Hypnea spp.,
Laurencia spp., Jania spp., Amphiroa spp. and Gracilaria mammillaris.

Smaller amounts of other reds, as well as scattered clumps of
Sargassum spp., Colpomenia sinuosa and other Phaeophyta with
occasional small Chlorophyta species are also present.

Zone 2 lies at O to about 30 cm below m.l.w.sp. and slopes
gently shoreward, averaging about 15 cm in depth. The substrate in
this zone is also dominantly coralline algae, though an occasional
Porites astreoides is also present. Zone 2 is chiefly populated by
Halimeda opuntia, Laurencia papillosa and Gelidiella acerosa, though
Pterocladia americana and Jania spp. are also important.

Below zone 2, the algal ridge continues to slope shoreward,
tending to form open-backed basins behind each boiler or lobe with
depths of about 30 to 60 cm. These basins are designated as zone 3.
Here, Porites astreoides occupies about 15% of the surface area, and
there are also scattered sand pockets in the coralline substrate.

The flora here is similar to the second zone, but Laurencia papillosa
and Halimeda opuntia are reduced and the total biomass of fleshy algae
is one third less than in the second zone.

The vertical seaward faces of the algal ridge at about 1 to 1.5m
depth were designated zone 4. This is a relatively smooth coralline
surface with a light cover of fleshy algae of many small species, the
most obvious being DiGtyopteris delicatula. All quadrats from zone 4
were taken centered at about O.5 m below m.l.w.sp.

Zone 5 occurs on the nearly flat and irregular seaward margin of
the fore-ridge carbonate pavement at depths of 3.5 to 4.5 m. The
substrate in this zone is a largely cor-algal pavement of dead corals
cemented together by crustose corallines, foraminifera and probably
submarine cementation of sediment. A few live corals, Porites
astreoides, Porites porites, and Siderastrea spp., are also present.
The dominant algae are Halimeda opuntia, Dictyota divaricata,
Amphiroa tribulus and Halimeda tuna.

Zone 6 is also predominantly a cor-algal pavement, but about 11%
of the surface is covered by the same living corals that characterize
zone 5, Porites astreoides being especially important. This zone
occurs around the sides and shoreward margins of the ridge lobes at
depths; of7 0.'5: to; 1:2)5*:me The most abundant algae here are Halimeda
opuntia, Dictyopteris delicatula, Dictyota divaricata, Dictyota
dentata, Sargassum vulgare and Jania spp., but numerous other species
also occur in small amounts.

Zone 7 lies in the shallow water (0.5 - 1.0 m) near shore
landward of the algal ridge, where a band of carbonate and terrigenous
cobbles and pebbles occurs along much of the shore of Boiler Bay.
Some corallines and corals also occur here, but in rather small amounts
In this band, the dominant algal species are Jania adherens, Padina
sanctae-crucis, Sargassum vulgare, Halimeda opuntia and Cladophoropsis
membranacea.

Zone 8 occurs on the beachrock which runs parallel to the shore
in the western and eastern sections of Boiler Bay. This zone is
frequently well above mean low water, and since it is partly protected
from wave action by the algal ridge, it is somewhat more subject to
desiccation and temperature and salinity extremes. However, our
quadrats were taken below mean low water springs at depths of about
OF to, 0.5m. Much of zone 8 is characterized by a turf of Hypnea
musciformis, Laurencia papillosa and other Rhodophyta. However, at
the east end of Boiler Bay, the Chlorophyta dominate this zone,
particularly Halimeda incrassata, Halimeda opuntia and Dictyosphaeria
cavernosa. At the west end of the bay, the breachrock is covered
predominantly with Sargassum vulgare with considerable amounts of
Sargassum platycarpum, Dictyota dentata and Chaetomorpha linum.
Because of this difference in the two areas of beachrock and our
inadequate sampling for the zone, we do not feel that we have
properly characterized the algal populations of zone 8.

In each of the above zones, quadrats were subjectively chosen
from several algal ridge lobes and their associated pavements as
being typical in algal cover for that particular zone. Macroscopic
benthic algae were removed as well as chunks of substrate from within
a 0.25 m area. The substrate was carefully examined for the smaller
species and all algae collected was sorted, identified and weighed for
wet biomass. Taylor (1960) was used extensively for identifications.
Skeletal carbonate for Jania, Amphiroa, Halimeda and Pencillus was
calculated using the data of Vinogradov (1953). The standing crops
of crustose coralline, algal symbionts and boring algae are not
included in this data.

Upstream-downstream dissolved oxygen analysis was applied to
Reference Reef, one lobe of the Boiler Bay algal ridge (Figs. 2, 5 &
ILL) A Sampling was accomplished at low water during spring tides when
quiet weather permitted a gentle flow of seawater over the ridge with
each wave crest. At that time, zone 1 is exposed except during a
wave crest and zone 2 is exposed with each wave trough. The water
flows southwestward over Reference Reef into the back-ridge lagoon.
The boundary of the flow across the ridge (see figure 5), and the flow
rates, were measured with standard float bottles. Flow volumes were
calculated through section B-B' of the effluent channel. Because of
the restricted area involved, the conjunction of spring tides, small
waves and light ENE winds was required to provide a measurable
channelization of effluent and a well defined area of measurement, as
well as to reduce the possibility of significant atmospheric oxygen
input from breaking waves. There was therefore often a considerable
waiting period between water sample collections. Dissolved oxygen
measurements were accomplished using standard Winkler techniques
(Strickland and Parsons, 1968).

Oxygen exchange was also studied on 100 cm? segments of coralline
and corals with their attached algal cover, which had been removed from

231-339 0 -'77 - 6

Boiller Bay jJUSt prior to testing. Polarographic electrodes were used
with a precision blood oxygen instrument (Radiometer model no. PHM-27)
and a salt water chamber maintained at + O.1L°C (see) Adey, 1973) 2° Full!
sunlight (early April) was employed, and following testing with an
entire sample, the fleshy algae were removed and the coral or
coralline substrate was tested again.

Results

A list of the algal species commonly encountered in each quadrat
and their standing crops is given in tables 1 and 2. The standing
crops of all algae for the Boiler Bay algal ridge are shown in figure 6
according to zone. The total number of species encountered and the
number of quadrats are also indicated. Fleshy algae are treated
separately from those with carbonate skeletons. Summary graphs of
the mean standing crops, relative to zone, exclusive of rare or
infrequent species, are given in figures 7 through 9. Those quadrats
taken from Robin, Beach and Isaacs Ridges on the south shore are
summarized in figures 1OA and 1OB.

Winkler oxygen values obtained for the Reference Reef sampling

stations shown in figure 5 are given in figure ll. On each sampling
occasion, daytime oxygen levels increased as water flowed over the
ridge and nighttime levels decreased. The oxygen thus produced or

consumed by the ridge organisms and net and gross productivity in g
carbon/m*/day (translated by g O5/m?* + O23, Ryther;,, 1956)?"are
calculated in table 3. Day and nighttime upstream samples were not
taken at the same point, and the possible effect of this is discussed
below.

Another estimate of the productivity of Reference Reef was
determined from the laboratory polarographic electrode study of small
samples. The rate of productivity for each zone of Reference Reef
was calculated as the product of the total surface area of that Zone
times the photosynthetic and respiratory rate derived in the laboratory
from the 100 cm? segment from the same zone. These calculations are
shown in table 4.

Our range of gross productivity for Reference Reef is compared
with other studies in table 5.

Discussion

The Waikiki Beach study (Doty, 1968) of algal standing crops

included algal ridge crest and back ridge or reef flat areas. For
comparison, we have plotted the Waikiki data for November 1967 in
figure 12. This month shows about an average level of standing crop

for the 17 month period (see Doty, 1971).

In terms of total algal standing crop (not including crustose
coralline, borers or symbionts), from the algal ridge crest back into

the lagoon, Waikiki and Boiler Bay are quite similar. The peak of
standing crop occurs on the ridge crest in both cases and this is
markedly reduced on the back ridge pavements. The shallower Waikiki
back ridge pavement has a higher standing crop than that of Boiler Bay.
However, the most striking difference is an apparent lack of a beach
rock and its accompanying secondary peak of standing crop at Waikiki.
Even in the total number of macroscopic species present, these two
areas are remarkably similar, Boiler Bay averaging about 33 species/
zone (see figure 6) and Waikiki about 37 species/zone, the difference
being easily attributable to collecting pressure (see figure 14).

In detail, however, the two areas are remarkably dissimilar. On
the Boiler Bay algal ridge, the red algae dominate strongly, forming
48-62% of the standing crop. (If the crustose corallines were to be
included, the dominance would be considerably greater.) This high
level of red algal standing crop is not concentrated in a few species,
but in over eight species of Laurencia, Hypnea, Gracilaria, Gelidiella,
Pterocladia, Jania and Amphiroa. On the pavements surrounding the
ridge (zone 6) and on the fore-beach cobbles (zone 7) this dominance
is reduced, but still the red algae make up 20-40% of the standing
crop. It is only in some areas of the beach rock (zone 8) that the
red algae are largely replaced by the green alga Halimeda or the
brown alga Sargassum.

In contrast, a number of species of Laurencia, Hypnea and
Gracilaria are present on the Waikiki reef, but none of these exceed
a mean of 80 g/m? in any one zone. Although Spyridia filamentosa
reaches mean values of nearly 200 g/m, only Acanthophora spicifera
reaches dominant levels. However, virtually all of the red algal
standing crop that is present occurs on the back ridge flat, especially
near shore. The 23 species of red algae in the quadrats on the
Waikiki algal ridge proper formed only .04% of the standing crop in
November, 1967, while in Boiler Bay's equivalent zones, 31 species (in
quadrats) formed 41% of the standing crop.

Green algae are not important on the algal ridge crests in either
Boiler Bay or Waikiki Beach. However, on the back and fore-ridge
Pavements and near shore in both areas, the greens become conspicuous.
The 30-40% of green standing crop in back ridge and beach rock at
Boiler Bay were mostly Halimeda spp. lowever, Caulerpa, Cladophoropsts,
Valonta and Dietyosphaeria occur at mean levels of 20-160 g/m2.

The 20% of the standing crop in back ridge and shore Waikiki contains
considerable Halimeda, but nearly equal amounts of Dictyosphaeria
cavernosa. Ulva reticulata, near shore, is also important, perhaps due
to higher pollution levels.

In contrast to the situation in Boiler Bay, the Waikiki ridge
standing crop is almost entirely composed of brown algae: three species
of Sargassum. These three species occur in varying amounts across
the reef flat to the shore, and it is only in the zone 30 meters from
shore, where Sargassum polyphyllum and Acanthophora spicifera share
most of the biomass, that the brown algal dominance of the standing

crop is challenged. At least three species of Sargassum occur on the
Boiler Bay ridge, but only in a single quadrat did they occur at levels
of more than a few grams. Sargassum is generally more important on
the beach rock and Padina sanctae-crucis on the cobble zone near shore.
Otherwise, Dilophus, Dictyopteris and Dictyota are all important minor
elements on the pavements around the Boiler Bay ridges and may
occasionally form blooms. In Hawaii, Dilophus does not occur, and

at Waikiki relatively few Dictyota and Dictyopteris occur as compared
to other Hawaiian areas (I.A. Abbott, personal communication).

In the context of these striking differences, it is interesting
to examine the standing crop of the exposed algal ridges on the south
shore of St. Croix. Green algae are virtually absent from the
highest of these ridges as they are at Waikiki. Also, the brown
algal standing crop, largely in the form of Sargassum vulgare, exceeds
that of the red algae which consists mostly of three species of
Laurencia. In the back ridge zones 2 and 3, the Sargassum drops
rapidly in abundance. Even though it tends to be replaced by
Dictyopteris, red algae (still mostly Laurencia spp.) begin to
dominate again. This suggests that on algal ridges subject to very
high wave energies, reds may be replaced by Sargassum. The
considerably higher and more exposed algal ridges on the southeast
shore of Martinique (studies in progress) show a similar relationship.
There, the highest ridge tops, at > 1mm.l.w.sp., are nearly bare,
while the low crests and surrounding highly turbulent pavements have

high standing crops of Sargassum. While Waikiki is not directly
exposed to the trade wind, as are the southeastern St. Croix ridges,
it does often receive a very large South Pacific swell. Boiler Bay,

on the other hand is largely blocked by the northern Virgin Islands
and rarely receives the equivalent large swells from occasional
northerly winds developed from passage of continental fronts
("northers").

Adey (1974) discusses the relationship between exposure, wave
action and algal ridge heights. The highest parts of the more
exposed St. Croix ridges reach 40-50 cm above mean low water springs,
with a mean spring tide range of about 30 cm. During some summer
neap tide cycles, the ridge zone 1 can be 20-30 cm above sea level for
four to six hours mid-day. Desiccation and heating are likely to be
critical on occasional quiet days and, along with mechanical stress
during more turbulent weather, probably accounts for the obvious
bareness of the higher part of the ridges. These factors probably
also account for the difference in standing crop decreasing from
3200 g/m at Boiler Bay ridge crests with an average height of +14 cm
to 2680 g/m on the south shore ridges averaging +30 cm high. ite
seems likely that Sargassum species are better able to withstand both
the turbulent water conditions and perhaps occasional periods of
limited desiccation and heating.

In recent years, it has generally been considered that benthic
algal standing crops are considerably lower in tropical than in
temperate to boreal areas (see e.g., Taylor, 1960). This has been

variously attributed to low ambient water nutrients, high insolation
levels, lack of beds of brown algae or a shift of standing crop to
crustose corallines, which are not usually included in standing crop
analysis (Bakus, 1969). (Note that Adey and Macintyre (1973) point
out that crustose corallines are probably as important in terms of
bottom coverage in subarctic-boreal waters as they are in the tropics.)
In our studies of the Boiler Bay algal ridge and that of Doty at
Waikiki, average wet standing crops up to 3 -3.5 kg/m? were found,
with individual quadrats ranging up to 5 kg/m2. On several
considerably larger algal ridges and pavements that we are now
studying in Martinique, algal standing crops are nearly twice as high.
Typical lower intertidal Ascophyllum, Fucus and Laminaria stands in
Norway have a standing crop of about 3.5 kg/m (Baardseth, 1970).
Upper sublittoral Laminaria hyperborea stands in the British Isles
average 5.0 - 6.5 kg/m* (Kain, 1971). An especially rich Gigartina
stellata stand in the lowest intertidal and infralittoral in Maine
achieved nearly 10 kg/m2 in the peak summer season (Burns and
Mathieson, 1972). Thus, while perhaps temperate-boreal standing
crops of larger algae tend to be higher than those in the tropics,
algal ridge and beachrock standing crops (those in turbulent zones)
can be similar, and, generally, in the seas around the older parts of
the higher, volcanic eastern Caribbean islands (Adey and Burke, 1976)
algal standing crops of reef structures are quite equivalent to or
perhaps larger than the average for northern shores. While brown
algae are less important than reds in the flora of the Boiler Bay
ridge, Sargassum, Dictyota and Dictyopteris are major components.
Similarly Sargassum actually exceeds the red algae in biomass on the
south shore St. Croix ridges while at Waikiki and Martinique, it is
highly dominant. Perhaps the main question to which our attention
should be directed is not why tropical standing crops of larger benthic
algae are so low, but why they only achieve high levels in special
situations.

The importance of grazing in the shallow tropical marine
environment has attracted considerable interest in recent years.
Randall (1961) contrasted the luxuriant intertidal algal crop in
Hawaii with the low stubble of the upper sublittoral and attributed it
to fish grazing (Acanthuridae, Scaridae and Pomacentridae) . The
conspicuous vegetation-free halo surrounding tropical reef areas has
also been shown to result from either fish (Earle, 1972; Randall,
1965) or echinoid (Ogden, et al., 1973) grazing, and removal of
urchins from reef areas can result in massive increases of algal
standing crops (Sammarco, et al., 1974). Grazing by urchins in
northern waters can also be extremely important in limiting standing
crop (Jones and Kain, 1967; Paine and Vadas, 1969). In the northern
North Atlantic, Adey is familiar with extended areas, virtually bare
of fleshy algae dominated by echinoids and coralline algae, and
Lebednik (personal communication) has seen similar areas in the
Aleutian Islands. Mead (1970) has suggested that the main group of
fish grazers, the percoid fishes, which are tropical and Cenozoic in

10

evolution are largely responsible for the present reduction of the
sublittoral algal "forests" in the tropics. LEVES di EEICUuLe to
escape the logic of this conclusion - an algal ridge or shallow
pavement is a special, turbulent water environment which is usually

inaccessible to grazing fish and Diadema. A few invertebrates
(mostly crabs, snails, limpets and chitons) do graze in this zone,
although their effectiveness is apparently limited. Echinometra is

often abundant on algal ridges and beachrock, however, the wave
energy is apparently generally sufficient to largely confine the
echinoids to their holes and to feeding on drift (Ogden, Abbott and
Abbott, 1974).

Below the shallow highly turbulent levels of the ridges and
pavements, the grazing marks of parrot fish, urchins and gastropods
are often strikingly apparent on the typically bare coralline or
rubble surfaces. In Caribbean waters, grazing, especially by parrot
fish, is critically important in the colonization and carbonate
buildup of the crustose corallines themselves (Adey and Vassar, 1975;
Steneck and Adey, 1976), and probably is an important factor
restricting algal ridges to the most exposed windward shores.

Our research group has spent a considerable amount of time in
daytime snorkeling around the algal ridges in Boiler Bay. Our
collective casual observations on the intensity of fish grazing are
as follows: zones 1 and 2 on the ridge, zone 8 on the beachrock and
perhaps to a lesser extent, zone 7 near shore are rarely grazed by
larger fish. These zones are either exposed in wave troughs or
continuously washed by waves. Zone 3, the algal ridge bowl is
periodically grazed, sometimes heavily depending on sea conditions.
Zones 4 and 6, on the other hand, are heavily and consistently grazed.
Not only are these zones easily reached by grazing fish, but cover for
the fish is abundant in the form of holes in the pavements and coral
structures, especially Acropora palmata. Zone 5, the deep pavement,
is the zone over which we have greatest disagreement concerning fish
grazing pressure. Generally, we do agree that it is less than the
pavement zone 6 because of lack of cover for the fish. Periodically,
however, it is probably massively grazed, especially by schools of
tangs and parrot fish. Figure 13 shows this subjective index of
grazing potential plotted against standing crop. Generally there is
indicated a strong inverse relationship between the ability of fish to
graze effectively and the standing crop of fleshy and filamentous
algae.

The upright calcareous algae, on the other hand, show no
apparent relationship between grazing pressure and standing crop.
Zone 1, no fish grazing, and zone 4, intense grazing, have about the
same level of calcareous standing crop. The only striking feature of
the plot is the very high standing crop of calcareous algae on the
deep fore-ridge pavement, zone 5, as compared to the other zones.
This largely results from the abundance of only two species, Halimeda
opuntia and to a lesser extent Amphiroa tribulus. Halimeda opuntia has

JLab

secondary peaks in zones 2 and 6 and the reason for this distribution
pattern is obscure.

In figure 14, the total number of non-calcified algal species for
each zone are plotted as a function of the number of quadrats taken
(i.e., total area collected for collecting pressure). The Waikiki
data of Doty are also included and are converted to 0.25 m? quadrats
on the basis of total area per zone (see Fig. 12). The relationship
indicated further suggests that the larger number of species taken at
Waikiki was a result of greater collecting pressure. Ibigi Sesleqbbets ILS),
the Boiler Bay data are plotted, species number/zone as a function of
quadrat number/zone. Since species number is a function of collecting
pressure, a correction factor is thus derived as shown to correct all
zones to 10 quadrats. (Note that a plot of number of species
occurring at mean levels of > lg/.25 m* is not or only slightly
quadrat dependent in the range that we worked.) Thus, in figure 16,
the total number of non-calcified algal species, corrected to 10
quadrats, as well as the number with a standing crop > 1 g/.25 m7,
is plotted as a function of Zone. Both measures show a consistent
drop in number shoreward.

Proceeding shoreward, daytime temperatures and suspended sediment
generally increase markedly in Boiler Bay, a feature very obvious to
the swimmer. Equivalent night temperature drops are to be expected,
especially in the winter. These physical characteristics are likely
responsible for the shoreward drop in species number.

Figure 17 indicates the variations in number of species from the
mean total species curve (Fig. 16) for Boiler Bay. Zones 3, 4 and 6
have four to six species above mean levels. These are zones of
heavy to intermediate grazing pressure, suggesting that due to a
reduction in space competition, grazing has increased richness. On
the other hand, the poorly grazed zones, especially 1 and 2, which
have a high standing crop are poor in the rarer species. Zone 8 is
the chief anomaly in the pattern. If lack of fish grazing is the
controlling factor, and this is indicated by a relatively high biomass,
one would expect total species number to be low. Ogden, Abbott, and
Abbott (1974) found 20% fewer species on the west beachrock in Boiler
Bay as compared to the offlying ridge crests. Their extensive search
for species should represent a large number of quadrats. We suggest
that our species number for zone 8 is too high, probably as a result
of insufficient sampling.

In table 5, our range of gross productivity values for Reference

Reef are compared with other areas. As is perhaps to be expected, our
values for a dominantly algal area are somewhat higher than those
previously given for coral reef communities. Also as can be seen in

table 4, algal ridge zones 1 and 2 are responsible for 55% of the area
productivity, and in terms of those zones alone, the productivity is
about 30 g C/m*/day. The contribution of crustose coralline algae is

2

indicated as 12.1 g C/m2/day. This is probably higher than normal,
since on the Boiler Bay ridge, the coralline surface is usually shaded
by the fleshy algal cover. In any case, it is considerably higher
than the previous figure of 1.5 g c/m* /day given by Marsh (1970).

Because of the critical importance of tide level to our process
of taking data on the ridge, it was not possible to take successive
oxygen readings at different light levels during a single day. This
could however be done on different days as the time of low water
springs changes. Presumably our daytime values being taken near noon
are maximum levels. On the other hand, Marsh (1970) indicated that
in ridge corallines, photosynthesis levels are constant for 10 hours
of the day. If this is true for other ridge algae, then our upper
value of gross productivity (Table 5) is high by about 15%. The
productivity of these ridges and especially the higher ridges of the
eastern islands (Adey and Burke, 1976) should be examined in further
detail.

Conclusions

The intertidal wave-washed crustose coralline algal ridges of
St. Croix, Virgin Islands develop a dense standing crop of fleshy
algae and have a gross productivity which is high for the tropical
reef environment. In caged areas and areas with considerably greater
wave action, especially in the presumably more eutrophic waters around
the higher volcanic islands such as Martinique, the standing crop is
often denser and also extends to greater depths. In St. Croix, where
wave energy is moderate, the heavy coverage of fleshy algae is greatly
reduced with depth away from the ridge crests. Thus, lack of intense
grazing pressure under turbulent water situations is probably
responsible for the rich algal area on the ridges. In a relatively
low-turbulence coral dominated reef environment, the maturity of the
tropical ecosystem with extensive filling of grazing niches, especially
by fish and echinoderms, results in a generally reduced primary
productivity. Cenozoic tropical reefs are characterized by coralline
algae and hermatypic schleractinian corals. The domination of this
combination perhaps resulted from the evolution of grazing bony fish
and some echinoid species. Algal ridges, and in some cases exposed
beachrock, are specialized turbulent water situations in a carbonate
environment where grazing niches cannot be filled either by fish or
by the key grazer Diadema antillarum.

Given favorable conditions of marine salinity and low levels of
marine erosion and suspended sediment, the frequency and extent of
fleshy algal-dominated reef environments, as well as the nature of the
flora developed, is thus largely dependent upon the strength and
constancy of the accompanying trade: wind and sea. Since well-
developed sublittoral algal pavements are especially prominent around
the older parts of the higher eastern Caribbean islands, it is
suggested that more eutrophic conditions, with greater algal growth
rates, are also critical.

3

Acknowledgements

Many people helped us in the course of this study, chief among
them being our own associates J.M. Vassar, P.J. Adey and R.S. Steneck.
N. Buckman-Ogden and J. Ogden, West Indies Laboratory, St. Croix both
read the manuscript and offered suggestions for its improvement.

I.A. Abbott deserves special thanks for both help with identification
problems and for critically reading the manuscript.

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& R. Burke. 1976. Holocene algal ridges and barrier reef
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& I. Macintyre. 1973. Crustose coralline algae: a re-
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edema Vassar.un 1975. Succession and accretion rates in
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14

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Fig. 5. Reference reef showing algal ridge zone
locations and water sampling sites—Q9.
See figure 11 for longitudinal (AA') and
effluent (BB") channel sections. B'B"
is the boundary of the water mass treated,
as determined by float bottles. The
arrows on BB" indicate the direction of
current flow.

Zones 1-3 as figs. 3 & 4.
Tp, Diploria spp.
Porites astreoides
Porites porites
Sideastrea siderea
Carbonate blocks
Algal nodules

Terrigenous stones

JARBLEAON

Thalassia

231-339 O-'77-7

Fig. 6B. Mean wet standing crop as a function of zone. Algae with
carbonate skeletons (Jania, lialimeda, Amphiroa and Penicillis)
are shown separately in terms of organic biomass and skeletal
Carbonate. Species number and the number of quadrats taken
per zone are indicated.

Carbonate
skeletons

(B)

~
a

=
@

Organics in
caicifed algae

standing crop (wet) g/,2 -102
is]
o

=
»

Fleshy | algae . \

0
total no. species 36 47 37 35 42 38

No. species wgt. > 1gm 12 16 20 12 13 18
no. 0.25 m2 quadranis/ 6 8 10 8 6 5
zone

(A)

Inner Lagoon

Outer Lagoon

A

depth (meters)

-4

) 10 20 30 40 50 60 me

Fig. GA. Generalized bathymetric transect from the shore across the
Boiler Bay algal ridge to the outer lagoon, showing the depths
and positions of the algal zones.

A. tribulus

Amphiroa spp. Nt me:
’
'
'
Na |

CALCAREOUS REDS
5
°
total
'_

Red Algae Standing Crop (Dmn2)
o s 3 8

Coelothrix irregularis 4

Centroceras clavulatum
hrysymenia sp.

Ea

Pterocladia americana Se

Gelidiella acerosa ray
< :

Gracilaria mammillaris
= Laurencia spp. ae +L. gemmifera L. Intricata
> a
c eC
n /
Ww 1
=
a

Laurencia papillosa H. spinella

Hypnea spp. \ 7

H. cervicornis
zone = 41 2 3 6 7 8

Fig. 7. Distribution and standing crop of the major species of red algae
in the algal ridge area of Boiler Bay.

-Aeq ZSTTOG JO eaze ehptazA TehTe syyj ut oebhte
usezb Jo setoeds zofew ey} Jo doz. Hutpueqs pue uotyanqtajsta °9 °bta

Siz 9 © Geek v Greets

(eae eg end sec ae ST]
ee ellaeydsodjoig

i ae eae eaey

| 4 sisdosoydope|9
edisajnes

Ee SAGE ‘ere

<= Se Se

au

e1uojen

(o)
o

epewleH

PN ‘HH
a =a epowley

Bun} ‘H
ie ee

ejyessesoul “Hy

(74/6) aeBjy uaain - --doi9 Sulpuers

fo)
Gite ee g
©
00r
008
O'9vC~e aalsaloivo
ooz

*Xeq AeTTog Jo eeze ohptz Tebte ey} UT
eebtTe umoriq Jo satoeds zofCeu ay3 go dozo 5utpuezs pue uotynqtzysta °*6 °*bta

Lv

mae.
winiyjesedAjod ‘s

5}5N19-@8}DUeS eUlped

sueuseyje snydojig

Bynyesijep
8110}d04A}91Gg

e}eJ8AIP &30A}91G

eyejuep e30A}91G

(7/6) eeBly umolg ---doiDd Bulpueys

BYeONID e304}91G
esonuis

ooPr

008

Total Browns

Brown Algae

9/m?)

~

300 300
Skeletal carbonate
Organics in
200 yi 200 calcified algae
5 =]
©
ee
100 — 100
iS
°
=
0, a
Dictyota dentata
Dictyota divaricata
fe Hypnea spinelia
€ Laurencia
> gemmifera \
tf \
Dictyopteris © \
delicatula ®
Dilophus guineesis =
Laurencia intricata
5
©
x
Sargassum hystrix
2800. |
Skeletal carbonate Laurencia obtusa
E '
2400 zone 1 1 2

Total Wet Weight

Generalized
South Shore Algal Ridge

Organics in calcified algae

i)
So
{=}
°o

1600

f=)

Algal Ridge

depth (meters)
a WN

Outer Pavement Lagoon

Fig. 10. Distribution and standing crop of fleshy algae on several of
the high algal ridges on southeastern St. Croix (see Table 2).

1.6 g of Halimeda opuntia was found in zone 2 of Beach ridge
and this is not shown in the diagram.

A - Brown algae
B - Red algae

Fig. 11. Day (x) and night (0) change of dissolved oxygen concentration
across Reference Reef. See figure 5 for the location of
sections AA' and BB’. (5) indicates the number of water
samples. Date and time of sampling is indicated at the right.

10.00_
eo G) 4fg, 1230
(4) 44, 1230
(5) i
—~, 9.00_ (6) 44, 1230
o
= (4) (6)
—Z
c
® 8.00_ ”)
re.)
>
<
Oo
9
S$ 700_
ro)
o (5)
a (5)

6.00_ (4) (8) 12%, 2230
(5) 12f, 0200

5) 1/q 0530

. eae eel et eee

Reference

Reef

Porites astreoides

meters

. 10 20 30
B' B
1
Effluent
Channel

2
0 5 10 meters

Fig. 12. Standing, crops of algae
at Waikiki Beach, November 1967.
The numbers under the somewhat
arbitrary zone breakdown at the
bottom of the diagram refer to:
(no. spp.) total number of species
collected, (sp. > 1 g) number of
species with a standing crop >1l
g/.25m2, (no. quad.) equivalent
number of 0.25 m2 quadrats.

Ruger LS

Algal standing crop as a func-
tion of ease of fish grazing (per
zone). x - fleshy and filamentous
algae, o - algae with carbonate
skeletons (Jania, Halimeda, Amphi-
roa, Penicillis).

Fig. 14.

Total number of species found/
zone (in quadrats) as a function of
the number of quadrats taken.

x - Boiler Bay, o - Waikiki data
(Doty, 1969) converted to 0.25 m2
quadrats. Zero was used as a
theoretical data point.

3200,

2400) x
\ i ~ brown algae
iN i tif
1600)
x Se sit
x7 wre /
ax
800 red algae ~ \
a INS = GeO
= UM ie re %
\ A eos green algae
3200) a
a
{o)
«x
© 2400) ‘\ is
g ae ! Se
a | aeons ;
S 160 yey Fleshy Algae
=
a x
80)
Halimeda

carbonate skeleton

fle

zone

no. spp
sp. >1g
no. quad.

depth (m)

(17) (23)
5.4

200 100

distance from shore (m)

Algal Ridge SHORE

10

easier

fish grazing

very difficult

standing crop wet weight 9 An2

total no. species “one

10 15
no. of quadrats taken “one

IDE ys ESI

Number of species
per zone, (x) total and
(o). > 1 g/.25m2, as a
function of quadrat
number (area collected
or collecting pressure)
in Boiler Bay. The

correction factor is
derived as shown (arrow)
to adjust all zones to
a collection pressure
of 10 quadrats.

for 10 quadrats
increase total

species number

by:

4 5 6 7
no. quadrats taken ~ one

Hagin Gy.

SSS aa i
é DN (X) total number of species
2
3 \x’ |, corrected to 10 quadrats (after
See see text Fig. 15), and (0) species with
3 ye mes standing crops (wet) of 1
~ Sneey ee a /0.25 :
5 a mr ———=p——-0. g/0.25m“, as a function of zone
E 10 on the Boiler Bay ridge (see
i<j Fig. 6).
zone: 5 412 3 6 “6
10
=19
0)
o 8
8 xx
Berge 17. Tu 6/*
eale
Deviation of total species §| 5 x x
number from the mean (corrected «£! 4 wa i ;
to 10 quadrats) as a function SURG P
of ease of fish grazing. = Ny
51 AX
a0 1

US Ue 8 10

ae Ec tab a2
deviation of total species nO from mean
zone

Weights less than 1 gram
Plants occurring as single small specimens

List of species, frequency of occurrence in quadrats, and wet standing

crop (g/.25m2), Boiler Bay algal ridge.

indicated by notation "tr".

Table l.

are not listed but appear in species number in figure 12.

Zone and quadrat

— 1A 1B 1c 1D 1E
Caulerpa racemosa 24.7 4.4
Cladophoropsis membranacea 19.4 88.7
Dictyosphaeria cavernosa 5.4
Halimeda incrassata
Halimeda opuntia 2.0 24.5
Halimeda tuna
Penicillis capitatus
Udotea spp.
Valonia spp. 3.4
Colpomenia sinuosa 8.5 10.8 VAS 9.6 SiD
Dictyota ciliolata
Dictyota dentata tr
Dictyota divaricata H39R5
Dictyopteris delicatula 82.2 ty Ex 59)
Dilophus alternans 39/20 59
Padina sanctae-crucis ‘Ex, Z3s{0) 16.2 2-55 1.5
Sargassum Vulgare 50753 83.4 107.9
Sargassum platYcarpum
Sargassum polyceratium 369.6 4.6
Amphiroa spp. tr eg tr tr tr
Centroceras clavulatum tr
Chrysymenia sp. 13.0
Coelothrix irregularis
Galaxaura spp.
Gelidiella acerosa 19.6
Gracilaria mammillaris 16.3 tr 200.2 40.8 Vics}
Hypnea cervicornis tx:
Hypnea musciformis 160.0 349.0 90.6 124.0 246.8
Hypnea spinella tr Lop
Jania spp. 222.4 9.7 aus 181.0 337.4
Laurencia intricata
Laurencia gemmifera 64.3 43.9 64.0 8.2
Laurencia papillosa 64.3 43.9 UZ Bi, 64.0 337.4

Pterocladia americana

36.5

tr

tr

tr

jong

34.0

14

187.4

70.6

165.2

68.3

tr

31.6

tr

183.0

94.6

158.8

67.3

IHU 65)

80.

85.

tr

1x5

Er

34.

17.7

tr

20.1

104.5

icy

7s

147.3

475.6

tr

tr

19/55

34.2

13.6

1535

156.8

185.0

164.8

tr

295.8

ae

tr

223.4

tr

164.0

400.1

TABLE 1 (CONT. 2F 2G 2H 3a 3B 3c

Caulerpa racemosa

Cladophoropsis membranacea

Dictyosphaeria cavernosa

Halimeda incrassata

Halimeda opuntia 54.7 35.0 554.6 187.8 59.0 133.5
Halimeda tuna 2.9 tr

Penicillis capitatus

Udotea spp.

Valonia spp. 12 4.4 0.3
Colpomenia sinuosa

Dictyota ciliolata

Dictyota dentata

Dictyota divaricata 36.9 16.7 46.5
Dictyopteris delicatula 7.8 tr tr

Dilophus alternans 30.1 59.5
Padina sanctae-crucis re tr
Sargassum vulgare 2.9

Sargassum platycarpum

Sargassum polyceratium

Amphiroa spp. tr tr tr
Centroceras clavulatum 6.1 150)
Chrysymenia sp.

Coelothrix irregularis 6.5 4.3

Galaxaura spp.

Gelidiella acerosa 384.8 33.8 214.8 166.6 165.3
Gracilaria mammillaris 70.0 Bir2 6.3 59) 14.3 225
Hypnea cervicornis 53

Hypnea musciformis 1.0

Hypnea spinella 31.2

Jania spp. 192.4 ne ee 21.3 69.0
Laurencia intricata tx tr

Laurencia gemmifera Pe tr

Laurencia papillosa 40.4 PESO) 85.6 4305 106.4 10.0

Pterocladia americana 25:40 3553 14.4 190.2 V8)

3D 3E 3F 44 4B 4c 4D 4E 4F 4G

136.2 24.0
tr 4.0
tr 3.0 ty
91.3(?)
102.5 188.8 26.4 tr 19) fe 177.6
tr 28.0 tr 67.0 6.3 90.7
1.6
4.3 13.0
itr 0.7 tr tx
tr 18.8 1.9 0.9 tr
tr tr 60.6 10.9
131.0
46.5 110.3 58.1
109.1 78.4 118.2 3.4 25.5 172.5
59.5 141.4 34.2 Ts 3
0.6 3.2 1.5 2.1 tr tr tr
5.1 Bee
18.8
2.6 3.9
0.3 173 3.5 4.2
tr 24.6 tr
16.9 5.4
tr 43.0 3.0 fee tr 3.5 17.6
Wy Aaa 54.6 4.0 Tal) GZ} 8.6 3e2
Talioat Dell tr tr 15.6
2.5
7.6 49.3 58.0 7/0) ARO 69.6 46.7 2ive3
tr 40.5 tr 4.7 tr 100.0 tr
tr tr tr .
19.1 98.4 38.0 Vi58 tr Ex

N
@
=
=

TABLE 1 (conT.) 4H 5A

5B 6A 6B 6D 7A
Caulerpa racemosa 9.0 26.3 136
Cladophoropsis membranacea 2.5
Dictyosphaeria cavernosa 2.0 2533 3.4
Halimeda incrassata 18 14,1
Halimeda opuntia 496.0 318.2 61.6 65.2 224.3 107.6 70.0
Halimeda tuna 46.7
Penicillis capitatus 8.4 5152 15 yo 2.0
Udotea spp. oa: 20.0 1.4
Valonia spp. 17 er
Colpomenia sinuosa 1.0 1.6
Dictyota ciliolata
Dictyota dentata 234.7 286.3 4.5
Dictyota divaricata tr 145.7 31.0
Dictyopteris delicatula tr 77.4 84.3
Dilophus alternans 99.4 tr tr
Padina sanctae-crucis 3.1 3.7 552
Sargassum vulgare tr 14.8 26.2 30.1
Sargassum platycarpum 222 4.6 tr

Sargassum polyceratium

Amphiroa spp.

Centroceras clavulatum

Chrysymenia sp. 0.3

Coelothrix irregularis

Galaxaura spp. 14.2
Gelidiella acerosa 107.3
Gracilaria mammillaris 1.2

Hypnea cervicornis

Hypnea musciformis

Hypnea spinella 2S
Jania spp. 78.0 73.4
Laurencia intricata 83.0
Laurencia gemmifera CY,
Laurencia papillosa tr
Pterocladia americana tr

25:15 14.3
H625:9)
1.3
59.6 6 72.30
IL e@
Ex

19) 2

tr

16.0

268.0

25.4 90.5 41.3 tr 22
253 0.8 0.6 VO
6.0 5.4 ‘er TEA: 836.9
229.6 28.4 109.8 86.1 100.3 238.7
25 137 5.3 2.4 15.3
9.3 19.3
25 0.6 3.8 fees
1.0
tex algal
TDAll 5.4 42.4 85.4 (ere 124.5 0.7 1.0
Toa 63.0 199.0
4.3 2.5
S}58) 529.1 Died 233.4 8:3
5.3 95.5 NS 5® 60.2 381.7
G7, Tez tr 9.0 130.5
104.9 tr 301.9
263
76.6 8.7 30.0
14.3 0.3 0.5
3825
5.3 300.0 123.9 204.0 121.4 8.9 121.3 ieee 7.4
119.5

TABLE 1 (cont) 12B 12¢ 12D 12E 12F

Caulerpa racemosa
Cladophoropsis membranacea
Dictyosphaeria cavernosa

Halimeda incrassata

tr 43.1 62.2
Halimeda opuntia 318.0 420.9 632.6 413.5 628.0
Halimeda tuna Boe 45.8 2.4 158.8 50.7
Penicillis capitatus far 2188 8.6
Udotea spp. 3.8 alas} 1.4
Valonia spp. 6.4 17.8

a al a i a

Colpomenia sinuosa 0.7 1.4
Dictyota ciliolata
Dictyota dentata
Dictyota divaricata 270.0 306.0 69.2 219.0 23252
Dictyopteris delicatula 6.8 1.9 Sas 8.2 2.3
Dilophus alternans
Padina sanctae-crucis 0.4
Sargassum vulgare 3.0
Sargassum platycarpum
Sargassum polyceratium
Amphiroa spp. 157.4 201 s1 252152
Centroceras clavulatum
Chrysymenia sp.
Coelothrix irregularis 0.9
Galaxaura spp. 44.8 1452 13.7 2.2
Gelidiella acerosa 0.8 3.7 4.0 15.4 4.7
Gracilaria mammillaris 0.3
Hypnea cervicornis tr
Hypnea musciformis
Hypnea spinella
Jania spp. 38.3 40.2
Laurencia intricata 0.8 0.3 0.6 tr

Laurencia gemmifera
Laurencia papillosa

Pterocladia americana

Table 2. South Shore Algal Ridges. , 4:1 Rebin

Isaac Beach Inner Outer Isaac Beach Isaac
1 i} 1 1 2 2 3

Caulerpa racemosa

Cladophoropsis membranaceae

Dictyosphaeria cavernosa

Halimeda incrassata

Halimeda opuntia 16
Halimeda tuna

Penicillis capitatus

Udotea spp.

Valonia spp.

Colpomenia sinuosa

Dictyota ciliolata

Dictyota dentata tz 22.8 ibs}
Dictyota divaricata 376.5 6.4
Dictyopteris delicatula 71.6 44.9 5 tr 122.6 101.5
Dilophus alternans 44.4 47.9

Padina sanctae-crucis

Sargassum vulgare sfillsy/ 843.0 290.7

Sargassum platycarpum

Sargassum polyceratium

Amphiroa spp. 137.1
Centroceras clavulatum 4.5
Chrysymenia sp.

Coelothrix irregularis

Galaxaura spp.

Gelidiella acerosa

Gracilaria mammillaris

Hypnea cervicornis

Hypnea musciformis

Hypnea spinella 49.0 58.2 60.0 IW sy-sah 3.0
Jania spp. tr 31.6
Laurencia intricata BAT CS) 156.6
Laurencia gemmifera tr Gr

Laurencia papillosa

Laurencia microcladia 295.0 Iie S} 240.9 15.4 2955 4.4

Laurencia obtusa 224.0 er 82.6 9.3 ilalE}S) 287.1

231-339 O-'77-8

Table 3. Productivity of Reference Reef derived from upstream-downstream
analysis of oxygen concentration (see figures 5, 11).

Oxygen determined by the Winkler method.

C produced

BoP

EGS}

Date Time 40, conc 05 produced 05 produced
(+ mg05/L) (mg/m? /hr) (g05/m?/12 hr) (gC/m2/12 hr)
Apr. 18,573 1230 0.88 7.56 x 103 90.7
Apr 17 Gien al 280 1.36 14.26 x 103 AGALe
Apr. 20) 85 73°* 1230 0.69 5.93 x 103 eee

MEAN NET PRODUCED, DAYTIME

Date Time A 05 conc O05 consumed O05 consumed
(— mg02/L) — (mg02/m*/hr) —(g0,/m?/12 hr)

July 93°73 0530 0.597 5.12 x 103 61.44
Dee, 82573 2230 0.197 1.66 x 103 19.92
Dec. 212), 1.73. ~. 10200 0.883 2.93 x 103 35.16

MEAN CONSUMED NIGHTTIME

NET PRODUCTION (24 hr)

GROSS PRODUCTION

2d4e3

33\03;

C consumed
(gC/m2/12 hr)

18.43
5.98
10.55

11.7 gC/m2

21.6 gC/m2

45.0 gC/m2

EEE

Table 4. (opposite page)

Productivity of Reference reef determined from small samples in
laboratory aquaria, using polarographic electrodes. The productivity
of the carbonate crusts with their attached fleshy algae was determined.
The larger algae were then removed and the productivity of the remain-
ing carbonate crust with its coralline crusts and boring algae was
redetermined. Porites astreoides occupies approximately 16% of the
surface area of the zone 6 in the "bowl" of Reference reef. The produc-

tivity of the remaining area of this zone was determined as a

ratio of

its standing crop to the standing crops found in zones 1, 2 and 3.

Table 4.

Collection Site

prod. resp. gross prod. resp. gross
zones 1 & 2 g0>/m*/hr 5.9 -4.6 10.5 2.3 = 22 4.5
Landing reef gC/m2 /hr 1.8 -1.4 a2 .69 -.66 1.35
gC/m2/12 hr 21.6 -16.8 38.4 8.3 7.9 16.2
Zones 1 & 2 g05/m*/hr 4.4 31.7 8.1 2.3 =2,2 4.5
Stick reef gC/m*/hr es tol 2.4 69 -.45 lesa
gC/m2/12 hr 15.6 -13.2 28.8 8.3 -5.4 IL) 7)
Zones 1 & 2 g0>/m*/hr 4.0 =2.3 Gea
Sand reef gC/m*/hr MG? -0.69 1.9
gC/m?/12 hr 14.4 “3.8 22.8
zone 3 g05/m2/hr Bye -3.2 6.4 .6 -1.2 1.8
Sand reef gC/m2/hr .96 ~.96 1.86 .18 =. 36 .54
gC/m2/12 hr lie so0 = ie 23.04 Deals =4, 32 6.48
Zone 6 g0>/m2/hr te 7/ -l1.1 2.8 1) 5 1b gC/m*/day
Porites astreoides gC/m2/hr 5 Syl -.33 . 84
only gC/m?/12 hr 6.12 -3.96 iL) sl

Gross production for zones 1 and 2: 30 gC/m*/day (mean) x 170.7 m2 (area zones 1 & 2) = 5121.3 gmC/day

Gross production for zone 3: 23.04 gC/m*/day (mean) x 62.7 m2 (area zone 3) = 1444.8 gmc
Gross production for P. astreoides in zone 6: 10.1 gC/m2/day (mean) x 155.6 m2 (area

zone 6) x 16% coverage P. astreoides =" -251)..5
Coral pavement in zone 6: 20.5 gC/m2 /day x 155.6 m2 x 84% coverage = 2679.0

9497/389 m2 =
PYM SIS) gmc /m2/day

Table 5. Comparison of the productivity of the algal ridge lobe,
Reference reef, with other reef systems.

Source & Area

This Study, Reference reef

Kanwisher & Wainwright, 1967

Florida, Corals

Kohn & Helfrich, 1957
Hawaiian reef community

Odum & Odum, 1955
reef community

Gordon & Kelly
Hawaiian reef community

Helfrich & Townsley, 1961
Hawaii, Eniwetok, P. Rico
Coral Reef

Gross Productivity gC/m?/day

24-45

261 -VOR2

VOS8=S207

4.9-32

ATOLL RESEARCH BULLETIN
NO. 212

PRELIMINARY OBSERVATIONS ON THE ALGAE,
CORALS, AND FISHES INHABITING THE SUNKEN FERRY
“FUJIKAWA MARU” IN TRUK LAGOON

by Roy T. Tsuda, Steven S. Amesbury, and Steven C. Moras

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

May 1977

PRELIMINARY OBSERVATIONS ON THE ALGAE,
CORALS, AND FISHES INHABITING THE SUNKEN FERRY
“FUJIKAWA MARU” IN TRUK LAGOON !

by Roy T. Tsuda,? Steven S. Amesbury,* and Steven C. Moras 2

Introduction

Of all the sunken ships in the Truk Lagoon, the armed Japanese
aircraft ferry "Fujikawa Maru" is by far the center of attraction to
SCUBA divers. The ferry, sunk on Feb. 17, 1944 during World War II,
is located about 1 km off the southwestern tip of Eten Island and can
easily be located since the masts rise above the surface of the water
and are clearly visible. The ferry is 439 ft. long and 58 ft. wide
with a tonnage of 6,938 (Ronald D. Strong, personal communication).
It lies upright on the barren silty bottom in 90 ft. (28.3 m) of
water; the main deck is 50 ft. (15.7 m) below the surface of the
water.

The "Fujikawa Maru," as well as the other sunken ships in the
lagoon, are in essence artificial reefs whose organismal components
represent a climax community that has become dominant after a 30
year period. The only (1972) who included the algal species collected
from the ferry (Station 13) on June 14, 1970.

L Contributions No. 67, University of Guam Marine Laboratory.

2
The Marine Laboratory, University of Guam, P.O. Box EK, Agana,
Guam 96910.

Agricultural Experiment Station, University of Guam, P.O. Box EK,
Agana Guam 96910.

(Manuscript received June 1975 -- Eds.)

The checklist of the algae, corals, and fishes reported below
represent observations made on two dives on March 30, 1975. Although
the number of dives is few, it should be pointed out that the sole
purpose of the dives was to record the species present. Each of the
three authors was responsible for a certain group of organisms -
Tsuda (algae), Amesbury (fishes), and Moras (corals). The soft
corals which represent a dominant component of the ferry are not
included since none of us is familiar with their taxonomy.

Checklist

The algae (21 spp.) are listed under their respective divisions;
the corals (26 spp.) and the fishes (34 spp.) are listed under their
respective families. The five species of algae collected on June 14,
1970 and not seen in 1975 are also included and are preceded by
asterisks.

Algae

Cyanophyta
Microcoleus lyngbyaceus (Kltz.) Crouan

Chlorophyta
Caulerpa ambigua Okamura
Caulerpa brachypus Harvey
Caulerpa filtcotdes Yamada
*Caulerpa lentilitfera J. Ag.
Caulerpa racemosa (Forskal) J. Ag.
*Dictyosphaerta cavernosa (Forskal) Boerg.
Halimeda coptosa Goreau & Graham
Haltmedo dtscotdea Decaisne
*Haltmeda gtgas Taylor
Haltmeda tnerassata (Ellis) Lamx.
Haltmeda macrophysa Askenasy
Halimeda opuntia (L.) Lamx.
Tydemania expedititonts Weber van Bosse
Udotea geppit Yamada

Phaeophyta
*Dietyopterts repens (Okamura) Boerg.
Dietyota bartayresti Lamx.
Lobophora vartegata (Lamx.) Womersley
Padina jonestt Tsuda
Turbtnarta ornata (Turn.) J. Ag.

Rhodophyta
*Champta compressa Harv.

Corals

Antipathia
Cirrhtpathes angutna Dana

Scleractinia
Acroporidae
Aeropora deltcatula (Brooks)
Astreopora sp.
Monttpora erythrae Marenzeller
Monttpora verrucosa (Lamarck)

Agariciidae
Pachyserts spectosa (Dana)

Caryophylliidae
Plerogyra sp.

Dendrophylliidae
Tubastraea aurea (Quoy & Gaimard)

Faviidae
Favia favus (Forskal)
Favia spectosa Dana
Favttes abdita (Ellis & Solander)
Favites favosa (Ellis & Solander)
Leptastrea tmmersa Klunzinger
Leptastrea purpurea (Dana)
Platygyra lamellina (Ehrenberg)

Fungiidae
Fungia fungttes (Linnaeus)

Mussidae
Lobophyllta corymbosa (Forskal)
Lobophyllia costata (Dana)
Symphyllia nobilis (Dana)
Symphyllta sp.

Pectiniidae
Peetinta lacinitata (Milne-Edwards & Haime)

Pocilloporidae
Poetllopora damtcornts (Linnaeus)
Poetllopora eydouxt Milne-Edwards & Haime
Sertatopora angulata Klunzinger

Poritidae
Porttes lutea Milne-Edwards & Haime
Porites sp.

Fishes

Acanthuridae
Acanthurus nigrofuscus Forskal
Acanthurus ntgrorts Cuvier & Valenciennes
Ctenochaetus striatus (Quoy & Gaimard)
Naso unteronts (Forskal)
Zebrasoma scopas (Cuvier)
Zebrasoma veltferum (Bloch)

Balistidae
Baltstapus undulatus (Mungo Park)

Blenniidae
Metacanthus atrodorsalts (Gunther)

Chaetodontidae
Centropyge btcolor (Bloch)
Chaetodon auriga Forskal
Chaetodon kleintt Bloch
Hentochus acumtnatus (Linnaeus)

Cirrhitidae
Cirrhttus ptnnulatus (Bloch & Schneider)
Paracirrhites forstert (Bloch & Schneider)

Gobiidae
Ambtlygobtus albtmaculatus (Ruppell)

Labridae
Chetlinus fascetatus (Bloch)
Eptbulus tnsidiator (Pallas)
Gomphosus varius Lacepede
Haltehoeres hoevent (Bleeker)
Thalassoma lutescens (Lay & Bennett)

Lutjanidae
Caesto caerulaureus Lacepede
Pterocaesto sp.

Scolopsts cancellatus (Cuvier & Valenciennes)

Mullidae
Parupeneus pleurosttgma (Bennett)

Pomacentridae
Abudefduf glaueus (Cuvier & Valenciennes)
Chromis caeruleus (Cuvier & Valenciennes)
Chromts dtmtdtatus (Klunzinger)
Dascyllus aruanus (Linnaeus)
Daseyllus retteulatus (Richardson)

Daseyllus trimaculatus (Ruppell)
Pomacentrus pavo (Bloch)

Scaridae
Searus sordidus Forskal

Siganidae
Stganus argenteus (Quoy & Gaimard)

Zanclidae
Zanelus cornutus (Linnaeus)

Discussion

A vertical zonation pattern is evident on the masts which rise
above the surface. The shallow water Poctllopora eydouxt (coral)
and Turbinaria ornata (brown alga) were only found on the masts at
the surface. Clouds of Pomacentrus pavo and Chromts caeruleus were
also observed around the masts.

The substratum on the deck is composed of fine and coarse frag-
ments of Haltmeda which have accumulated over the past 30 years.
Although corals cannot settle here, this substratum provides an ideal
habitat for the massive holdfasts of the green algae Udotea geppit
and Haltmeda tnerassata.

The bulkhead provides an ideal habitat for mats of the flabellate
form of Tydemanta expeditionis, Caulerpa filtcotdes, and the straggly
Haltmeda coptosa. Lobophora variegata encrusts the bulkhead of the
upper deck. Lobophyllta costata and Symphyllita sp. were the dominant
corals on the various superstructures of the ferry with some colonies
reaching 40 cm in diameter. The pomacentrids were the most numerous
of the resident fish species. Besides those pomacentrids observed
around the masts, schools of Dascyllus species were seen around the
coral growths of the ferry. Acanthurids were quite common though
never in large schools.

Except for the epiphytic Champta compressa on Haltmeda, members
of the red algae were conspicuously absent. Likewise, populations
of chaetodontids were sparse. Few piscivorous fishes were observed
on the ferry. Considerably more were seen on another ship, a sunken
destroyer off Dublon, which was resting on a coral substrate which
supported a large fish fauna of its own. Schools of roving fishes
(Caesto caerulaureus and Pterocaesto sp., and the rabbitfish Stganus
argenteus) were also very abundant, but this is probably quite
variable over time.

The more than sixty sunken ships (Stewart, 1972) in the Truk
lagoon present a unique opportunity for the study of reef community
structure. The time available for colonization of the artificial

reefs is known for each of the ships. The variation in size of the
ships and their position with respect to water depth, substrate type,
distance from one another, and distance from natural reef areas
provide a series of natural experiments on the effect of a variety
of environmental factors on reef community development. It is

hoped that this checklist will serve as a start for further studies
in this unique natural laboratory which is protected by local law.

Literature Cited

Stewart, W. H. 1972. Historical and geographic tourist map of the
ghost fleet in the Truk Lagoon, Eastern Caroline Islands, Pacific
Ocean, with notes on the ocean and World War II. Trust Territory
of the Pacific Islands, Publications and Printing Division.

Tsuda, R. T. 1972. Some marine benthic algae from Truk and Kuop,
Caroline Islands. Atoll Res. Bull. 155: 1-10.

ATOLL RESEARCH BULLETIN
NO. 213

CHEMISTRY OF FRESHWATER POOLS ON ALDABRA
by A. Donaldson and B. A. Whitton

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

May 1977

Aldabra, showing location of 20 pools studied.

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CHEMISTRY OF FRESHWATER POOLS ON ALDABRA
by A. Donaldson and B. A. Whitton !

ABSTRACT

An account is given for 20 freshwater pools on Aldabra of some
of their physical and chemical parameters likely to be of particular
biological significance. The majority of these pools were permanent
throughout the wet season studied, but dry for the remainder of the
year. They were in general characterized by moderate to high levels
of both phosphate and ammonia. Some pools were chosen for more
detailed study of changes through both the whole season and individual
days. Levels of Na, K, Mg, Ca, Cl tended to increase through the part
of the season studied, whereas those of phosphate and ammonia showed
no such clear trend. Diurnal cycles were evident for various
parameters such as level of dissolved oxygen, and in at least one case
Ca and possibly also Mg. It is suggested that the algae Oedogonium
and Plectonema gloeophilum may play a major role in bringing about such
diurnal changes in Ca.

INTRODUCTION

Although freshwaters usually cover only a small fraction of the
total surface of atolls, they do provide a habitat of particular

ecological interest. Further, they are probably the easiest habitat
for which it is possible to obtain accurate data on a wide range of
chemical parameters of biological significance. Apart from comments

on salinity, there are apparently no reports in the literature on the

1 Department of Botany, University of Durham, Durham, England

Manuscript received 8 July 1975 -- Eds.)

chemistry of such pools. The following account deals with
representative freshwater pools on Aldabra during the wet season of
1972/1973.

The four islands (Fig. 1) of the Aldabra group (9° 2aeSi 46° 20° E)
are formed of coral reefs elevated 5 to 10 m above present sea-level.
They are situated in a relatively dry part of the southwest Indian
Ocean, with a mean annual rainfall of 1070 mm (see both Stoddart,
197la and Stoddart and Mole, in press). Further background
information on their geology, geography and ecology is given in the
volume edited by Westoll and Stoddart (1971), while Braithwaite et al.
(1973) have provided a detailed account of the islands' geological
history. Baker (1963) commented briefly on the phosphate content of
rocks and soils. He reported that guano is rare, but that it does
occur in small quantities on West Island. Small pockets of residual
soil also occur locally on this island, and their composition ranges
from 5.2 - 12.6% P. Pale yellowish brown soils in the Cinq Cases
region are low in phosphate, rarely exceeding 0.87% P. All but two
of the pools reported in the present survey are from either West
Island or the Cing Cases region. Baker also described a small area
of phosphatic rocks containing between 15.7% and 19% P, but these could
not be included in our study since freshwater pools were absent in
theaxy) walcainalty,.

McKenzie (1971) recognized five main categories of terrestrial
aquatic niche (for Entomostraca) on Aldabra: permanent freshwater ~
reservoirs, temporary rainwater rock-holes, brackish pools, tide
controlled pools, highly saline pools not obviously tide controlled.
The salinity range for permanent freshwater reservoirs fell between
O.6 and 9.4%o0 and that for temporary rockholes between O.1 and
VOiR-Dy Siar. Our study deals largely with sites which fall into
McKenzie's category of temporary rainwater rockholes. The pools
chosen were however mostly permanent throughout the wet season of
1972/73, though nearly all were dry for the remainder of the year
(Table 1). They were all at the lower end of the salinity range.

The account first summarizes data on the chemistry of pool waters
(but not muds). It then discusses features of their chemistry which
provide clues to the factors influencing the pools which are themselves
of importance in influencing organisms associated with the pools. A
more detailed account of their algal vegetation is included in another
paper following on the present one.

METHODS

Data summarized in Table 1

Grid references. These have been devised as follows. Either the
pool itself, or the nearest obvious landmark, was first located on

the reduced lay-down of the June 1960 aerial photographs (D.O.S. (P.M.
SEY) Aldabra West, Aldabra East, 1969). The position on these
photographs was then related to the two nearest points (such as sites
with bench-marks) findable both on the aerial photographs and the
outline map of Aldabra which includes both grid lines and bench marks
(D.0.S. 304 - series Y852, 1964). Using these marker points, the
position of the pool was determined on the outline map, thus giving
the grid reference.

Geological stratigraphy. The types of rock were established, by
relating the grid reference to the map given by Braithwaite et al.
(1973), and not by direct observation on the ground.

Vegetational types. The names used are based on those given by
Fosberg (1971).

The authors hold a detailed guide to the location of these pools,
together with a system of nomenclature for pools on Aldabra. They
would be pleased to supply copies to any interested.

Water analyses

Provided that a pool was deep enough, probe measurements and a
sample were taken at O.2 m depth. If they were shallower, then the
sample was taken half-way between the surface and the bottom. It was
filtered immediately through a No. 2 SINTA funnel into a 250 ml
polythene bottle. A No. 2 funnel is quoted by the manufacturers as
having a maximum pore diameter of 40-50 um. Although many algae had
one or more dimensions smaller than this, nevertheless the SINTA
funnel held back the great bulk of the algal standing crop.

Part of the filtrate was used immediately for analyses with the
aid of HACH portable kits, while the remainder was stored in the
polythene bottle at 10 1G apart from the times between collection and
return to camp, and during transit to U.K.

ifs 0.D-,5 — Optical density measurements were made only on the
stored samplee, and are probably the least reliable of the parameters
recorded.

2: Temperatures. Use of thermistor incorporated with oxygen probe
and/or mercury thermometers.

3. pH. Use of PYE UNICAM portable meter model 293.

4. Dissolved oxygen. Use of Lakelands Instrument Co. portable meter,
with Mackereth type electrode.

5-8. Na, K, Mg, Ca were all measured after return to U.K. Analar
HCl was added to the samples and measurements made by atomic absorption
spectroscopy (PERKIN-ELMER 403). With the possible exception of the
lowest K levels, the errors associated with storage may be assumed to
be small.

231-339 O-'77-9

4
De Glee Use of argentometric titration.

HOS 2: Orthosphosphate, polyphosphate and organic phosphate.
Use of HACH portable kits. Levels below the minimum detectable were
recorded only for polyphosphate (0.01 mg 17lp) .

The so-called polyphosphate fraction may possibly be heterogeneous,
as it is that part which gives the orthophosphate colourimetric
reaction only after hydrolysis with H2S0q4. If any orthophosphate-
containing material which passes through the SINTA funnel should not
be soluble in the acid provided with the HACH kits (final c. 1.8), then
it would be included in the polyphosphate rather than the
orthophosphate fraction.

13-15. (NH3 + NHg)-N, NO2-N, NO3-N. Use of HACH portable kits.

It seems possible that the values indicated for nitrite and nitrate
were underestimates due to interference from other substances in the
pools. Urea, for instance, if present, would cause a marked inter-
ference. Samples from several sites where it was thought probable
that interference would occur were checked by the method of additions
and no such interference was in fact found. As it was impractical to
carry this out for every analysis, the data summarized here for
nitrite and nitrate should be treated with caution.

Representative rock samples were taken from the sides of the pools.
Small amounts (¢. 0.2 g) were subsequently removed from immediately
below the algal layer (epilithic, or endolithic, if latter present), and
analyzed for Mg, Ca and inorganic phosphate soluble in cold 25%, HCl.

LOCATION OF POOLS STUDIED AND SAMPLING PROGRAMME

The location of the pools studied is shown in Fig. 1, examples are
illustrated in Figs. 2 - 6, and environmental details summarized in
Table 1. Total rainfall was exceptionally high during the 1972/73 wet
season (1 Dec - 25 May: 1414 mm) and some of the pools listed in Table
l as permanent through the wet season probably hold water only inter-
mittently during wet seasons with low rainfalls. All the pools
studied were free of obvious tidal influence.

The pools were sampled for their water chemistry, and in most
cases also for rock samples. For practical reasons it was not
possible to adopt for the water chemistry survey a rigorous programme
of visits to pools, time of day of sampling, or measurements of all
parameters for all samples. A summary of the individual analyses
carried out is given in Table 2:

Programme A was used to establish the main features of the pools,
and possible correlations between individual ions. This consisted of
54 samples from 19 different pools. Where samples were taken from a
Single pool on more than one occasion (Table 2), these were in all

cases separated by at least two weeks. In most cases there was about
17 weeks between the first and last sample. Most samples in A were
taken between O800 and 1100 h.

Programme B was used to establish the changes in individual
parameters over 24 h. It involved a further 82 samples in addition
to those in A. Pools Wl, W2, W4, W5, W1LO were all studied over full
24 h cycles, while Tl and T2 were studied from early morning to late
evening.

In addition to these sampling programmes, individual measurements
for temperature, pH and dissolved oxygen were made both on other
occasions, in these same pools, and also in other pools. Where
results from these are included, this is mentioned specifically.

RESULTS AND DISCUSSION

Individual parameters

Data concerning individual parameters are summarized in Tables 3

and 4, and those on diurnal cycles in Table 5. The relationship
between various pairs of ions is shown in Fig. 7, and examples of
diurnal cycles in Fig. 8. A wide range of factors may be expected
to influence the chemistry of a pool (Fig. 9). The following

discussion includes the evidence available for establishing that
particular factors are of quantitative significance.

Ike Optical density. Water in pools with decaying leaves sometimes
appeared brown in colour, this being especially evident if Casuarina
"needles' were present. In the present survey, even after storage,

the range of values for O.D.429 nm from pools near Casuarina was
Clearly different from pools away from Casuarina (0.146 + 0.055,
Hey O.O48) +. 0.025, n = 15).

Dre Temperature. Depth profiles were carried out in the deeper pools
on West Island on various dates and various times of day. During the
day-time there was usually a slight decrease in temperature with
increasing depth, the maximum recorded difference being 3.6 C (in Wl

with maximum depth of 0.69 m, at 1520 h on 8 March). A very slight
increase however sometimes occurred immediately above the surface of
the bottom mud (e.g. in Wl). At night-time the pools were more or

less isothernal, with minor differences in temperature not exceeding
iL S@NKeHe

a. pH. The lowest value recorded for pool water (at O.2 m depth),
pH 6.2, is slightly above the minimum of pH 5.9 found by Trudgill
(1970) for soil under Casuarina forest.

4. Dissolved oxygen. Among the relatively few measurements taken
just before dawn, no pool became completely free of dissolved oxygen.

6

However it seems probable that in the deeper and more sheltered pools,
the water immediately above the bottom sediments sometimes becomes
anaerobic, and possibly even that occasionally a whole pool may become

anaerobic. The night-time level in pool W2 fell to 0.5 mg 15 atnoe2m
depth (Fig. 8), while the deeper pools often showed a very marked
decrease in oxygen with depth during daytime. The maximum such

decrease found was a drop of 7.7 mg 171 (in wo, dropping from 9.1 mg
1-1 at the surface to 1.4 mg 171 at the maximum depth of,02/73) mat
1430 h on 19 Feb.).

5-8. Cations. As seawater has lower K/Na and higher Mg/Ca ratios
than found in any of these freshwater pools, it would be expected that
the more a terrestrial pools is subject to the influence of added
seawater, then the nearer its ratios for these ions would approach
those in seawater. The pools deviating most markedly from lines of
best fit drawn for K v. Na and Mg v. Ca (Fig.: 7) are:

high K/Na T1>W4>CC1O>CC11 ... W6<W9<W7<CC9 high Na/K
high Mg/Ca CC9>W1>CC1lO>CC12>CC1l ..5. W2<W7<T2<W3 high Ca/Mg

Only CC9 (Bassin Flamant) combines both low K/Na with high Mg/Ca.
This suggests the possibility either that this large pool is
influenced by seawater more directly than any other pool, or that
complicating factors are less important here than elsewhere McKenzie
(1971) has suggested that freshwater in this area forms a lens floating
on top of seawater. However other factors might also be important in
determining cation ratios in this pool. CC9 already has a much higher
cation level than many of the other pools studied, so any addition, as
from animal excreta, will have less effect on the overall ratios than
it would in pools with lower cation levels. Further, in spite of the
considerable animal activity associated with CC9, it seems probable
that visiting animals are of much less quantitative importance in
relation to the volume of the pool than they are for smaller pools
such as :T'2’.

Apart from CC9, there was no obvious relationship between K/Na
and Mg/Ca, suggesting that other factors such as local rock or animal
behaviour probably play an important role in controlling K levels. Of
the four pools with the highest K/Na ratios, Tl, CClO and CCll are in
fact subject to very high tortoise activity, and W4 to frequent visits
by crabs (Table 1). If excreta from these animals is adding
relatively more K than Na, this is presumably a reflection of high
K/Na levels in the vegetation on which the animals feed, since
selection excretion of K in preference to Na is most unlikely.

Where pools were sampled on several different occasions, Na and K
levels increased through the season, roughly parallel to increases in
Giles This was probably due largely to evaporation, but as suggested
above animal excreta may also sometimes make a significant contribution
to the increasing levels. . Data for three occasions in W2 when the
depth of water was almost the same are summarized in Table 6. The

increase in K, as a fraction of seawater, was four times as great as
that for Na and Cl, suggesting that bird excreta in this pool (Table 1)
may be an important source of K.

If there were no factor ever leading to a loss of Na, K and Cl
from pools, they would become ever more saline. However probably all
pools are capable of filling up their whole catchment area and over-
flowing to other small catchment areas after long periods of rain.
However, during the period summarized in Table 6, W2 was never diluted

by rainwater sufficiently for such overflow to occur. In shallow
pools lacking any well defined rim there is probably also some loss of
bottom detritus due to wind erosion in the dry season. However the

sides of W2 are sufficiently steep to make this explanation unlikely
here:

Among the four pools whose Mg and Ca levels were followed in
detail over a day's cycle (Table 5, Fig. 8) one (W7) showed a clear
diurnal cycle for Ca and a possible one for Mg, two, Wl (Fig. 8) and
W5, showed an erratic distribution of values suggesting that diurnal
cycles might be being masked by some other effect, while one (W2)
showed no indication of cyclical change. This last had the least
diurnal range in pH.

Pools (Wl and W7) both had massive growths of Oedogonium filaments
and Plectonema gloeophilum flocs at the time of survey, whereas these

were sparse or absent in W2 and W5. Both these algae in Wl and W7
had a heavy calcareous encrustation, so presumably were important
agents helping to remove Ca from solution. A diurnal cycle such as

found in W7 might result from a combination of day-time removal of Ca
by deposition around the surface of the algae, and a night-time
release of Ca resulting from the higher levels of COz present due to
respiration in the pools.

2) Chloride. The increasing Cl levels found through the season
suggest that data on Cl levels in pools immediately prior to drying
out would be of great interest, since any organism present will
presumably suffer extremely high osmotic levels, and this might well
prove one of the most critical factors in their survival ata
particular pool.

MO=i2 i Phosphate. Fractions attributed to orthophosphate,
polyphosphate and organic phosphate by the HACH kit methods were all
well represented in the pools, and usually all three were present at
levels likely to be of significance to organisms able to use these
particular forms of phosphate. Examples occurred of each of the three
fractions being the most abundant, but in general orthophosphate and
Organic phosphate were more abundant than polyphosphate.

Although levels showed considerable variation, W4 and W7 were
Clearly characterized as low (ortho-) phosphate pools, while W2 showed
very high levels. In contrast to the four cations measured, there

8

was no obvious tendency for concentrations of any of the phosphate
fractions to increase through the season.

13 AS ic Inorganic combined nitrogen. The Aldabra pools always
carried moderate to very high levels of ammonia (Table 4). It seems
likely that nitrite and nitrate levels were low in comparison, but due
to the problem of interference by reducing substances, this is still
uncertain. For instance, 10 mg urea 171 causes an apparent reduction
of 50% in a solution of O.5 mg KNO3-N 11, The results obtained for
nitrite and nitrate have therefore been omitted from Tables 3, 4 and
5, but are summarized here as they do provide some clues to the
situation in the pools. Only 5 out of 54 analyses in programme A gave
detectable values for these ions. The values recorded were, for
NO»5-N: minimum, < 0.003; maximum, O.52, mean including W2, 0.008, mean
excluding W2, < 0.003; for NO3-N: minimum, < 0.01, maximum, 0.56, mean
including W2, 0.12, mean excluding W2, < O.O1 mg ibe a In one case
(W2, Fig. 8), however, there was an apparent diurnal cycle of these
ions, with the oxidized forms reaching a peak in the afternoon. Even
then, the indicated nitrogen present in these oxidized forms represen-
ted only 3.5% total inorganic combined nitrogen.

As with orthophosphate, there was no tendency for ammonia levels
in particular pools to increase through the season.

Rock analyses

Data for these are summarized in Table 7. As only 28 analyses
were carried out, and little attempt made to separate recent sedimentary
material from older rock, it is not possible to make any detailed
comparison between chemistries and rock analyses. A few preliminary
observations do however suggest that such a survey might prove of
considerable interest. Two rock samples from Wl showed by far the
highest Mg/Ca ratio. Similarly the highest percentage PO4-P was
recorded from rock below W2, the pool with the highest P levels in the
water.

GENERAL DISCUSSION

Dynamics of soluble phosphate and nitrogenous compounds

The most striking features of the freshwater pools on Aldabra are
the high levels of both dissolved phosphate and inorganic combined
nitrogen often present, and the fact that the latter is apparently
represented: largely by ammonia-N. The main source of both phosphate
and ammonia is presumably from the excreta of animals such as crabs,
birds and tortoises. The high levels of organic phosphate indicate
the importance of such excreta. As many of these animals carry out
the bulk of their feeding away from the pools, addition of their
excreta to the pools will bring about, during the wet season, a net
transfer of nutrients from surrounding ecosystems to the water.

Some 60-87% of the total nitrogenous waste material in marine
and freshwater crustacea is in the form of ammonia (Lockwood, 1968).
In bird faeces most nitrogen occurs as excreted uric acid and amino
compounds contained in the undigested food residues (Ganning and Wulff,
LIGY)|. Copeman and Dillman (1937) found that 85% of a guano sample
in contact with water was converted to ammonia in 4 days. The large
quantities of ammonia usually present in the pools are therefore
probably in part due to direct excretion of ammonia, and part to
microbial breakdown of uric acid and other organic nitrogen compounds.

It is widely assumed in the literature that in an aerobic
environment ammonia will become oxidized quite rapidly as a result of
microbial activity, first to nitrite and then to nitrate. It seems
unlikely that the apparent near absence of such a process in pools on
Aldabra can be explained entirely by the interference in analytical
techniques discussed above. Ganning and Wulff (1969) reported a
Similar strange lack of nitrification in brackish water rockpools
receiving much bird excreta by the Baltic Sea, and these authors
suggested that some inhibitory substances might be present in the pools
hindering oxidation of the ammonia. Although they apparently made no
allowance for chemical interference in their analyses of nitrite and
nitrate, they did find these ions to be relatively abundant at some
seasons of the year, the overwhelming predominance of ammonia occurring
only during late summer.

Neither phosphate nor ammonia showed any obvious tendency to
increase in a particular pool during the season. As the algal
standing crops also did not show any marked increase during the season
(see next paper), accumulation by these organisms is unlikely to be a
major cause of net removal of these nutrients. The very high pH values
occurring during the afternoon in some pools are probably a significant
cause of the loss of NHg-N, as a result of the ammonia - water
equilibrium, NH4t + H20%NH3 + H30t, being displaced towards the right-
hand side. An odour of ammonia was sometimes detectable in the
vicinity of several pools on Aldabra. Removal of soluble phosphate
is less easily explained, though probably precipitation of calcium
phosphate is one significant cause.

Comparisons with waters in other regions

Several examples exist in the literature of waters with levels of
phosphate and inorganic combined’ nitrogen rather similar to those of
the pools on Aldabra, excluding pool W2. Howmiller (1969) suggested
that the hypersaline Arcturus Lake on Genovesa Island, Galapagos, is an
outstanding example of guanotrophy. The nutrient levels reported for
this lake were 0.46 mg 1-1 PO, and 0.45 mg 1-1 NO3-N; ammonia was at a
low level in comparison with nitrate. The rockpools described from
the Baltic by Ganning and Wulff (1969) showed variation from one to
another and also throughout the year; the maximum levels reported were
0.731 mg 1-1 PO4g-P and 0.325 mg 1-1 NHy-N.

It is difficult to find any reports of waters carrying the levels
of phosphate and inorganic combined nitrogen found in pool W2, and which

10

are not polluted by sewage or some other human activity. A pond in
Hyderabad studied by Seenayya (1971) provides a fairly close parallel,
but this pond (Golkonda Pond) is heavily polluted by sewage.

ACKNOWLEDGEMENTS

We are most grateful to the Royal Society and the Natural
Environment Research Council for financial support and encouragement
in our research on Aldabra, and for the help in collecting data
provided by, among others, L.U. Mole, F.W. Topliffe, J.R. Wilson and
H. Charles, for helpful discussion with Dr. S. Trudgill, for analyses
by W. Simon, T. Brett, P.J. Say and Mrs. G.A. Walker, and for. typing
by Mrs. V. Evans.

REFERENCES

Baker, B.H. NODS Geology and mineral resources of the
Seychelles Archipelago. Mem. geol. Surv..Kenya 3: 1-140:

Braithwaite, C.J.R., Taylor, J.D. and Kennedy, W.J. IQ YSs
The evolution of an atoll: the depositional and erosional
histony of Aldabva.) Phil.) Trans. ‘R- sSOC.. Bn26Os 50/52 Or

Copeman, P.R. v.d. R. and Dillman, F.J. TOBE Changes in the
composition of guano during storage. Tic AGE Cie OC Ce es
IL Rossy e

Fosberg, F.R. OTA Preliminary survey of Aldabra vegetation.

Phil Trans. wisn SOG. BiaZ6Os, 25-2258

Ganning, B. and Wulff, F. 1969. The effects of bird droppings
on chemical and biological dynamics in brackish water rockpools.
Oikos 20: 274-286.

Howmiller, R.P. 1969. Studies on some inland waters of the
Galapagos. ECOLOGY) O21) (3-00):

Lockwood, A.P.M. 1968. Aspects of the Physiology of Crustacea.
328 pp. Oliver & Boyd, London & Edinburgh.

McKenzie, K.G. 1971 ¢ Entomostraca of Aldabra, with special
reference to the genus Heterocypris (Crustacea, Ostracoda).
Phil. SLANS. wise SOC, Be 260%, 207 —2984

Seenayya, G. OFA Ecological studies on the plankton of certain
freshwater ponds of Hyderabad - India. I. Physico-chemical
complexes. Hydrobiologia 37: 7-31.

Stoddanty: Dare 197l1la. Scientific studies at Aldabra and

neighbouring islands. Phid ma TLansingwh. MSOC VBE ZOO OO

ALI

Stoddart, D.R. ILS} 7/AU 0}, Place names of Aldabra. Phighs “rans.
Rew SOCe BB 260s) 6Si-652)-

Stoddart, D.R. and Mole, L.U. (in press) Climate of Aldabra Atoll.

Sverdrup, H.U., Johnson, M.W. and Fleming, R.H. 1946. The Oceans,
their physics, chemistry and general biology. 1087 pp.
Prentice-Hall, N.Y.

niretuclepalilil; Sout. 1972. Process Studies of limestone erosion in
littoral and terrestrial environments, with special reference
to Aldabra Atoll, Indian Ocean. Ph.D. Thesis, University of
Bristol, England.

Westoll, T.S. and Stoddart, D.R. (eds) 1971. A Discussion of
the results of the Royal Society Expedition to Aldabra 1967-68.
Piss rash Reto OC sb) 260) o> 4e pple

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Table 3. Summary of data on physical and chemical parameters for water
in pools. Mean values are based on programme A, minimum and maximum

ones on programme B.

Values added in brackets are extremes found

during spot readings taken anywhere in freshwaters on Aldabra.

Concentrations are in mg 1-l.

detectable values are treated as zero).

0-D- 455

temperature

orthophosphate-P
"bolyphosphate"-P

organic
phosphate-P

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minimum maximum

0.021

OnZ73

DANSE 340) 372 (41-2)

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Table 7. Analyses of rock samples taken from sides Of pools.
Values given are for material soluble ine 25%, Hel Vacs a percentage of
the total rock, including soluble component.

min. max. mean
Mg Of22 IoD 0.543
Ca 27.6 SS eal DN ATE

Ie) 0.01 OL.L7. 0.05%

ILOjn AT

Pool Wl at beginning of wet season.
of floating Oedogonium.

Note massive

flocs

Pal

22

ldaKop o)5

Pool Wl near end of wet season, showing its steep sides
exposed as pool dried out. Marker in this (and subsequent
figures) is 1m long.

Fig. 4. Pool W2, when almost full. The surrounding white,
terrestrial areas are ones which lack the normal
Tolypothrix byssoidea algal cover.

Pad

1DaIC De Pool W4. Note sharp boundary between the terrestrial rock
which is never submerged, and which appears dark due to
growths of Tolypothrix byssoidea, and the now exposed upper
part of the pool which has a cover of Calothrix parietina.

24

Bp Che Ove

Pool W7, when relatively low. Note flocs of Closterium

floating on the surface, and the black mud already deposited
as the pool dries out.

Fig. 7. Relationship between various pairs of ions studied in
programme A:

1000 [

10 10-0 100-0 1000-0
Na mg I-1

100-0

1-0 | J

10-0 1000 1000-0
Ca mg I-!

|
| a) log K v log Na, regression coefficient = 0.9, correlation
| coefficient = 0.9

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coefficient = 0.5

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.
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= 0.5 correlation coefficient = 0.7

a) Log’ N vlog ortho PO4-P, regression coefficient = 1.8,
correlation coefficient = 0.8

Fig. 8. Diurnal cycles of various parameters in pools Wl and W2
(on 5/6 June 1973)

280 are
274 30:7
wa 268 205 w2
262 28-3
256 Temperature 271
°c
250 25:9
244 247
: 238 23:5
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ATOLL RESEARCH BULLETIN
NO. 214

- OBSERVATION ON REDOX POTENTIAL
IN FRESHWATER POOLS ON ALDABRA

by B. A. Whitton and M. Potts

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

May 1977

OBSERVATION ON REDOX POTENTIAL
IN FRESHWATER POOLS ON ALDABRA

by B. A. Whitton and M. Potts !

ABSTRACT

In comparison with most values reported in the literature for
freshwaters, small freshwater pools on Aldabra were found always to
have values for redox potential which were low, the great majority
lying below + 200 mV at a standard pH of 7.0, using a correction of
58 mV per unit pH difference from the standard pH. Values were
especially low in pools in Casuarina forest, where readings below
+ 50 mV were recorded frequently. Typical freshwater pools on Aldabra
combine a well oxygenated environment with low redox potential, an
environmental combination which has received little investigation in
the diterature.

INTRODUCTION

In a previous study of the chemistry of freshwater pools on
Aldabra (Donaldson and Whitton, 1976), it was found that most of the
inorganic nitrogen present was apparently in the form of ammonia
rather than nitrite or nitrate. This occurred in spite of the fact
that the pools were often highly super-saturated with oxygen by daytime.
The only other observations in the literature reported a similar
(apparent) loack of nitrification are those made by Ganning and Wulff
(1969) in brackish rockpools by the Baltic Sea. These latter authors
suggested that some inhibitory substances might be present in the pools
which hindered oxidation of ammonia.

Since the previous account of the pools on Aldabra, the present
authors have had the opportunity to collect further data from the atoll
which would help account for any lack of nitrification. A full report
of these will be published elsewhere, but a summary of observations on
redox potential in the pools is given here in order that it may be read
together with the previous account. These observations were all made
during the period December 1974 - January 1975.

1 Department of Botany, University of Durham, Durham, England.
(Manuscript received 8 July 1975 --Eds.)

METHODS

The system used for naming the pools has been described in the
previous account (Donaldson and Whitton, 1976).

Measurements of redox potential were made using a PYE UNICAM
portable meter model 293 with PYE UNICAM combined redox electrodes,
these consisting of a platinum indicator electrode and a silver /
Silver chloride reference system. Cleaning and buffering of the
electrode was made at frequent intervals during sampling. The Pt
indicator electrode was first cleaned by rubbing gently with emery
paper and then immersing in concentrated, chlorine-free HNO3. It was
then rinsed and calibrated against a redox buffer. Measurements of
pH were taken at the same time as redox potential, using a similar
meter, but witha pH electrode. Data on redox potential is reported
here both\as a direct’ reading, (Eh): and one Vcorrected) "to pHi 7 Or (hiayr
Due to wide diurnal fluctuations in pH, it is essential to make some
initial comparison of results with redox potential values corrected to
a standard pH value. The correction value of 58 mV per unit pH used
in the earlier literature was used here. Where simple tests of adding
acid or alkali to samples of pool water weré carried out, these did in
fact indicate that this was a reasonable choice of correction value.

It was unfortunately not possible to take measurements of dissolved
oxygen simultaneously with those of redox potential due to loss of
equipment on route to the atoll. However observations on ammonia,
nitrite, nitrate, phosphate and algal vegetation indicated that the
behaviour of the pools was in general very similar during the present
period of study to that during the previous period.

RESULTS

Of the 20 pools whose chemistry was described by Donaldson and
Whitton (1976), six were chosen for detailed study of redox potential
changes with depth and time on 17 January 1975. Part of the data
obtained is summarized in Table 1. As the sediment surface tends to
be both ill-defined and a region of rapid change, not too much attention
should be paid to small differences between readings taken immediately
above the sediment surface.

In addition to these readings, measurements were taken in other
small pools on 17 January, and also in a range of pools on other dates.
These somewhat extended the upper range of redox potential values, the
highest being a.small pool filled with Oedogonium:

te pH Eh Eh7
0950 h 2829 iO + 245 mV + 274 mV
All freshwater pools found with an Eh7 value in the water near the

surface of the pool lower than + 50 mV (including W7, W8, W9) were
associated with the Casuarina Forest, and had waters coloured a pale

brown.

DISCUSSION

These data are obviously fragmentary, and any conclusions should
be treated with caution until a more intensive study has been carried
out on them. Nevertheless they do indicate the probability of several
features of interest.

ie All the pools showed marked variations in redox potential during
the day. As repeated readings at any one time were consistent, and as
the pattern of changes was quite different in different pools, it seems
unlikely that these variations were associated with any sort of
instrument error. It may be pointed out that the pool which showed an
increase between successive readings during the day, W2, is also the
pool for which Donaldson and Whitton (1976) reported a late afternoon
peak in nitrite and nitrate values.

Die In comparison with data for soils and lake sediments, the
literature on values for redox potential in freshwaters is rather
sparse. It would however seem clear from the literature available

(Baas-Becking et al., 1955; Hutchinson, 1957) that in comparison with
most oxygenated waters, the values for pools on Aldabra are low, and
those for the pools in the Casuarina Forest remarkably low.

Bye If freshwaters behave in a manner similar to soils (Pearsall, 1938;
Reddy and Patrick 1975), then the redox potential measurements recorded
for the majority of Aldabra pools correspond with environments which do
not favour nitrification. If the values quoted by Pearsall (1938) are
corrected to Eh7 values, then he found that with only three exceptions,
all soils studied where nitrate predominated over ammonia had an Eh7
greater than + 234 mV, and soils lacking nitrates had an Eh7 less than
204 mV. The three exceptions found by Pearsall were all soils
receiving drainage from stream waters containing nitrate.

These observations would provide an explanation for the rarity of
detectable nitrate in Aldabra pools. In most instances the redox
potential values correspond to an environment which is sufficiently
reducing that nitrification would not be expected to occur. Further,
pool sediments were always found to be markedly reducing, and as the
sediments are in many cases disturbed frequently by the activity of
crabs, any chemical changes taking place in the sediments may be
expected to have a marked effect on the chemistry of the water above
the sediment.

It is less clear what are the probable agents responsible for
bringing about these low redox potential measurements. The fact that
all the pools in the Casuarina Forest had low values suggests that. the
brown materials leaching from the debris of "needle" and other fallen
parts of the tree may here be partly responsible. However the
variations taking place during a single day within one pool suggest
that some other quite different factors must play a role. Disturbance

of the sediments and release of excreta by crabs would seem likely such
FaCEOwRS!.

ACKNOWLEDGEMENTS

We are most grateful to the Royal Society and the Natural
Environment Research Council for financial support.

REFERENCES
Baas Becking, L. G. M., Kaplan, I. R. and Moore, D’. 1960. Limits
of the natural environment in terms of pH and oxidation-
reduction potential. JW GeOL es G8 QAZ=284\,
Donaldson, A. and Whitton, B. A. 1976. Chemistry of freshwater pools
in Aldabra Atoll. Atoll Research Bulletin, this issue.
Ganning, B. and Wulff, F. 1969. The effect of bird droppings on

chemical and biological dynamics in brackish water rockpools.
OLTKOS 120) 274-286.

Hutchinson, G..E. OSs A Treatise on Limnology. 1. Geography,
Physics and Chemistry. 1015 pp. John Wiley Sonsiine a INiay 2,
U.S.A.

Pearsall, W. H. 1938. The soil complex in relation to plant
communities 1. Oxidation-reduction potentials in soils.
ey HCO V6 a LOOM y

Reddy) Ke Rai vand yPalteraa chy Wai, Witan LOW y. Effect of alternate
aerobic and anaerobic conditions on redox potential, organic
matter decomposition and nitrogen loss in a flooded soil. Soil

Biol. Biochem. 7: 87-94.

RYE OIC —S eI Ono Cm COL OUC> SINE IE6C, V6 OVG= OB n/c sSe Coe OSL caOnomec oO mCOL-EGUa OO mee OIC O9C—s Onin OF OS8T-OT8T

IGi= GN= SLO 6G GEG Rim GUS E UE CUG= Aol EPS) BrCl CXS VAIS Ol4e yh IPS — BIG ord ARI) is OV EES Sie) Os SSbI-SZPT
GIs GIS OGG ERG VES C= GSS Wiehe WI SiS SEOC) EEG — Gi Sr SIGE Chekd= hie CIES 9) eeks esi (als GUS EE O€OT-0780
Goce ea qusuTpes

JO 90eyJANS MOTEeq uM OOT

EU NI C/E POG GE Corse. loli} Eee - “Che O7= OBL SESE t= On OV6SSisse SCOLEOIt SOLS tac Sit= Orla. Val Bae OS8T-OT8T
CE= Git OYE*S) SOs Guy da GOTO AS te. 76g = teh So SSE: f3}5(0\7 [Et (Achim Gath KO isl SSS (ONES {EAS GS SIGS ONES GG SSPI-SCPT
GShr Sloss SiGe S) GARG ES Siar = Clan Gi (E85. hse @ Silden AGES GEG ELGr SOE OGmEnGm OG LG OB == Oa Le S8 Cave Gy tO SiGe Cass O£OT-O0Z280

queutpes Fo soezans ZeAO

Gv Os SEE OG -Gb= Cll On} GeOG iAP Ose Cl)e/h anes: - Achiae Oi (MONE AS ksi Sine CSG) Cede Clear Oshiae Che (<as OS8T-OT8T
UVa OSs ie ee ee On OV CGO Lm Lem L a O9- lOSiees Gail, i6Gllt Gila Cl sOlniGulem lot G8= GEG) CeO7 —SGilt OOLE G6eL iyaige SSVT-SCPL
Cfgar - @Olshs BITS) OSA hehe -eYene WE SS) eke avbne Sear Clg>S) (Gg = tise Aline GSMs ING “OAS (orsiing Ste a/6 t339KA (eS (lo) ia Seats). Sh O€OT-O7Z80
[ood jo soezins mMojTaq um Oz
‘ua ua ud a Ate? Wer gist ‘ua ug Hd 3 ‘ua ug Hd al ‘ug ua Hd 34 Lue Ue = Hae 2 u = Z ie Xe
6M 8M LM DM €M ZM Kep JO outz pue food ut uotz1Tsod
(9x09 38eS) sTood xts z0J senTeA (AW) TeTQUeqZOd xopez pue Hd ‘(D, ) sanqezoedusy, T eTqeL

231-339 O-'77 - 1

ATOLL RESEARCH BULLETIN
NO. 215

ALGAL FLORA OF FRESHWATER
HABITATS ON ALDABRA

by A. Donaldson and B. A. Whitton

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

May 1977

°3X0] UT peuUOCTqjUeU seoeTd FO uoTRIeDOT Hutmous ‘erzgqeptw ec Ota

ALGAL FLORA OF FRESHWATER
HABITATS ON ALDABRA

by A. Donaldson and B. A. Whitton !

ABSTRACT

An account is given of the algal flora of freshwater habitats on
Aldabra, together with ecological notes for each species. The distri-
bution of these species in 20 pools chosen for more detailed study is
summarized. Based on their occurrence within these 20 pools, the most
widespread species are: Lyngbya sp., < 1 um (20); Calothrix parietina
(15); Phormidium mucicola (15); lLyngbya nordgardhii (14); Oscillatoria
pseudogeminata (14); Phacus orbicularis (13).

INTRODUCTION

Although freshwaters cover only a small part of the surface of
Aldabra, they nevertheless provide a wide variety of habitats for algae.
They range, for instance, from shallow depressions holding water for only
a few hours to pools permanent throughout the wet season. The physical
and chemical parameters of the water may also show considerable varia-
tion, as has already been described in the previous paper (Donaldson and
Whitton, 1976). Preliminary observations (Whitton, 1971) indicated
that the algal flora of freshwater pools was sometimes quite rich in
species, so this habitat was chosen for more detailed study in the
survey carried out during the wet season. The following account
summarizes the floristic observations made during this survey.

METHODS

Location of pools and sampling programme

Observations were made on pools on West Island, Middle Island and
South Island. The pools chosen for especially detailed sampling

1 Department of Botany, University of Durham, Durham, England
Manuscript received 8 July 1975 -- Eds-)

2

included the 20 whose nomenclature and chemistry were described by
Donaldson and Whitton (1976), together with a further 79 mostly in the
Takamaka, Cing Cases and Anse Cédres regions. The maximum level of
Cl recorded in the chemical survey of pools was 1260 mg 17! in cc9

(+t =,:2.3% 5 salinity).

It seems probable that almost all the other pools used for
observations were, at any particular stage of drying out, less saline
than, €¢9). However it also seems probable that most pools reach high
salinity values towards the end of the wet season. CC9, recorded by
us aS 2.3%0, was reported to have a salinity of 1-5%0 by McKenzie (1971).

Unless stated otherwise, records were made during the wet season,
November 1972 - Junel973. A brief comparison is however added of
observations made in December 1968 - January 1969.

The 20 pools listed in Table 1 were sampled on the following
number of occasions: WL, 6; 9 W2;, Se WS), Far JW4) (Si WS), 7 el era
Wi, (Si OWS), Ses WO aes Ss WIO2,. hs sWwiOS),, dys) Ml Si 2) e-em CCA ESE
CCS; S72 (CCB 2 wi CO 3i35 Cel}, ike COINS eA cle male Although
only a few details are included here, quantitative plankton counts were
made for pools Wl, W2, W3, W4, W5, W6, W7 and W9.

Samples were taken from each obvious microhabitat within a pool.
The principle microhabitats (together with abbreviations used throughout
the text) were as follows:

F floating at or near the surface (including neuston)
P plankton

Ep epilichire

En endolithic

El epipelic

Et epiphytes

A aufwuchs, not including true epiphytes.

Other microhabitats occurring occasionally included dead wood, leaves
and structures resulting from man's activities.

The species recorded are all based on direct microscopy observa-
tions. A much fuller list would no doubt have resulted if culture
techniques had been used, since under these conditions not only might
a few of the rarer plankton species have had an opportunity to develop,
but probably also many of the widespread terrestrial species.

Taxonomic

The eukaryotes named have been identified using the floras standard
for the respective groups. Although it has so far not proved possible
to name the remaining forms, this is in most cases for lack of repro-
ductive stage or because the groups require specialist help. The
authors can supply further details of these forms to any workers dealing
with algae on other atolls. The separation of the two Closterium forms

termed here sp. A and sp. B did however cause difficulty. Sometimes
the two forms were quite distinct, whereas on other occasions material
was present which appeared intermediate.

The problems involved in naming the blue-green algae (Myxophyta)
were rather different. There are several quite different approaches
available in the literature for treating these organisms, but none are
ideal for producing a descriptive account of field populations including
a wide range of forms. The method used in the preparation of this |
account was as follows. As far as possible, a binomial was allocated
using the conventional floras, in particular those of Geitler (1932),
Desikachary (1959) and Starmach (1966). For some genera where the
limits between species as originally described appear to be based almost
entirely on rather arbitrary size (trichome width) limits, we have
divided up the overall size range in a more systematic manner, following
the practice already adopted by Golubié (1967). We have then used the
conventional binomial which corresponds most closely to the modified
size limits. These hatter are added after the binomial and the author
responsible for the original description. The genera treated in this
Manner are: Aphanocapsa, Aphanothece, Chroococcus, Gloeocapsa,Gloeothece.

The computer numbers for each species are those used in a recording
system produced by B. M. Diaz, N. T. H. Holmes, M. K. Hughes and the
authors. Further details of this system are available on request to
the authors.

RESULTS
Taxonomic

The list of photosynthetic organisms found in pools is summarized
in the next section. It consists entirely of algae with the exception
of one aquatic monocotyledon, Naias graminea. A few other angiosperms
were however occasionally partially submerged in transient pools, this
being true especially of the (?) introduced herbs Portulaca oleracea
and Stachytarpheta jamaicensis on West Island.

Algal species which are more typically terrestrial are included
within the lists for some pools. This is because some rocks are
alternatively submerged and exposed as the water level rises and falls
during the wet season.

For convenience all items in the floristic list are termed species
in this account, although in a few cases varieties are included, while
in others several different forms have been lumped together.

Rllowmisitac Wisi

MYXOPHYTA

010215 Anabaena variabilis Kutz. DANG

4

010216

010531
010532
010533
010534
©10535
010536
010602
010631
010632

010902

010905
010910
010911
011302

011534

011538

011539:

011540

011541

011811

011903

A. ambigua Rao. 126 Widespread, and sometimes abundant,
especially at the beginning of the wet season; later formed
spores which dropped on to the bottom.

Aphanocapsa fusco-lutea Hansg. > 1 < 2 um. IDA aml, Ne
A. montana Cramer. > 2 < 4 um. HE 5 7NG

A. grevillei (Hass.) Rabenh. > 4 < 6 um. Perle aA
A. roeseana de Bary. > 6 < 8 um. Peeler Aye

A). testacea (A. Br.) Nagi > 84m. IDA ade ONG

A. delicatissima W. et G. S. West. < 1 um. IPL, VNC
Aphanothece pallida (Kutz.) Rabenh. B, sp.

Aphanothece saxicola Nag. < 2 um. Po Ra. Ae

A microspora (Menegh.) Rabenh. > 2 < 4 um. LIB as dhl. JANG
Calothrix braunii Born. et Flah. nea Ne Widespread, especial-

ly among floating flocs of other algae; at least probably
just growth stage of C. parietina.

C. elenkinii Kossinskaja. Ep.
C. marchica Lemm. Ns
G. uparietina Thuret. “Bp. Main cover of most rocks in pools.

Chlorogloea microcystoides Geitler. Ep.

Chroccoccus turgidus (Ktutz.) Nag. > 8 < 16 um, lamellate
sheath. Ep, A. Widespread.

C. minutus (Kiutz.) Nag. > 4 < 6 um, non-lamellate sheath.
Pe Ely Ae

C. membranicus (Menegh.) Nag. > 6 < 8 um, non-lamellate sheath.
Be eEds Epi ae

C. turicensis (Nag.) Hansg. > 8 < 16 um, non-lamellate sheath.
Dahl, Ep, A.

C. spelaeus Ercegovic. > 16 < 32 um, non-lamellate sheath.
ISP ATH esa oy TN

Cylindrospermum muscicola. El, Ep, A.

Dactylococopsis raphidioides Hansg. PR: Rare; recorded only
from West Is.

011904

012201

013107

012631

012640

012641

012642

012647

O22731

012732

©12733

012801

012802

012901

OL3S13

013601

013602

013604

013801

D. rupestris Hansg. 12) Recorded in only two pools. Abundant
iin, (SILL.

Entophysalis granulosa Kutz. Ep.

Hapalosiphon welwitschii W. et West. EM Ep.

Gloeocapsa montana Kitz. > 2 < 4 um, sheath colourless. A a sL
Ep, A.

G. dermochroa Nag. > 2 < 4 um, sheath yellow-brown. 15 dail
Ep, A.

G. kutzingiana Nag. > 4 < 6 um, sheath yellow-brown. 12a oy
Ep, A.

G. muralis Kutz. > 6 < 8 um, sheath yellow-brown. IDF AL Aoi
A.

G. sanguinea (Ag.) Kitz. > 4 < 8 um, sheath red-violet-blue.

Ep. Predominantly terrestrial, but found occasionally
in pools.
Gloeothece violacea Rabenh. < 2 um. Ep.
G. palea (Ktitz) Rabenh. > 2 < 4 um. IO) pa tnll 5 VN
G. repestris (Lyngb.) Bornet > 4 < 6 um. ldjoyye tad le 7Ne Mainly

restricted to terrestrial habitats, but occasional records
for pools, where it can form macroscopic colonies overlying
rocks.

Gloeotrichia echinulata (J. E. Smith) P. Richter. Et.

G. ghosei R. N. Singh. Et. Forms large brown lobed colonies
on Chara.

Gomphosphaeria aponina Kutz. Pe Rare.

Homoeothrix varians Geitler. Bt. Frequent epiphyte on

Gongrosira, and sometimes also on dead wood.

Hyella caespitosa Born. et Flah. En.

H. fontana Huber et Jadin. En. Especially abundant in pools
W5 and W6é.

H. balani Lehmann. En.

Johannesbaptistia pellucida (Dickie) Taylor et Drouet. P.

Found only in the more brackish pools, where it sometimes
formed large floating clumps.

6

014101

014201

014202

014203

014204

014205

014206

014207

014208

014209

014210

014211

014212

014214

014215

014231

014232

014603

014631

014632

014633

014634

Lithonema adriaticum Erceg. En. Recorded only from W2.
Lyngbya aestuarii Liebm. Dp IB. Very abundant in CC9

L. allorgei Fremy. ILA ING

L. aerugineo-coerulea (Kutz. ) Gom. ee VLA AN,

L. confervoides Ag. F, PB, A«

L. digueti Gom. Pl hl AS Widespread.

L. epiphytica Hieron. Bite Frequently recorded epiphyte on

Plectonema gloeophilum and Oedogonium.,.

hieronymusii Lemm. 1D AB Wanlbes Forms olive-green floating
mats in pools in Casuarina Forest; eaten by Cardisoma.

kutzingii Schmidle. Et. Occasional epiphyte on Oedogonium
limnetica Lemm. Ey, Ps Widespread.

majuscula Harvey. ING doy, Jobe

martensiana Menegh. (Hoy tI NE Widespread, and often

abundant, either among other algae, or forming nearly
unigal sheets.

Lyngbya nordgardhii Wille. Ep. Widespread.

Tie

pusilla (Rabh.) Hansg. Bt. Frequent epiphyte of Plectonema
gloeophilum, Oedogonium and Pithophora.
rigidula (Kutz.) Hansg. Bites Frequent epiphyte of

Plectonema gloeophilum, Oedogonium and Pithophora.

Lyngbya sp., not above, < 1 um. lab NG Probably = L. erebi

W. and G. S. West.

Lyngbya sp., not above, > 1 < 2 um. El, A:

Merismopedia trolleri Bachmann. El. South’ Is. oni:

M. minima G. Beck. < 1 um. Bl. South Is. only.

M. toni seni Lemm. > 1°<-2- ym. 1a South is*q only:

M. punctata Meyen. > 2 < 4 wm. El. A record from pool W2

is only one for any species of this genus away from South Is.

glauca (Ehrenb.) Nag. > 4 < 6 um. Bis South Is. only.

014707

014801

014802

014902

015201

015202

015207

015210

OU521:3

015218

015504

015505

015506

015508

015509

015510

OlS512

015514

Microchaetetenera Thuret. Ep. Generally rare, but large
growths developed in the research station reservoirs and in
the gutters of the solar stills.

Microcoleus chthonoplastes Thuret. Ep, El.

M. sociatus W. et G. S. West. Idjop IHIEG Occasional, and never
abundant.

Microcystis flos-aquae (Wittr.) Kirchn. P. South Is. only,

where it sometimes forms dense blooms.

Nostoc carneum Ag. BP. For most of the wet season present
in only small amounts, but towards the end of wet season
produced large floating colonies in some West Is. and Middle
SEDO SIs

N. commune Vaucher. 1M lehdk A species characteristic of
depressions which receive frequent re-wetting and drying out
during the wet season; not a typical component of vegetation
of pools holding water for long periods.

N. microscopicum Carm.

N. punctiforme (KUtz.) Hariot. re ED ype Ne Occurs mostly
on rocks in pools but occasionally also planktonic in the
vicinity of Chara and Naias.

N. sphaericum Vaucher. Bl. Forms characteristic spherical
colonies up to 20 mm diameter in the Cinq Cases region;
shallow depressions which have collected water may have their
bottom almost covered with these colonies.

N. piscinale Kttz. Po Pe ily. Ac

Oscillatoria amphibia Ag. El, A.
O. angusta Koppe. El, A.
O. animalis Ag. IM, Tek 7Ne Often the first alga to appear

when a pool is wetted, when it forms green sheets over the
mud, which eventually rise as the pool deepens; eaten by
Cardisoma.

O. brevis (Kutz) Gom. Fug nl, ae

O. chalybea (Mertens) Gom.

O. chlorina Kutz. bap ae Ne

O. claricentrosa Gardner. HL, 7Nc

O. geminata Menegh. Idk A INe

8

O15 Sul:
015518
O15 520

0175523

015538

015585
015586
015587
015588
015589
015590
015591
015592
015593
015594
015595
015596
015597
015598
015701
015704
015705

015707

limosa Ag. le 2NG
mougeoti Kutz. iL Ne
proboscoidea Gom. Tal NG

pseudogeminata Schmidle. El, A. Widespread, and recorded
frequently throughout the wet season.

OsecilTatoriay Spi. ,1 2252) 4m. 1 Ba dal An exceptionally large

form for Oscillatoria; although it was suspected of being
a growth stage of Lyngbya majuscula, it was not possible to
establish this. Restricted to South Island, and most
abundant in the more brackish pools, where it forms sheets
which lift of the bottom mud, and are eaten by Cardisoma
and tortoises.

Oscillatoria tenuis Ag. aL AN 7Nc

O.

O.

(O)-

acuta Bruhl et Biswas. El, A.
acuminata Gom. Tl ONG
acutissima Kuff. ill, NG
amphigranulata Woronich. ple Vite
guttulata van Goor. Pele Ate
jasorvensis Vouk. ilps NG
koetlitzi Fritsch. ub Ne
lacustris (Kleb.) Geitler. FA
limnetica Lemm. Ieee lib ep Ne

nigra Vaucher. Wm NG

obscura Bruhl et Biswas. a NG
okeni Ag. Ei Ate

tambi van Goor. eo ANe

Phormidium africanum Gom. Ep, A.

BR.

Ee

Pe

foveolarum Gom. Ep, A.
hendersonii Lemmn. Bp, as

jenkelianum Schmidle. Ep, A.

015708

015769

015770

O157 71.

OU57'7 2

015773

015802

015806

015807

015815

015817

015818

015901

015903

©1593.

016101

016103

016104

016301

016602

016604

016801

016901

P. mucicola Huber-Pestalozzi et Naumann. Ep, A. Frequently
present in the mucilage of other species.

P. bohneri Schmidle. Ep, A.
P. corium Gom. Ep, A.

P. molle Gom. Ep, A.

P. usterii Schmidle. Ep, A.
Px fragile Ep, A.

Plectonema boryanum Gom. i, Ep Ae Frequent both as epilith
and forming a network of filaments among floating algae.

P. gloephilum Borzi. 1 18D) p) Wl 7Ns Abundant, especially in
West Is. pools where it often formed pink globular floating
lumps which sometimes became the dominant alga.

P. gracillimum (Zopf.) Hansg. A liege lb ZANe

P. tomasinianum Bornet. nA lake

P. notatum Schmidle. Ep, EL, A.

P. puteale (Kirchn.) Hansg. op Ile Ne

Pleurocapsa aurantiaca Geitler. HO WILE Nc

P. amethystea Kold. Ep, Hl, A Recorded only from pool Wl.
P. minor Hansg. IMD Wey ANG Widespread and often abundant.
Pseudanabaena catenata Lauterb. la lyAwWANS

P. schmidlei Jaag. ladbp No

P. schmidlei Jaag var. gracilis Skuja. Duly ANE

Radaisia cornua Sauv. Ep.

Schizothrix arenaria (Berk.) Gom. Ep. Occasional in pools on

West Is., rare elsewhere.

S. calcicola (Ag.) Gom. Ep.
Siphononema polonicum Geitler. Ep. Only one record, from W2.
Spirulina subsalsa Oerstd. El, A. Most abundant in South Is.

pools, especially the more brackish ones where it sometimes
became the dominant species.

10

016933

017432

017435

017436

017437

017451

017602

017606

017801

018033

018101

030201

030202

030205

030206

030207

030249

030401

030501

030502

Spirulina gigantea Schmidle > 2 < 4. um.

Synechococcus cedrorum Sauv. > 2 < 4 um. Bg Aw

S. aeruginosus Nag. > 8 < 16 um. lajoy, | tb Ne

S. maior Schroeter. > 16 < 32 um. NG

S. maior Schroeter var. maximum Elenkin et Hollerbach. > 32 um.
Synechococcus < 2um. mL ING

Tolypothrix byssoidea (Berk.) Kirchn. Ep. One of the main

terrestrial algae of Aldabra: pool records probably in large
part due to inwash of terrestrial material.

tT. Gisterta, Kutz. Ep; Bl vA. Ocassional. Truly aquatic
species.
Westiellopsis prolifica Janet. Ep. Restricted to moist rock

just above maximum water level of pools.

Xenococcus kerneri Hansg. puters Rare. Epiphyte of Oedogonium
sp. and Portulaca sp.

Chroococcopsis gigantea Geitler. Wo, IN Especially abundant
in pool W2.

EUGLENOPHYTA

Euglena acus Ehrb. P, Bl, “A. A common species, especially
on West Is.; dominant in W2 in the wet season.

E. gracilis Klebs. Piece A One of the most widespread
species on Aldabra.

E. spirogyra Ehrb. 1 lbs Only 2 records, both from W2.

E. minuta Presc. Pepi rle AC. Widespread, but most abundant
in West Is. pools.

Euglena oxyuris Schmarda. Pye EA aA
Euglena spp., not above. PL oy ee
Lepocinclis ovum (Ehrenb.) Lemm. IDs Abundant in West Is.

pools, rare elsewhere.
Phacus caudatus Htib

Phacus orbicularis Hub. Pe bib An One of the most widespread
species, usually present throughout the wet season.

uma

030601 Trachelonomas hispida (Perty) Stein emend. Deflandre var.

coronata Lemm. 12)

030602 Trachelomonas volvocina Ehrb. pt Formed nearly unialgal
blooms in W5 and W6 for a short period, occasional records
elsewhere.

CRYPTOPHYTA
049931 cryptomonad < 8 wm long blue-green. 12s
049932 cryptomonad, > 8 < 16. um long, blue-green. Pe
049956 cryptomonad, > 16 < 32 um long, brown. De
049959 cryptomonad, > 16 um long, green. P.
PYRROPHYTA

059969 dinoflagellate A. ID Wie Occurs both as epiphyte on

Oedogonium and free-living. Recorded only on West Is.
059970 dinoflagellate B. P. Recorded only towards end of wet
season. Widespread, but never abundant.

XANTiNWOPHYTA
061150 Goniochloris sp. P.

062050 Akanthochloris sp. P, A.

CHRYSOPHYTA
081450 Synura sp. P. Rare.
BACILLARIOPHYTA

Centrales. Records for centric diatoms on South Is. only
090750 Chaetoceros sp.
099151 centric diatom < 8 um diameter. Pi As
099152 centric diatom > 8 < 16 um diameter. Bn NG

Pennales
100250 Amphora spp. El, A. At least two distinct species.
101901 Nitzschia acicularis W. Smith. Pi Ae Only records from

South ise

V2

101904 WN. palea (Kutz.) W. Smith. DP ide, ae By far the most
widespread freshwater diatom.

109950 pennate diatom, other spp. Pe ml eNe

CONJUGATOPHYTA
120269 Closterium sp. NP iy TAL ING Very abundant on West Is., where
it forms conspicuous growths - less abundant on other islands.
120270 Closterium sp. B. EF Bly 2B
120271 Closterium cynthia De. Not. Ps, oe

120272 Closterium sp. D. By a iN

120369 Cosmarium blyttii Wille. Pi, As Widespread, but not forming

large populations. Present throughout wet season.

120370 Cosmarium polygonum (Na&g.) Arch. D> VAY

120306 Cosmarium subcostatum Nordst. IDS Ne

120372 Cosmarium tinctum Ralfs. BAS

120373 Cosmarium trachydermum West and G. S. West. Jer Ne.

121451 Mougeotia sp., cs 7 um P. Only record in water-catchment drum,
Middle Is.

VW22154 \ Spirogyra sp., >. 32 s 48 um, 1 .chloroplast,~ non “episicater
I aisle Found only on South Is.

122469 Staurastrum polymorphum Bréb. P, -A.
122470 Staurastrum sp. Be ioe gNe
122471 Staurastrum sp. on 1D) JING
122472 Staurastrum sp. De P, A.
CHLOROPHYTA
Volvocales
130201 ane globosa Korshik. Pp, Recorded only for one pool

near Anse Cédres.

130402 Chlamydomonas globosa Snow. Pr Widespread, especially on
West Is. and Middle Is.

130450 Chlamydomonas sp. 12h

183)

130901 Eudorina elegans Ehrb. Pee Widespread, especially on
West Is. and Middle Is.

131150 Gonium sp.

131601 Phacotus lenticularis (Ehrb.) Stein. Pie Ax Widespread and
often abundant.

132601 Pleodorina californica Shaw. P. Only one record, from
South Is.

Chlorococcales

140201 Ankistrodesmus acicularis (A. Br.) Korshik. P.

140205 A. braunii Brunnth. P.

140206 A. falcatus (Corda) Ralfs. P.

140207 A. longissimus Lemm. Wille. P.

140208 A. minutissimus Korshik. ie Occasionally the dominant
species jn the plankton. Maximum population recorded =
3.6 x 10 cells/ml on 5 Feb 1973 in pool W4.

140209 A. mucosus Korshik. Pie AS

140210 A. pseudomirabilis Korshik. Ps Often abundant.

140402 Characium ornithocephalum A. Br. Et. Rare.

140403 Characium strictum A. Biss He. Rare, except for one sample
from W5/W6.

140501 Chlorella ellipsoidea Gerneck P, A.

140504 C. vulgaris Beyer. Pp, A.

140508 C. mucosa Korshik. Pi, As

141003 Coelastrum cambricum Arch. var. rugosum Rich. P.

141009 Coelastrum microporum Nag. P.

141501 Dimorphococcus lunatus A. Bie De Recorded only on South
Is., where it occasionally formed blooms.

142102 Golenkinia radiata Chod. Digs Je Recorded only on South Is.,
where it is widespread.

142401 Kentrosphaera bristolae G. M. Smith. lade, Ne

142650 Micractinium sp. Py

231-339 O - '77 - 12

14

142750

142902

142907

142910

143103

143501

143501

143513

143513

143508

143510

143510

143510

144102

144103

144104

144403

144605

144703

150150

150250

S202

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Nautococcus sp. ig De Forms distinctive yellow-green surface

scums on some West Is. pools.
Oocystis crassa Wittr. 1D ING
O. parva We et Gs So Wesit: P, A.
O. pusilla Hansg. Pye As

Pediastrum boryanum P, A.

Scenedesmus acuminatus (Lag.) Chodat. 1 78
S. acuminatus (Lag.) Chodat var. biSeriatus Reinh. Pil:
S. bijugatus (Turp.) Lagerheim. Dy wAe Widespread, and often

quite abundant.

S. bijugatus (Turp.) Lagerheim var. alternans (Reinsch) Hansg.

Deane

Sep Obiiquuss, (Turps) skutzs Pi, A

S. quadricauda (Turp.) Bréb. PReAZ

S. quadricauda (Turp.) Bréb. var. abundans Kirchn. De

S. quadricauda (Turp.) Bréb. var. setosus Kirchn. Pies

Tetraedron incus (Teil.) G. M. Smith. Pa,

T. minimum (A. Br.) Hansg. Digt Ay

T. muticum (A. Br.) Hansg. P, A.

Treubaria triappendiculata Bern. Dips

Lagerheimia genevensis Chodat. P. Recorded only in Jan. 1969.
Sorastrum spinulosum Nag. Be

Ulotrichales &Chaetophorales
Apatococcus sp.

Aphanochaete sp. Biter
Hormidium rivulare Kutz. El.

Endoderma reineckei Schmidle.. Et. Almost restricted to being
epiphytic on Pithophora.

15

152901 Gongrosira debaryana Rabh. Ep, El: Widespread, and usually
abundant.

152949 Gongrosira sp. A. Ep, El:

154435 Stichococcus bacillaris Nag. Ep. Rare.

154603 Trentepohlia iolithus (L.) Wallr. Ep. Widespread. When
pools dry down, this species is visually obvious as a pink
or Orange covering.

154604 T. odorata (Wiggers) Wittrock. Ep.

154734 Ulothrix tenerrima Kutz. Ep. El.

15995 Chaetophorales spp. other El.

160731 Oedogonium sp., < 8 um. INF 1S); Lb ies

160732 Oedogonium sp., > 8 um < 12 um. Be Bp ably, Pht:

160733 Oedogonium sp., > 12 < 16 um. IY, INO II We.

160734 Oedogonium sp., > 16 < 20 um. 1, IO), dll Wiee,

160801 Pithophora oedogonia (Mont.) Wittr. By Et. Widespread, but
seldom abundant. Sometimes used by White-eye as almost
exclusive nest material.

161050 Rhizoclonium sp. By eit.

CHAROPHYTA
170106 Chara zeylanica Kl. ex Willd. var. diaphana (Meyer) R. D. Wood
ANGIOSPERM
251602 Naias graminea Del.
Distribution within 20 pools also studied for water chemistry
The distribution of species within the 20 pools described by
Donaldson and Whitton (1976) is shown in Table 1, while some of the
observations on these pools are summarized in Table 2.
GENERAL OBSERVATIONS
Geographical

Some species were found only on West Is., while others only on
South Is. The following are some examples:

16

West Is. (all restricted *tojyonly onesor vai few" poolls))

Gomphonema aponina (not var. multiplex), Siphononema polonicum,
Euglena spirogyra, dinoflagellate A, Akanthochloris sp., Characium
ornithocephalum, C. strictum.

South Is.

widespread species:

Microcystis flos-aquae, Spirogyra sp., Staurastrum spp., Coelastrum
cambricum, Dimorphocaccus lunatus, Golenkinia radiata, Tetraedron
incus, Naias graminea

locally frequent or abundant species:

Gloeotrichia ghosei, Nostoc sphaericum, Oscillatoria sp. > 32 um,
Chara zeylanica

restricted to only one or a few pools:

Johannesbaptistia pellucida, cryptomonad (> 16 um, long green),
Carteria globosa.

In a few cases the reasons for the distribution of a species being
restricted are fairly clear. For instance Gloeotrichia ghosei occurs
only as an epiphyte on Chara, which latter has itself a restricted
distribution. It is worth commenting that the first three species
listed as restricted to West Is. were found only in W2, the most
eutrophic of any pool studied on Aldabra. The four species listed as
locally frequent or abundant all had the focus of their distribution in
the region of CC9 (Bassin Flamant).

It is difficult to suggest any obvious factor of pool morphology
or chemistry which might inhibit the spread to West Is. of some of the
species widespread on South Is. Microcystis and Spirogyra, which are
visually very obvious, were certainly quite absent from West Is. in both
IGS and 19i7/3\.

Seasonality

Examples of species which produced by far their largest crops
early in the season are: Oscillatoria animalis, Closterium sp. A,
Eudorina elegans, Oedogonium spp., Pithophora oedogonia. The Pithophora
tended to form spores which dropped into the mud, and which presumably
did not germinate until the pool dried out and became re-wetted.
Examples of species which were evident only towards the end of the
season are: Gloeotrichia ghosei, Trachelomonas volvocina, dinoflagellate
Be

Influence of specific environmental factors on pool flora

(a) Pools which were intermittent rather than permanent through the wet
season tended to have a relatively sparse flora, as is shown for W3, W10
and. €€ll in Table 2. On the other hand the total surface area of the
pool appears to have rather little influence on the richness of its
flora, as can be seen by comparing Table 1 of Donaldson and Whitton
(1976) with Table 2 of this paper. For example, W4, the pool for which
the largest number of species were recorded, has a maximum surface area
of only -about 3.5 m2.

7

Macroscopic colonies of Nostoc were almost always obvious in
transient pools. Although these colonies occurred in various morpho-
logical forms, it seems probable that these may all be referred to one
of the three binomials, N. commune, N. microscopicum and N. sphaericum.
The other important algal components of such pools were mostly species
of Lyngbya, Phormidium and Plectonema. Transient pools had few
flagellates, and never became green with floating algal flocs on the
surface.

(b) Although permanent throughout the 1972/73 wet season, two other
pools, W102 and W103, both had a very restricted flora (Table 2). Both
these pools were dominated throughout most of the period by blooms of
Euglena gracilis.

(c) Several species were restricted to the more saline of the pools
included in this survey e.g. Johannesbaptistia pellucida, Spirulina
subsalsa.

Grazing

Planktonic algae sometimes showed a rapid fall in population
density, and in some such cases observations strongly suggested that
grazing by planktonic animals was responsible. For instance, the rapid
development of a population of ostracods was probably responsible for
the destruction of a dense Ankistrodesmus falcatus population in W2.

Larger algae were on many occasions observed to be eaten by crabs.
For instance grazing by Cardisoma carniflex appeared to have a clear
impact on the floating mats of Oscillatoria animalis which developed
early in the season in W2, while this species was frequently observed
to eat Spirogyra and Chara in the Bassin Flamant region. Nostoc
colonies may possibly be less vulnerable to grazing by crabs than these
other species, but Cardisoma carniflex was seen to eat N. commune in W110.
Geocrapsis stormi was seen to eat Oscillatoria animalis and Oedogonium
in Wl.

As already noted by Grubb (1971) tortoises also often eat some of
the larger algae. This occurred for instance, with the flocs of
Oscillatoria sp. > 32 um and Spirogyra in CC5 and CClO. During
observations made on CC1O throughout one day it became clear that
tortoise grazing had an obvious impact on growths of both species.
Tortoises were not seen to eat Nostoc colonies, though these were often
readily available.

Comparison of 1968/69 observations with 1972/73 observations

Although most of the observations reported in this paper were made
during the 1972/73 survey floristic notes were also made during the
1968/69 survey. The former survey was much less extensive both in time
and in number of pools studied, and only about half the number of species
listed for 1972/73 were found *in 1968/69. Three species were recorded
only in the earlier survey: Wollea bharadwajae, ?Lobomonas sp.,
Lagerheimia genevensis. (Of these, the material recorded as Wollea

18

bharadwajae may possibly have been only a form of Anabaena ambigua
growing in the Cing Cases region).

Pools Wl, Tl and T2 were studied in both surveys, and the floras
recorded in both surveys tended to be rather similar. In Tl, five
species were recorded in the two samplings made in 1968/69 but not in the
three of 1972/73, while 11 species were recorded in 1972/73 but not
1968/69. However almost as many differences were recorded between
indvidual surveys made within either of the two seasons.

ACKNOWLEDGEMENTS

We are most grateful to the Royal Society and the Natural
Environment Research Council for financial support, and to Mrs. V. Evans
for typing this paper.

REFERENCES
Desikachary, T. V. 1959\. = Cyanophy ta’. Indian Council Agricultural
Research.
Donaldson, A. and Whitton, B. A. 1976. Chemistry of freshwater pools

on Aldabra Atoll. Atoll Research Bulletin, this issue.

Geitler, L. 1932. Cyanophyceae. In: Rabenhorst's Kryptogamen-Flora,
Vole a4): Akademische Verlagsgesellschaft, Leipzig.

Golltubice,. 7S. 1967. Algenvegetation der Felsen. Ein d6kologische
Algenstudie im dinarischen Karstgebiet. Die Binnengewadsser
23 MBS E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart,
Germany.

Grubb, P. aS) 7/ake The growth, ecology and population structure of
giant tortoises on Aladabra. . Phil. Trans. R. Soc: B 260: 327-372.

McKenzie, K. G. OW ale Entomostraca of Aldabra, with special
reference to the genus Heterocypris (Crustacea, Ostracoda).
Phil. Trans. R. Soc. B 260: 257-298.

Starmach, K. 1966. Cyanophyta-Sinice, Glaucophyta-Glaukofity
(Polish): Flora slodkowodna Polski, Pol. Acad. Sci. 2; 1-807.

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26

Summary of data concerning distribution of algae within

Table 2.

the 20 pools chosen for most detailed study.

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ATOLL RESEARCH BULLETIN
NO. 216

TERRESTRIAL AND FRESHWATER ALGAE
OF THREE WESTERN INDIAN OCEAN ISLANDS
(ASTOVE, FARQUHAR AND ST. PIERRE)

by B. A. Whitton and A. Donaldson

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

May 1977

TERRESTRIAL AND FRESHWATER ALGAE
OF THREE WESTERN INDIAN OCEAN ISLANDS
(ASTOVE, FARQUHAR AND ST. PIERRE)

by B. A. Whitton and A. Donaldson !

Introduction

In contrast to the many studies of the marine algae of tropical
limestone islands, the terrestrial and freshwater algae have received
little attention. A preliminary account for Aldabra (Whitton, 1969)
is apparently the only description of such algae from the Western
Indian Ocean. This account emphasized the abundance of blue-green
algae, and it was suggested that these might be of considerable ecologi-
cal significance. The following note provides comparative observations
made during brief visits (by B.A.W.) to Astove (10°6'S, 47°45'E),
Harguhar Atoll (1O°1L1l"S) and SE. Pierre (9°19"S, 50°43"E during
Matkehr—s Apia 1: 19173

The ecology and vascular plant species of Astove and Farquhar
Atoll have recently been described in some detail (Astove: Bayne et
al., 1970; Fosberg & Renvoize, 1970b; Farquhar Atoll: Stoddart &
Poore, 1970; Fosberg & Renvoize, 1970a), while the data known for
St. Pierre have been summarized by Vesey-Fitzgerald (1942), Stoddart
(1967) and Piggott (1969). The reader is referred to these accounts
for general environmental information. Samples were either studied
while still fresh, or dried down and re-wetted before viewing. In
addition a few samples from each island were used as a source of
inocula for subsequent growth in the laboratory using a range of media.

Observations

As the visits to each of these islands lasted only a few hours,
the accounts are obviously very incomplete. Nevertheless it was
rapidly obvious that while all three islands resembled Aldabra (Whitton,
1969) in having blue-green algae constituting the bulk of the algal
vegetation, the overall abundance varied markedly between them, being
most abundant on St. Pierre and least abundant on Farquhar. The
species records are summarized in Table lI.

1 Department of Botany, University of Durham, Durham, England

(Manuscript received June 1974--Eds.)

231-339 O - '77 - 13

Astove

(Visited 3 April 1973). The algal vegetation of the western side
of this elevated atoll (the only region visited) rather closely resem-
bled that of West Island, Aldabra. The rocks had a general cover of
Tolypothrix byssoidea, with Gloeocapsa sanguinea almost always closely
associated with it. Tolypothrix byssoidea also sometimes formed small
black patches on bare sand, and in such cases the principal alga
associated with it was Schizothrix arenaria. Conspicuous growths of
(typical) Nostoc commune colonies were sometimes present on the lime-
stone, while N. commune var. flagelliforme was frequent in the coconut
plantations. Small freshwater pools were quite common, especially
towards the lagoon. These were mostly 2-4 m diameter and reached (at
the time of visit) up to 0.6 m deep. These had an attached algal
cover, a plankton, and usually also a partial surface cover of flocs or

filaments buoyant as a result of associated oxygen bubbles. Species
almost always present at the sides of these pools included: Calothrix
parietina, Lyngbya spp., Gongrosira sp. and Oedogonium sp. The plank-

ton consisted of varying proportions of blue-green algae (including
Anabaena variabilis, Phacotus lenticularis, Chlorella sp., Scenedesmus
quadricauda, Tetraedron minimum and Euglena sp. Floating flocs mostly
consisted of Closterium sp., Oedogonium sp., Spirogyra sp.

Only one species was recorded on Astove which has so far (the

authors, unpublished) not been found on Aldabra. This was Stigonema
hormoides, which was locally frequent in an area about 1 km south of
the settlement, occuring on shaded concavities on rocks. As this

species is easily visible to the naked eye, and relatively distinctive,
its presence on Astove may represent a genuine difference from Aldabra.

Farguhar Atoll

(Visited 16 March and 6 April 1973; North Island only). The
algal vegetation was relatively inconspicuous. This was no doubt
largely due to the general cover of coconut trees over most of the
island, and to the rarity of standing freshwater. Nevertheless it
seems possible that these are not the only explanations since typical
Nostoc commune colonies were not recorded anywhere, and N. commune var.
flagelliforme occurred only very rarely. Exposed rock surfaces did
however resemble those of Astove in having a thin cover of Tolypothrix
byssoidea and Gloeocapsa sanguinea.

One small area of freshwater was found, apparently identical with
the marshy area referred to by Stoddart and Poore (1970, p.16). Its
general appearance and algal species composition was rather similar to
that of a saline pool, so it seems possible that it may not remain fresh
water throughout the year. This pool overlay a dark-coloured fine silt,
and lacked exposed rock surfaces. The only algae found were blue-green
algae, these mostly occurring as floating granules or mucilaginous lumps.
Such flocs taken from deeper layers in the pool contained many purple
photosynthetic bacteria, which together with some forms of Aphanocapsa
were often sufficient to colour the granules pink. The blue-green
algae included: Aphanocopsa spp., Chroococcus minutus, Synechococcus

elongatus, Lyngbya spp.
St. Pierre

(Visited 17 March 1973). This island is sufficiently small (417
acres) for an impression of the whole area to have been obtained during
the visit, which occurred a year after mining for guano had finally been
completed.

Observations on the algal flora of St. Pierre are of particular
interest in view of the high levels of phosphate likely still to be
available in many microhabitats. In spite of the large literature
published in recent years on the effect of phosphate eutrophication in
freshwaters, there have been no previous accounts of the algae of
naturally occurring very phosphate-rich environments.

At the time of the visit there was much standing water, with
hundreds of small freshwater pools. About one-sixth of the island is
covered with young trees of Casuarina (up to 4m high). In addition
there were about 20 much older trees of this species and 5 coconut
palms. The open part of the island is deeply dissected by thousands
of crevices. The solitary Pisonia grandis found in 1960 by Piggott
(1969) is apparently dead, and the angiosperm vegetation of this region
consisted of Ipomoea and various herbs, especially Stachytarpheta
jamaicensis, Turnera ulmifolia, Vernonia cinerea and Lippia nodiflora
(all names based on field observations only).

Both blue-green algae and lichens were very conspicuous. As with
the other islands, the more exposed parts of the rock were dominated by
Tolypothrix byssoidea. Epilithic lichens were much more abundant here
than on any of the other islands (including Aldabra). Crustose lichens
were especially abundant on the lips of the rocks adjacent to fissures.
Small blue-green alga containing lichens were also often evident,
especially where the covering of free-living epilithic blue-green algae
was particularly thick. Recent observations have shown that the latter
phenomenon occurs also on Aldabra.

No large Nostoc commune colonies were seen, but a thin film of
Nostoc was common in rock concavities. The small pools were sometimes
completely filled by growths of Rhizoclonium, mixed with numerous small
blue-green algal colonies. In the young Casuarina woodland,
Oscillatoria animalis was abundant. Samples from this region cultured
on agar with mineral medium developed, in addition to algal colonies, a
particularly rapid and vigorous growth of actinomycetes.

Discussion

Although Table I is merely a preliminary list of the algal floras
of these atolls, nevertheless they probably give a fair indication of
the relative proportions of the main algal groups represented. Blue-
green algae are by far the most important group, and free-living
eukaryotic algae are largely restricted to the pools. Tolypothrix

byssoidea was the most widespread alga on all three islands, being

ubiquitous on rocks and widespread on sand. This species has a
similar distribution both on Aldabra (the authors, unpublished) and the
Bikini wsVands! = (Gaylor, = 1950) The Indian Ocean islands do however

show a marked contrast to the latter in that neither Porphyrosiphon
nor Symploca have so far been found on them, whereas Taylor reports
them to be major constituents of the vegetation of sand on Bikini.

Acknowledgements

This survey was carried out while the authors were in receipt of a
grant from the Royal Society and Natural Environment Research Council
for research on Aldabra. The authors are most grateful to both these

bodies and to Dr. D. R. Stoddart for a guide to the literature about
St. Pierre.

TABLE 1. Terrestrial and freshwater algal floras

Astove Farquhar St. Pierre

MYXOPHYTA
Chroococcales
Aphanocapsa fusco-lutea Hansg. + + +
A. grevillei (Hass.) Rabenh. +
A. muscicola(Menegh.) Wille + + +
Aphanothece castagneii (Bréb.) Rabenh. +
A. pallida (Kltitz.) Rabenh. + + +
Gloeocapsa magma (Bréb.) Hollerbach.
emend. + +
G. Sanguinea (Ag.) Kitz. + + +
Gloeothece rupestris (Lyngbye) Barnet +
Chroococcus minutus (Ktitz.) Nag. +
C. turgidus (Ktitz.) Nag. + + +
C. turicensis (Ndag.) Hansg. + + +
Synechococcus aeruginosus Nag. +
S. elongatus Nag.
Chamaesiphonales
Pleurocapsa minor Hansg. em. Geitler + ot
Hyella fontana Huber et Jadin + +
Hormogonales
Hapalosiphon welwitschii W. et G.S. West +
Westiellopsis prolifica Janet + + +
Stigonema hormoides (KlUtz.) Born.
et Flah. oF +
Calothrix marchica Lemm. + +
C. parietina Thuret +
Gloeotrichia sp. +
Microchaete tenera Thuret +
Plectonema boryanum Gomont ap + +
P. gloeophilum Borzi +
P. gracillimum (Zopf.) Hansg. +
P. notatum Schmidle +
P. terebrans Born. et Flah. ap
Tolypothrix byssoidea (Hass.) Kirchn. + + +
ne conus, KUeZ + + +
Cylindrospermum muscicola Ktitz. +
Nostoc carneum Ag. + a +
N. commune Vaucher ar af +
N. commune Vaucher var. flagelliforme
(Berk et Curtis) Born. et Flah. + +

N. microscopicum Carm. sec. Harvey,
in Hook + + +

6
TABLE 1. (Cont=)
Astove Farquhar St. Pierre

N. paludosum Kitz.

N. piscinale Kltz. + +
N. punctiforme (Ktitz.) Hariot

N. piscinale Ktitz. + +
N. sphaericum Vaucher

Anabaena variabilis Klutz +
Oscillatoria animalis Ag. +

O. pseudogeminata G. Schmid

O. tenuis Ag.

Phormidium africanum Lemm.

P. angustissimum W. et G. S. West

Pp oeecnue a
Lyngbya allorgei Frémy
L. digueti Gom. + +
L. erebi W. et G. S. West +
L. martensiana Menegh ; +

L. nordgardhii Wille

L. perelegans Lemm +
Schizothrix arenaria (Berk.) Gom +

S. calcicola (Ag.) Gom +

+
+

+
+
t+etttteteeteeteetett

CONJUGATOPHYTA

Spirogyra sp.

Closterium sp.

Cosmarium subcostatum Nordst.
Cosmarium sp.

+++ 4+

CHLOROPHYTA

Phacotus lenticularis (Ehr.) Stein
Ankistrodesmus minutissimus Korschik.
Chlorella sp.

Scenedesmus quadricauda (Turp.) Bréb.
Tetraedron minimum (A.Br.) Hansg.
Gongrosira sp.

Trentepohlia iolithus (L.) Wallroth +

++ tet

Oedogonium sp. (? p.) +r +
Rhizoclonium sp. c. 40 um broad +
EUGLENOPHYTA
Euglena sp. c. 30 um long iF
Euglena sp. c. 75 um long a
XANTHOPHYTA
+

Akanthochloris sp.

TABLE Ls. (Conte)

Astove Farquhar St. Pierre

BACILLARIOPHYTA
Nitzschia palea (Klitz.) W. Smith +
Nitzschia sp. +

Total 43 23 Sil

References

Bayne), (Ga dre, aCogani,, “BH, wbt among, vA. Wier, EecaZ era. aire yn GIsUOlyneeare
Hutson, As; Poore, M2 BaD. ;) stoddart; Deak. sande tay lomnadmuDE
1970 Geography and ecology of Astove. Atoll. Res. Bulls 36:
88-99%.

Desikachary, T. V. 1959 Cyanophyta. 686 pp. Indian Council of
Agricultural Research, New Delhi, India.

Fosberg, F. R. and Renvoize, S. A. 1970a Plants of Farquhar Atoll.
Atoll Res). Bullet! 363) 2 i= Sys

Fosberg, F. R. and Renvoize, S. A. 1970b Plants of Astove Island.
Atoll Resi Bull. 336s a VOM aia.

Geitler, L. 1932 Cyanophyceae. In: Rabenhorst's Kryptogamen-Flora,
Volk way Akademische Verlagsgesellschaft, Leipzig.

Piggott, C. J. 1969 A Report on a-Visit to the Outer Islands of
Seychelles between October and November ]960. Land Resources
Division of Overseas Surveys, Tolworth, Surrey, England.

Stoddart, D. R. 1967 Summary of the ecology of coral islands north of
Madagascar (excluding Aldabra). Atoll Res’. Bull). VUSs 5S8—ol.

Stoddart, D. R. and Poore, M. E. D. 1970 Geography and ecology of
Farquhar Atoll. Atoll Resi! (Bulls (36: \7=26:

Taylor, W. R. 1950 Plants of Bikini and other northern Marshall Islands.
Ann Arbor, Michigan: University Press.

Vesey-Fitzgerald, L. D. E. F. 1942 Further studies on the vegetation on
islands in the Indian Ocean. Ue TECOUS, SOc) peal GO.

Whitton, B. A. 1969 Terrestrial and freshwater algae of Aldabra.
Rhine rams oo (Rien SOCsy Onde Be ZOOs i249 —2Obis

ATOLL RESEARCH BULLETIN
NO. 217

TERRESTRIAL AND SWAMP ALGAE FROM THREE ISLANDS
IN THE CHAGOS ARCHIPELAGO, INDIAN OCEAN

by B. A. Whitton, A. Donaldson, D. J. Bellamy and C. Sheppard

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

May 1977

TERRESTRIAL AND SWAMP ALGAE FROM THREE ISLANDS
IN THE CHAGOS ARCHIPELAGO, INDIAN OCEAN

by B. A. Whitton, A. Donaldson, D. J. Bellamy and C. Sheppard !

INTRODUCTION

During visits to the Chagos group of islands by two of the
AlLehORS (OD) Wie Be an wWantlanyielOHS); Dewi. E. Cen Se eErOnnwanuary to
March 1975), several collections were made of conspicuous algal growths
from terrestrial and swamp habitats. As these growths provide some
interesting similarities and differences from those reported previously
from other islands in the Indian Ocean (Donaldson & Whitton, 1976;
Whitton & Donaldson, 1976), a brief account of their floristic
composition is given here.

The Chagos Islands (6°S, 71°W) form part of the British Indian
Ocean Territory. The samples described here were taken from three of
these islands, Egmont, Eagle and Danger. Egmont Is. is one of the
atolls which lie around the periphery of the Chagos archipelago, while
Eagle Is. and Danger Is. may be considered as being part of the rim of
a very large former atoll, now almost entirely submerged. No rainfall
records exist for any of these three islands, but stations have at times
existed on three of the other peripheral islands, Diego Garcia, Salomon
Is. and Peros Banhos, the most recent being those from Peros Banhos.
Here it is estimated that during 1950-1966 there occurred a mean annual
rainfall of 3999 mm (Stoddart, 1971). Seasonal changes in
precipitation were relatively small, with the lowest values being in
May and June.

METHODS

Samples were dried soon after collection and then sealed in
individual packets. They were viewed after re-wetting with distilled
water. In certain cases simple cultural techniques were used either
to help confirm identifications or to demonstrate the presence of
further species. Species noted only after culture are noted below.

Department of Botany, University of Durham, Durham, England.
(Manuscript received 8 July 1975 --Eds.)

The species categories used in the various tables of results are
based on these used in an algal computer recording system held at
Durham, and described further in Donaldson and Whitton (1976). The
size ranges given are standard ranges used in our system; the full
range of these widths have not necessarily been found in these samples
from the Chagos Is.

RESULTS
Eagle Is.

Almost 50 different samples were collected here during 1975.
Three quite different types of algal community were represented, two
from clearings in the coconut plantations, and two from Typha swamp
regions. Their species compositions are compared in Table 3.

A. Thin olive-brown sheets spreading across debris overlying sand in
a clearing; dominated by Nostoc commune.

B. Flattened to hemispherical lumps, reaching a maximum height of
9mm, lying over grey or pale brown sand in coconut clearings;
frequent. The main blue-green algal layer is only 1 mm thick,
the remainder of the lumps being largely filled with sand.
However there is usually a zone about 2 mm thick below the main
blue-green algal layer which has a faint green tinge, this being
due to moss protonema and Lyngbya martensiana. Some of the
filaments of Tolypothrix byssoidea are moderately calcified,
while those of Lyngbya martensiana and Schizothrix arenaria are
also sometimes calcified.

(en Algal layer forming patches in more or less open areas within a
swamp community dominated by Typha sp. and Eleocharis sp. These
algal layers are probably completely submerged after heavy
rainfall, but at the time of collection formed a dark grey slimy
layer overlying a dark humus; moss shoots are also mixed with the
algal layer. On drying out, the algal layer turns to a grey-
green colour, as a result of the highly calcified layer around
the sheaths of the dominant alga, Tolypothrix byssoidea.

Die + Continuous layer of blue-green algae overlying dense brown
fibrous mass, near a mangrove swamp. Algal layer about 3 mm
thick, gelatinous, crumbly, irregular on surface. Obvious
vertical zonation (Table 4). Some, but not all, of the dominant
alga, Tolypothrix byssoidea, is calcified.

Egmont

Samples of mat were taken from a barachois community both in 1973
and: .19)75.. The water associated with this community presumably varies
considerably in salinity according to the pattern of rainfall in the
previous few weeks; at the time of study in 1975 it was slightly
brackish (Na, 236 mg 1).

The detailed composition of the mat is somewhat variable, but
always had a layer dominated by Oscillatoriaceae overlying a layer of
purple photosynthetic bacteria. The single sample of mat returned in
1973 was much thicker than any of the 10 samples returned in 1975, the
photosynthetic layer reaching a depth of 10 mm and with obvious vertical
zonations of the various blue-green algae present. Details of this
mat profile are summarized in Table l. The composition of this mat
rather closely resembles one already described for rock pools on the
Florida Keys, U.S.A., even though these latter are intertidal (Fischer
and Golubit¢, in Ginsburg, 1967) and termed by Golubié (1973) a
"stratified community mat in intertidal rock pool'.

The samples returned in 1975 were much thinner, with the blue-
green algal layer about 3-4 mm thick. A consolidated species list is
given in Table 2.

Danger Is.

All terrestrial algal crusts were dominated by Tolypothrix
byssoidea. Other species represented are listed in Table 5.

DISCUSSION

All these terrestrial and swamp algal communities are dominated by
blue-green algae, and only in the Typha - Eleocharis swamp on Eagle Is.
are eukaryotic algae represented by more than a few scattered cells.
Three out of the six different types of algal community described from
these three islands are dominated by Tolypothrix byssoidea, though the
structure and composition of all three examples are each rather
different. The extent to which this alga becomes calcified varies
markedly, with the most highly calcified material occurring in the
Typha - Eleocharis swamp. During the more detailed studies of the
authors on Aldabra (Donaldson & Whitton, 1976; A. D. and B. A. W.,
unpublished data), no sample of Tolypothrix byssoidea has ever been
found which approaches this swamp community in extent of calcification.

REFERENCES

Donaldson, A. and Whitton, B. A. 1976. Algal flora of freshwater
pools on Aldabra Atoll. Atoll Research Bulletin, this issue.

Ginsburg, R. N. 1967. Stromatolites. Science, N. Y. 157: 339-440.

Golubic, S; ONS The relationship between blue-green algae and
carbonate deposits. INNS), Carae, INS Gs Re Mobley.) ES NG Gaels)
The biology of Blue-green Algae. 676 pp., pp. 434-472.
Blackwell Scientific Publications, Oxford.

Sieeelolenceie, Ie. Is) ALS)\7/be Rainfall on Indian Ocean Ocean coral islands.
Atoll Research Bulletin 147: 1-21.

4

Whitton, B. A. and Donaldson, A. 1976. Terrestrial and freshwater
algae of three Western Indian Ocean islands. Atoll Research
Bulletin, =thismus sues

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Table 2. Consolidated list of algal species from barachois mat
community, Egmont Is.

010531 Aphanocapsa fusco-lutea Hansg. > 1 < 2 yum
010532 A. montana Cramer > 2 < 4 um

010632 Aphanothece sp. > 2 < 4 um

OLO902. Calothrix braunii Born. et Pillah.

010911 C. parietina Thuret

011538 Chroococcus minutus (Klitz.) Nag. > 4 < 6 um, non-lamellate
sheath

O11534) \€. turgidus (Kiltz.) (Nag. > 8 < 16 um, lamellate sheath
011540 C. turicensis (Nag.) Hansg. * 8 < 16 m, non-lamellate sheath
012201 Entophysalis granulosa Ktitz.

013602 Hyella fontana Htiber et Jadin

014205 Lyngbya digueti Gom.

015202 Nostoc commune Vaucher

015702 Phormidium angustissimum W. et G.S. West

015704 P. foveolarum Gom.

015802 Plectonema boryanum Gom.

015931 Pleurocapsa minor Hansg. < 4 um

015932" Pleurecapsa)sp.<> 4 <.6 jm

016631 Schizothix sp. < 1 um

016931 Spirulina subtillissima < 1 pm

Table 3. Species composition of four different algal communities,
Eagle Is. The details given are generalizations based on a range of
samples, and for composites where vertical zonation is apparent.
Organisms seen only after cultures are indicated by +, rather than a
frequency score.

A BEC Dp)
010531 Aphanocapsa fusco-lutea > 1 < 2 um 2
010902 Calothrix braunii its
010950 Calothrix sp. +
011534 Chroococcus turgidus > 8 < 16 um,
lamellate sheath ay ile
011541 C. spelaeus > 16 < 32 um non-lamellate sheaths 2
012640 Gloeocapsa dermachroa > 2 < 4 um sheaths
: yellow-brown Ds
012645 Gloeocapsa sp. > 2 < 4 um, sheath violet iL
012732 Gloeothece palea > 2 < 4 um LE 5
012733 G. rupestris > 4 < 6 um 3
013107 Hapalosiphon welwitschii + +
014202 Lyngbya allorgei i} tt
014211 L.. martensiana Bets
014252 Lyngbya sp. > 1 < 2 um 2
014801 Microcoleus chthonoplastes ys eS
015202 Nostoc commune 5 Be 2
015218 WN. piscinale +
015211 WN. punctiforme 2:
015704 Phormidium foveolatum + +
015511 Oscillatoria claricentrosa 2
015802 Plectonema boryanum ch:
015932 Pleurocapsa sp. < 4 um iL
016602 Schizothrix arenaria 3 eo eS
017601 Tolypothrix byssoidea 2) 35> bee US
017606 TT. distorta AF ee 2
062050 Akanthochloris sp. iE
120250 Closterium sp. i
122150 Spirogyra sp. 2
palmelloid green alga (c. 13 um diameter) 2
160732 Oedogonium sp. > 8 < 12 um il
160801 Pithophora oedogonia 2
239950 unknown moss (es) 2S

231-339 O -'77 - 14

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Table 5. Species list for algal crusts from Danger Is.

Aphanocapsa montana > 2 < 4 um

Gloeocapsa dermochroa > 2 < 4 um, sheath yellow-brown
G. kutzingiana > 4 < 6 um, sheath yellow-brown
Lyngbya martensiana

Microcoleus chthonoplastes

Nostoc commune

Plectonema boryanum

P. notatum

Pleurocapsa Sp. < 4 um

Schizothrix sp. (c. 4.5 um wide)

Tolypothrix byssoidea

= 1” " Ried os ca
SOLUS SE ce De

ATOLL RESEARCH BULLETIN
NO. 218

THE HOLOCENE REEF SYSTEMS
OF EASTERN MARTINIQUE, FRENCH WEST INDIES

by Walter H. Adey, Patricia J. Adey, Randolph Burke
and Leslie Kaufman

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

May 1977

a leat
y Ts :

THE HOLOCENE REEF SYSTEMS
OF EASTERN MARTINIQUE, FRENCH WEST INDIES

by Walter H. Adey,! Patricia J. Adey, Randolph Burke 2
and Leslie Kaufman 2

ABSTRACT

The windward east coast of Martinique ranges from high, eroding
Pleistocene volcanic cliffs in the north, to relatively low, early
Tertiary, embayed shores partly capped by Pleistocene limestones, in the
southeast. An extensive shallow carbonate shelf is developed off the
central part of this coast. The types of Holocene reefs found on
Martinique correspond to the pattern that is characteristic for Lesser
Antillean islands: the volcanic northern coast is barren; massive algal
ridges have developed on benches cut in the limestones and other early
Tertiary rocks of the southeast coast; on the central east coast, where
the shelf is well developed, an extensive and double bank barrier system
has formed. Parts of the bank barrier reef system are reaching the
mature stage typical of high energy coasts and have algal ridge caps
built primarily by crustose corallines.

The bank barrier reefs of Martinique are unusual in that the corals
have been replaced on the crests by a carbonate pavement covered with
dense, diverse stands of fleshy algae, Sargassum spp. being especially
common. The term "fleshy algal pavement" has been applied to this type
of reef. Based on upstream-downstream measurements of dissolved oxygen,
a winter productivity level of 33 g07/m2/day indicates that these ex-
tensive reefs with their dense plant cover may be the most productive
in the eastern Caribbean.

Though transitional and less extensive, reefs with fleshy algal
pavements also occur on Grenada, St. Lucia, southeastern Basse Terre,
and Nevis. All of these island shores are locally characterized by high
sea water turbidities, presumably resulting from run-off, indicative of
eutrophic conditions. However, where algal pavements are well developed,

1/7 Smithsonian Institution, Washington, D.C. 20560
2/ University of South Florida, St. Petersburg, Florida 33701
3/ Johns Hopkins University, Baltimore, Maryland 21218

Dire

the reef surface also generally lacks extensive large scale porosity and
large populations of Diadema antillarum. Large parrot fish of the genus
Scarus are also very poorly represented, although this may result from a
general lack of filamentous algal turfs normally characterizing their
diet.

Based on Cl4 dating of cores, we have concluded that about 600 years
B.P., the Martinique inner bank barriers shifted from Acropora palmata
to fleshy algal domination. We postulate that the surfaces of these
reefs developed into pavements by the filling of the interstitial reef
spaces by either gradual or catastrophic means. This eliminated the
large scale porosity required by algal grazerssuch as Diadema (and per-
haps some parrot fish) in this high energy area.

In the reefs of the Grenadines where the surrounding waters are
quite clear, Similar carbonate pavements occur in the high energy areas,
but lack a fleshy algal cover. We conclude that flat, featureless car-
bonate surfaces in areas of strong wave or current energy are conducive
to the development of stands of fleshy algae. Especially where nutrient
levels are high, massive stands of fleshy algae can become dominant on
the reefs.

INTRODUCTION

Recent studies of Caribbean coral reefs, especially those of Adey
and Burke, 1976, Macintyre and Glynn, 1974, and Adey, 1975, have demon-
strated the existence of extensive reef structures with considerable
accumulations of Holocene carbonate. In addition, massive algal ridges
have been found in numerous localities throughout the Lesser Antilles
(see Adey and Burke, 1.c.). While there are undoubtedly fewer species
of reef organisms in the West Indies than in the Indo-Pacific, this is
probably largely due to the great difference in geographical area and to
the relative isolation of the Caribbean during the Neogene. There is no
field evidence for climatic inhibition of the rate of Holocene or earlier
carbonate reef build-up in the Caribbean, a conclusion that correlates
with new evidence for long term stability of Caribbean sea temperatures
(Clamap,- 19,76).

Research on living Lesser Antillean reefs has not been extensive.
Prior to the recent work out of West Indies Laboratory, St. Croix (see
W.I.L. Special Publications, beginning 1972), and the reconnaissance
study of Lewis (1975) in the Grenadines, most studies had been done in
areas of very limited Holocene reef development. Areas such as the east
coast of Martinique had been considered devoid of reefs, in part due to
the fact that the structures found in the area did not correspond to the
popular conception of what a reef should be. The waters around Martini-
que tend to be turbid, with visibilities ranging from three to twelve
meters, depending upon wave activity. Coral growth on the reefs is re-
stricted, and has commonly been replaced on the crest by carbonate
pavement covered with a heavy, diverse growth of fleshy algae, which we
refer to as "fleshy algal pavement". Fish biomass is reduced as compared

316

to that found on the rich Caribbean Acropora palmata reefs. In spite of
their unusual morphology, it would appear that Martinigque's Holocene
carbonate reef systems are among the most massive and extensive in the
eastern Caribbean and biologically they may be among the most productive
in the West Indies.

In the island arc extending from the Virgin Islands to Grenada,
approximately eight hundred kilometers of windward coast is available
for potential reef development (not including Barbados). As part of the
present study, a reconnaissance survey of this coast has been made by
boat and plane (Adey and Burke, 1976). As a result of these observations,
the windward island shores of the Lesser Antillean island arc are found
to fall into four categories: (1) barren, terrigenous (volcanic) sedi-
ments, 30% of total shoreline; (2) barren, non-volcanic, 14%; (3) bank
barrier reefs on carbonate shelves at depths of 12-25 m, 38%; (4) reefs
on shallow (<10 m) pre-Holocene benches, 18%. Approximately 52% of the
bank barrier reef shores are dominated by fleshy algal pavement reefs
such as found on Martinique, 36% by Acropora palmata reefs, and 12% by
an A. palmata-algal ridge transition type characterized by crustose
corallines and Millepora. Algal ridges in all stages of development
occur in some areas as scattered caps on the bank barriers. Shallow
benches, which occur primarily on the shores of limestone islands, are
dominated by high algal ridges in exposed areas, and in more protected
areas by A. palmata and coralline-Millepora reefs. In category (2)
listed above are found some limestone island shores which are steeply
cliffed, unbenched and without shallow reefs (5% of total), as well as
some shores where the carbonate shelves are deep (>30 m) and also lack
well developed reefs (9% of total).

Adey and Burke (1976) discuss the distribution of these reef types
throughout the Lesser Antilles. These same authors (in ms.) present a
model for control of Holocene reef development by the height of pre-
Holocene shelves or benches, wave energy and water turbidity. The young
northeastern coast of Martinique is characterized by steep cliffs cut in
volcanic sediments, a narrow steeply slooping shelf and very minimal reef
development. The older eastern and southeastern coasts, on the other
hand, have a well developed shelf and are rich in Holocene reefs.
Martinique is located in a peak of trade wind energy and constancy in
the eastern Caribbean, and an index of wind strength and constancy shows
a reduction to the north as well as the south (Adey and Burke, in ms.).
East to southeastern Martinique is characterized primarily by bank bar-
rier reefs with well developed fleshy algal stands on their crests. The
southernmost part of the coast is dominated by a bench algal ridge sys-
tem. These areas and their reef biotas are described below.

All of the authors listed took part in both the field work and the
preparation of this paper. However the various sections were largely
the responsibility of the following individuals: geological structure
and history - Adey and Burke; corals - Burke; algae and productivity -
Adey and Adey; fish - Kaufman.

DESCRIPTION OF THE REEFS

The following description includes the reefs from Presqu'ile de la
Caravelle to Baie des Anglais (Fig. 1). It is based on three aerial re-
connaissance flights, extensive examination, including coring and diving,
at four sites (Vauclin Point reef, Cayes de Pinsonelle algal ridge,

Islet de Ramville reef and Islet Lezard algal ridge) and numerous short
visits to other points.

The reef between Passe du Vauclin and Cap Ferré has not been ex-
plored by diving. Aerial reconnaissance indicates that it is dominated
by a fleshy algal pavement and with coralline-Millepora mounds on the
crest in the north and a high algal ridge in the south, northeast of
Point Macré. Thus, it is intermediate in character between the inner and
outer systems that will be described below. Aerial reconnaissance also
indicates that fleshy algal dominated reefs, with scattered coralline-
Millepora mounds, exist near shore to the north of Caravelle, as far as
Pte. Lahoussaye. This is apparently equivalent to the inner system to
the south. Further to seaward, extending northwestward of Caravelle, a
deeper ridge-like structure also occurs. This could be either a develo-
ping Holocene bank structure or a Pleistocene equivalent of the shelf
edge features. We have not examined either of these areas in the water.

The major Holocene reefs of Martinique lie on the east coast be-
tween Presqu'ile de la Caravelle and the southeastern corner of the
island. North of Passe du Vauclin they are arranged in a double (inner
and outer) bank-barrier system. South of Cap Ferré, a single bench
algal ridge system is predominant.

The double reef system, from Passe du Vauclin to Presqu'ile de la
Caravelle, morphologically appears to be a set of bank-barriers developed
during the middle Holocene atop the carbonate island shelf (Fig. 2).

Bank barriers, which are common on many eastern Caribbean carbonate
shelves, where depths are 15-25 m, typically consist of a coral or coral-
line framework cap (5-10 m thick) overlying a sand, rubble and head
coral base. Adey and Burke (1976) suggest that these formed as "bars",
largely during mid-Holocene, by coral head trapping of carbonate sand
and rubble. Although the inner reef group in Martinique, which extends
irregularly from Pte. du Vauclin to Baie du Galion, is rather atypical

in form, paralleling the shore in a manner that is more characteristic
of "bench" reefs, a series of holes bored in the Vauclin Point reef con-
sistently penetrated the lower limit of the Acropora palmata facies at
depths begween 4-5 meters (Fig. 3), below which carbonate sand with
occasional coral rubble is the major component. In the deepest hole in
Vauclin Point reef drilling character abruptly became very hard at 14 m.
We were unable to obtain core at this point, but this suggests a basement
contact. It may be that the particular orientation and embayed nature
of the shoreline has allowed the formation of a bank, based upon a bench.
Such circumstances would require the bench to be relatively deep, at
least 14 meters. Profiles across the Martinique shelf suggest that it
shoals landward to such a degree that, if such a bench is present, it
does not rise far above the shelf proper. These inner reefs may be bank
barriers formed against an abrupt change of slope at the contact of the
carbonate shelf and the volcanic rocks forming the shore.

The outer system is also unusual for the Lesser Antilles in that
the apparent basement is relatively deep (25-33 m). This may result
from unusually massive sand and rubble buildup on an embayed and north-
south oriented coastline. Another possibility is the presence of a
Pleistocene reef structure lying under the outer system. Our deepest
core at Pinsonelle (Fig. 5) extended to only 13 m, however repeated
drillings through the bank barrier in St. Croix (Adey and Burke, in
prep.) as well as the minimum bank thickness at Vauclin reef (Fig. 3)
and the shelf profile (Fig. 2), suggest that such an old reef core is
not likely. Although other double bank-barrier systems are known in the
Lesser Antilles (e.g., World's End and Horseshoe Reefs in the Grena-
dines), the Martinique system is the most extensive of these.

Profiles across several other points on the inner reefs (Figs. 5-7)
are generally similar to that of Vauclin reef. Although it is not obvi-
ous on figure 1, it can be seen on the standard marine chart (HO 1009)
that Sans Souci (inner), Pte. de la Prairie and especially Ramville Is-
land reefs are more exposed than Vauclin Point reef. As a result, these
reefs tend to be shallower near the crest, and both Ramville and Pte. de
la Prairie have numerous low algal mounds and spurs (incipient algal
ridges). These features are particularly apparent from the air, espe-
Cially during low tides, but can be very difficult to see from a boat
unless the sea is calm.

The inner reef system apparently developed extensive Acropora
palmata caps over the bank-barrier foundation in the late Holocene.
Large samples, extracted from the "clean" cores of several A. palmata
arms, from the top and bottom of this facies in Vauclin reef gave C
dates of 560 and 1670 years B.P. respectively. Most of the A. palmata
excavated in a 3 m2 hole dug to a depth of 2 m into the reef crest
(Fig. 3) appeared to be in growth position. Today this coral only domi-
nates a narrow and irregular band on the front faces of these reefs just
below the zone of heavy algal growth.

Coring on the very high energy ramparts of Pinsonelle algal ridge
(Fig. 4), indicates that crustose corallines have dominated the frame-
works of the algal ridges on this system for at least two thousand years;
a single cl4 date from a narrow A. palmata band at a depth of 4-5 m was
2110 years B.P. Although we cored only in an emergent area on the outer
system, the similarity in position and appearance of these structures
would indicate that the results obtained at Pinsonelle are valid general-
ly. It would be especially desirable to obtain a coring through to the
Pleistocene shelf in an area where no algal structures have reached sur-
face level, such as at Cayes de San Souci (see Fig. 8). However, there
is little reason to believe that the basement in such cases is any shal-
lower than at Pinsonelle. At Passe du Brigot an algal ridge is developed
on both sides of the channel, and a section across the north ridge
(Fig. 9) is very similar to that at Pinsonelle both with regard to depth
profile and animal and plant populations (Fig. 10).

A single hole was drilled in the southern Martinique "bench-ridge
system" at Lezard Is. (Fig. 11). At eight and one half meters below the
surface of the ridge, drilling resistance increased markedly, suggesting

6.

a transition from predominantly crustose coralline material to rock base-
ment. However, a drill rod broke at this point, and we were unable to
retrieve the bit and obtain core from the base of the hole. Our inter-
pretation of these structures as "bench type" is based primarily on their
location near shore, very shallow lagoons, and association with several
predominantly limestone islands upon which wave cut terraces could have
been formed at a period of lower sea level (see Adey and Burke, 1976).

A similar drilling pattern was encountered in the bench algal ridge at
Chateaux Point, Guadeloupe, where limestone basement was obtained with
great difficulty from 7 m down.

The present eastern Martinique reef complex is characterized by an
overwhelming dominance of large standing crops of fleshy benthic algae
in crest and upper fore-reef areas, especially of the genera Sargassum
and Gracilaria. This characteristic is also shared by some of the reefs
of southeastern Basse Terre, St. Lucia, Grenada, and Nevis (see Adey and
Burke, 1l.c.). Otherwise, such extensive fleshy algal reefs do not gen-
erally occur on Lesser Antillean islands except locally on sheltered
and low "old" algal ridges (Connor and Adey, 1976). (Note the apparent
fleshy algal covered pavement on high energy eastern Curacao (Van den
Hoek et al., 1975.) The reasons for this somewhat unique algal domi-
nance is surely due in part to the limitation of grazing by fish (Scari-
dae and Acanthuridae) and echinoids (Diadema antillarum) by the high
wave energy. However, Similarly shaped reefs with about equivalent wave
action (see Adey and Burke, in ms.) in the Grenadines lack a fleshy algal
cover. Through the Lesser Antilles, there appears to be a correlation
of algal pavements with high turbidity; all of this is discussed below.

CORALS

P. J. Roos, in his 1971 study of the stony corals of the Netherland
Antilles referred to a private collection of twelve species from Martin-
ique made by P. W. Hummelinck. Eleven of these species were found during
the course of the present study, Solenastrea bourni being the only ex-
ception, which was perhaps overlooked due to its similarity to the more
abundant Montastrea annularis. Twenty-two additional species, including
four major carbonate-producing Hydrozoans, have been identified and are
listed in table 1. Their distribution along several inner and outer
bank barrier transects is shown in figures 5-10.

Coral diversity on the lower fore-reef zones of the inner system is
rather high, but total populations do appear to be generally low relative
to comparable reef surfaces in St. Croix, Antigua or the Grenadines.

The number of scleractinian species observed is about fifty percent of
that reported from the intensive investigations in Jamaica (Goreau and
Wells, 1967). However, as Roos (l.c.) noted "...there is considerable
variation in important (diagnostic) characteristics...and great varia-
tion in growth form (of many of the corals and therefore) the great
number of species of Agaricia in Goreau and Wells (1967) has to be
interpreted with care." We conclude that this caution must be applied
to some other genera as well. Thus, no attempt has been made to
differentiate beyond the generic level for numerous morphologies of

as

Agaricia spp., Mycetophyllia spp., and what are apparently Colpophyllia
spp. The list of coral species presented here (Table 1) could probably
be increased by thirty to forty percent if taxonomic distinctions were
clearer.

Inside the inner reef system water circulation and turbulence are
restricted, creating conditions suitable for well developed lagoonal
communities. Many of the inner bays have extensive mangrove populations,
and coral production is low. However, in the lagoon area immediately
behind the inner reefs, extensive beds of Thalassia testudinum and
Syringodium filiforme as well as a complementary benthic algal popula-
tion (see below) are found. The primary corals found here are Manicina
areolata, and four species of the family Poritidae, Porites astreoides,
Porites divaricata, Porites furcata, and Porites porites, the latter
three species being most common. Also, associated with these are the two
species of the family Siderastreidae (Siderastrea radians and Sideras-
trea siderea), which inhabit all other zones of the reef as well.
Occasional scattered colonies of Diploria clivosa, Diploria strigosa and
Oculina valenciennesi (near shore) are also common. An extremely deli-
cate form of the Hydrozoan Millepora alcicornis is abundant in the
shallow seagrass zone adjacent to the immediate back reef. Apparently
this species prefers these shoal areas, which are characterized by a
considerable current but low turbulence. This is also a common location
for the formation of circumrotary corals (free nodules) by both Sideras-
trean species.

Except for the reef crest, which is locally dominated by the Hydro-
zoans M. complanata, and M. sgquarrosa, the back and upper fore reefs are
characterized by large standing crops of fleshy algae and generally only
a few scattered corals.

The area of coral dominance on the inner reef system is restricted
to the fore reef and extends from just beneath the zone of heavy algal
growth at a depth of 3-4 meters, to the sand at 10-15 meters. Because
of the turbid water and the resultant reduction of light penetration, the
corals within this zone represent a community usually found within a
much wider depth range, and many colonies have assumed the plate-like
form usually associated with deep water, although these are generally of
less fragile construction.

Below the algal zone, the abundance of the various species of the
order Faviidae increases with depth and then decreases again near the
base of the deep fore reef. The members of the family Faviinae, includ-
ing Favia fragum, D. clivosa, D. strigosa, Diploria labyrinthiformis and
Colpophyllia natans, are common over the entire reef, although the latter
two species are more common on the fore reef. The family Montastreinae
(represented by Montastrea annularis and Montastrea cavernosa) is gener-
ally more common near the break in the fore reef slope and on the deep
fore reef. Although there are some marked species preferences in the
upper and lower portions of the fore reef slope (Acropora palmata above
and D. labyrinthiformis and Colpophyllia spp. below), the most striking
change in coral populations is in their morphologies.

The general trend of coral morphology, changing with increasing
depth from crusts to heads to foliose, is similar for most major corals
at all locations carefully examined on Martinique. The trend, however,
is especially marked in Diploria spp., Montastrea annularis and Porites
astreoides. Primarily encrusting forms of Diploria clivosa and Diploria
strigosa and hummocky mounds of P. astreoides are found in the areas
dominated by fleshy algae near the outer part of the algal pavement.
Further seaward, they assume a hemispherical shape and are locally asso-
Ciated with patches of Acropora palmata. On the steeper fore reef, most
corals gradually adopt the foliose morphology. Foliose Diploria spp.,
M. annularis, P. astreoides and Colpophyllia spp. shingle wide areas of
the reef face and associated pillars, from the break in the fore reef
slope to the base of the deep fore reef, over a depth range of up to ten
meters.

Additional components occurring near the base of the deep fore reef
community include Scolymia lacera, Meandrina meandrites, Mycetophyllia
spp., and Siderastrea siderea.

On the outer reef system (Figs. 4, 8-10), the scleractinian corals
are concentrated along the deep fore reef and also occur locally in back
reef (reef flat) areas behind the algal ridges. The deeper fore reef
community seems to be rather homogenous from our observations of three
different localities, Pinsonelle, Cayes du Vauclin and Cayes de San Souci.
Where the slope steepens markedly, there is a sharp decrease in fleshy
algae and a corresponding increase, in’ the, frequency of) coralssaetne
Hydrozoan Millepora squarrosa, and three Diploria spp., especially D.
labyrinthiformis, occasional A. palmata, along with the secondary corals
(e.g., Favia spp., Siderastrea spp., P. astreoides) dominate the transi-
tion across the slope break. A gradual increase in’ M. annularis, Colpo-
phyllia spp., Montastrea cavernosa, and large S. siderea then occurs
down the reef face. These coral communities also occur on the sides of
the deep sand channel& and patches that are scattered over the fleshy
algal pavements.

The back reef areas behind the algal ridges of the outer system are
much more complex in that large patches of less energy tolerant species
(A. palmata, P. porites) are locally very abundant. The back reef slope
(sand cliff) is mostly occupied by seagrass communities, though well de-
veloped faviinid reefs of M. annularis, Diploria spp. with abundant
alcyonarians occur locally. Occasional A. palmata colonies also occur

in this area.

There are a few major Caribbean coral species that are relatively
rare in the Martinique reef system. Agaricia agaricites, which charac-—
teristically shingles vast areas on deep reefs in Jamaica and shallow
areas in Belize, is rather uncommon here. Similarly, Acropora cervicor-
nis, which is moderately common on fore reefs of the Grenadines, Guadel-
oupe, Antigua, and the Virgin Islands and is a mid-reef framework builder
in the central and western Caribbean, was found living only at the base
of a patch reef in Havre du Robert.

BENTHIC ALGAE

As described above, on Martinique the crustose corallines are the
main builders of the algal ridge caps on the outer reef system, the near-
shore bench ridges in the south, and the incipient ridges forming on the
crest and upper fore reef along the inner system. They also form a thin
crust over much of the surface of the algal pavements of both the outer
and inner reef systems. The taxonomy and ecology of crustose corallines
in the Caribbean is presently under study (WHA), and, as many of the
species are as yet undescribed, only a brief treatment of the dominant
elements is given here.

In the deeper zones of the reefs throughout the Caribbean, the crus-
tose corallines are poorly represented, although the genera Paragonioli-
thon (n. gen.), Neogoniolithon, Hydrolithon and Tenarea do occur. However,
in the shallower zones that are dominated by Acropora palmata and Mille-
pora complanata, the corallines typically encrust large portions of the
usually abundant dead coral substrate. Porolithon pachydermum, Neogoni-
olithon megacarpum (n. sp.) and Lithophyllum congestum are dominant in
well lighted areas, and Neogoniolithon accretum, Hydrolithon borgesenii
and sometimes Lithothamnium ruptile and Paragoniolithon solubile are
important in shaded situations (see Adey and Vassar, 1975; Van den Hoek
et al., 1975). On rubble fragments scattered around the corals and
patches, Hydrolithon borgesenii and sometimes Neogoniolithon mamillare
are usually the dominant encrusters. The coralline populations on the
inner reef system of Martinique is generally typical in the narrow A.
palmata band and deeper fore-reef Zones.

Although some Caribbean algal ridges are characterized by frameworks
of the anastomosing branching type of L. congestum (Steneck and Adey,
1976), this has not been found to be the situation on Martinique and
Guadeloupe, perhaps due to physical damage to this plant caused by higher
energy levels. Although L. congestum occurs in this area, the ridges
appear to be built largely by Porolithon pachydermum with a considerable
admixture of Millepora.

On the fleshy algal pavements (which occupy the zones ordinarily
dominated by A. palmata in other areas) the crustose coralline population
is quite unusual beneath the heavy plant cover. Here Neogoniolithon n.
sp., Mesophyllum n. sp., Lithophyllum n. sp. and Archeolithothamnium n.
sp., all shade plants, are abundant. Most of the plants usually associa-
ted with A. palmata, with the addition of Archeolithothamnium dimotun,
also occur on these pavements, but are much less abundant than those
listed above. All of the species, except Archeolithothamnium n. sp.,
have been found in scattered localities elsewhere in the Caribbean, where
they occur locally under fleshy algal patches. The Archeolithothamnium
n. sp., a large, coarsely branched plant, is presently known only from
Martinique.

The fleshy algae dominating the crest of Martinique reefs are both
diverse and abundant. The Vauclin Point pavement, with over 100 species
of macro-algae and an average standing crop of about 4 kg/m? (wet), is
richer than any previously described equivalent tropical area (see Connor
and Adey, 1976). (Van den Hoek et al., 1975, reported 142 algal species

10.

from a transect across the shore in south west Curacao. However, the
transect included both shore and deeper zones. The maximum number in

that study for a community-zone was 55. An equivalent number for the
back algal pavement in this study would be over 90.) A series of 0.25 m2
areas were collected in transects across Vauclin Point reef and Pinsonelle
algal ridge. Standing crop measurements of the dominant species found are
shown in figures 12 and 13. Numbers of quadrats, species encountered,
frequency and standing crop by species are given in table 2.

On the Vauclin Point pavement, the maximum standing crops of fleshy
algae are found on the upper fore reef (seaward of the surf zone) and on
the back reef areas. There is a marked decrease in the standing crop in
the surf zone, mostly due to a reduction of Sargassum platycarpum. On
the algal ridge at Pinsonelle, the reduction of standing crop in the very
high energy zone (which is also largely intertidal) is much more marked.
Here, except for a small amount of Sargassum vulgare, virtually all of
the species occurring on the fore and back ridge pavements are absent,
and several Laurencia species dominate.

The Sargassum species are the major elements on the Martiniquan reef
and ridge crests, although several Gracilaria, Dictyopteris, Dictyota and
Laurencia species also occur in abundance. Floating Sargassum is fre-
quent on these shores during periods of rough seas, suggesting that it is
torn off by wave action (the relationship between wave action and algal
standing crops over several years was described by Doty (197la) fora
Hawaiian algal pavement). On the other hand, heavy wave activity seems
often to be associated with the presence of dense algal growth, perhaps
serving to inhibit the predation of Diadema antillarum and parrot and
surgeonfishes (mainly Sparisoma rubripinne and Acanthurus bahianus) .
Thus, Sargassum appears to reach high densities within a relatively nar-
row range of wave energy. Large algal stands do occur intertidally (on
algal ridges) in relatively quiet St. Croix, but in Martinique, they ex-
tend down to 3-4 m on the medium-high energy inner reefs, and reach
10-15 m on the very high energy outer system.

With adequate protection, D. antillarum can probably survive long
periods of intense wave action without moving from shelter. Within the
area of dense algal growth behind the crest at Vauclin Point reef are
several bare depressions occupied by large groups of Diadema. These
typically cover an area of about 3 x 5 meters and are approximately 50 cm
deep, sharply delineated at the upper rim by the surrounding algal forest.
Apparently the urchins are kept within these patches by the force of cur-
rent and swell as well as the whipping action of the long Sargassum fronds.
During periods of especially intense wave activity these patchy Diadema
seem to be restricted even more narrowly and pioneer algal growth begins
to encroach upon the bare carbonate areas. As conditions return to normal,
however, this new growth quickly disappears. Areas adjacent to the barren
Diadema depressions, when cleared of their heavy algal growth are usually
kept clear by Diadema grazing when wave energy stays at moderate levels.
However, under rough conditions these areas are slowly re-covered with
fleshy algae. It would appear that if wave action were to diminish
greatly for a considerable length of time, the Diadema already present on
the reef could decimate the algal stand. Therefore consistency of wave

1b

action would seem to be as critical as intensity. This might provide one
explanation for the extent of the algal pavements that are present on
Martinique, located as it is in the zone of maximum trade wind constarcy.

Our studies have indicated that three factors are related to the
presence of sublittoral fleshy algal pavements: strong wave energy,
smooth carbonate pavement and turbid waters. In addition to the inhibi-
tion of grazing activities that has been discussed above, it is also
likely that wave energy plays an important role in the formation of
smooth carbonate pavements by reduction of the coral population through
breakage and the filling of the reef matrix with transported sediment
and debris. Once such a smooth pavement is formed, grazing is further
reduced because holes and niches used for protection by reef organisms
are drastically reduced in number. However the combination of high wave
energy and smooth carbonate pavement is usually not enough in itself to
stimulate the formation of heavy fleshy algal growth, as is evidenced in
many reefs in the Grenadines where the water is quite clear. It would
seem that turbid water, probably rich in nutrients, is also required for
the general heavy growth of fleshy algae, as has been observed by Adey
and Burke (1976).

As has been discussed above, the crests of Martinique reefs were not
always covered by fleshy algal pavements, but were dominated in the past
by Acropora palmata, although it is probable that the water turbidity
and energy conditions were not significantly different during that period
than they are today. It seems likely that the changeover occurred when
the A. palmata reef grew to near sea level and gradually was transformed
by wave energy into a carbonate pavement upon which fleshy algae could
flourish in the turbid water.

PRODUCTIVITY

Coral reefs are biologically among the most productive systems known.
Oxygen exchange rates (based on changes in oxygen concentrations in water
flowing over reefs and transformed to organic carbon), have been used to
obtain gross figures of about 5 to 25g C/m2/day. In a typical Caribbean
A. palmata reef environment, living corals and coralline algae cover a
relatively small proportion of the total reef surface. Usually, much of
the total area at any one time consists of dead coral substrate with a
short, dense turf of small filamentous algae, blue-greens often being
abundant. An algal ridge or pavement with a large standing crop of
fleshy algae might be expected to be more productive than a typical coral
reef; Connor and Adey (1976) suggested this in the study of an algal
ridge lobe on St. Croix, where values of 24 to 45 C/m2/day were obtained.

Preliminary oxygen exchange studies were undertaken on both Vauclin
Point reef and Pinsonelle algal ridge, using the Winkler method for mea-
suring dissolved oxygen. At Vauclin Point reef, the breaker zone begins
approximately one third of the way back from the seaward margin of the
algal pavement. For each of the 15 sets of oxygen data taken, five
samples were collected: seaward of the reef; beginning of the algal Zone;

231-339 O-'77 - 15

2

beginning of the breaker zone; end of the breaker zone; at the landward
limit of the pavement. By plotting these data as a function of position
and measuring water flow (float bottles) over the reef, it was possible
to determine gain or loss of oxygen in the breaker zone and the net ex-
change of oxygen-between reef surface and the water in terms of g/m2/hr.
Dissolved oxygen values of 6 to 7.2 mg/l were found across the Vauclin
Point reef and mid-winter results for both clear and partly cloudy
(overcast) days are shown in figure 14. From these data, a gross pro-
ductivity for the clear day of approximately 33g 05/m2/day or mlOg
C/m2/day was determined. This is well below that obtained by Connor and
Adey (l.c.) for the St. Croix algal ridge in April. However, the marked
difference between the cloudy and the bright days suggests that, even at
this latitude, daily and seasonal differences may be considerable.
Summer productivity probably is considerably higher.

Only a single series of oxygen samples were taken across the Pin-
sonelle algal ridge, on the second of January. The oxygen concentrations
rose from 6.8mg/l in front, to nearly 8.lmg/1 immediately behind the
ridge pavement system. About two-thirds of this rise occurred on the
rather short back ridge algal pavement, indicating that this very shallow
area is considerably more productive than the deeper Vauclin Point pave-
ments. This further suggests strong limitation of productivity with de-
creasing light in sublittoral reef systems. Adey at al. (studies in
progress) found gross oxygen production in Rod Bay, St. Croix during
November to be 17g 05/m2/day for a 1m deep mixed coral and algae back-
reef and 13g 09/m2/day on a typical Acropora palmata-algal turf fore-reef
with an average depth of about 4m. (Very similar levels of 17 and 10g
O5/m2/day, respectively, were also predicted using species productivity
values of Doty (1971b) and the standing crops of algal turfs typical of
the reef.) Thus, for approximately the same depth (lm), productivity on
the Martinique algal pavement was twice that of the back reef on St.
Croix. This agrees with the theoretical consideration that while fila-
mentous algae generally show 10-20 times greater productivity per unit
weight than the more massive algae (Doty, l.c.), the standing crops of
massive fleshy algae on these algal pavements are 20-30 times greater
than that of filamentous algal turfs in a "coral" reef.

These data suggest that with very extensive algal pavements at
depths of 1-2 m, the reef systems-of Martinique are probably the most
productive in the eastern Caribbean.

FISHES

The fish assemblages of the algal-dominated Martinique inner reef
system differ markedly from those of typical A. palmata and mixed A.
palmata-coralline-Millepora reefs of the eastern Caribbean. The fishes
associated with coral and bare carbonate areas are similar to those of
the mixed reefs, although certain conspicuous species are rare or ab-
sent. The fleshy algal pavements are heavily populated with typical
grass bed populations usually alien to the mixed and A. palmata reefs.
This contrast in fish assemblages may be explained by the general

Ste

disruption of small scale habitats as the formerly Acropora palmata
dominated reefs of Martinique acquired their fleshy algal pavement caps.

During February-April 1975, about one hundred hours between 0900 and
1900 were spent observing fishes at five general locations on Martiniquan
reefs: Ramville Island, Sans-Souci channel and Pte. du Vauclin on the
inner bank barrier system; at Lezard algal ridge on the southern bench
system; and on the more protected parts of Loup Garou on the outer bank
barrier system. Assemblages were described by snorkeling or SCUBA diving
within a broadly defined habitat (e.g., fleshy algal pavement) and record-
ing the relative abundances of the fishes seen. Data were recorded with
time from the beginning of each observation period, allowing the construc-
tion of sampling effort (time) /species number curves. From these curves
it was determined that even in the most heterogenous environments, where
frequent crossing of broad habitat boundaries complicated data recording,
fifty to sixty minutes was usually sufficient to record over ninety per-
cent of the visible species. The required time was somewhat longer in
regions of high wave energy and turbidity. The major patterns discussed
here include only the most abundant species. These were often observed
within the first thirty seconds, without significant change in relative
abundance estimate over several hours of observation. The relative
merits and drawbacks of visual population estimates of reef fishes have
been discussed by Bardach (1959) and reviewed by McVey (1972) and
Gunderman and Popper (1975).

Table 3 lists all species encountered during the Martinigue study,
grouped in order of greatest relative abundance. Table 4 compares the
relative abundance of several characteristic fishes on fourteen eastern
Caribbean reefs, including Martinique. The reefs have been categorized
on the basis of dominant surface features; Union Island, Goyave and Nevis
reefs include patches of two categories.

As seen from Table 4, compared to mixed A. palmata-coralline-
Millepora reefs, “pure stands" of A. palmata had fewer parrotfishes and
were missing many other species. Randall (1967) noticed smaller popula-

tions of fishes on "Luxuriant reefs of Acropora palmata..." and attri-
buted this pattern to "...limited food for fishes which feed on attached

marine life or organisms therein," in spite of the excellent shelter
provided by the continuous coral cover. Our experience on Rod Bay, Reef,
St. Croix, where living A. palmata occupied less than 30% of the surface
and algal turfs are responsible for 80% of the productivity, further
suggests that grazing fish may be food limited on extensive "pure stands"
of A. palmata.

The Martiniquan reefs differ even more dramatically from those with
both patchy A. palmata and coralline-Millepora pavement. Many species
are missing from Martinique and even Sparisoma viride and Scarus vetula
have a very restricted distribution. However the reefs support a much
greater proportion of grass bed fishes such as the bucktooth parrotfish,
mutton hamlet, and blackear and dwarf wrasses (occurrence in grass beds
cited from Randall, 1968; Bohlke and Chaplin, 1968). This peculiarity of
the fish populations is almost certainly not biogeographical, since all
of the absent species are found on reefs both to the north and south, and

14.

no special biogeographical boundaries fall through this area. Further-
more, at the reef at Goyave, Gaudeloupe (which is apparently in the
process of developing from a typical Acropora palmata reef into one with
a fleshy algal cover) there are adjacent patches of fleshy algal pavement
and A. palmata surrounded by coralline-Millepora pavement. The assem-
blage on the fleshy algal pavement is similar to that of Martinique,
while the assemblage on portions dominated by A. palmata is typical of
this type of reef.

From Table 4 it can be seen that the Martiniquan fish assemblages
resemble those of other reefs with extensive fleshy algal pavements much
more than those with extensive A. palmata or mixed patches of A. palmata
and coralline-Millepora pavement, even where they occur on the same is-
land. This suggests that the fish assemblages on fleshy algal pavement
reefs are due to either structural or food resource characteristics of
these reefs which differ from those Of/A-palmata ‘or ‘mixed reetsr

The large Scarus spp. and Sparisoma viride are characteristically
grazers and scrapers of carbonate surfaces rather than browsers on the
larger fleshy algae. (Their main food: in ‘ai/typical A. palmata reek mils
probably the abundant algal turfs growing on dead coral (Randall, 1967),
often the most significant primary producers in this environment (Adey,
studies in progress). It is possible that pavement reefs with dense
fleshy algal stands do not have sufficient food resources of the proper
type to maintain these big turf grazers.

Most parrot fishes form herds in open water, a behavior which is
probably facilitated by bright color patterns (Hamilton, in press).
When encountering the snorkeler (and presumably any other potential dan-
ger) large Scarus spp. and S. viride disappear beneath ledges or among
interlocking arms of A. palmata. Since reefs dominated by fleshy algal
pavements usually lack this shelter as well as grazeable algal turfs,
these fishes would be forced to adopt rather different habits to survive
there as adults. Apparently this has not occurred for any of the larger
parrotfishes excepting the yellowtail, Sparisoma rubripinne. When en-
countered away from shelter, this species assumes a mottled, cryptic
color pattern and nestles down against the algal cover. On the pavements
at Martinique, Nevis and Guadeloupe it was usually observed browsing on
fleshy algae rather than grazing on the small amount of turf-covered
coralline pavement that was available. Ten S. rubripinne were shot for
stomach analyses. Stomach contents were difficult to identify to species,
however most were filled with fleshy fragments. Thus S. rubripinne
seems much better suited to life on the fleshy algal pavements .than’S.
viride or any of the large Scarus spp.

Sparisoma radians is also capable of cryptic coloration and behavior
on fleshy algal pavement, as well as feeding on the algae growing there.
Randall (1967) found it feeding primarily on Thalassia in grass beds.

The mutton hamlet and dwarf and blackear wrasses are cryptic carnivores
that are capable of matching any mottled green or brownish substrate.
Since they are abundant both in grass beds and on fleshy algal pavement,
they apparently do not prey exclusively on organisms peculiar to one or
the other of these habitats.

IS} 6

Many fishes which can reach great abundance on all reef types are
of extremely localized distribution on fleshy algal pavements. For ex-
ample, it has been demonstrated that juvenile Eupomacentrus planifrons
(threespot damselfish) inhabit live, branching coral ("bushes") which
the fish kills as it matures (studies in progress - Kaufman). Adults
prefer dead corals of similar morphology, on which they "farm" small
fleshy algae by chasing herbivores from a territory. This study was
performed at Jamaica and St. Croix, but similar patterns of micro-
distribution for this and other species were observed throughout the
eastern Caribbean and the Florida Keys. In Martinique, juvenile E.
planifrons were fairly abundant on back-reef Porites porites shoals and
fissured rocks, but nowhere else. Adults were restricted almost entirely
to small patches of A. palmata on the fore-reef, where they defended
territories on dead coral.

Observations on the local distributions of many other species tend
to support the hypothesis that differences in fish assemblages between
reefs dominated by fleshy algal pavements and those dominated by living
A. palmata and/or algal turf-covered coralline-Millepora pavements are
primarily due to differences in either or both habitat structure and
food resource availability. The reefs of Martinique assumed their pre-
sent character approximately 600 years B.P., before which the reefs were
dominantly A. palmata. Presumably, the fish assemblage at that time was
as different from that found today as are those of existing A. palmata
reefs. In assessing the historical effects of fishing on local popula-
tions, one should note that major changes in the distribution and abun-
dance of Caribbean reef fishes probably have occurred as the reefs
themselves have matured. In many respects, these changes may be inde-
pendent of man's activities, and are probably still in progress today.

ACKNOWLEDGMENTS

The authors are grateful to the Martinique Prefectural Government,
which freely gave the permission, visas and customs clearances required
to pursue this study. The American Consulate in Fort de France, and
especially Vice Consul Thomas L. Randall, Jr. helped us in numerous
ways. Special thanks are due to Raymond and Margarite Asselin of
Vauclin, who allowed us to work out of their cottage on Petite Grenade
Island. Claude and Jacqueline DeLeuze of Vauclin provided very welcome
assistance in the form of equipment loan and help in locating necessary
facilities. In addition to the authors, Judith Connor and Michael Vassar,
assistants to the Coralline Program, also participated in many aspects of
the investigation. John Ogden and Lee Gerhard of West Indies Laboratory,
St. Croix and Patrick Colin of the University of Puerto Rico read the
manuscript and offered valuable suggestions for its improvement. The
project was supported by the Smithsonian Research Awards Program and
Smithsonian Secretary's Fluid Research Fund.

TG.
REFERENCES

Adey, W. 1975. The algal ridges and coral reefs of St. Croix: their
structure and Holocene development. Atoll Research Bull. 187:1-67.

Adey, W. and J. M. Vassar. 1975. Succession and accretion rates in
Caribbean crustose corallines., Phycologia 14:55-69.

Adey, W. and R. Burke. 1976. Holocene bioherms (algal ridges and bank
barrier reefs) of the eastern Caribbean. Geol. Soc. Am. Bull.

Adey, W. and R. Burke. In Ms. Holocene bioherms of the Lesser
Antilles: geological control of development. A.A.P.G. Mem.7th
Caribb. Geol. Congr.

Bardach, J. 1959. The summer standing crop of fish on a shallow
Bermuda reef. Limnology Oceanogr. 4:77-85.

Bohl keys Drei uranGy Grey. C.i1Ge Chaplin. 1968. Fishes of the Bahamas and
Adjacent Tropical Waters. Acad. Sci. Philadelphia. 771 pp.

Climap Project Members. 1976. The surface of the ice-age earth.
Science 191 :1131-1137.

Connor, J. and W. Adey. 1976. The benthic algal composition, standing
crop and productivity of a Caribbean algal ridge. Atoll Research
Buda.

Doty, M. S. 197la. Antecedent event influence on benthic marine algal
standing crops in Hawaii. Mar. Biol. Ecology 6:161-166.

Doty, M. S. 1971lb. The productivity of benthic frondose algae at
Waikiki Beach, 1967-1968.» ;Hawaid»Bot.. Sci. Paper 22: 1-119).

Goreau, L. and J. W. Wells. 1967. The shallow water scleractinia of
Jamaica: revised list of species and their vertical distribution
range.) Bull. .Mar-Sea .1/j:442=453..

Gunderman, N. D. and D. Popper. 1975. Some aspects of recolonization
of coral rocks in Eilat, Gulf of Aquaba by fish populations after
poisoning. Mar. .Biol., 30:L09-117.

Hamilton, F. In press. Functional coloration of coral reef fishes.
Cited in. Coral..reef, ecology,’ C. U...Smith., (ed<),.

Lewis, J. 1975. A preliminary description of the coral reefs of the
Tobago Cays, Grenadines, West Indies. Atoll Res. Bull. 178:1-9.

Macintyre, I. 1972. Submerged reefs.of ‘the eastern Caribbean... Am.
Assoc. Petr. Geol. Bull. 56:720-736.

Macintyre, I. and P. Glynn. 1974. Internal structure and developmental
stages of a modern Caribbean fringe reef, Galeta Point, Panama.
VIIth Caribbean Geol. Conference. Abstracts p. 40.

Vr.

Mevey, J. P- ~1972. “Ecology of the’ Pokai, Bay artificial reef, Oahu.
Univ. Hawaii Contr. 114-16000133461. 29 pp.

RanGalele Wis Hie LOGI). Food habits of reef fishes of the West Indies.
Cited in: Proceedings of the International Conference on Tropical
Oceanography. 847 pp. University of Miami, Miami, Fla.

Randall, J. E. 1968. Caribbean reef fishes. T.F.H. Publication, Inc.
Jersey City. 318 pp.

Roos, P. J. 1971. The stony corals of the Netherlands Antilles.
p. 1-108. in Studies on the Fauna of Curacao and other Caribbean
islands. No. 371. P. W. Hummelinck, ed., The Hague.

Steneck, R. and W. Adey. 1976. The role of environment in control of
morphology in Lithophyllum congestum, a Caribbean algal ridge
builder. Botanica Marina.

Van den Hoek, C., A. Cortel-Breeman and J. Wanders. 1975. Algal
zonation in the fringing coral reef of Curacao, Netherlands
Antilles, in relation to zonation of corals and gorgonians.
Aquatic Botany 1:269-308.

Presqu’ile de
la Caravelle

Pinsonnelle
Ridge

Cayes de
Sans Souci

Ridge
coral back reef
ee

algal ridge

fleshy algal pavement

coral fore reef Passe

du
algal mounds Vauclin

Cincipient algal ridge)

Anglais — Ferré
algal ridges
L

Scale - km

“Baie des Anglais

laakeps lke

|. Cabrit

Holocene bioherms, on eastern Martinique. Transects: H - Hardy Is.; Li) =
Lezard Is. (Fig. 11); Brigot Ridge - Cayes du Vauclin (Fig. 9); C = Cayes

de ‘San Souci (Fig. 8); PB = Pinsonelile algal ridge (rigs. 4, 10) 7 Vo- Vauclin
Point. (Fig. 3);;S —:San Souci channel) (Fig:.../7); ‘Sp. = Pointe de Laybraamaaie

(Fig. 7); R- Ramville Is. reef (Fig. 5)

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———
y 3 ¢ l

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TT

Fig. 3. Section across Vauclin Point reef, V on figure Te

coralline crust with fleshy algae
150 m

Np E> ~ PNSbreake: zone

SRG EEE, 8M
; Of REE AC SKZ2
Bre OMS OSS
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=10 S77 Acropora palmata fe eons
= Y Porites porites : ei alae
o Q head corals | ? Been) ses
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carbonate sand with core
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a eS A ged
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crustose coralline
Thalassia

core holes

head }
corals

=|
Vv
“¥ fleshy algae, dominantly Sargassum
AE
% foliose

Fig. 4. Section across Pinsonelle algal ridge on the outer reef system,
Pon figures.

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Rubble

Fig 2 6.

Sans Souci Channel

(inner system)

meters 200 a Bo iahd See” (Oc "

Profile and surface zonation across San Souci Channel reef. Section

S on figure 1. Corals: (1) Porites porites. (2) Montastrea annularis,
Diploria clivosa, D. strigosa. (3) P. furcata. (4) P. astreoides,
Dichocoenia stokesii, Manicina areolata, Favia fragum, M. annularis.

(5) Millepora spp., Acropora palmata. Corallines: Neogoniolithon
westindianum. Pavement: qdrazed coralline encrusted. Mat: algal

stabilized sediment.

SoS R OES
, 00 8 00 8 o 2 ° 0° 96° 2,0°0 0%

o,0° ° so on eo1e Ss Ne On" °,sCorals (ie

mean low water

Pte. de la Prairie (inner system)

10
meters 50 100 150 200
Fig. 7. Profile and zonation across Pte. de la Prairie reef.

Section Sp on
figure 1. Corals: (1) Porites) porites, P< astreoides, Siderastrea
siderea, S. radians, Manicina areolata, Favia fragum. (2) Diploria
spp. (3) Montastrea annularis. (4) Acropora palmata. (5) P.

astreoides. (6) Millepora squarosa. (7) P. porites, P. furcata,
Mycetophyllia sp., Agaricia sp.

; Bite ath
pee He ee a eS ES

y

uA
<

& Sponges

Rubble

SEAWARD
Se ee Eee

:

Cayes de Sans Souci

(outer system)

meters 500 1000

Fig. 8. wvrofile and zonation across Cayes de San Souci. Section C on

figurewl 4) Corals:. (1) Daploria ‘labyrinthiftormmis, D. clivosa,
Millepora spp., Montastrea annularis, Agaricia agaricites. (2)

Isophyllia sp. Mycetophyllia sp., Porites astreoides. (3) Favia

fragum, Diploria sp., Siderastrea siderea, P. astreoides. —.(4)
Acropora palmata. (5) deep reef community growing on carbonate
spurs. Corallines: lLithophyllum congestum, Neogoniolithon sp.

Sediment

mean low water

Brigot Ridge

Cayes du Vauclin (outer system)

meters 200 400 600 800

imal) 4 SIP

Profile and zonation, Cayes de Vauclin (Brigot). Marked Brigot

Ridge on figure 1. Corals: (1) Millepora sp. and Porites astreoides.
(2) Millepora sp., Diploria clivosa, upper section; Siderastrea
siderea, Montastrea annularis, Isophyllia sp., lower section.
Coralline algae, about 75% living - Lithophyllum congestum, Neo oni-
olithon megacarpum, Neogoniolithon spp., Porolithon pachydermum.

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Table 1. Corals Recorded from the Reefs of Martinique

ANTHOZOA
Scleractinia

Acroporidae

Acropora cervicornis

A. palmata

Agariciidael
Agaricia agaricites

Poritidae
Porites astreoides
P. porites
P. divaricata
P. furcata

Siderastreidae
Siderastrea radians
s. siderea

Faviidae
Montastreinae2

Montastrea annularis

M. cavernosa

Faviinae
Favia fraguum
Diploria strigosa
D. clivosa
D. labyrinthiformis
Manicina areolata
Colpophyllia natans3

Rhizangiidae
Astrangia solitaris
Phyllangia americana

Oculinidae
Oculina valenciennesi

Trochosmilidae
Meandrina meandrites

Dendrogyra cylindrus

Dichocoenia stokesii

Mussidae
Scolymia lacera
Mussa angulosa
Isophyllastrea rigida
Isophyllia sinuosa
Mycetophyllia lamarckiana

Cariophylliidae
Eusmilia fastigata

HYDROZOA

Milleporina

Milleporidae
Millepora alcicornis4
M. squarrosa
M. complanata

Stylasterina

Stylastridae
Stylaster roseus

1. Three additional morphologies were observed.
2. Solenastrea bournoni was observed by P. W. Hummelinck but we did

not encounter it.

3. At least three different morphologies were observed.
4. Extremely variable morphologies were encountered including a long,
thin, fingered variety and a wrinkled vase-forming variety,

apparently ecovariants.

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Typical habitat:

Table 3.

Martinique Fish, Grouped by Relative Abundance

(T) indicates uncertainty in identification.

Class I

Jenkinsia lamprotaenia (T)
Holocentrus ascensionis

H. rufus

Atherinomorus stipes
Allanetta harringtonensis
Planktivorous Carangid (T)
Ocyurus chrysurus

Haemulon chrysargyreum

H. sciurus

Mulloidichthys martinicus
Pempheris schomburgki
Kyphosus sectatrix (T)
Eupomacentrus fuscus

E. partitus
Microspathodon chrysurus
Abudefduf saxatilis
Chromis multilineatus
Halichoeres bivittatus

H. radians

H. maculipinna

Thalassoma bifasciatum

H. poeyi

Sparisoma rubripinne

S. radians

Scarus croicensis
Ophioblennius atlanticus
Labrisomus nuchipinnis et al.
Malacoctenus aurolineatus
Malacoctenus macropus
Malacoctenus erdmanni
other Malacoctenus spp.
Acanthurus coeruleus

A. bahianus

A. chirurgus
Coryphopterus glaucofrenum
ee nyalinusi et adv. (1)

Dwarf Herring
Longjaw Squirrelfish
Common Squirrelfish
Hardhead Silversides
Reef Silversides

(on Pte. du Vauclin) ~
Yellowtail Snapper
Smallmouth Grunt
Bluestriped Grunt
Yellow Goatfish
Copper Sweeper
Bermuda Chub

Dusky Damselfish
Bicolor Damselfish
Jewelfish

Sergeant Major

Grey Chromis
Slippery Dick
Puddingwife

Clown Wrasse
Bluehead Wrasse
Blackear Wrasse
Yellowtail Parrotfish
Bucktooth Parrotfish
Striped Parrotfish
Redlip Blenny

Hairy Blenny
Goldline Blenny

Rosy Blenny

Imitator Blenny

Blue Tang

Ocean Surgeonfish
Doctorfish
Bridled Goby
Glass Gobies

i ee

ee * *

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Table 3 (con't.)

37
38
39
40
41
42
43
44
45
46
47
48
49
50
52
52
53
54
55
56
5if,
58
59
60
ol
62

63
64
65
66
67
68
69
70
7
72
a3
74
5)
76
Vil.
78
79
80
81
82
83

Synodus intermedius
Aulostomus maculatus
Adioryx coruscus
Myripristes jacobus
Cephalopholis fulva
Alphestes afer
Hypoplectrus puella
Serranus tigrinis
Priacanthus cruentatus
Lutjanus griseus

L. apodus

Haemulon flavolineatum
H. plumieri
Eupomacentrus planifrons
E. variabilis

E. leucostictus
Doratonotus megalepis
Hemipteronotus splendens
Sparisoma viride

S. chrysopterum
Gnatholepis thompsoni
Gobiosoma evelynae
Balistes vetula
Cantherhines pullus
Canthigaster rostrata
Diodon holocanthus

Class II

Sand Diver
Trumpetfish

Reef Squirrelfish
Blackbar Soldierfish
Coney

Mutton Hamlet
Barred Hamlet
Harlequin Bass
Glasseye Snapper
Grey Snapper
Schoolmaster
French Grunt
White Grunt

Yellow Damselfish, 3-spot

Cocoa Damselfish
Beaugregory

Dwarf Wrasse

Green Razorfish
Stoplight Parrotfish
Redtail Parrotfish
Goldspot Goby
Sharknose Goby
Queen Triggerfish
Tail-Light Filefish
Sharpnose Puffer
Spiny Puffer

Class III
Synodus synodus Rockspear
Echidna catenata Chain moray
Strongylura spp. Needlefish

Hemiramphus brasiliensis
Adioryx vexillarius
Sphyraena barracuda
Epinephelus guttatus
E. adscensionis

Gramma loreto

Rypticus sapnoaceous
Caranx ruber
Trachinotus goodei
Scomberomorus regalis
S. cavalla

Lutjanus mahogony
Haemulon aurolineatum
Odontoscion dentax
Equetus acuminatus
Pseudopeneus maculatus
Gerres cinereus (T)
Bothus lunatus

Ballyhoo

Dusky Squirrelfish
Great Barracuda
Red Hind

Rock Hind

Royal Gramma
Soapfish

Barjack

Palometa

Cero

King Mackeral
Mahogony Snapper
Tomtate

Reef Croaker
Cubbyu

Spotted Goatfish
Yellowfin mojarra
Peacock Flounder

FR

Pad

x mm Mm Mm OM OM

x

x

mm mM OM MOM

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xm mM Mm MM

39).

Table 3 (con't.) Class nin (conte) FR AP BR
84 Scorpaena plumieri Spotted Scorpionfish x

85 S. grandicornis Grass Scorpionfish x

86 Chaetodon capistratus Foureye Butterflyfish x x x
87 Ge striatus Banded Butterfly x x
88 Bodianus rufus Spanish Hogfish x

89 Hemipteronotus novacula Pearly Razorfish x

90 Sparisoma aurofrenatum Redband Parrotfish x Si
OW Lactophryis triqueter Smooth Trunkfish x x
92 Sphaeroides greelyi Caribbean Puffer x
98 S. spengleri Bandtail Puffer x
94 Diodon hystrix Porcupinefish x x

Class IV

95 Dasyatis americana Stingray

96 Gymnothorax moringa Spotted Moray

97 G. vicinus Purplemouth Moray

98 Apogon maculatus Flamefish

99 Dactylopteris volitans Flying Gurnard
100 Scarus vetula Queen Parrotfish

Class V

101 Trachinotus falcatus Permit
102 Caranx hippos Crevalle Jack
103 Pempheris poeyi Shortfin Sweeper
104 Pomacanthus paru French Angelfish

105 Eupomacentrus mellis Honey Gregory

106 Halichoeres garnoti Yellowhead Wrasse

Lagoon, Sand, Grass

Near the Reef

107 Ophichthus ophis Spotted Snake Eel

108 Myrichthys oculatus Goldspotted Snake Eel
109 Hippocampus reidi Slender Seahorse

110 Mugil curema Mullet

data Hypoplectrus nigricans Black Hamlet

WN Monocanthus ciliatus Fringed Filefish

dal) M. tuckeri Slender Filefish

114 Antennarius multiocellatus Longlure Frogfish

Cryptic within carbonate pavement

115 Stathmonotus spp.
116 Enneanectes pectoralis

A. Palmatay

A. palmata coralline-Millepora

fleshy algal pavement

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Hypoplectrus chlorosus (+) a ees >. Always Seen LOS
Pomacanthus paru Sine
parisoma ne oripinne oe oe cee a wae ee ee ee eoe eee eoo0e eoe e@ooe eoe eee ‘ateve Frequently Seen 2— 9
alichoeres poeyi eles eiomeverers 2007 CSO ee ae eee
parisoma radians ae So Ie ** Occasional 1
Doratonotus megalepis ee net on ao
° Rare 1

Table 4.

Fish species contrasting most in relative abundance on different reef types. Parentheses
indicate the presence of a species on small patches of A. palmata near or surrounded by
coralline-Millepora pavement.

(ee)
=

ATOLL RESEARCH BULLETIN
NO. 219

ISLAND NEWS AND COMMENT

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

May 1977

ISLAND NEWS AND COMMENT

Again we have material on hand for a substantial News and
Comment number, but have not had time to edit much of it for pub-
lication. We do not want to delay publication of the present
issue, nor do we want to delay further a number of reviews of books
and other publications that may interest our readers. Hence we
will include such items as are ready in the present short number.

Rather than including extensive bibliographic material in
Island News and Comment, we hope in the future to devote an entire
issue to bibliographic articles noting and commenting on island
literature. We are, hence, very much interested in receiving re-
prints and other papers dealing with islands, as well as references
to such works if copies are not available.

We also welcome informative or critical reviews of signifi-
cant books or other publications on islands or reefs. If they
are typed in the format used for reviews in this issue the editors'
work will be facilitated.

NEWS

THIRD INTERNATIONAL CORAL REEF SYMPOSIUM: Under joint sponorship
of the University of Miami, the U.S. Geological Survey, and the
Smithsonian Institution, a third of the series of symposia started
at Mandapam Camp, India and continued on the S.S. Marco Polo on the
Great Barrier Reef, Australia,will be held at the University of
Miami, in Miami, Florida, May 23 - 27, 1977, with a reception to be
held on the evening of May 22. The second circular, being mailed
as this is written, will give authoritative details.

Before and after the symposium, field trips are being planned
and with leaders as follows:

Jamaica Jeremy Woodley

Barbados Noel James & Colin Stearn

Bermuda Wolfgang Sterrer

Bahamas Conrad Gebelein

Grand Cayman Harry Roberts & Clyde Moore

Belize James Miller & Ian Macintyre

Martinique & Walter Adey & Robert Dill
St. Growx

Panama Ross Robertson & Peter Glynn

Florida Keys
& Florida Bay Gene Shinn
Dry Tortugas Gary Davis

231-339 O-'77 -17

Whether or not any particular one of these trips will take place
will depend on whether sufficient people indicate that they will partic-
ipate. The trips must be self-supporting, as there is no available source
of funds to subsidize them.

The chairman of the organizing committee is:

Dr. R. N. Ginsburg
Fisher Island Station
Miami Beach, Fla. 33139

MID-PACIFIC MARINE LABORATORY: The U.S. Energy Research and Development
Administration (Division of Biomedical and Environmental Research) sup-
ports the operation of the Mid-Pacific Marine Laboratory on Enewetak
Atoll as an extension of the Hawaii Institute of Marine Biology (University
of Hawaii). The laboratory has recently been reorganized and expanded to
a year-round operation, with special interest in programs related to the
distribution of radionuclides and other materials in an atoll environment,
the cycling of these materials through the atoll ecosystem, and the pos-
sible implications of these processes to man. Researchers interested in
undertaking studies at Enewetak should contact Dr. Stephen V. Smith,
Director, MPML, P. 0. Box 1346, Kaneohe, Hawaii, 96744.

FILM AVAILABLE ON THE ECOLOGY AND ETHOLOGY OF CORAL REEF FISHES: Dr.
Ernst S. Reese, Department of Zoology and the Hawaii Institute of Marine
Biology of the University of Hawaii, has completed an educational docu-
mentary film on the social behavior and related aspects of ecology of
butterflyfishes of the Family Chaetodontidae. This is a professional
quality film in color, 16mm with sound. The film was made over the past
8 years during studies carried out in Hawaii, at Eniwetok Atoll in the
Marshall Islands, and at Heron Island on the Great Barrier Reef. The
film shows the feeding and agonistic behavior of 15 species. Each species
belongs to one of three feeding "guilds" (coral feeders, omnivores,
plankton feeders) which in turn determines the space-related behavior of
each species. The purpose of the film was to examine how one group of
fishes successfully coexists and shares the resouces of space and food on
the reef. There is an intimate relationship between the patterns of
social behavior and the ecology of each species which has resulted in

the different strategies of resource utilization. A detailed study guide
is available with the film. For a brochure and further information on
rental, preview or purchase, please write to Dr. Ernst S. Reese, Hawaii
Institute of Marine Biology, P.O. Box 1346, Kaneohe, Hawaii 96744.

SECOND SMITHSONIAN PHOENIX ISLAND EXPEDITION: As a follow-up to their
1973 expedition to the Phoenix Islands, Roger Clapp, F. R. Fosberg,

and D. R. Stoddart revisited the Phoenix Islands in March and April,
1975, courtesy of the SAMTEC Project of the U.S. Air Force. The official
purpose of the visit was to assess the environmental effects of Air Force
activities on Enderbury and Hull Islands. Scientific observations were
made in various fields and a series of topographic profiles were levelled
across various transects of the islets. Fortunately it was possible to

visit all eight of the atolls, so the reconnaissance study of the
Phoenix Group was essentially completed. Reports on several aspects are
being prepared for publication in a future issue of ARB.

COMMENT

THE PROBLEM OF DEFINITION OF THE TERM "ATOLL": I was recently asked
about definitions of atolls and especially about what to call Cayo Sal,
of the Venezuelan coast. This is a somewhat edited version of my reply,
which may be of interest to some readers.

This raises a question of terminology which probably never can be
resolved except arbitrarily, and then certainly not to the satisfaction
of all groups of users. The difficulty is that different groups tend to
write definitions to satisfy different requirements and it is often not
possible to formulate a definition that suits all purposes equally well.

For geologists the essential characteristics seem to be that the
material must be reef limestone or at least calcareous skeletal debris
and that the reefs and islets must surround or partially surround a
shallow body of water termed a lagoon. Some geologists insist that the
structure rise from deep water, others are willing to accept an essen-
tially morphological definition, though they argue endlessly about the
origin of the essential form.

The definition quoted to you by Dr. Bryan was formulated by
ecologists, essentially for use by ecologists. It differs from other
definitions in that it also includes islands which do not have lagoons
but are emergent though essentially sea-level reef-limestone structures.
It excludes those that are elevated more than a very few meters above
mean low-tide level, and it also excludes such islands as are closely
associated with high islands or continental land, or are separated from
such higher land only by shallow water. Bryan has used the 100 fathom
line as separating shallow from deep water. I would probably use a
Shallower limit, such as 100 m. However, this illustrates one of the
principal problems with definitions of natural phenomena. They usually
do not exhibit sharp boundaries and tend to be continua, with gentle to
sharp gradients. There seems to be no easy solution to this problem.
The only one that at all satisfies me is to make the definition apply to
the main body or "core" of the phenomenon and be arbitrary about just
where in the transitional area the boundary should be recognized.

I realize that this will not satisfy anyone who simply wants an
authoritative definition. Some years ago we had a project to compile an
"Oxford Dictionary" type glossary of coral reef terms. ‘Several of us
participated in:the work, with Prof. Rhodes W. Fairbridge as editor. It
was never published. Now the manuscript is being updated by a committee
on terminology set up at the Second International Coral Reef Symposium in
1973, under chairmanship of David R. Stoddart. Hopefully this will see
the light of day sometime soon. However, it will merely bring out the
true demensions of the problem, but will not solve it.

piLereirac

Really, the only thing I can suggest is that you either use the
term atoll and define it to your satisfaction, or, alternatively, dodge
the question by calling Cayo Sal an "atoll-like island."

F. R. Fosberg

AN_ IRRESPONSIBLE SCIENTIFIC EXPEDITION: From August 22 to October 21
an expedition, called Line Islands Expedition, mounted by the Gilbert
and Ellice Islands Government which toured the islands in the central
Pacific which are or will possibly become part of the new independent
Gilbert and Ellice Islands nation. The purpose was to assess the
economic possibilities of the islands and to collect scientific infor-
mation that may be of use in administering these islands by the new
government. The personnel of the expedition were officials of the
government and, in addition, an ornithologist from Hamburg, Germany and
a graduate student from Hawaii interested in corals.

The islands visited were Christmas, Fanning, Washington, Malden,
Starbuck, Caroline, Flint, Vostok, Phoenix, Sydney, Hull, and Gardner.
Reports were submitted to the government by the principal members of the
expedition, either general, or on the specific subjects covered by cer-
tain members, which were duplicated but appear to be unedited. Their
scientific content will not be commented on here. Perhaps it will appear
later in a more finished state.

What is of immediate interest is information in several of these
reports on the unfortunate behavior of crew members of the ship Teraaka,
during their visits to the islands. What transpired was apparently an
orgy of slaughter of all sea-birds that they could lay hands on--nest-
ing birds that trustingly do not leave their nests at the approach of
man. In some instances almost whole populations were killed, to some
extent for food, but mostly just for "sport".

Furthermore, while some of these birds were being cooked, the
thick dry "peat" under the Pisonia forest on Vostok Island was set on
fire. An attempt was made to put out the fire, but it was evidently
not completely extinguised, and was recently reported as still burning.
Vostok is, or was, a perfect gem of an island unaltered by man. It was
a perfect, functioning relatively simple natural ecosystem, a notable
sea-bird rookery, a natural laboratory for the study of phosphate rock
formation processes and other biological interactions and environmental
relations. It was a prime candidate for preservation as an interna-
tionally recognized “island for science." Now, if the report of the
persistence of the fire is true, it will be reduced to a barren and
lifeless waste, a desert island in the worst sense of the term.

If this is an example of the way the citizens of the new nation
behave, right under the eyes of their government officials, the future
of any of the non-human inhabitants of these central Pacific islands
looks dim indeed. And if the biota and the total biological diversity
of these island ecosystems become progressively more and more depleted,
the long-term future of the islanders also looks very dim.

We regard it as a shame that such occurrences took place on a
scientific expedition. We concur fully with the recommendations of
Dr. Grossmann, the expedition ornithologist, on pages 16-18 of his re-
port, as minimum steps to be taken. We agree that education in conser-
vation principles in the schools is the only long-term effective solution
to such problems. It is obvious that the education of the present
generation of adults has been woefully deficient in these matters.

REVIEWS :

Carlquist, S. 1974. Island Biology. 1-660, Columbia University Press,
N. Y. and London, $25.00. No one seriously interested in the scientific

aspects of islands can very well afford not to have this book immediately
available. Not only are few aspects of island biology neglected, but
there are few where significant new information and ideas are not pre-
sented. A new starting point is provided for the study of many important
aspects of this subject. Although there may be many areas where others
will differ from the views expressed by Carlquist, one cannot afford to
neglect to examine his data, ideas, and conclusions critically.

He has been fortunate to have opportunities to amass a vast amount of
actual experience and data on islands and island-like areas throughout
the world. Where he has known of or suspected interesting insular
phenomena he has gone. His institution, Claremont Graduate School, is to
be congratulated on giving him the freedom to do this, as is his mother
for making his early insular explorations possible.

To avoid an encyclopedia-sized work, he has adopted the approach of
only mentioning, not repeating, the detail given already in his own
extensive publications and in readily available works of others. He has
not, of course, omitted points necessary to support the ideas expressed
here, but has as nearly as practical made this a work presenting his own
newly discovered or recorded data and the new ideas and conclusions to
which they have led him. Although casting a wide net for important facts
and ideas in the works of others, and giving full credit for these, this
is essentially his own work and seems fully internally coherent and
consistant--amazing for a 660 page book.

Logically the first chapter deals with dispersal--how the ancestors
of island species may have reached the islands. This is a vexed subject
on which there has been much speculation based mostly on purely circum-
stantial evidence. Carlquist's chapter is no exception to this. It is
perhaps the weakest chapter in the book. It ascribes great effectiveness
to bird transport, on the basis of fleshiness of fruit and such features,
and admits little importance for wind. Familiarity with typhoon-strength
winds and knowledge of "jet streams'' suggests to me that wind may be a
very important agent even for dispersal of plants that are obviously
specially equipped to be carried around locally by birds or other agencies.
Long distance dispersal must, under any circumstances, be an extremely
infrequent and unusual event. This is perhaps not sufficiently stressed
in the chapter on dispersal, which in most respects is very thorough.

One of the features of island evolution that has been most discussed,
and that Carlquist devotes seven chapters to, is the so-called "adaptive
radiation." This is the phenomenon of differentiation of populations of
a group to enable them to occupy a diversity of habitats or ecological
niches, developing distinctive adaptive features and behavior patterns
accordingly. The first of the seven chapters on this subject, discusses
in detail the general features and implications of the concept, and is
certainly the most adequate and thought-provoking treatment of the
subject that I have seen. In particular, it offers an explanation of why
adaptive radiation seems so conspicuously a feature of oceanic island
biotas. This he relates to the "disharmonic" nature of these biotas,
the incomplete and unbalanced representation of the broad spectrum of
families normally found in continental areas. Thus lessened competition,
or evolutionary vacua, permit expression of evolutionary potentials in
a particular group that would be prevented by prior effective occupancy
of most of the niches by members of other groups. This principle, while
implicit in some previous discussions of adaptive radiation, is here
brought out with admirable clarity and with abundant illustrative examples
and details It is a major accomplishment of the book. The other six
chapters on the subject are detailed expositions of the occurrence of
adaptive radiation in the principal geographical regions where it is
conspicuous.

The following chapters are, with one exception, devoted to classes
of features prominent in island biotas that are not, or at least not
obviously, the result of adaptive radiation or clear responses to habitat
factors. These include woodiness in usually herbaceous plant groups;
gigantism; types of loss of, or reduced, dispersability such as flight-
lessness, reduction of obvious dispersal mechanisms, increase in size of
propagules, and other less striking but possibly equally pertinent
phenomena; peculiar developments in reproductive biology; reduction,
modification or loss of appendages in animals; and other topics related
to or correlated with insularity.

Chapter 15 is a lengthy essay on equatorial highlands and mountains
showing how they partake of the nature of islands, sharing some of the
features discussed earlier in the book for true islands. An impressive
amount of information is brought together under this head. In this, as
here and there in previous chapters, one occasionally gets an impression
of strained interpretations, but these are sometimes most thought-provoking.

Although the coverage of the whole subject of insular phenomena is by
far the most thorough ever written, it is done in such a fashion as to be
challenging. Further work and discussion are provoked and stimulated
rather than cut off. A tremendous field, full of problems, is laid out.
Reference to previous documentation is extensive. Each chapter has a
large list of cited references. The material in the whole work is made
readily accessible by an index of biological names used and by a remarkably
detailed subject index.

Lest the above generally laudatory comments lull uncritical readers
into accepting everything in the book as gospel, a few particular points

i/

may be indicated where erroneous statements have crept in or where there

is room for strong difference of opinion: Page 8, the Tahitian cotton

is not endemic, even though it has been called Gossypium taitense. P. 11,
the last two sentences seem to be somewhat contradicted by the discussion
of geckos in the last paragraph on p. 8. Pp. 31 and 38, the date of the
Fosberg Gouldia reference should be 1936. P. 63, Peperomia does not have
"relatively large seeds." P. 70, rather than Boerhavia diffusa the Pacific
Island Boerhavia is mostly B. repens, which is notably polymorphic species.
The suggestion that pantropical strand species lack variation only indicates
that slight attention has been paid to them, probably because of the wide-
spread idea that they lack variation. P. 80, Oeno Atoll is not a raised
atoll. P. 86, Rhipsalis definitely shows differentiation, at least in
Madagascar. P. 131, the expression "most mesic" suggests that mesic is
synonymous with wet, rather than intermediate between xeric (dry) and
hydric (wet). Pp. 212-213, the statement that geological sequences of
islands from old to young are not present in southern Polynesia is defi
nitely erroneous, as such sequences are very clear in the Society, Cook

and Austral island groups. P. 359, there are many lowland arborescent
plants in the Galapagos, at least, that are not stem-succulents, e.g.
Croton, Bursera, Acacia, Pisonia. P. 450, an exception to the statement
that Pacific Bidens have not advanced far toward arborescence is

B. hendersonensis which is truly a small tree, some meters tall and with

a strong, hard woody trunk. P. 480, contrary to the statement that neither
fruits nor seeds of Sapindus are known to be capable of flotation, seeds

of Sapindus saponaria are not uncommon in tropical beach drift. P. 483,
Pleomele, a weak segregate from Dracaena, is definitely not an endemic
Hawaiian genus, but includes species in other areas.

The book is generally attractively put together, with abundant
illustrations, as well as excellent drawings. Many of the author's well-
known superb photographs are included, but the quality of the reproduction
of many of them is disappointing. The captions of plates 14.14, 14.15,
and 14.17 are transposed. The lines are rather long for comfortable
reading.

In spite of such minor criticisms, Island Biology is a tremendous
achievement and establishes Carlquist as probably the greatest name in

the contemporary island biological field.

F, R. Fosberg

Wood, E. J. F. and R. E. Johannes. Tropical Marine Pollution. 192 pp.
Elsevier Oceanography Series 12, Elsevier, Amsterdam, Oxford, N. Y.

1975. $26.95. It is not often that a review of a major subject can come
close to covering all that is known about the subject. That the editors
of this book could not only succeed in doing this, but include one major
original research report, is an indication of how meager our knowledge of
tropical marine pollution is. The paucity of information, however, is not
an indication that the subject is unimportant. Enormous populations in
tropical countries and on tropical islands live in immediate proximity to
the sea and are dependent on it for part of their nutrition. The biota

of the tropical oceans are enormous and diverse. The growing threat of
death from man-caused pollution hangs over all tropical marine and maritime
plants and animals, including man.

Why tropical marine pollution? Johannes, in his introductory chapter,
deals with this question and shows clearly that knowledge derived from
temperate situations cannot necessarily be extrapolated to tropical ones.
Tropical organisms are usually living much nearer their lethal temperature
threshold than are those in temperate waters, and that other physiological
tolerances are also narrower in the tropics. He shows in tabular form
a long list of comparisons of pertinent features of tropical organisms
with corresponding temperate ones.

The other chapters in the book are:

Chapter 1. INTRODUCTION: MARINE COMMUNITIES RESPOND DIFFERENTLY TO
POLLUTION IN THE TROPICS THAN AT HIGHER LATITUDES (R. E.
Johannes and Susan B. Betzer)

Chapter 2. POLLUTION AND DEGRADATION OF CORAL REEF COMMUNITIES (R. E.
Johannes)

Chapter 3. THE RESPONSE OF MANGROVES TO MAN-INDUCED ENVIRONMENTAL
STRESS (William E. Odum and R. E. Johannes)

Chapter 4. TROPICAL SEA GRASS ECOSYSTEMS AND POLLUTION (J. C. Zieman)

Chapter 5. EFFECTS OF THERMAL POLLUTION ON TROPICAL-TYPE ESTUARIES,
WITH EMPHASIS ON BISCAYNE BAY, FLORIDA (J. C. Zieman and
E. J. Ferguson Wood)

Chapter 6. BIOLOGICAL IMPACT OF A LARGE-SCALE DESALINATION PLANT AT

KEY WEST, FLORIDA (Richard H. Chesher)

9

From our viewpoint Johannes" own masterful treatment of the impact
of pollution on coral reefs is the core of the book. Probably no one
would have been nearly as capable of writing this account as Bob Johannes.
Anyone concerned with the future of coral reefs will find it of great
interest.

The chapters on mangroves, by Odum and Johannes and on sea-grass
ecosystems by Zieman will also be of concern to many of our readers.

Suffice it to say that the tropical marine ecosystems are already in
trouble, and that the trouble will surely get worse as long as the sea
is used as the world's cesspool.

I have only one criticism of the editing that is worth mentioning.
The habit of abbreviating generic names in binomials is a great nuisance
and source of confusion when the full binomial has not been spelled out
at least on the same page, preferably in the same paragraph. It is
particularly bad when several generic names are abbreviated which begin
with the same initial letter.

The book is expensive, but is essential to any tropical marine
biologist. It has also the virtue of being readable (except parts of
Chesher's chapter) and interesting.

F. R. Fosberg

Johannes, R. E., et al. The metabolism of some coral reef communities;
RD A IN A NE NS SS eS

a_ team study of nutrient and energy flux at Eniwetok. BioScience 22(9):
541-543. 1972. The high biological productivity of atoll coral reefs

bathed by nutrient-poor oceanic waters has been long appreciated but

poorly understood. This preliminary report heralds some of the interesting
results from a formidable team effort to illumine some of the processes
that maintain a coral reef ecosystem.

From their base aboard the research vessel Alpha Helix of the Scripps
Institute of Oceanography in May-June, 1971, a team of scientists was
able to simultaneously monitor a large variety of changes in chemical,
biological and physical properties of sea water flowing over the windward
reef at Eniwetok Atoll near the old transect laid out by Odum and Odum in
their pioneering reef ecosystem study (Ecol. Monogr. 24: 291-320, 1955).

Nitrogen fixation proceeds at very high rates in the system, result-
ing in a net export of that element from the reef. N fixation in turn is
allied to efficient utilization of phosphorus, very little of which is
lost from the benthic reef community to the water. Phosphorus, it is
suggested, must be recycled very efficiently between photosynthetic
organisms and heterotrophic ones. In this context it is interesting that
the presence of endozoic algae in the tissue of corals and other reef
organisms is directly correlated with low release of dissolved organic
and inorganic phosphorus.

Bryce Decker

ee Te

10

King, Warren B., ed. Pelagic studies of seabirds in the Central and
Eastern Pacific Ocean: Smithsonian’ Contr. Zool: 158; 1=277% 1974:

Those who have seen the concentrations of seabirds in island rookeries
during breeding seasons and their almost total absence on the same islands
during other parts of the year can scarcely help wondering where they go.
For some species this information has been very scanty, indeed, in the
past. The Pacific Ocean Biological Survey Program of the Smithsonian
provided excellent opportunities to work on this problem, and this publi-
cation is a beginning toward making the results of these observations
available. It includes seven papers treating the Sooty Fern, the Wedge-
tailed Sheawater, the Black-footed and Laysan Albatrosses, 18 species of
Storm Petrels, and the Red-tailed Tropic Bird in detail. The papers are
abundantly illustrated with maps, diagrams and tables, and the volume
contains a short, by no means exhaustive, list of pertinent references to
previous work. Those interested will await the future numbers of the series
that will treat other bird species.

F, R. Fosberg

McGregor, C. The Great Barrier Reef. Time-Life Books, Amsterdam, 1-184.
1974. This is a beautifully illustrated account of one of the most

remarkable and magnificent features on the face of the earth. It can be
highly recommended to the non-technical person who has an intelligent
interest in the unusual features of the world and an appreciation for
outstanding natural beauty. The author emphasizes the fact that even a

1260 mile-long complex of reefs, channels, lagoons, and island is threatened
with exploitation and degradation at the hands of man in this greedy age.

F. R. Fosberg

TANE vols. 17: 1-212, 1971; 18: 1-200, 1972. This handsome little
journal is published by the Auckland University Field Club, and carries
articles on New Zealand natural history, with much material on the .off-
shore islands around New Zealand. It is especially dedicated, though not
restricted, to the publication of articles written by students, reporting
their own original field studies and observations. Within this framework
the articles maintain a very high standard and are both well written and
well edited. The content runs the gamut of outdoor observational natural
history, including archaeology.

These two recently received volumes are of special interest in that
each reports the results of a student-organized expedition, or annual
field camp, to a small island. Vol. 17 contains 8 articles resulting
from the 1970 Whale Island field camp, as well as articles on other small
islands and several on mainland observations. Vol. 18 is largely taken
up by 15 articles resulting from the 1971 Red Mercury Island field camp.

One cannot commend too highly this type of training in field biology.
It at the same time introduces beginning biologists to serious scientific
study, with results valuable to science, and provides them an incentive
to maintain a quality in their observations high enough to stand the
scrutiny of their scientific colleagues normally directed to published

ila

work. The results, as shown by these articles, are very worth-while,

The subscription price of TANE is at present $1.00 per volume.
Correspondence should be addressed to:

TANE

c/o The Botany Department
University of Auckland
Private Bag, Auckland, N. Z.

F. R. Fosberg

12

SHORT ARTICLE

Remarks on the Botany and Statistics
of the Bahama Islands

by William T. Gillis
NATURAL HISTORY OF THE BAHAMA ISLANDS

The Bahama Islands consists of nearly 30 major islands, 661 smaller
ones called cays (pronounced "keys''), and nearly 2400 rocks, jutting over
a shallow platform with ocean deeps surrounding it. The total area is
over 4,400 square miles. The islands stretch in a giant arc 760 miles
long from northwest to southeast, with greatest width about 150 miles
(from Gun Cay to Man-of-War Cay, Abaco). The archipelago straddles the
Tropic of Cancer with Georgetown, Great Exuma lying just a few miles
north of ‘this line. -— The span i's’ from .20° 55" to 27° 30° INe maiteeand
from 71° 10' to 79° 20' W. Long. which places the islands in the trade
wind belt.

Topography is somewhat monotonous, the highest point on any of the
islands being Mt. Elvernia on Cat Island with an elevation of 210 feet.
Much of the island group (e.g., the western side of Andros, the southern
half of Middle Caicos, etc.) is known as "swash" and is subject to over-
flow by the sea in very high tides and in storms. Large salt-water lakes
at sea level occur on several islands, there appearing to be no connection
with the sea except likely underground. There are only two so-called
rivers in the whole archipelago, both on Andros. Other seemingly fresh-
water lakes are high in calcium ion concentration.

The Bahamas enjoy a climate of the tropical savanna type (Aw in the
Koppen system), with no frost ever recorded. Rainfall decreases southward
from about 50-60 inches in the north to less than 20 inches in Inagua.
Hurricanes are likely to strike some part of the islands every few years.
Because of the limestone substrate's presence close to the surface, only
about 35,000 acres of land are presently under cultivation. A dairy herd
is maintained on Eleuthera. An abandoned sugar mill stands on Abaco.
Native pine has been harvested on the northern islands but not touched
commercially in the disjunct area in the Caicos group (it has been bull-
dozed there however).

The flora consists of about 1000 species of vascular plants, about
30 of which are endemic. The vegetation types appear to be fairly simple,
consisting mainly of pineland, coppices (not unlike the hammocks of
southern Florida), scrublands of shrubs and small trees (called simply
"bush" in the islands), strands and mangroves. The floristic relation-
ships of the flora and vegetation are generally with Florida, Cuba,
Hispaniola, and to a lesser extent, Puerto Rico and Yucatan.

[Supporting papers for a talk given at the annual meeting of the Associa-
tion for Tropical Biology at the Smithsonian Institution, December 1974,
that we thought might interest some of our readers.--Eds. ]

13

There are two political units within the geographical region called
the Bahamas: the Commonwealth of the Bahamas (which received independence
within the British Commonwealth on 10 July 1973), and the Turks and Caicos
Islands, a Crown Colony of Great Britain. Prior to Jamaican independence,
the Turks and Caicos were administered as part of Jamaica. Since 1848,
they have not been affiliated with the Bahama Islands politically. From
April 1972 until a few months ago, there were no commercial transportation
connections between these two political units!

(prepared December 1974)
PHYTOGEOGRAPHIC HYPOTHESES RELATING TO BAHAMA FLORA

1. All plants of the Bahamas have had a source area outside the Bahamas.
For endemics, the progenitors were from outside the region, and are
probably missing from the region now.

2. Bahama plants which are also in Florida are chiefly restricted to the
Florida Keys, or at least south of Lake Okeechobee. A few species
have entered the Bahamas from Florida. These are restricted to the
northern islands (e.g. Asters, poison ivy).

3. Plants which do not cross the Gulf Stream from the Bahamas into
Florida by and large have no special adaptation for wind dispersal,
nor are eaten by birds, nor have any special niche in South Florida.

4. Most new plants in the flora (that is, have entered the region in
recent years) have probably been introduced by wind (especially
hurricanes), by rafting, or by birds. Some inadvertent introduction by
activities of man is probably also at work.

5. Endemism increases with the size of islands (banks), lessening of
rainfall, and decreasing latitude.

6. San Salvador has a greater affinity with Hispaniola in its fauna and
flora than can be explained by proximity.

7. Each island has a distinctive assemblage of plants, with different
dominants on each island. Exception: Abaco and Grand Bahama.

8. Toward the south, the following trends are noted:
Strand plants of the north move into upland thickets.
Species develop broader leaves.
More species occur that have gray, dead-looking foliage.

There is a tendency for the plants present to be more drought-
resistant.

14

9. Anomalous situations:
Why is Grand Turk's flora so similar to that of South Caicos?
What is the cause of the present distribution of pine?

What prevents the Cuban element in the flora of South Andros from
crossing into Mangrove Cay or North Andros?

Why does the "southern element" in the flora not move up
the Exuma Cays to New Providence?

(Prepared 1974)
PLANT DISTRIBUTION IN THE BAHAMA ISLANDS

Cosmopolitan species - mangroves
strand plants

Bahamas —- Cuba

South Andros - Cuba - Catalpa punctata
Pseudocarpidium wrightii
Callicarpa hitchcockii
Phoradendron trinervium
Crossopetalum aquifolium

Bahamas -— Yucatun - Mimosa bahamensis
Bahamas — Hispaniola

San Salvador - Hispaniola - Croton discolor
Zanthoxylum bifoliolatum

Aberrant - Chamaesyce vaginulata
' Limonium bahamense
Nashia inaguense
Caesalpinia murifructa

Bahamas - Florida

(a) widespread in Florida
(b) southern tip of peninsula or Florida Keys

Plants on Bimini and/or Grand Bahama and not in Florida:

Thouinia discolor Haematoxylum campechianum
Diospyros crassinervis Rhachicallis americana
Tabebuia bahamensis

Phialanthus myrtilloides Cassia lineata

Triopteris jamaicensis Buxus bahamensis
Stigmaphyllon sagraeanum Erythroxylum spp.

Grimmeodendron eglandulosum

Endemic - 2

1 species of Vernonia
1 species of Marsilea
1 species of Lobelia
1

species of Cynanchum

species of Agave (Section Bahamana)

Cuba —- Cay Sal - Chamaesyce centunculoides

APPROXIMATE SIZES OF THE MAJOR ISLANDS

Abaco

Acklins

Andros

Berry Islands

Bimini

Cat

Crooked

Eleuthera

Exuma

Exuma Cays and Little Exuma
Grand Bahama

Inagua (incl. Little Inagua)
Long

Mayaguana

New Providence

Ragged

Rum Cay

San Salvador

Cay Sal and Lobos

Dimensions

105

SY

Td

43

19

90

40

73

40

64

26

21

ifah

15

X 7 miles

X 4 miles

X

22 miles

7

14 miles

4

5

7

miles

miles

miles

miles

miles

miles

miles

miles

miles

miles

Square Miles

372
12.
2,300

uy

30
60

2

(Lands and Surveys Dept., Ministry of Development, Commonwealth
of the Bahamas, Statistical Abstract, 1970).

a5

16

POPULATION OF THE BAHAMAS

Abaco 6550
Acklins 936
Andros 8,845
Berry Islands 443

Bimini (and Cay Lobos) 1503

Cat 2657:
Crooked 689
Eleuthera 9,468
Exuma Sh OW,
Grand Bahama 25,859
Harbour Island and
Spanish Wells BETS

Inagua 1,109
Long Cay (Fortune) 26
Long Island 3,861
Mayaguana 581
New Providence TOU 503
Ragged 208
Rum Cay 80
San Salvador 776
TOTAL 168,812 (1970 Census)

(Dept. of Statistics, Cabinet Office, Commonwealth of
the Bahamas, Statistical Abstract, 1970)

* The population of these islands is also included in the figure for
Eleuthera. The grand total does not count this figure twice.

1)

List of Atoll Research Bulletin
titles issued to date

To avoid the correspondence resulting from numerous requests for titles
of all ARB numbers issued to date we are listing them here. We also take
this opportunity to inform those interested that numbers 1-117 are entirely
exhausted or almost so. A few copies of some of them are left but will
only be sent to fill gaps in sets in libraries of institutions actively
engaged in research on islands. A $10 per number charge plus a $5 issue
handling charge will be assessed for supplying these, the money to be
paid into the Smithsonian Institution ARB fund.

1. Basic information papers, by various authors. 1-25, Sept. 10, 1951.

2. Symposium on coral atoll research, by various authors. 1-14,
Sept. LO.) 1951,

3. Vertebrate ecology of Arno Atoll, Marshall Islands, by J. T. Marshall,
jhe,» dle srl (leien TES) Tile )bule

4. Marine zoology study of Arno Atoll, Marshall Islands, by R. W. Hiatt
and, D. Strasbune. =13. Oct. 15, 1951.

Saotne Soils of Arno Atoll, Marshall’ Islands, by E. L. Stone, Jr. 1-56,
None. 5)9 LOSI.

6. The agriculture of Arno Atoll, Marshall Islands, by E. L. Stone, Jr.
146, Novs' 15, 1951:

7. The plants of Arno Atoll, Marshall Islands, by D. Anderson. 1-4,
PavAs, Nov. Lo, 951.

8. The hydrology of Arno Atoll, Marshall Islands, by D. C. Cox. 1-29,
Bece (5) L951.

9. The coral reefs of Arno Atoll, Marshall Islands, by J. W. Wells.
it—4 Deed 155° 1951.

10. Anthropology-geography study of Arno Atoll, Marshall Islands, by
L. Mason, J. Tobin and G. Wade. 1-21, Sept. 1, 1952.

11. Land tenure in the Marshall Islands, by J. Tobin. 1-36, Sept. 1, 1952.

12. Preliminary report on geology and marine environment of Onotoa Atoll,
Ciibent islands, byub. wienGloud. Jnr. W773. Dee. WS, 1952.

13. Preliminary report on marine biology study of Onotoa Atoll, Gilbert
islands. by A. H. “Banner, and) J. Es Randall: 1-625 "Dec. 15, 1952.

14. Description of Kayangel Atoll, Palau Islands, by J. L. Gressitt.
iG Dees elas L952

231-339 O - '77 - 18

18

15a

m6.

7s

iS).

1S)

20.

Zils

Lis

BBN.

24.

2s

26.

Zl.

28%

Zo).

30.

SARE

32%,

The insect life of Arno, by R. L. Usinger and I. La Rivers. 1-28,
April 30, ° 1953).

The land vegetation of Arno Atoll, Marshall Islands, by W. H.
Hatheway. 1-68, April 30, 1953.

Handbook for atoll research, by various authors, edited by) Ea Re.
Fosberg and Marie-Héléne Sachet. 1-129, Mary,1 55+ 1953.

Ichtyological field data of Raroia Atoll, Tuamotu Archipelago, by
R. R. Harry. 1-190,. July. 31,. 1953.

Check list. of: atolls, by E. H.. Bryan, Jr...,1-38, Sept... 30, 19537

Health report of Kapingamarangi, by R.; Es ;Miller,...1-42,..Sept.u30,
1953%. ;

Notes on Ngaruangl and Kayangel Atolls, Palau Islands, by J. L.
Gressitt. lp, sept. 30, pLO5 30

Summary of information atoll soils, by E. L. Stone, Jr. 1-4, Sept.
305.1953" ;

Vegetation of Central Pacific Atolls, a brief summary, by F. R.
Rosberg. ~ 1-76. Sept... 30, 1953.

Enumeration of the decapod and stomatopod Crustacea from Pacific
coral islands, by L. B. Holthuis. 1-66, Nov. 15, 1953.

Bryophytes from Arno Atoll, Marshall Islands, by H. A. Miller and
Mas Doty.” T-0> Nov. 15, 1953.

Scorpions on coral atolls, by M.-H. Sachet. 1-10, Nov. L5H Q5ar
Nutrition study in Micronesia, by M. Murai. 1-239, Jan. 31, 1954.

Preliminary report on land animals at Onotoa Atoll, Gilbert Islands,
by E. T. Moul. 1-28, May 31, 1954.

A summary of information on Rose Atoll, by M.-H. Sachet. 1-25,
May 31, 1954.

The hydrology of the Northern Marshall Islands, by T. Arnow. 1-7,
May 31, 1954.

Expedition to Raroia, Tuamotus, Part 1. Expedition to Raroia, Tuamotus,
by N. D. Newell. 1-12; Part 2. Physical characteristics of Raroia,

by N. D. Newell. 13-21; Part 3. General map of Raroia Atoll, by N. D.
Newell. Nove 30, 1954.

Raroian Culture, Part 1. Economy of Raroia Atoll, Tuamotu Archipelago,
by B. Danielsson. 1-91; Part 2. Native topographical terms in Rarota,

;
I

5) 34

34.

a5

36.

ear

38.

39.

40.

4l.

42.

AS

44.

45.

19

Tuamotus, by B. Danielsson. 92-96; Part 3. Native terminology of
the coconut palm in Raroia Atoll, by B. Danielsson. 97-99; Part 4.
Bird names in Raroia Atoll, by B. Danielsson and A. Natua. - 100-101;
Part 5. Check list of the native names of fishes of Raroia Atoll,
by B. Danielsson. 102-109, Nov. 30, 1954.

Floristics and plant ecology of Raroia Atoll, Tuamotus, Part 1.
Floristic and ecological notes on Raroia, by M. S. Doty. 1-41; Part
2. Ecological and floristic notes on the Myxophyta of Raroia, by

J. Newhouse. 42-54; Part 3. Ecological and floristic notes on the
Bryophyta of Raroia, by H. A. Miller and M. S. Doty. 55-56; Part 4.
Ecological and floristic notes on the Pteridophyta of Raroia, by K.
Wilson. 57-58, Nov. 30, 1954.

Animal ecology of Raroia Atoll, Tuamotus, Part 1. Ecological notes on
the mollusks and other animals of Raroia, by J. P. E. Morrison.

1-18; Part 2. Notes on the birds of Raroia, by J. P. E. Morrison,
19-26, Nov. 30, 1954.

Interrelationship of the organisms on Raroia aside from man, by M. S.
Doty and J.P. E. Morrison. 1-61, Nov. 30, 1954.

Reefs and sedimentary processes of Raroia, by N. D. Newell. 1-35,
Nov.’ 30, 1954.

Pumice and other extraneous volcanic materials on coral atolls, by
Meri. sachet. (1-27, May 15, 1955.

Northern Marshall Islands Expedition, 1951-1952. Narrative by F. R.
Fosberg. 1-36, May 15, 1955.

Northern Marshall Islands Expedition, 1951-1952. Land biota:
Vascular plants, by F. R. Fosberg. 1-22, May 15, 1955.

Bryophytes collected by F. R. Fosberg in the Marshall Islands, by
BoA. Midler.) 4, May 15, 1955.

Canton Island, South Pacific, by 0. Degener and E. Gillaspy. 1-51,
Aves * 15.5. ° 2955*

The insects and certain other arthropods of Canton Island, by R. H.
Van Zwaluwenburg. 1-11, Aug. 15, 1955.

The natural vegetation of Canton Island, an equatorial Pacific atoll,
by W. H. Hatheway. 1-9, Aug. 15, 1955.

The hydrology of Ifalik Atoll, Western Caroline Islands, by T. Arnow.
TES, “Aug. Loy 1955).

A partial list of the plants of the Midway Islands by J. A. Neff and
Be A. “DUMOMe .! LL eAne. 1 Sg S 5.

[i oe OF on Pes cell Es el es et a

N

20

46.

47.

48.

49.

50%.

a1.

a2.

Diss

54.

De

56.

57.

58.

59.

60.

6i.

62;

63%

Conspicuous features of organic reefs, by J..I. Tracey, Jr., P. E.
Cloud,. Jr..and K. 0. Emery.,. 1-3; Aug..15, 1955:

Fishes of the Gilbert Islands, by J. E. Randall. 1-243, Aug. 15,
HO55%

The geography of Kapingamarangi Atoll in the Eastern Carolines, by
Herold J. Wiens. 1-86, 1- [7], June 30, 1956.

Bioecology of Kapingamarangi Atoll, Caroline Islands: Terrestrial
aspects, by William A. Niering. 1-32, June 30, 1956.

Geology of Kapingamarangi Atoll, Caroline Islands, by Fdwin D. McKee.
1-38, June 30, 1956.

Observations on French Frigate Shoals, February 1956, by Arthur
Svihla. sl=2, Sept. 15,21957.:

Zonation of corals on Japtan Reef, Eniwetok Atoll, by Eugene P. Odum
and Howard T. Odum. 1-3, Sept. 15, 1957.

Slicks on ocean surface downwind from coral reefs, by F. R. Fosberg.
1-4, Sept. 15, 1957.

Field notes on atolls visited in the Marshalls, 1956, by Herold J.
Wiens. 1-23, Sept. 15, 1957.

Agricultural notes on the Southern Marshall Islands, 1952, by W. H.
Hatheway. 1-9, Sept. 15, 1957.

Atolls visited during the first year of the Pacific Islands Rat
Ecology Project,. by J.T. Marshall, i Jr... L-11,;:Sept..15.,.4957¢

Preliminary report on the flora of Onotoa Atoll, Gilbert Islands, by
Edwin T.. Moul. 1-48, Sept. 15,.1957.

The Maldive Islands, Indian Ocean, by F. R. Fosberg. 1-37, Sept.
155 £95 Te

Report on the Gilbert Islands: Some aspects of human ecology, by
Rene L.A. -Catala. . 1-187, -0ct./ 31, ,1957.

Climate and Meteorology of the Gilbert Islands, by M.-H. Sachet.
1-4, Octs 31,1957.

Long-term effects of radioactive fallout on plants? by F. R. Fosberg.
ded. Maye 15.1959).

Health and sanitationsurvey of Arno Atoll, by J. D. Milhurn. 1-7,
May -15,, 1959.

Report on a visit to the Chesterfield Islands, September, 1957, by
by-F.. Cohic. 1-11, May 15,1959.

64.

65.

66.

67.

68.

69.

70.

WA

Pie

733

74.

ipe\e

Vo

Lhe

1S

79.

80.

Zi

Canton Island, South Pacific (Resurvey of 1958), by Otto Degener and
Isa Degener. 1-24, May 15, 1959.

Some marine algae from Canton Atoll, by E. Yale Dawson. 1-6, May
Pie V95or

Notes on the geography and natural history of Wake Island, by E. H.
Bryan, Jr, l=—22), May sl5,7:1959.

Vegetation and flora of Wake Island, by F. R. Fosberg. 1-20, May
Ey, BLOS 9.

Additional records of phanerogams from the northern Marshall Islands,
by! FE. R. Fosberg.. 1-9), May 15, 1959.

Contribution to a German reef-terminology, by Georg Scheer, 1-4,
May 15, 1959.

Atoll news and comment, Editors. 1-7, May 15, 1959.

Microclimatic observations at Eniwetok, by David I. Blumenstock and
Daniel F. Rex, with a special section on Vegetation by Irwin E. Lane.
tix. 158, June 30), 1960:

Report on Tarawa Atoll, Gilbert Islands, by Edwin Doran, Jr. 1-54+24,
Vee. /5, 1960.

Some aspects of Agriculture on Tarawa Atoll, Gilbert Islands, by
R. R. Mason. AIOE OGE. USA 1960.

Birds of the Gilbert and Ellice Islands Colony, by Peter Child, 1-38,
Det e5) L960.

A report on Typhoon Effects upon Jaluit Atoll edited, by David I.
Blumenstock. 1-105, April 15, 1961.

Observations on Puluwat and Gaferut, Caroline Islands, by William A.
Niering. 1-10, December 31, 1961. Historical and climatic
information on Gaferut Island, by Marie-Héléne Sachet. 11-15,

Dec. 315, L9G

A check list of marine algae from Ifaluk Atoll, Caroline Islands,
by Isabella A. Abbott. 1-5, Dec. 31, 1961.

Narrative report of botanical field work on Kure Island, 3 October
1959) to 9 October 1959s ebyytorace, RB (Clay./ 1-4;9oDee. 31591961"

Botanical observations on Leeward Hawaiian Atolls, by Charles H.
Lamoureux. 1-10, Dec. 31, 1961.

The tropical coral reef as a biotope, by Sebastian A. Gerlach, 1-6,
Decne Sa oaks e

22

81.

82.

832

84,

BDis

86.

87.

88.

89.

90%

Ws

923

93%

94.

95%.

96%

97.

987.

Qualitative description of the coral atoll ecosystem, by F. R.
Fosberg.. te l—bas -Dece,, Siamio6il:

|
|
|

Heron Island, Capricorn Group, Australia, by F. R. Fosberg, R. F.
Thorne and J. M. Moulton. 1-4, 5-13, 15-16, Dec. 31, 1961.

Notes on-some of the Seychelles Islands, Indian Ocean, by C. J.
Pregote,,/ 1-10, Dec. 7314 1961.

Atoll News and Comments. Editors, 1-14, Dec. 31, 1961.

Land tenure in the Pacific - A symposium of the Tenth Pacific Science
Congress convened by Edwin Doran, Jr. 1-60, Dec. 31, 1961.

Geography and land ecology of Clipperton Island, by Marie-Héléne
Sachet. 1-115,-Feb. 28, 1962.

Three Caribbean atolls: Turneffe Islands, Lighthouse Reef, and
Glover's Reef, British Honduras,. by D. R. Stoddart. 1-151, June
305219625

Coral Islands, by Charles Darwin, with introduction, map and
remarks by D. R. Stoddart. 1-20, Dec. 15, 1962.

Geophysical observations on Christmas Island, by John Northrop.
P=2ea,Dec. “155. 1962:

Plants of Christmas Island, by Alvin K. Chock and Dean C. Hamilton,
Jee: :1l=7,5. Dees 155.1962:

Central subsidence. A new theory of atoll formation, by Hans Hass.
1=4,))Dec.115,° 1962;

Vascular plants recorded from Jaluit Atoll, by F. R. Fosberg and
M.=H. ‘Sachet:| -.1-39,, Dec. ' 155 429622

A brief study of the cays of Arrecife Alacran, a Mexican atoll, by
F. Re Fosberg.. 1-25; Decs:i15, 11962.

Atoll news and comments, Editors, 1-19, Dec. 15, 1962.

Effects of Hurricane Hattie on the British Honduras Reefs and Cays,
October 30-31, 19615: by’ D. R.-Stoddart, +1=142),\May 15 }A1963n

Some aspects of the meteorology of the tropical Pacific viewed from
an atoll, by Ronald L. Lavoie. 1-80, May 15, 1963.

The flora and vegetation of Laysan Island, by Charles H. Lamoureux.
1-14, Nov. 15, 1963.

Insects and other invertebrates from Laysan Island, by George D.
Butler, Jr., and Robert L. Usinger. 1-30, Nov. 15, 1963.

aor

100.

LOT:

102;

TOA

104.

105.

106.

1075

108.

109.

UO),

at.

ie

1 fal 8

ha

Ae5 I.

116.

23

Notes on the Wedge-tailed Shearwater at Heron Island, Great Barrier
Reef, Australia, by A. 0. Gross, J. M. Moulton, and C. E. Huntington.
1-11, November 15, 1963.

Atoll news and comments, Editors, 1-16, November 15, 1963.

Notes on Indian Ocean atolls visited by the Yale Seychelles
Expedition, by Alan J. Kohn. 1-12, Sept. 30, 1964.

Notes on reef habitats and gastropod molluscs of a lagoon island
at North Male Atoll, Maldives, by Alan J. Kohn. 1-5, Sept. 30, 1964.

Observations on the birds of French Frigate Shoal and Kure Atoll,
by Miklos D.F, Udvardy and Richard E. Warner. 1-4, Sept. 30, 1964.

Carbonate sediments of Half Moon Cay, British Honduras, by D. R.
Stoddart. 1-16, Sept. 30, 1964.

Floristic report on the marine benthic algae of selected islands
in the Gilbert Group, by Roy T. Tsuda. 1-13, Sept. 30, 1964.

New records of Halimeda and Udotea for the Pacific area, by Edwin
EeMouls , fu, Sept. 30, 1964.

Place names on Nukuoro Atoll, by Vern Carroll, 1-11, Sept. 30, 1964.

Atoll News and Comment, Issued by The Pacific Science Board 1-8,
Sept. 30, 1964.

A preliminary list of the algal flora of the Midway Islands, by
Richard G. Buggeln, 1-11, July 15, 1965.

Marine algae from Laysan Island with additional notes on the
vascular flora, by Roy T. Tsuda. 1-37, July 15, 1965.

An annotated bibliography of recent papers on corals and coral reefs,
by John D. Milliman. 1-58, July 15, 1965.

Atoll News and Comment, Issued by The Pacific Science Board. 1-14,
July 1551 1965

Terrestrial Sediments and Soils of the Northern Marshall Islands,
by F. Raymond Fosberg and Dorothy Carroll. 1-156, Dec. 31, 1965.

Northern Marshall Islands land biota: Birds, by F. R. Fosberg.
1-35, March 31, 1966.

Marine benthic algae from the Leeward Hawaiian Group, by Roy T.
Tsuda. 1-13, March 31, 1966.

Reef studies at Addu Atoll, Maldive Islands, edited by David R.
Stoddart. 1-122, March 31, 1966.

=z

oad

4
¢
C
;

LRI@GOWITtIiTinat

CRLITLIOMAIIA AL

N

Pa

as

miuirkn

re rw A

AlInrr;~imri

24

JLILY/ 6

Nos.

Atoll News and Comments, Issued by The Pacific Science Board, 1-8,
March 31, 1966.

1-117 were issued by the National Academy of Science--N.R.C. The

following issued under the sponsorship of the Smithsonian Institution,
Washington, D. C.

118:
9).
120.

IW A1EG
W726

23).
124.
123%
126).
IEZ TEE
128

AST

130.

ANS WES

Ecology of Aldabra Atoll, Indian Ocean, edited by David R. Stoddart.
1=T4l. Nov. (5, 19677

Atoll News and Comment. 1-6, Nov. 15, 1967.

A Record of Benthic Marine Algae from Johnston Atoll by Richard G.
Buggeln and Roy T. Tsuda. 1-20, March 30, 1969.

The Algae of Kapingamarangi Atoll, Caroline Islands. Part l.
Checklist of the Cyanophyta, Chlorophyta and Phaeophyta by Jan
Newhouse. 1-7, March 30, 1969;

Marine Toxins from the Pacific II. The Contamination of Wake
Island Lagoon by Albert H. Banner, Judd C. Nevenzel and Webster R.
Hudgins. 1-9, March 30, 1969.

Wake Island Vegetation and Flora, 1961-1963 by F. R. Fosberg and
M.-H. Sachet. 1-15, March 30, 1969.

Ecology of Terrestrial Arthropods on the Tokelau Atolls by Alden D.
Hickley. 1-18, March 30, 1969.

Reconnaissance Geomorphology of Rangiroa Atoll, Tuamotu Archipelago
by D. R. Stoddart with List of Vascular Flora of Rangiroa by
Marie-Héléne Sachet. 1-44, March 30, 1969.

Island News and Comment. 1-19, March 30, 1969,

Ornithology of the Marshall and Gilbert Islands by A. Binion
Amerson, Jr. May 28, 1969.

Notes on Birds Observed in the Comoros on Behalf of the Smithsonian
Institution by A. D. Forbes-Watson. 1-23, Aug. 15, 1969.

Four Southwestern Caribbean Atolls: Courtown Cays, Albuquerque
Cays, Roncador Bank and Serrana Bank by John D, Milliman. 1-26,
Aug. 15, 1969.

A Botanical Description of Big Pelican Cay, a Little Known Island
off the South Coast of Jamaica by C. D. Adams. 1-10, Aug. 15, 1969.

Post-Hurricane Changes on the British Honduras Reefs and Cays:
Re-Survey of 1965 by D. R. Stoddart. 1-25, Aug. 15, 1969.

IL 3.

US\36

134.

sie

E36.

1S

138.

dbo ie)

140.

141.

142.

143.

144 e

145.

146.

147.

148.

25

Plants of Satawal Island, Caroline Islands by F. R. Fosberg. 1-13,
Augie 5) L969"

A Collection of Plants from Fais, Caroline Islands by F. R. Fosberg
and Michael Evans. 1-15, Aug. 15, 1969.

Plants Collected on Islands in the Western Indian Ocean During a
Cruise of the M. F. R. V. "Manihine," Sept.-Oct. 1967 by M. D.
Gwynne and D. Wood. 1-15, Aug. 15, 1969.

Island News and Comment. 1-17, Aug. 15, 1969.

Coral Islands of the Western Indian Ocean by D. R. Stoddart. 1-224,
Aue 207 1970

Carbonate Sand Cays of Alacran Reef, Yucatan, Mexico: Sediments by
Robert L. Folk and Augustus S. Cotera. 1-16, Feb. 16, 1971.

The Vertebrate Fauna and the Vegetation of East Plana Cay, Bahama
Islands by Barrett C. Clough and George Fulk. 1-17, Feb. 16, 1971.

The Island of Anegada and its Flora by W. G. D'Arcy. 1-21,
Feb. 16, 2971"

Inshore Marine Habitats of Some Continental Islands in the Fastern
Indian Ocean by Alan J. Kohn. 1-29, Feb. 16, 1971.

The Distribution of Shallow-Water Stony Corals at Minicoy Atoll in
the Indian Ocean with a Check-List of Species by C. S. Gopinadha
Pilate ott 2 heb. 16. LOZ

The Uninhabited Cays of the Capricorn Group, Great Barrier Reef,
Australia by SB. Domm.. 1-27, Keb. 16, 1971.

The Safe Use of Open Boats in the Coral Reef Environment by S. B.
Domm. 1-10, Feb. 16, 1971.

The Vascular Flora and Terrestrial Vertebrates of Vostok Island,
South-Central Pacific by Roger B. Clapp and Fred C. Sibley. 1-10,
Inloys allay) Ale)7/Abe

Notes on the Vascular Flora and Terrestrial Vertebrates of Caroline
Atoll, Southern Line Islands by Roger B. Clapp and Fred C. Sibley.
te. Reb. 16, LO715

Records of Mallophaga from Pacific Birds by A. Binion Amerson, Jr.
and) Ky ¢.. Emerson. 1-S05 Feb. 16, 1971.

Rainfall on Indian Ocean Coral Islands by D. R. Stoddart. 1-21,
Bebe! Om 29)7 Ar

Island News and Comment. 1-38, Feb. 16, 1971.

26

149,

150).

5.

1G

153).

154.

NB) 36

156.

ois

158.

159.

160.

161.

162.

163.

164.

165%.

166.

Geography and Ecology of Diego Garcia Atoll, Chagos Archipelago
edited by D. R. Stoddart and J. D. Taylor. 1-237, Aug. 27, 1971.

The Natural History of French Frigate Shoals Northwestern Hawaiian
Islands, by A. Binion Amerson, Jr. 1-383, Dec. 20, 1971.

Bacterial counts in surface open waters of Eniwetok Atoll, Marshall
Islands, by Louis H. DiSalvo. 1-3, Dec. 31, 1972.

Marine studies on the North Coast of Jamaica, edited by Gerald J.
Bakus. 1-6, Dec. 31, 1972.

Fish diversity on a coral reef in the Virgin Islands, by Michael J.
Risk. 1-4, Dec. 31, 1972.

Recolonization of a population of supratidal fishes at Eniwetok
Atoll, Marshall Islands, by William A. Bussing. 1-4, Dec. 31, 1972.

Some marine benthic algae from Truk and Kuop, Caroline Islands, by
Roy Tl. Tsuda... I-10; Dees 31,7 1972;

Additional records of marine benthic algae from Yap, Western Caroline
Islands, by Roy T. Tsuda and Mary S. Belk, 1-5, Dec. 31, 1972.

Carbonate lagoon and beach sediments of Tarawa Atoll, Gilbert Islands,
by Jon N. Weber and Peter M. J. Woodhead. 1-21, Dec. 31, 1972.

Birds seen at sea and on an island in the Cargados Carajos Shoals,
by R. Pocklington, P. R. Willis and M. Palmieri. 1-8, Dec. 31, 1972.

Geomorphology and vegetation of Iles Glorieuses, by R. Battistini
and G. Cremers. 1-10, Dec. 31, 1972.

Reef islands of Rarotonga, by D. R. Stoddart,.With list of vascular
Plants by F. R. Fosberg. 1-14, Dec. 31, 1972.

South Indian sand cays, by D. R. Stoddart and F. R. Fosberg. 1-16,
Dec.. 31, 1972.

Island News and Comment. 1-26, Dec. 31, 1972.

The natural history of Gardner Pinnacles, Northwestern Hawaiian
Islands, by Roger B. Clapp. 1-25, Dec. 31, 1972.

The natural history of Kure Atoll, Northwestern Hawaiian Islands,
by Paul W. Woodward. 1-318, Dec. 31, 1972.

Central Western Indian Ocean Bibliography, by A. J. Peters and
Jj. ¥F. CG. Lionnet.. 1-322, May, 2,'1973;

Crown of thorns (Acanthaster planci) plagues: The natural causes
theory, by Peter J. Vine. 1-10, Nov. 23, 1973.

67:

168.

169.

170.

LAA

£72.

Th3s

174.

WTS.

L7G.

1 ae

17S

el.

hie 0)

181.

OZ.

27

A study of some aspects of the crown-of-thorns starfish (Acanthaster
planci) infestations of reefs of Australia's Great Barrier Reef,
by R. Endean and W. Stablum. 1-62, Nov. 23, 1973.

The apparent extent of recovery of reefs of Australia's Great
Barrier Reef devastated by the crown-of-thorns starfish, by R.

Endean and W. Stablum. 1-26, Nov. 23, 1973.

Investigations of Acanthaster planci in Southeastern Polynesia during

1970-1971, by Dennis M. Devaney and John E. Randall...1-23, Nov. 23, 1973.

Population levels of Acanthaster planci in the Mariana and Caroline
Islands 1969-1972, by James A. Marsh, Jr., and Roy T. Tsuda. 1-16,
NOU 2S al Sic

The natural history of Laysan Island, Northwestern Hawaiian Islands,
by, Charles A. Ely and: Roger B./Clapp.. -1=361,.Dec.. 31,.1973.

Comparative Investigations of Tropical Reef Ecosystems: Background
for an integrated coral reef program. Edited by Marie-Heléne Sachet
and Arthur .1.. Dahl. .1-169,..Dee.: 15,1974.

Preliminary checklist of the marine benthic plants from Glover's
Reef, British Honduras, by Roy T. Tsuda and Clinton J. Dawes. 1-13,
Dee. 15, 1974.

The Natural History of Pearl and Hermes Reef, Northwestern Hawaiian
Islands, by A. Binion Amerson, Jr., Roger B. Clapp, and William 0.
hire, The 91-306. Dee..31, “LOA4.

Observations on the birds of Diego Garcia, Chagos Archipelago, with
notes on other vertebrates, by A. M. Hutson. 1-25, Jan. 15, 1975.

The birds of the Iles Glorieuses, by C. W. Benson, H. H. Beamish,
G. Jouanine J. Salvan, and o¢. BE. Watson... 1-34, Jans 15,. 19/75,

Fulgoroidea from Aldabra, Astove, and Cosmoledo Atolls, collected
by the Royal Society Expedition 1967-68 (Hemiptera-Homoptera), by
Mey D., Webb. 1-10, Jan. 15, 1975.

A preliminary description of the coral reefs of the Tobago Cays,
Grenadines, West Indies, by John B. Lewis. 1-9, Jan. 15, 1975.

Marine zonation and ecology of Cocos Island, off Central America,
by Gerald J. Bakus. 1-9, Jan. 15, 1975.

Biogeography of reptiles on some of the islands and cays of Eastern
Papua-New Guinea, by Harold Heatwole. 1-32, Jan. 15, 1975.

Sandiicays om longatapu.eby Da R. Stoddart. 1-8, Jan-.d5.~ L975.

The murine rodents Rattus rattus, exulans and norvegicus as avian
predators, by FE.) 1. Norman. 1-13, Jan. 15, 1975.

i
|-
U

ALIAAL

N

28

83%

184.

185).

186.

187.

188.

189.

t90.

Ode

1:92"

1193".

194,

Moye,

196%.

IESE

OSs

199.

Ducie Atoll: Its history, physiography and biota, by Harold A.
Rehder and John E. Randall. 1-40, Jan. 15, 1975.

Marine turtles in the Phoenix Islands, by George H. Balazs. 1-7,
Jans elo skool.

Island News and Comment. 1-40, Jan. 15, 1975.

The Natural History of Lisianski Island, Northwestern Hawaiian
Islands. 1-196, Feb. 15, 1975.

The Algal Ridges and Coral Reefs of St. Croix their structure and
Holocene development, by Walter H. Adey. 1-67, Aug. 6, NLS)

Anegada Island: Vegetation and Flora by W. 6G. D'Arcy... LL=39)
JN Gers SOHAL) YS)a

The Natural History of Namoluk Atoll, Eastern Caroline Islands by
Mac Marshall with identifications of vascular flora by F. R.
Fosberg. 1-53, Aug. 6, 1975.

Almost-Atoll of Aitutaki: Reef Studies In The Cook Islands, South
Pacific, Edited by D. R. Stoddart and P. E. Gibbs. 1-158, Aug. 13,
17D.

Bibliography of The Natural History of the Bahama Islands by
William T. Gillis, Roger Byrne, and Wyman Harrison. 1-123, Aug. 20,
1975.

The Natural History of Johnston Atoll, Central Pacific Ocean, by
A. Binion Amerson and Philip C. Shelton, Dec. 19/6.

A photographic survey down the seaward reef-front of Aldabra Atoll,
by Edward A. Drew, in press.

Topography and coral distribution of Bushy and Redbill Islands and
surrounding reef, Great Barrier Reef, Queensland, by Carden C.

Wallace and E. R. Lovell, in press.

Coral Cays of the Capricorn and Bunker Groups, Great Barrier Reef
Province, Australia, by P. G. Flood, in press.

Submarine cementation of grainstone fabric, St. Croix, U. S. Virgin
Islands, by Lee C. Gerhard, in press.

Christmas Island (Pacific Ocean): Reconnaissance geologic
observations by Mark J. Valencia, in press.

Notes on the vertebrate fauna of Tongareva Atoll, by Roger B. Clapp,
in press.

Observations on vegetation of blue-faced booby colonies on

ZON
Cosmoledo Atoll, Western Indian Ocean, by Mary E. Gillham, in press.

200. Vegetation of sea and shore-bird colonies on Aldabra Atoll, by
Mary E. Gillham, in press.

201. Life history notes on some Aldabran land birds by C. B. Frith,
in press.

202. Climate of Aldabra Atoll, by D. R. Stoddard and L. U. Mole, in press.

203. Annotated check list of corals in the Mascarene Archipelago, Indian
Ocean, by Gerard Faure, in press.

204. Annotated check list of octocorallia in the Mascarene Archipelago,
Indian Ocean, by Gérard Faure, in press.

205. The hosts of the Coral-Associated Indo-West-Pacific Pontoniine
shrimps, by A. J. Bruce, in press.

206. The Natural History of Necker Island, Northwestern Hawaiian Islands,
by Roger B. Clapp and Eugene Kridler, in press.

207. The Natural History of Nihoa Island, Northwestern Hawaiian Islands,
by Roger B. Clapp, Eugene Kridler, and Robert R. Fleet, in press.

208. Notes on plants of the genus Caulerpa in the Herbarium of Maxwell
S. Doty at the University of Hawaii by Wm. Randolph Taylor, in press.

209. Marine algae of the Te Vega 1965 expedition in the Western Pacific
Ocean, by Wm. Randolph Taylor, in press.

210. Marine algae known from the Maldive Islands, by H. E. Hackett, in
press.

211. The benthic algal composition standing crop, and productivity of a
Caribbean algal ridge, by Judith L. Connor and Walter H. Adey, in
press.

212. Preliminary observations on the algae, corals, and fishes inhabiting
the sunken ferry "Fujikawa Maru" in Truk Lagoon, by Roy T. Tsuda,

Steven S. Amesbury, and Steven C. Moras, in press.

213. Chemistry of freshwater pools on Aldabra, by A. Donaldson and B. A,
Whitton, in press.

214. Observation on redox potential in freshwater pools on Aldabra, by
B. A. Whitton and M. Potts, in press.

215. Algal flora of freshwater habitats on Aldabra, by A. Donaldson and
B. A. Whitton, in press.

216. Terrestrial and freshwater algae of three Western Indian Ocean

30

ZANT

208.

219)

Islands (Astove, Farquhar and St. Pierre), by B. A. Whitton and
A. Donaldson, in press.

Terrestrial and swamp algae from three islands in the Chagos
Archipelago, Indian Ocean, by B. A. Whitton, A. Donaldson, D. J.
Bellamy, and C. Sheppard, in press.

The Holocene reef systems of eastern Martinique, French West
Indies, by Walter H. Adey, Patricia J. Adey, Randolph Burke and

Leslie Kaufman, in press.

Island News and Comment, in press.

U. S. GOVERNMENT PRINTING OFFICE : 1977 O - 231-339

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