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Nos. 252-255
e :

ae”
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fe, NY

ATOLL RESEARCH
BULLETIN

July 1981

252. Bird and Denis Islands, 254. Natural History of Raine
Seychelles Island, Great Barrier Reef
by D. R. Stoddart and F. R. by D. R. Stoddart, P. E. Gibbs,
Fosberg and D. Hopley

253. Topographic and Floristic 255. Short Original Articles
Change, Dry Tortugas, Florida, by various authors
1904-1977
by D. R. Stoddart and F. R.
Fosberg

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

July 1981

ACKNOWLEDGEMENT

The Atoll Research Bulletin is issued by the Smithsonian
InstheutiLone vasa park Of ats activity in) tropical biology, to place
on record information on the biota of tropical islands and reefs,
and on the environment that supports the biota. The Bulletin is
supported by the National Museum of Natural History and is produced
and distributed by the Smithsonian Press. The editing is done by
members of the Museum staff and by Dr. D. R. Stoddart.

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

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

Editors

ete HOSIbD ence:
Ian G,. MacIntyre
M.-H. Sachet

Smithsonian Institution
Washington, D.C. 20560

Do Ike SeoOclaice

Department of Geography
University of Cambridge
Downing Place
Cambridge, England

ota! Neg Ad fines i or thie Wels

Pit ri » ia PNB Hh meh): th y ne ui ai hd } we itu : Aolteo aan
‘ rs = er sph! a
~ \ , (ft

ATOLL RESEARCH BULLETIN
NO. 252

BIRD AND DENIS ISLANDS, SEYCHELLES

by D. R. Stoddart and F. R. Fosberg

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

July 1981

Contents

ee Geography and ecology of Bird Island, Seychelles

Introduction

Morphology and structure
Climate

Vegetation

Flora

Invertebrates

Reptiles

Mammals

Birds

History

Dis Plants recorded from Bird Island

3% Geography and ecology of Denis Island, Seychelles

Introduction

Morphology and structure
Climate

Vegetation

Flora

Invertebrates

Reptiles

Mammals

Birds

History

ae Plants recorded from Denis Island

Se References

Manuscript received May 1980 --Eds.

page

OoowoounwsP FE

Pe
=! ©)

N —
On Ww

WWWWWWNHNNDY
iS) [al ©) ©) ©) Sy Ql Gin

es)
OV

PS
OV

veal

List of Figures

The Seychelles Bank
Bird Island in 1976
Beach sediment at Bird Island
Denis Island in 1977

Monthly rainfall at Denis Island,
1971-1962

List of Tables

Scientific studies at Bird Island
Characteristics of Bird Island beach sands

Monthly rainfall at Bird Island, 1951-1962

Key to the literature on insects collected at

Bird Island
Scientific studies at Denis Island

Monthly and annual rainfall records at
Denis Island

following
following
following

following

following

page

page

page

page

page

page

LAL

aja

thal

50

50

26

28

Os

ASL

2s

List of Plates
Bird Island: Suriana zone on the northeast
shore

Bird Island: Pisonia and Cordia woodland
with Suriana on the northeast shore

Bird Island: Tournefortia parkland in the
northeast

Bird Island: tree-like Tournefortia in the
northeast

Bird Island: pioneer sedges and Scaevola on the
east shore

Bird Island: pioneer Ipomoea pes-caprae on the
east shore

Bird Island: pioneer sedges, Scaevola and
Tournefortia on the northeast shore

Bird Island: airstrip from the southeast

Denis Island: phosphate cliffs with Casuarina
woodland, southwest shore

Denis Island: phosphate cliffs at Muraille
Bon Dieu.

Denis Island: detail of phosphate cliffs at
Muraille Bon Dieu

Denis Island: Typha swamp

atta

following page 11

following page 50

BIRD AND DENIS ISLANDS, SEYCHELLES

by D. R. Stoddart and F. R. Fosberg

1. Geography and ecology of Bird Island, Seychelles

INTRODUCTION

Bird Island is one of two small coral islands on the northern
margin of the Seychelles Bank, western Indian Ocean. It is situated
in latitude 3°43'S and longitude 55°13'E, 98 km northnorthwest of Mahe,
the main granitic island of the Seychelles (Figure 1). J.C.F. Fryer
spent one week there in 1908 and later contributed the basic account
of the island (Fryer 1910). Subsequently it has been the subject of
brief visits by geologists, botanists and zoologists (Table 1), but no
comprehensive account of the island appeared until the recent major
paper by Feare (1979), based on observations during a year's residence
during a study of the Sooty Tern colony.

The purpose of the present paper is to present information on the
terrestrial botany of the island, and to call attention to some early
sources on its natural history (notably the visit by H.M.S. Alert in
1882) not mentioned by Feare. Detailed information given by Feare,
especially on the birds, will not be repeated here, and the reader is
referred to his paper for a more comprehensive account.

MORPHOLOGY AND STRUCTURE

The island is 1800 m long and a maximum of 830 m wide (Figure 2);
Coppinger (1885) gives dimensions only half as great, but both
Horsburgh (1809) and Fryer (1910) give approximately similar figures.
Figure 2 is based on a compass traverse map made in March 1976, with
additional detail from air photographs, and yields an area of 82.5 ha.
No elevations have been determined on the island, but the maximum
height is unlikely to be more than 4 m above mean sea level. Tidal range
at springs is 1.2 m and at neaps O.9 m.

Bird consists entirely of reef-derived carbonate sands and fine
gravels, with small quantities of drift pumice. Beaches are wide and
extensive around the entire coast, except for some erosion and cliffing
at the southern point. In March 1976 there was a broad unvegetated
spit, composed of several beach ridges, at the northeast end of the
island, but this is probably a largely seasonal phenomenon and varies

Table 1.
Date

1771 August

1808
1822

1882 4 March

1891

1908 24 July
- 1 August

LOSS

1952" 2iesepe
LI5S5

1960 14-15
September
1962 March
1963 4 Nov.
1369 #2

1970 14 July

1972 June,
August-November

1973 May-October

1974 July

1976 25-27
March

LOT]

Visitor

Lt W. Robinson
Lt D. Thomas,
Eagle
Hirondelle

F. Moresby
R.W. Coppinger

H.M.S. Alert

Lt C.H. Simpson

Lt F.M. Leake
H.M.S. Stork

JAC srs vhGy.erT

D. Vesey-FitzGerald
E.S. Brown

M. Ridley, R. Percy

C.J. ePiggotct

B.H. Baker
C. Jeffrey
R. Bailey

F.L. Lambrecht

J. Procter

Crue weare

D.R. Stoddart

L.G. Nikiforov
et al.

Scientific studies at Bird Island

Subject

Charting

Wrecked
Survey
General

collecting

Charting

General survey
and collecting

Birds
Insects
Birds
Coconuts
Geology
Botany
Birds
Insects
Botany

Birds,
general ecology

Botany,
general ecology

Geology and
sediments

Publication

Dalrymple 1780

Froberville 1848
Moresby 1842

Coppinger 1885;

Bowdler Sharpe
1884

Hemsley 1885

Admiralty Chart

Fryer 1908, 1910

Vesey-FitzGerald
1941

Mattingly and
Brown 1955

Ridley and Percy
LOS,

Piggott 1968, 1969
Baker 1963

Jeffrey 1962
Bailey (1967)
Lambrecht 1971

Procter 1970

Feare 1973a,
LOVSby LO SeloiiGay
lOV6b:, (OVO C RLY
Feare and High
LOW

This paper

Nikiforov et al.
1978

in location with the monsoons from the northeast to the northwest sides
(cf. the trace of the 1960 shore from air photographs in Figure 2).
Piggott (1968, 53) concluded from the existence of this spit that the
island was increasing in size, but this seems unlikely.

The beaches face a reef flat 300 m wide, which dries at low
springs, on the southeast side, but the western beaches overlook
deeper water. Table 2 gives characteristics of beach and berm
sediment samples from sites around the coast, based on sieving at
0.25¢ intervals, and Figure 3 gives cumulative curves and sample
locations. The beach samples (from mid-beach) are generally rather
coarser than the beach crest samples, though this is only at all
pronounced on the leeward coast. AE ches samples Mie wane ther coarse
to medium sand size range, are well to moderately well sorted, exhibit
little skewness, and are platykurtic to mesokurtic.

Nikiforov et al. (1978, 46-61) measured the height of beach ridges
at the northeast point reaching successively 2-3, 3.1 and 3.4 m from
sea to the vegetated island surface (datum not stated). Holes were
drilled in the second and third of these units to depths of 6.75 m;
subsurface sediment samples were mainly coarse-grained and poorly
sorted calcareous sands with shell and mollusc fragments. The centre
of the island was also drilled to a depth of 5.2 m below the surface;
the sediments were unsorted carbonate materials with near-surface
cementation. Nikiforov et al. (1978, 53) interpret this cemented
layer as an ancient beachrock, dated at 2500-2700 years B.P., but from
their description it appears to be a phosphate rock.

Phosphate rock covers much of the central part of Bird Island.
Figure 2 shows the distribution of this material according to Baker
(1963, 46-48) and Piggott (1968). The phosphorite has maximum
thicknesses of 1-2 m, and wells in it penetrate to the water table at
depthsiot 175-2) m: The rock is brown and speckled, well-cemented and
dense at the surface but crumbly below. The upper surface is
intricate and fractured, with many loose blocks, while the lower limit
of the rock is characteristically broadly lobate and irregular. A
sample of the phosphorite has been analysed by T.P. Scoffin, who finds
P905 28.2% and CaO 52.7%. He comments: "The rock is extensively
phosphatized and grains are hard to identify. There are some
unaltered coral grains but most grains are completely or partially

replaced. Those that are partly replaced have undergone centripetal
phosphatization at their rims. There is a dense brown phosphatic fine
grained or amorphous crust on the grains. This crust is not banded.

One grain shows unaltered carbonate cement needles in skeletal chambers.
There are bladed grains showing undulose extinction which may be broken
pieces of an earlier phosphate cement.” The surface guano overlying
the cemented material has been largely removed for export. Piggott
(1968, 39) describes the soil overlying the phosphorites as Jemo
Series, in contrast to the Shioya Series of the rest of the island.

Table 2. Characteristics of Bird Island beach sands

Sample Location Position @% mean ¢@% median oO Sk K

I I G

1 West Bay Beach 1.44 1.40 0.78 0.05 0.83

2 Berm O78 0.81 0.62 -0.04 1.04

3 Northeast Beach 1.56 1.46 0.79 OSLO 535

Bay
4 Berm 15S) 1b abs} 0.69 0.16 1{O©
5 Southeast Beach ies sh eat 0.48 0.20 1.08
Bay

6 Berm 1.68 ARS AZ 0.50 -0.08 0.96

7 Southwest Beach 0.92 0.83 0.60 O.19 ALS AL)
Point

8 Berm 0.93 0.92 0.48 0.06 OZ

9 Northeast Beach 159 1.08 0.60 ON 7 7/ O24
Point

10 Berm LAS es (OVS (5;7/ 0.05 ab ILI.

aLaL Northwest Beach T55 1.40 O4/3 OMS 0.88
coast

ED. Berm 0.89 0.92 0.78 -0.04 1.08

CLIMATE

Bird Island falls within the general Seychelles régime of an
extended period of dry Southeast Trade winds from April to November,
and a wet season of northwesterly winds and calms during December to
March. Rainfall records are available only for 1961 and 1962,
totalling 2358 and 1589 mm respectively; the monthly figures show
considerable variation between the two years (Table 3). The incidence
of rainfall in 1961-62 was very similar to that of Denis Island
(Table 5), though with marked divergences in December 1961 to February
1962. Feare (1979) also measured rainfall, recording 703 mm between
15 August and 5 November 1972.

Table 3. Monthly rainfall at Bird Island, 1961-1962 (mm)

Year Jan Feb Malaka AIG May Jun Jul Aug Sep Oct

LO GH GasOA.0/ WSCC S) elA2 SO) )/ 3h LoS NESS Opal AealesS MOM OZ 635

IQ62Z- BW55S) P2453) Zito) MWeeoil QIASS S555 A2IYoS USssl ZHO@ ~ Zo.)

Nov Dec Year

WIG ACI My 200%: 285 je

IQS TNA .8) ILA}5.5) LEKSI3}5 5)

Jan Feb Mar Apr May Jun Jul Aug Sep Oct

MeanwuEZIO.Gme o> Ono 2 non tOVol “N84 2958 Lagie 29455 123.5

Nov Dec Year

Main 29057 USA wWS7aigs

Source: Stoddart (1971)

VEGETATION

Horsburgh (1809, 126), in the first description available, says
that Bird Island "is very low, with small bushes on it", and all later
nineteenth century accounts are broadly similar. "Le seul végétal qui
pousse sur cette terre désolée, est le veloutier [Scaevola taccada]",
comments Froberville (1848, 99). Moresby (1942, 12) found it "a small
sandy island with a few shrubs on it" in about 1822, and Pelly (1865,
231) puts it in the class of "low sandy islands, sprinkled with scrubby
brushwood". In 1882 Coppinger (1885, 211) supplied more detail during
the visit of H.M.S. Alert. He found a "dense thicket of bushes, which
forms a fringe around the margin of the island" (presumably mainly
Scaevola taccada); a central area with "scrubby grass ... [and] several
introduced plants gone wild"; and "two or perhaps three Casuarina".

By the time of Fryer's visit in 1908 there was still a narrow
marginal belt of Scaevola and Tournefortia argentea bushes about 10 m
wide; a bare area with practically no vegetation, comprising the Sooty
Tern colony; rough ground on the phosphate rock, with herbs and grasses;
and a small coconut plantation extending east to west across the centre
of the island. The composition of the herbaceous ground cover was not
recorded, though Fryer's collections included Cassytha filiformis,
Ipomoea pes-caprae and Tribulus cistoides; Boerhavia repens and

Portulaca cleracea were probably also widespread. A few years before
Fryer's visit, a visiting magistrate in 1903 had enumerated 6000
coconuts, mostly young, with "a few filaos" [Casuarina] and three acres
(1.2 ha) of maize (Seychelles Archives, C/SS/74(1), 139). In spite
of this general impression of a virtually treeless landscape, however,
it is worth noting that in 1908 Fryer recorded both Calophyllum
inophyllum and Terminalia catappa, together with Hernandia sonora and
Morinda citrifolia, neither of which have been seen since (though the
Morinda is very probably present).

Nevertheless, it is clear that the vegetation today differs
greatly from that of the period before 1908. The whole island is now
covered with coconut woodland, managed as a plantation, with peripheral
tall Casuarina and a littoral hedge of Scaevola taccada. Feare (1979)
distinguished seven physiographic and vegetational units, broadly
congruent with the following:

1. Littoral hedge, dominated almost exclusively by Scaevola taccada
and extending almost continuously along the west and southeast coasts.
The hedge is about 5 m tall and 10-25 m wide. Its inner margin is
often marked by trees of Guettarda speciosa, and along the southeast
shore it is fronted for about 200 m by a low zone of Suriana maritima
(Plate 1).

2. Littoral woodland of Thespesia populnea, found on the northeast
coast where the shore is aggrading and where, as Feare (1979) states,
the phosphatic sandstones outcrop. Pisonia grandis and Cordia
subcordata are also present in this unit (Plate 2), but are rare.
Thespesia is also found on the cliffed eroding shore at the south
point.

3. Casuarina woodland. This has been planted as a windbreak inside
the Scaevola hedge, especially in the southwest and eastern parts of
the island and around the settlement. Many of the trees are 25-30 m
tall, but in recent years many have been felled for use in
construction.

4, Coconut woodland. A tall closed woodland of coconut palms, with
a ground cover which varies with substrate. On sand the surface is
occupied by grasses, Lippia and other low herbs; on phosphatic
sandstone by tall Kalanchoe pinnata, Nephrolepis and Carica papaya as

well as a diverse low herbaceous flora. Small gardens are scattered
through the plantations. Weeds such as Achyranthes aspera and
Phyllanthus casticum are ubiquitous. In 1962; according) to: Piggott

(1968, 41) the plantations yielded 40 tons of copra per annum, with
10-40 nuts per tree.

5. Tournefortia parkland (Plates 3 and 4). In the northwest part of
the island, where the Sooty Terns nest, there is an open area of
Boerhavia repens and Portulaca oleracea, dominated by mature trees of
Tournefortia argentea and occasional shrubs of Scaevola and Suriana.

The Tournefortia trees are 4-5 m tall. The vegetation of this area
is now managed to promote the breeding of the terns (Feare 1976).

6. Pioneer herbaceous vegetation (Plates 5, 6 and 7). Eades) aks
extensive in the northeast bay and on the northeast spit. ihe alts
dominated by Ipomoea pes-caprae, grasses and sedges, and small shrubs
of Scaevola and Suriana. Terns also nest here, though in lower
densities.

7. Airstrip herbaceous vegetation, with a low mat of Lippia nodiflora
and other herbs and grasses (Plate 8).

8. Settlement, with a diverse collection of introduced weeds and
decorative plants (including Tabebuia heterophylla, massive Ficus
benghalensis, and Calophyllum inophyllum), and crop plants. Many of

these were introduced at an early date. Coppinger (1885) recorded
coconuts, Casuarina, cotton, sugarcane, papaya, yam and gourd in 1882;
Fryer (1910) added maize and tobacco in 1908. Conversely plants such

as Catharanthus roseus were not recorded until 1962, and Stachytarpheta
jamaicensis and Ricinus communis not until 1970, though presumably
introduced considerably earlier.

The extent of the phosphorite on Bird Island raises questions
about the possible former extent of Pisonia grandis woodland, in view
of Fosberg's hypothesis of the role of acid Pisonia humus in the
precipitation of phosphate cement. Pisonia is now rare on Bird (only
a single tree was seen in 1976), but Feare (1979) drew attention to
the existence of litter and humus beneath Scaevola, Tournefortia,
Suriana and Cyperus, suggesting that these may substitute for Pisonia
in the phosphate reaction. Collections were made of litter from
beneath eight species of trees and shrubs in 1976, and pH was measured
over a 27 day period in the laboratory. pH levels were stable during
this period, and averaged as follows:

Scaevola BIg) Casuarina 6.4
Guettarda Bye Tournefortia 6.6
Suriana 6.0 Pisonia Gig
Thespesia Ges Cordia ia

None of these dry-season litter samples resembled the deep humus found
under Pisonia woodland elsewhere, however, and it is unlikely that such
high pH levels would lead to phosphate precipitation. These
measurements do not, of course, resolve the question of the former
existence of Pisonia woodland on Bird Island; but if it ever did exist
it had clearly disappeared before 1882.

FLORA

The flora of Bird Island, excluding sea-grasses, totals 101
species, about half introduced. Of these, 43 were recorded by
Coppinger or Fryer in or before 1908; Jeffrey recorded 7 in 1961-62,
Procter 37 in 1970, Feare 55 in 1972-74, and the present study 62.

The flora includes sixteen tree species, but of these probably only
Pisonia, Thespesia, Cordia, Tournefortia and Guettarda are native, and
some of these may have colonised naturally since the transformation of
the island in the early twentieth century. Shrubs are represented
only by Scaevola and Suriana, both indigenous. It should be noted
that both Fryer (1908, 1910) and Summerhayes (1931) misidentified
Suriana as Pemphis acidula, as did Procter (1970); Pemphis is not
found on the islands of the Seychelles Bank, nor indeed is it found in
the western Indian Ocean islands north of St Joseph Atoll.
Christensen's (1912) record for Bird of Acrostichum aureum is almost
certainly a locality error, since A. speciosum is found on Denis Island
and is not otherwise recorded for Bird Island; and the same is clearly
true for Typha javanica, recorded for Bird by Summerhayes (1931).

INVERTEBRATES

Feare (1979) has recorded the presence during his stay on Bird
Island of land crabs, Myriapoda, Insecta and Arachnida. In addition,
Fryer made collections of insects and some other groups in 1908.
Table 4 keys the subsequent literature of insect determinations from
Bird. Only Lepidoptera, Coleoptera and Orthoptera are at all well
represented. However, the records of Diptera have been extended by
studies of the mosquitoes by Mattingly and Brown (1955) and Lambrecht
COHAN Aedes albopictus, A. albocephalus and Culex pipiens are all
biters and from time to time abundant. In addition, from Fryer's
collections, Budde-Lund (1912) records a terrestrial isopod, Hirst
(1911) five species of spider, and Hirst (1913) a scorpion.

REPTILES

Feare (1979) records nesting by Green and Hawksbill turtles.
It is worth noting that Dalrymple's chart of 1780, based on the Eagle
survey in 1771, is annotated "many turtle". There is now no means of
knowing either the size of the former nesting population or the scale
of the decline in numbers. Fryer also collected the two common
geckos, Hemidactylus frenatus and Phelsuma madagascariensis (Boulenger
1911); Feare (1979) suggests that the former may have been introduced.
Fryer did not collect the common skink Mabuia sechellensis.

MAMMALS

Bird Island was formerly known as Ile aux Vaches marines, though

Table 4. Key to the literature on insects collected at Bird Island

Family Number of species Source

Orthoptera : 9 Bolivar 1912, 1924
Dermaptera ilk isyogare ILE)iLO)

Odonata dk Campion 1913

Hemiptera 9 Distant 1913

Neuroptera 1 Needham 1913

Lepidoptera iLL Meyrick 1911; Fryer 1912
Coleoptera 13} Champion 1914, 1917; Gebien

ILGA29. Seorwe LIL 5 aALGa.se
Sal o~rrecly LQ iby

Hymenoptera 1 Forel 1912

Diptera @ Lamb 1912; Mattingly and
Brown 1955; Lambrecht 1971

in his paper Feare (1979) makes no mention of the former existence of
marine mammals there. Horsburgh (1809, 126-127) refers to "Sea Cows,
or Bird Island", and comments that "When this island was explored by
the Eagle Cruizer, from Bombay, in 1771, many sea-lions (probably
Manutees or large seals) were seen on the beach"; Dalrymple's chart
of 1780, from the same survey, is annotated "many Seals". Moresby
(1842) calls the island Ile aux Vaches. Froberville (1848, 99)
paraphrasing an account dating from the wreck of the Hirondelle in
1808, states: "Les vaches marines se trainent souvent sur les dunes
pour y dormir au soleil, et, dans cet état, on peut les entendre
souffler de bien loin: cet animal est peu redoutable malgré sa taille
énorme et ses dents menacantes; nos matelots en assommérent plusieurs
pendant notre séjour, mais nous n'en faisionspas beaucoup de cas, la
chair n'en étant pas supportable; une seule tortue nous faisait plus de
plaisir lorsque nous parvenions 4 la surprendre". lasaieng (CUS iKo)s aly)
commented that "there is no doubt the island was the haunt of the
species of dugong known as Vache marine, which unfortunately has long
ceased to exist". No such animal was recorded by the Alert in 1882,
or since that time. In discussing these and other records, I have
concluded that the records referred to seals of unknown species rather
than to dugongs (Stoddart 1972).

10

BIRDS

Early accounts agree on the great size of the bird populations.
In 1771 the Eagle found "birds innumerable" (Horsburgh 1809, 127).
The first collections and observations on birds were made by Coppinger
in 1882; subsequently Fryer made observations in 1908 which were
utilised by Feare (1979).

Coppinger (1885, 212) specifically states that there were "no land
birds" at the time of his visit. Given the sparse vegetation and his
familiarity with land birds of the Seychelles islands, it is unlikely
that he would have failed to record them had they been present. By
the time of Fryer's visit the Madagascar Fody Foudia madagascariensis
and the Ground Dove Geopelia striata had been introduced. Coppinger
mentions "multitudes of white egrets" and collected two males of
Bubulcus ibis (Bowdler Sharpe 1884), the first record of this
presumably native species; Feare (1979) had taken Fryer's reference
to a "white heron" as the first mention of this species. The Mynah
Acridotheres tristis was first recorded by Feare (1979), and the date
of its introduction after Fryer's visit is not known.

Of the seabirds, Coppinger in 1882 found the ground"everywhere
excavated by burrows of petrels", and collected a male Puffinus
pacificus (Bowdler Sharpe 1884, as P. chlororhynchus); he also stated
that "many gannets are breeding on the island". Fryer found boobies
"breeding in small colonies at the edge of the goelette Sooty Tern

colony". He names the species Sula piscator (= S. sula?), as does
Gadow (1907), who comments that "The guano of Bird Island, Seychelles,
was principally formed by a great colony of this species". In

the 1930s Vesey-FitzGerald (1941) found both nesting Masked Boobies
Sula dactylatra and immature Red-footed Boobies S. sula. To further
complicate matters, Coppinger (1885, 213) refers to "a large brown
bird" which was probably a Brown Booby Sula leucogaster. According

to Feare (1979), all three species are occasionally seen today. There
must remain some doubt as to which were present and breeding in the
nineteenth century, though the absence of trees suggests that only the
ground-nesting Masked and Brown Boobies could have been numerous.

Other early seabird and shorebird records are of less interest.
Coppinger (1885) mentions "a great flock of frigate birds", and terns
in "great numbers"; he collected a Brown Noddy Anous stolidus (Bowdler
Sharpe 1884). Fryer (1910) also mentions the noddies nesting in
Scaevola and Tournefortia, and refers also to a "large grey tern" and
a gull. Of migrants, Coppinger mentions "great flocks of turnstones
and curlews" (Bowdler Sharpe 1884 cites skins of the former) and "a
few oyster catchers". Fryer lists turnstone, plover, curlew,
sanderling, whimbrel and stilt, as well as the moorhen Gallinula
chloropus, all of which are listed by Feare (1979).

Nothing is to be added to Feare's (1976) account of the Sooty Tern
colony. He suggests that this numbered over 1,000,000 pairs in 1907.

Tae

By the mid 1930s, with habitat change and interference, it had
declined to 65,000 pairs (Vesey-FitzGerald 1941) and by 1955 to

18,000 pairs (Ridley and Percy 1966). Thereafter it recovered to
120,000 pairs in 1966 (Ridley and Percy 1966) and to 395,000 pairs
occupying 117 ha by 1973) \(Peare 1976). Feare estimated that during

his study the birds laid some 300,000 eggs during the period 8-17 June.

In addition to the comprehensive list of vagrants and migrants
recorded by Feare (1979), there are subsequent records by Feare and
High (1977) and by Turner and Forbes-Watson (1976).

HISTORY

The early history of Bird Island is virtually unknown. The Eagle
called there and charted the island in 1771. The Hirondelle, a French
privateer with 180 people on board, was wrecked there in 1808.

"They proved water by sinking a pit in the sand, remain'd there 22 days
until they constructed a raft, on which a part arrived at Mahe"
(Moresby 1842). Thereafter, apart from Moresby's own visit in 1822,
there is no information before the visit of H.M.S. Alert sixty years
later.

Feare (1979) has outlined the major changes in the ecology of the
island brought about by guano mining and the introduction of the
coconut. The former began in 1895: before that, in 1882, Coppinger
(1885, 211) found only two negroes salting fish and birds. By early
February 1896 385 tons of guano had been removed, and by early April
500 tons (Seychelles Archives B/44.195, 233). During 1900-1905 a
total of 17,000 tons was exported, and the resource was apparently
exhausted by 1906 (Baker 1963). The population at this time (1903)
numbered 42 men, 17 women and 31 children (Seychelles Archives
C/SS/74(1), 139); subsequently it declined to a total of 12 in 1931,
but then increased as the plantations were established, to 25 in 1947
and 49 in 1960.

The plantations had the effect of drastically reducing the area
available for nesting seabirds, and during the general disturbance
introductions included rats, mice, donkeys, pigs, goats, cats and dogs.
A new phase was opened in 1971-73 by the clearing of the airstrips and
the opening of tourist accommodation, but this increased accessibility
has coincided with a new ecological awareness on the part of the owners
and in a considerable intensification of scientific studies.

ACKNOWLEDGEMENT

We thank M. Guy and Mme Marie-France Savy for their generous help
On) Bird eisiand) an 976s

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Bird Island in 1976

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66-66

Plo

poe Bi K: b 4 ae 4 7 +A a

Pl. 1. Bird Island: Surtana zone on the northeast shore

Pie.) wBiendtsilandipeeeson7a ward Cordta woodland with Surtana on
the northeast shore

Pl. 3. Bird Island: Tournefortta parkland in the northeast

WFO SF

Pl. 4. Bird Island: tree-like Tournefortia in the northeast

>
rad «

ie ia a ok Ee Sela

Pl. 5. Bird Island: pioneer sedges and Scaevola on the east shore

Pl. 6. Bird Island: pioneer Ipomoea pes-caprae on the east shore

Pl. 7. Bird Island: pioneer sedges, Secaevola and Tournefortta
on the northeast shore

Pl. 8. Bird Island: airstrip from the southeast

2. Plants recorded from Bird Island

POLYPODIACEAE

Acrostichum aureum L.
Fryer, in Christensen (1912) [in error for Denis Island? ]

Nephrolepis biserrata (Sw.) Schott
Procter 4040 (Mahe); Fryer, in Christensen (1912); Gardiner, in

Christensen (1912); Feare, sight.

Nephrolepis multiflora (Roxb.) Jarrett
Stoddart 7110 (US).

Polypodium scolopendria Burm.f.
Fryer, s.n. (K); Fryer, in Christensen (1912) (as P. phymatodes)
[ TYPHACEAE |
[Typha javanica Schnizl ex Rohrb.
Fryer, in Summerhayes (1931) (in error for Denis Island?) ]
POTAMOGE TONACEAE
Thalassodendron ciliatum (Forsk.) d. Hart.
Cymodocea ciliata (Forsk.) Ehrenb. ex Aschers.
Stoddart 7123 (US); Fryer, sight (1910) (as Cymodocea) .
GRAMINEAE

-*Cynodon dactylon (L.) Pers.
Stoddart 7112 (US); Procter, sight.

*Dactyloctenium aegyptium (L.) Willd.
Stoddart 7072 (US); Procter, sight; Feare, sight.

14

*Digitaria horizontalis Willd.
Procter 4049 (MAHE).

*Eleusine indica (L.) Gaertn.
Procter 4049 (MAHE); Coppinger, in Hemsley (1885); Feare, sight.

Enteropogon monostachyum K. Schum. ex Engl.
Procter, sight.

Enteropogon sechellenis (Barker) Dur. and Schinz
Stoddart 7064 (US), 7091 (US); Procter, sight.

Eragrostis subaequiglumis Renv.
Procter 4044 (MAHE); Stoddart 7074 (US).

*Eragrostis tenella (L.) Beauv. ex R. and S. var. tenella
Piggott, S.n. (K); Stoddart 708i) (US); Procter, sight.

Eragrostis tenella var. insularis Hubb.
Vesey-FitzGerald 5656 (K).

Lepturus repens (Forst.) R.Br.
Fryer, sight, in Summerhayes (1931).

*Panicum repens L.
Fryer, sight, in Summerhayes (1931) [locality as "Ile aux Vaches"].

Pennisetum polystachion (L.) Schult.
Fryer, no number, in Summerhayes (1931).

*Saccharum officinarum L.
Coppinger, sight (1885).

Sporobolus virginicus (L.) Kunth
Fryer, no number, in Summerhayes (1931); Procter, sight.

Stenotaphrum micranthum (Desv.) Hubb.
Stenotaphrum subulatum Trin.
Procter 4041 (MAHE); Stoddart 7073 (US), 7129 (US); Feare, sight.

*Zea mays L.
Fryer, sight (1910).
CYPERACEAE

Cyperus sp.
Stoddart 71207 (US);

Cyperus conglomeratus Rottb.
Procter 4037 (MAHE) (as C. pachyrhiza); Stoddart 7071 (US); Feare,
sight (as C. pachyrhiza).

15

Cyperus dubius Rottb.
Mariscus dubius (Rottb.)
Fryer 26 (K) (as Mariscus dregeanus); Procter 4038 (MAHE); Feare,
sight.

Cyperus ligularis L.
Mariscus ligularis (L.) Hutchinson
Fryer 25 (K) (as Mariscus rufus); Jeffrey 1201 (MAHE) (mis-
labelled: this specimen is from Denis Island); Stoddart 7087 (US).
PALMAE
*Cocos nucifera L.
Coppinger, sight (1885); Fryer, sight (1910); Piggott, sight
(1969); Procter, sight; Feare, sight; Stoddart, sight.

ARACEAE

*Alocasia macrorrhiza (L.) Schott.
Stoddart 7109 (US).

Colocasia esculenta (L.) Schott.

Stoddart 7/1 Sr (us).

LILIACEAE sensu latissimo
*Agave rigida Northrop var. sisalana Perr. ex Engelm.
Agave sisalana Perr.

Feare, sight; Stoddart, sight.
*Crinum sp.?

StoddarewAk6 (US).

MUSACEAE

*Musa sapientum L.

Feare, sight; Stoddart, sight.

MARANTACEAE

*Maranta arundinacea L.
Fryer, sight, in Summerhayes (1931); Procter, sight.

16

CASUARINACEAE

Casuarina equisetifolia L.
Stoddart 7107 (US), 7133 (US); Coppinger, sight (1885); Fryer,
sight (1910) ;' Piggott, sight’ (1969); Procter, sight; Feare, sight

MORACEAE

*Artocarpus altilis (Park.) Fosb.
Stoddart, sight (juvenile).

Ficus sp.
Stoddart 7098 (US); Feare, sight.

*Ficus benghalensis L.
Stoddart 7097 (US).

Ficus nautarum Baker
Fryer, no number, in Summerhayes (1931); Procter, sight.
AMARANTHACEAE

Achyranthes aspera L.
Procter, sight.

Achyranthes aspera cf. var. fruticosa Boerl.
E.S. Brown in 1952-1953 (BM).

Achyranthes aspera cf. var. velutina (H. and S.) Townsend
Stoddart 7079 (US); Feare, sight.

*Amaranthus dubius Mart. ex Thell.
Stoddart 71037 (US) ; 7127" (US).

*Amaranthus caudatus L.
Fryer, no number, in Summerhayes (1931).

*Amaranthus cleraceus L.
Feare, sight.
NYCTAGINACEAE

Boerhavia diffusa L. (sensu lato) (prob. = B. repens L.)
Fryer \32° (KK).

Boerhavia repens L. var.

Procter 4043 (MAHE) (var. maris-indica Fosb.); Stoddart 7083 (US);

Feare, sight.

iL)

*Bougainvillea glabra Choisy
Stoddart 71027 (US):

*Mirabilis jalapa L.
Stoddart 7094 (US), 7111 (US); Feare, sight.

Pisonia grandis R. Br.
Stoddart 7080 (US); Feare, sight.
PORTULACACEAE
Portulaca oleracea L. var. granulato-stellulata v. Poelln.
Stoddart 7088 (US); Coppinger, in Hemsley (1885); Procter, sight;
Feare, sight.
LAURACEAE
Cassytha filiformis L.
Stoddart 7077 (US); Fryer, sight (1910); Procter, sight; Feare,
sight.
HERNANDIACEAE
Hernandia sonora L.
Hernandia peltata Meissn.
Hernandia nymphaeaefolia (Presl) Kubitzki
Fryer, sight (1910) (as H. peltata).

CAPPARIDACEAE

*Cleome viscosa L.
Feare, Sight.

*Cleome gynandra L.
Gynandropsis gynandra (L.) Briq.
Procter 4046 (MAHE); Feare, sight.

MORINGACEAE

*Moringa Oleifera Lam.
Stoddart 7122 (US); Feare, sight.

18

CRASSULACEAE
*Kalanchoe pinnata (Lam.) Pers.
Stoddart 7084 (US); Procter, sight; Feare, sight.
LEGUMINOSAE
Caesalpinia bonduc (L.) Roxb.
Caesalpinia bonducella (L.) Flem.

Fryer 7 (K)i

Cassia occidentalis L.
Procter, Sight; Feare, sight.

Sesbania sericea (Willd.) Link
Stoddart 7105 (US); Coppinger, in Hemsley (1885) (as S. aculeata
Pers.).
ZYGOPHYLLACEAE
Tribulus cistoides L.
Stoddart 7130 (US); Fryer, sight (1910); Procter, sight (as
Tribulus sp.); Feare, sight.

SURIANACEAE

Suriana maritima L.
Stoddart 7069 (US); Coppinger, in Hemsley (1885); Feare, sight.

EUPHORBIACEAE
Acalypha indica L.
Fryer 27 (K); Procter 4045 (MAHE); Stoddart 7082 (US); Feare,
sight.

*Fuphorbia cyathophora Murr.
Fryer, no number, in Summerhayes (1931) (as E. heterophylla L.).

*Fuphorbia hirta L.
Stoddart 7114 (US); Procter, sight; Feare, sight.

Euphorbia prostrata Ait.
Procter, sight; Feare, sight.

*Pedilanthus tithymaloides (L.) Poit.
Stoddart 7117 (US).

iL

*Phyllanthus amarus Schum.
Stoddart 7/1e2'5> (US)

Phyllanthus casticum Willem.
Stoddart 7089 (US), 7113 (US); Piggott, sight (1969); Feare, sight.

Phyllanthus maderaspatanus L.
Stoddart 7062 (US).

*Ricinus communis L.
Stoddart 7063 (US); Procter, sight; Feare, sight.
SAPINDACEAE
*Cardiospermum halicacabum L.
Fryer 24 (K).
MALVACEAE
*Abutilon indicum (L.) Sweet
Stoddart 7106 (US), 7121 (US) ; Coppinger, in Hemsley (1885); Procter,
Sight (as Abutilon mauritianum); Feare, sight.
*Gossypium hirsutum L.
Stoddart 7104 (US); Coppinger, sight (1885); Procter, sight (as

Gossypium purpurascens); Feare, sight.

*Malvastrum coromandelianum (L.) Garcke
Procter, sight; Feare, sight.

*Sida acuta Burm. f£.
Feare, sight.

Sida parvifolia DC.
Stoddart 7124 (US).

Thespesia populnea (L.) Sol. ex Correa
Stoddart 7076 (US); Feare, sight.
GUTTIFERAE
Calophyllum inophyllum L.

Fryer 17 (K); Stoddart 7099 (US) (var. takamaka Fosb.); Feare,
sight.

20

TURNERACEAE
*Turnera ulmifolia L.
Stoddart 7067 (US); Procter, sight; Feare, sight.
PASSIFLORACEAE
*Passiflora suberosa L.
Stoddart 7090 (US); Fryer, no number, in Summerhayes (1931);
Procter, sight; Feare, sight.
CARICACEAE
*Carica papaya L.
Stoddart 7085 (US); Coppinger, sight (1885); Fryer, sight (1910);
Piggott, sight (1969); Procter, sight; Feare, sight.

CUCURBITACEAE

*Cucurbita sp.
Stoddart 7108 (US).

*Cucurbita cf. maxima Duchesne
Stoddart 7134 (us);

*Cucurbita moschata Duchesne
Procter, sight; Feare, sight.

[ LYTHRACEAE |

[Pemphis acidula Forst.
Fryer, sight, in Summerhayes (1931); Procter, sight (both in error
for Suriana maritima L.) |

COMBRETACEAE

Terminalia catappa L.
Fryer 16 (K).

APOCYNACEAE

*Catharanthus roseus (L.) G. Don
Vinca rosea L.
Lochnera rosea (L.) Spach
Fryer 29 (K) (as Lochnera rosea); Stoddart 7086 (US), 7101 (US);
Piggott, sight (1969); Feare, sight.

“al

*Plumeria sp.
Stoddart, sight (juvenile).

CONVOLVULACEAE

*Tpomoea batatas (L.) Lam.
Fryer, no number, in Summerhayes (1931).

Ipomoea macrantha R. and S.
Ipomoea tuba (Schlecht.) G. Don
Stoddart 7093 (US); Fryer, sight (1910); Feare, sight.

Ipomoea pes-caprae (L.) R. Br.
Stoddart 7075 (US) (var. brasiliensis (L.) v. Ooststr.); Fryer,
sight (1910); Procter, sight; Feare, sight.

BORAGINACEAE

Cordia subcordata Lam.
Stoddart 7078 (US); Feare, sight.

Tournefortia argentea L.f.
Messerschmidia argentea (L.f£.) Johnst.
Stoddart 7068 (US); Coppinger, in Hemsley (1885); Fryer, sight
(1910); Procter, sight; Feare, sight.

VERBENACEAE

Lippia nodiflora (L.) Michx.
Phyla nodiflora (L.) Greene
Fryer 31 (K); Procter 4042 (MAHE); Stoddart 7066 (US); Feare,
sight.

Stachytarpheta jamaicensis (L.) Vahl
Stoddart 7065 (US); Procter, sight; Feare, sight.

LABIATAE
*Plectanthrus sp.
Stoddart. 75) (0S).
SOLANACEAE
*Capsicum minimum Roxb. (= C. frutescens L.?)

Feare, sight.

22

*Datura metel L.
Stoddart 7095 (US); Procter, sight; Feare, sight.

*Nicotiana tabacum L.
Fryer, Sight (1910);

*Solanum melongena L.
Feare, sight.

Solanum nigrum L. (s.1.)
Stoddart 7126 (US); Feare, sight
SCROPHULARIACEAE
Sériga asitatica (L-) O05 Ktze.
Preyer 33. (K)i
BIGNONIACEAE

*Tabebuia heterophylla (L.) Hemsl.
Stoddart 7096 (US), 7100 (US); Feare, sight (as Tabebuia rosea).

ACANTHACEAE
Asystasia multiflora Kl.?
Stoddart 7119 (US).
*Asystasia gangetica (L.) T. Anders. (= Asystasia multiflora K1.?)

Procter, sight; Feare, sight.

RUBIACEAE

Guettarda speciosa L.
Stoddart 7092 (US), 7132,,(US).);. Feare,. sight.

Morinda citrifolia L.
Fryer 4 (K).

GOODENTACEAE

Scaevola taccada (Gaertn.) Roxb.

Stoddart 7070 (US); Fryer, sight (1910) (as Scaevola koenigii);
Piggott, sight (1969); Procter, sight (as Scaevola frutescens); Feare,
sight.

23

COMPOSITAE
Vernonia cinerea (L.) Less.
Stoddart 7061 (US); Fryer, no number, in Summerhayes (1931);
Procter, sight; Feare, sight.

SOURCES

Coppinger collection: listed by Hemsley (1885), pp. 16-17; additional
sight records in Coppinger (1885).

Fryer collection: Fryer's paper on Bird Island (1910) and his
manuscript diary; enumerations of flowering plants in Summerhayes
(1931) and of ferns in Christensen (1912).

Procter collections: Department of Agriculture, Mahe, and enumeration
of collections and sight records in 'List of plants recorded on
Bird Island on 14 July 1970', typescript.

Jeffrey collection: Department of Agriculture, Mahe.

Feare records: sight records cited in Feare (1979).

Stoddart collection: determinations by F.R. Fosberg.

A few miscellaneous collections in herbaria indicated.

Species marked * are judged to be introduced.

Some of the sight records may well be misidentifications.

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3. Geography and ecology of Denis Island, Seychelles

INTRODUCTION

Denis Island is located on the northern margin of the Seychelles
Bank in latitude 3°48'S. and longitude 55°40'E., 89 km northnortheast
of Mahe and 52 km east of Bird Island. It is 2000 m long, has a
maximum width in the north of 1380 m, and has a total area of 120 ha
(48 per cent larger than Bird Island) (Figure 4).

The island was discovered by and named after Denis de Trobriand,
in command of the Etoile, in August 1773, but the first full account
resulted from the visit of J.C.F. Fryer in August 1908 (Fryer 1910).
Subsequently the island was visited by B.H. Baker, C. Jeffrey and C.J.
Piggott in the early 1960s (Table 5), but the only connected account
of the natural history of Denis remains that of Fryer. The present
paper is based on a visit on 10-11 December 1977; it brings together
the existing literature, including unpublished material in the
Seychelles Archives, and also provides an account of the vegetation
and flora.

MORPHOLOGY AND STRUCTURE

Denis is an island formed entirely of carbonate sands which have
been extensively phosphatised. The beaches are narrow and apparently
stable; there are no pronounced spits or beach ridges, as at Bird
Island, and no storm deposits. The average surface elevation is
probably only about 2.5 m, rising to 3.5 m in the north and northeast,
though no instrumental determinations have been made. About 80 per
cent of the surface area is occupied by phosphate rock, totalling about
95 ha, and this reaches the coast to form irregular cliffs about 2m
high at the southwest corner (Muraille Bon Dieu) (Plates 9 and 10) and
‘at the pronounced indentation on the east coast (Sans Tache). As on
Bird Island, the upper surface of the phosphate rock is blocky and

irregular, with many loose fragments. The lower surface of
cementation is irregular and lobate, forming pillars and inlets in the
cliffs (Plate 11; also illustrated by Fryer 1910, Plate 2). This

morphology is identical to that of similar rocks at Raine Island, Great
Barrier Reef, where the lower surface is also exposed in cliffs.

26

Table 5. Scientific studies at Denis Island

Date Visitor Subject Publication

P75) U2 Aug). D. de Trobriand Discovery De Trobriand, in
Etiole Fauvel 1909

1906 29-31 W.F. Stephens Phosphate Stephens 1906a,

March 1906b

1908 2-8 J.C.F. Fryer General survey Fryer 1910

August and collecting

ODT, J. Hornell Collecting Burleigh 1979

1952 E.S. Brown Insects Mattingly and Brown

L955

1960 Sept Coding Piggott Coconuts Piggott 1968, 1969
B.H. Baker Geology Baker 1963

1962 March C. Jeffrey Botany Jeffrey 1962

1969? F.L. Lambrecht Insects Lambrecht 1971

LOW Te vO—n: D.R.. Stoddart Botany This paper

December General ecology

According to Piggott (1968, 55), the phosphate rock is more than 1.2 m
thick in the northeast but thinner elsewhere. Baker (1963, 41-46)
describes a double horizon of cementation in the northeast, in the
following section: 5-46 cm: phosphate rock, 30-90 cm: unconsolidated
sand, 18-30 cm: phosphate rock. According to Baker the phosphate rock
is thicker in the centre of the island, where it forms a single horizon.
The cemented material is overlain by guano, much of which has been
removed for export. Piggott (1969, 63) terms the superficial material
a Jemo Series soil, in contrast to the Shioya Series of the rest of the
island, and Baker (1963) gives total P7205 content of guano samples of
3165, 2852 andy ..8 per cent.

Towards the eastern side of the island there are elongate water-
filled depressions about 1 m below the general surface level. Tite aes
not known whether these are natural or whether they result from guano-
digging operations earlier this century. They contain fresh water and
hydromorphic soils, which are unusual on western Indian Ocean coral
islands. Piggott (1969, 63) gives the following profile:

O- 1:3) em Black clay loam with coarse soft crumb structure,
containing about 10 per cent P7205 and 20 per cent
organic matter.

DY

13-38 cm Brown phosphatic sandstone; very dark in the upper
part but getting pale below; the sand being fairly
coarse.

38-46 cm Coarse white sand, with the watertable at 46 cn.

In addition to the phosphate rock, beachrock outcrops extensively
along the southeast shore, where it was noted by Fryer in 1908.
Remnants of beachrock indicate this this beach was formerly longer, but
the coast itself seems to be a stable one. Beachrock also outcrops
intermittently along the shore of the west bay. Tidal movement of
sand suggests that the beachrock is much more extensive than the
outcrops at any time suggest.

As at Bird Island, surface reef is only extensive along the
eastern side of the island. The shoal on which Denis stands is very
much smaller than that of Bird, and the surrounding surface of the
Seychelles Bank is 55-75 m deep.

CLIMATE

Rainfall records were maintained at Denis Island from 1951 to
1962 (Table 6, Figure 5). The mean annual rainfall over 12 years was
1730 mm, with extremes ranging from 1176 to 2643 mm. In the two years
(1961-62) for which records were also kept at Bird Island, the Denis
Island rainfall was substantially greater (12 pér cent greater in 1961,
a wet year, and 19 per cent greater in 1962, an average year). Leas
interesting that the pattern of wet and dry years at Denis during
1951-1962 closely resembles that at D'Arros in the northern Amirantes
(Stoddart, Coe and Fosberg 1979), suggesting a general regional control
of rainfall variability. December and January are the wettest months
and July much the driest. All months show considerable variation in
rainfall totals from year to year, but Denis does not experience the
prolonged dry seasons and severe droughts which occur on coral islands
in the southern Seychelles.

VEGETATION

De Trobriand in 1773 described the vegetation of Denis Island as
follows: "quelques parties de l'isle sont coupées par des esp&ces de prairies
dont l'herbe parait trés bonne quelqu'autres petites portions sont d'une
terre assez séche melée de sable: environ la moitié de la surface de
l'isle est couverte d'assez gros arbres mais dont le bois m'a semblé
etre trop gros et trop spongieux pour etre propre a4 la construction des
‘vaisseaux" (in Fauvel 1909, 47). These trees may well have been
Pisonia grandis. Fryer (1910, 19) unfortunately concluded that "the
vegetation is of no interest, being all secondary in character and
dependent on the coconut cultivation to which its whole surface is
devoted". He did, however, distinguish a littoral hedge of Scaevola,
Tournefortia and Suriana (wrongly identified by both him and Summerhayes
(1931, 278) as Pemphis acidula), with Hibiscus tiliaceus. Trees

28

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29

mentioned by him included Casuarina, Calophyllum, Hernandia, Ficus
and Pisonia. He also noted the existence of Typha; a large fern
(probably Acrostichum rather than the Asplenium he mentions); and
Catharanthus, as well as bananas and vanilla. Piggott (1969, 59-65)
also remarks on the number of large trees in the east and centre of
the island (Calophyllum, Hernandia, Pisonia, Ficus, Terminalia), and
found both Catharanthus and Cassytha to be widespread.

Based on the visit in 1977 and the map and collections then made,
the following vegetation units may be distinguished;

1. Littoral hedge, dominated by Scaevola taccada but also with
Tournefortia argentea and Suriana maritima, often much covered with
Cassytha filiformis. On its landward side both Guettarda speciosa
and Hibiscus tiliaceus are common, and in places, especially on the
west and southeast sides, Scaevola is replaced by Guettarda. The
littoral hedge on the west side is narrower, less continuous and more
diverse, and in places (on the slightly eroding northeast shore and
where the phosphate rock outcrops) it is entirely lacking.

2. Casuarina woodland. Casuarina equisetifolia is extensively
planted as a windbreak along the greater part of the coastline (Plate
10), forming a zone 20-100 m wide, often with the trees set out in

regular rows. Many are tall and old and some are being removed for
timber.
3. Coconut woodland. This covers the entire centre of the island,

and many of the trees are about 80 years old. When the plantation
was begun on a commercial scale is not known, but it was probably
about 1890. By 1903 there were 20,000 trees yielding 50,000 nuts per
month (Seychelles Archives C/SS/74(1), 139), and by 1906 the yield was
60,000 nuts per month (Stephens 1906a). Piggott (1969) found the
plantation well managed, in spite of the age of the trees and the
effects of Rhinoceros Beetle and coccids, and he commented on the

high quality of the copra in spite of the poor quality of the nuts.
The plantations are still scattered with broadleaf trees - massive
Barringtonia and Calophyllum and smaller Tabebuia, Morinda, Terminalia
and Pisonia. The fern Polypodium and Kalanchoe form extensive and
dense ground cover on the phosphate areas; Catharanthus and other
weedy plants are widespread. Piles of coconut husks are covered
with the fern Nephrolepis.

4. Marsh vegetation. The inland freshwater marshes support tall and
impenetrable stands of the fern Acrostichum speciosum and the bulrush
Typha javanica (Plate 12). The latter is rare on coral islands; it

is common on high islands in the Seychelles, notably on Praslin and
' Curieuse, which are the closest high islands to Denis. Alocasia
macrorrhiza is cultivated in these marshes.

30

FLORA

Fryer in 1908 recorded a total of 29 species of plants (Fryer
1908, 1910; Summerhayes 1931); 17 species collected by Jeffrey in 1962
are in the Department of Agriculture Herbarium, Mahe; and the 1977
visit yielded 72 species. With occasional archival records the known
flora stands at 91 species, rather less than that of Bird Island, but
undoubtedly more remain to be recorded. It is apparent that many
species were introduced at Denis at an earlier date than at Bird
Island. Casuarina, Terminalia, lime and pumpkin were recorded by the
visiting magistrate as early as 1903 (Seychelles Archives C/SS/74(1),
139), and in 1908 Fryer (1908, 1910; Summerhayes 1931) recorded
vanilla, banana, cotton, Datura metel, Capsicum frutescens and
Solanum nigrum, as well as Kalanchoe and Stachytarpheta. The Agave,
Carica and Ricinus found in 1977 must also have been introduced at an
early stage.

INVERTEBRATES

With the exception of the mosquitoes, recently collected by
Mattingly and Brown (1955) and Lambrecht (1971), knowledge of the
insect fauna of Denis Island derives entirely from the collections of

Fryer in 1908. The known fauna of about 60 species is dominated, as
at Bird Island, by Coleoptera, Lepidoptera and Diptera. The
literature on Fryer's collections is keyed in Table 7. In addition,

Hirst (1911) records four species of spiders, and Hirst (1913) a
scorpion, all collected by Fryer.

REPTILES

When De Trobriand discovered Denis in 1773 he recorded it as
"généralement couverte de tortiies de terre et de mer" (in Fauvel 1909,
47). By 1908 Fryer (1910, 19) found only "a few introduced specimens"
of the Giant Land Tortoise Geochelone gigantea, and he commented that
turtles had become "very scarce"; turtle tracks were seen in 1977 on
the western beach, but there is no means of estimating the size of
either the past or the present population. Apart from De Trobriand's
description, there is no further eye-witness account of the Giant Land
Tortoise on Denis Island. In 1927, however, James Hornell collected
fossil tortoise eggs, presumably from phosphorites, on Denis. These
have now been radiocarbon dated by Burleigh (1979), and yielded an age
of 1308+85 years B.P., or in calendar terms approximately 680+100 A.D.
These eggs confirm De Trobriand's account of the existence of tortoises
on Denis Island, and the date, which predates the human occupation of
the Seychelles, establishes that they must have colonised the island
naturally and not been introduced.

Fryer also collected the skink Mabuia sechellensis and the gecko
Phelsuma madagascariensis, according to Boulenger (1911); by
implication (Fryer 1910, 19) he also collected the gecko Hemidactylus
frenatus.

Sil

Table 7. Key to the literature on insects collected at Denis Island

Family Number of species Source
Orthoptera ©) Bolivar 1912, 1924
Dermaptera 1 Burr 1910

Odonata 1 Campion 1913

Hemiptera 9 Distant 1913

Neuroptera a Needham 1913

Lepidoptera 12 Meyrick 1911; Fryer 1912
Coleoptera iat Arrow 1922; Champion 1914;

Gebien 1922; Maulik 1931; Scott
1912; Sicard 1912; Fleutiaux
1923

Hymenoptera 4 Cockerell 1912; Forel 1912;
Meade-Waldo 1912

Diptera 14 Edwards 1912; Lamb 1912, 1914,
1922; Lambrecht 1971; Mattingly
and Brown 1955; Stein 1910;
Theobald 1912; Bezzi 1923

MAMMALS

In 1773 De Trobriand also mentioned the presence of "vaches
marines" on Denis Island (Fauvel 1909, 47). There is no other mention
of these animals on the island, other than Fryer's (1910, 19) comment
that "the dugongs are extinct". As with the similar but more extensive
records at Bird Island, it is probable that these animals were seals
rather than dugongs (Stoddart 1972, 215).

BIRDS

De Trobriand in 1773 found the island covered with birds, "dont
plusieurs esp@éces inconnues A ceux de nous qui avons faits des
campagnes rares. Ces oiseaux étaient si peu accoutumés a voir des
hommes que nous en avons pris une trés grande quantité dans les arbres
et que nous en avons tué autant que nous avons voulu avec des batons";
he also speaks of "la prodigieuse quantité d'oiseaux de toutes
espéces" (in Fauvel 1909, 47).

32

The island lacks the enormous Sooty Tern colony of Bird Island,
and also the colony of Wedge-tailed Shearwaters. The two most
abundant seabirds, especially in the Casuarina, are the White Tern
Gygis alba and the Brown Noddy Anous stolidus. Fryer (1910)
recorded the former as common but did not mention the latter. The
Crested Tern Thalasseus bergii was also seen in 1977.

Fryer (1908, 1910) recorded the Souimanga Sunbird Nectarinia
sovimanga (as Cinnyris) in 1908, and this is the only native landbird.
All the other land birds are introduced. They include: Barred Ground
Dove Geopelia striata, introduced prior to Fryer's visit in 1908;

Mynah Acridotheres tristis, not mentioned by Fryer, and, as at Bird,

a more recent colonist; Madagascar Fody Foudia madagascariensis,
introduced to Mahe in 1879 and to Praslin in 1904, but already "common"
on Denis in 1908; Turtledover Streptopelia picturata, also present in
1908; Moorhen Gallinula chloropus, “extremely common" in the marshy
areas in 1908 and also in 1977; and escaped Gallus in 1908.

Migrants include Turnstone Arenaria interpres, common in 1977, and
Whimbrel Numenius phaeopus, collected in 1908 and seen in 1977.

There are no records of Tropicbirds, Boobies and Frigatebirds,
and they certainly do not nest. The absence of attention to the birds
of Denis by previous workers on Seychelles birds is remarkable; there
are no records for the island given by Vesey-FitzGerald (1940, 1941),
Gaymer et al. (1969), Loustau-Lalanne (1962, 1963), or Betts (1940).

HISTORY

At the time of its discovery in 1773, Denis was uninhabited and
there was no trace of any human occupation. The island was leased on
31 December 1815 to Captain Lesage (Unienville 1838, III), presumably
for fishing and turtling, but nothing at all is known of this period in
its history. It presumably became permanently occupied, and in about
1880 a 40 ft high lighthouse tower was erected. This collapsed on
10 December 1881. An iron tripod was then built to hold the light,
but this rusted badly, and a stone light was proposed to replace it in
1890. The light was rebuilt in August 1893 but Fryer (1908) found it
"somewhat primitive". The present cast-iron braced tower was built in
July 1908-September 1910 (Seychelles Archives: B/38, 434, 472; B/39,
168; B/41, -262; -B/42, 22) \289,. 304;4319,. 406; B/43,7.337, 365: B/Ae,
1/25, 2057 B/ 50S) In 1903 the population consisted of 27 men, 7
women and 2 children (Seychelles Archives C/SS/74(1), 139).

In 1906 W.F. Stephen made the first report on the guano deposits,
which he found to be more extensive than at Bird Island but untouched.
He estimated (1906a) that over half the island was covered with guano,
giving a total reserve of 25,000-30,000 tons. He took 30 samples, and
later analyses (1960b) gave phosphoric acid contents for the guano of
26.2, 31.3, 30.6 and 29.7 per cent, and.of: the: wunderlyingwrock 27.3;
24:5, 20,2, 19.0, and 20.7*> per? cent. Serious digging did not start

38

until 1929, however. Between 1929 and 1941 a total of 16,195 tons
were exported, mostly in the period 1929-1934 (Baker 1963). After the
guano digging ended, coconut palms were rapidly planted over the whole
island. Donkeys, pigs and poultry had been introduced by the early
1960s (Piggott 1969).

It is difficult to give any precise idea of the ecological history
of Denis Island, since it, like Bird, has clearly undergone profound
changes in recent centuries. The amount of phosphate rock must
indicate the existence of large seabird colonies within the past five
thousand years. These may have been tree-nesting species such as the
Red-footed Booby, if the island had Pisonia woodland on it, or ground-
nesting species such as the Masked and Brown Boobies, if it were
largely bare. The former existence of a large tortoise colony, which
would require adequate shade, and the presence of many broad-leaf trees
in the coconut woodland, as well as De Trobriand's account in 1773,
suggest that the island, unlike Bird, was wooded, and that vegetation
was not suppressed by ground-nesting birds in the numbers of the Bird
Island Sooty Terns. The occurrence of woodland would also account for
the existence of a native sunbird and the early introduction of other
landbirds. Presumably the native seabirds must have been largely
destroyed during the fishing, turtling and coconut growing of the
nineteenth century, and the tortoise itself probably became extinct
at a very early date. The very extensive guano digging and coconut
planting fifty years ago would have completed the ecological
transformation.

An airstrip was cut by the owners of the island in 1975-76, anda
small tourist hotel opened in 1978. This should give the opportunity
for more detailed scientific observations in the future.

ACKNOWLEDGEMENT

We thank M. Pierre Burckardt for making it possible to work on
Denis Island in 1977.

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4. Plants recorded from Denis Island

POLYPODIACEAE

Acrostichum speciosum Willd.
Stoddart 8109 (US).

Asplenium sp.?
Fryer (1910) (but Christensen 1912 records no ferns).

Nephrolepis cf. hirsutula (Forst.f.) Presl
Stoddart) 8083) (US); -

Polypodium scolopendria Burm.f.

Stoddart 8097 (US).

TYPHACEAE
Typha javanica Schnizl ex Rohrb.

Jeffrey 1209 (MAHE); Stoddart 8143 (US); Fryer (1908) (as Typha) ;
Fryer (1910) (as Typha angustata); Summerhayes (1931) (incorrectly
cited as from Bird Island).

POTAMOGE TONACEAE
Thalassodendron ciliatum (Forsk.) d. Hart.
Fryer (1910) (as Cymodocea).

GRAMINEAE

*Cenchrus echinatus L.
Jeffrey 1202 (MAHE); Stoddart 8140 (US).

Dactyloctenium ctenoides (Steud.) Bosser
Stoddart 8098 (US).

36

*ETAGrOStIS Ciliaris (ie) Rear.
Jeffrey 1199 (MAHE).

*Eragrostis tenella (L.) Beauv. var. tenella
Stoddart 8104 (US).

Eragrostis tenella (L.) Beauv. var. insularis Hubb.
Stoddart 8139 (US).

*Fleusine indica (L.) Gaertn.
Stoddart 8082 (US).

Enteropogon monostachyos (Vahl) S. and E.
Jeffrey 1200 (MAHE) (= E. sechellensis (Baker) Dur. and Schinz?)

Enteropogon sechellensis (Baker) Dur. and Schinz.
Stoddart 8087 (US).

*Panicum maximum Jacq.
Stoddart 8134 (US).

*Pennisetum polystachion (L.) Schult.
Stoddart 8102> (US)F, Silos: (US).

Sporobolus virginicus (L.) Kunth.
Stoddart 8150 (US).

Stenotaphrum dimidiatum (L.) Brong.
Stoddart 8094 (US).

Stenotaphrum micranthum (Des.) C.E. Hubb.
Jeffrey 1208 (MAHE).

*Tricholaena rosea Nees
Rhynchelytrum repens (Willd.) C.E. Hubb.
Stoddart 8101 (US).

CYPERACEAE

Cyperus conglomeratus Rottb.
Cyperus pachyrrhizus Nees ex Boeckl.
Pycreus odoratus (L.) Urban
Jeffrey 1205 (MAHE) (as var. effusus (Rottb.) Kuk.); Stoddart
8099 (US); Summerhayes (1931) (as Pycreus odoratus and as C.
pachuyrrhizus)

Cyperus dubius Rottb.
Stoddart 8122 (US).

37

Cyperus ligularis L.
Jeffrey 1201 (MAHE) (mislabelled as from Bird Island); Stoddart
8112 (US), 8144 (US).

*Cyperus rotundus L.
Stoddart 8142 (US).

Fimbristylis cymosa R.Br.
Stoddart 8121 (US).
PALMAE
*Cocos nucifera L.
Bayer n( IOS, lOO), sights) Piggott (1969)), sight; Stoddart, isight.
ARACEAE
*Alocasia macrorrhiza (L.) Schott
Stoddart 8110 (US).
LILIACEAE
*Agave rigida Northrop var. sisalana Perr. ex Engelm.
Agave sisalana Perr.
Stoddart, sight.
*Crinum sp.
Stoddart, sight.
MUSACEAE
*Musa sapientum L.
Stoddart, sight.
CASUARINACEAE
*Casuarina equisetifolia L.
Fryer (1910), sight; Piggott (1969), sight; Stoddart 8085 (US).
MORACEAE
Ficus sp.

Fryer (1908), sight (as “banyan"), (1910), sight; Piggett (1969),
sight.

38

*Ficus benghalensis L.
Stoddart 8141 (US).
AMARANTHACEAE

*Achyranthes aspera L. var.?
Jeffrey 1210 (MAHE).

*Achyranthes aspera L. cf. var. aspera
Stoddart 8086 (US).

*Amaranthus cf. dubius Mart. ex Thell.
Stoddart 8105 (US).

NYCTAGINACEAE

Boerhavia repens L.

Jeffrey 1206 (MAHE) (as Boerhavia coccinea Mill., but probably

this species); Stoddart 8093 (US).
Pisonia grandis R.Br.
Fryer (1908), sight (as 'Mapou'), (1910), sight (as Pisonia
calpidia); Piggott (1969), sight; Stoddart 8148 (US).
PORTULACACEAE
Portulaca oleracea L. var. granulato-stellulata v. Poelln.
Stoddart 8092 (US).
LAURACEAE
Cassytha filiformis L.
Piggott (1969), sight (as Cassytha); Stoddart, sight.
HERNANDIACEAE
Hernandia sonora L.
Fryer (1908) sight (as 'Bois blanc'), (1910), sight (as
Hernandia peltata); Piggott (1969), sight (as Hernandia peltata).

MORINGACEAE

*Moringa oleifera Lam.
Stoddart 8135 (US).

39

CRASSULACEAE

*Kalanchoe pinnata (Lam.) Pers.
Stoddart 8114 (US); Fryer, no number, in Summerhayes (1931) (as
Bryophyllum pinnatum Kunz:).

LEGUMINOSAE

*Canavalia gladiata DC.
Fryer, no number, in Summerhayes (1931).

*Cassia occidentalis L.
Stoddart 8106 (US), 8136 (US).

*Crotalaria verrucosa L.
Stoddart 8111 (US).

ZYGOPHYLLACEAE

*Tribulus terrestris L.
Fryer, no number, in Summerhayes (1931).

RUTACEAE

*Citrus aurantiifolia (Christm.) Swingle
Noted by Visiting Magistrate in 1903 (Seychelles Archives).

SURIANACEAE

Suriana maritima L.

Jeffrey 1207 (MAHE) (mislabelled Pemphis acidula Forst.) ;
Stoddart 8138 (US); Fryer (1910), sight (as Pemphis acidula); Piggott
(1969), sight; Fryer, no number, in Summerhayes (1931) (as Pemphis
acidula).

EUPHORBIACEAE

*Acalypha indica L.
Fryer, no number, in Summerhayes (1931).

_*Euphorbia cyathophora Murr.
Euphorbia heterophylla L.
Jeffrey 1204 (MAHE) (as Euphorbia heterophylla L.); Stoddart 8090
(US).

*Euphorbia hirta L.
Fryer, no number, in Summerhayes (1931); Stoddart 8084 (US).

40

Euphorbia microphylla Heyne ex Roth.
Fryer, no number, in Summerhayes (1931).

*Euphorbia prostrata Ait.
Fryer, no number, in Summerhayes (1931).

*Pedilanthus tithymaloides (L.) Poit.
Stoddart 8151 (US).

Phyllanthus maderaspatensis L.
Jeffrey 1198 (MAHE); Fryer, no number, in Summerhayes (1931).

*Ricinus communis L.
Stoddart 8125 (US).

MALVACEAE

*Abutilon indicum (L.) Sweet
Stoddart 8116 (US); Fryer, no number, in Summerhayes (1931).

*Gossypium hirsutum L.
Gossypium maxicanum Tod.
Fryer, no number, in Summerhayes (1931).

Hibiscus tiliaceus (L.) L.
Stoddart 8118 (US); Fryer (1908), sight (as 'Bois du var"), @910);
sight; Fryer, no number, in Summerhayes (1931).

*Sida acuta Burm.f.
Stoddart 8128 (US).

Sida parvifolia DC.
Jeffrey 1203 (MAHE); Stoddart 8088 (US).

Thespesia populnea (L.) Sol. ex Correa?
Stoddart.-8113 “ (US) .
GUTTIFERAE
Calophyllum inophyllum L. var. (probably all var. takamaka Fosb.)
Stoddart 8115 (US), 8119 (US); Fryer (1908), sight (as 'Takamaka'),
(1910)), Sight; Piggott. (1969)}iarsighte.
TURNERACEAE

*Turnera ulmifolia L.
Stoddart 8130 (US).

41

PASSTFLORACEAE
*Passiflora suberosa L.?
Stoddart 8146 (US).
CARICACEAE
*Carica papaya L.
Stoddart 8149 (US).
CUCURBITACEAE

*Cucurbita cf. maxima Duchesne
Stoddart, sight.

Melothria maderaspatana Cogn.
Fryer, no number, in Summerhayes (1931).

[ LYTHRACEAE |
[Pemphis acidula Forst.

Fryer (1910), sight (in error for Suriana maritima L.);
Summerhayes (1931) reporting Fryer's sight record. |

COMBRETACEAE

Terminalia catappa L.
Stoddart 8126 (US); Piggott (1969), sight.

BARRINGTONIACEAE

Barringtonia asiatica (L.) Kurz.
Stoddart 8147 (not retained).

APOCYNACEAE
*Catharanthus roseus (L.) G. Don

Stoddart 8124 (US), 8131 (US); Fryer (1910), sight (as Vinca
rosea); Piggott (1969), sight (as Catharanthus).

CONVOLVULACEAE

Ipomoea sp.
Pryern (1910), ‘sight.

42

*Tpomoea coccinea L.
Jeffrey 1211 (MAHE).

Ipomoea macrantha R. and S.
Stoddart 8152 (US).

*Tpomoea obscura (L.) Ker-Gawl
Stoddart 8145 (US).

Ipomoea pes-caprae (L.) R.Br. ssp. brasiliensis (L.) v. Oooststr.
Stoddart 8120 (US).
BORAGINACEAE

Cordia subcordata Lam.
Stoddart 8096 (US), 8117 (US).

Tournefortia argentea L.f.

Stoddart 8107 (US); Fryer (1908), sight (as 'Bois tabac"), (1910), sight
(as Tournefortia); Fryer, no number, in Summerhayes (1931); Piggott
(1969), sight (as Tournefortia).

VERBENACEAE

*Lippia nodiflora (L.) Michx.
Stoddart 8123 (US).

*Stachytarpheta jamaicensis (L.) Vahl
Stoddart 8129 (US); Fryer, no number, in Summerhayes (1931).
SOLANACEAE

*Capsicum frutescens L.
Fryer, no number, in Summerhayes (1931).

*Datura metel L.
Fryer, no number, in Summerhayes (1931).

Solanum nigrum L. var. americanum (Mill.) O.E. Schulz
Stoddart 8103 (US).
SCROPHULARIACEAE

Strigawastaticanm ln.
Stoddart 8089 (US).

43

BIGNONIACEAE
*Tabebuia heterophylla (L.) Hemsl.
Stoddast 8s3) (US):
ACANTHACEAE
Asystasia bojeriana Nees
Stoddane lJ) (US)e
RUBIACEAE

Guettarda speciosa L.
Stoddart 8091 (US).

Morinda citrifolia L.
Stoddart 8095 (US).

Spermacoce repens (DC.) Fosb. and Powell
Stoddart 8093 (US).

GOODENIACEAE

Scaevola taccada (Gaertn.) Roxb.
Stoddart 8137 (not retained); Fryer (1908), sight (as "Bois
veloutier').

COMPOSITAE

Vernonia cinerea (L.) Less, var. parviflora DC.
Stoddart 8132 (US).

Synedrella nodiflora (L.) Gaertn.
Stoddart 8100 (US).

SOURCES

Fryer collections and observations: Fryer's paper on Denis Island
(1910) and his manuscript diary (1908); enumerations of flowering
plants in Summerhayes (1931).

Piggott observations: Piggott (1969).
Jeffrey collections: Department of Agriculture, Mahe.

Stoddart collections: determinations by F.R. Fosberg.

Species marked * are judged to be introduced.

Some of the sight records may well be misidentifications.

Ghee REALE ene e) BN

| * pbwep Bis fener (aN
we Pady ,

icine ior ee

Avett \ wrt SeGET eh 6
athe aia) ete

pet 4 Le

ree iY : i @ hee %
PIOi.74 \ Vass ki +
hry 3 y TH} F
pared. (2.0) set bom ae
(30) GOLe teh
fa nue

iivoktes 2a oitee

Dr ea ts ee
a oa A 2) ae

fou val Bowoass PAY

5. References

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46

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47

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48

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50

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ornithology of the Seychelles Islands. JIbis, (14) 5: 518-531.

[]sana

mwmEroding shoreline

4+ Beach ridge crest

Inland phosphorite

Phosporite outcrop
at shore

YA Beachrock

— Road

P Pisonia

H Hibiscus

x Calophyllum

ED)

a
<

¥ Casuarina

oy

G Guettarda

Vay

€ Cordia

Ss Scaevola

© Tournefortia

Suriana

°
BS} Herb mat benchmark

[[]]}cocehut woodland

rn — Marsh and
ae standing water

{e) m 200

Fig. 4. Denis [stand in 1977,

6500

SOG,

AOO

30©

200

100

Was. oD)

Gas
D>
Ce

Monthly rainfall at Denis Island, 1971-1962

Pl. 9. Denis Island: phosphate cliffs with Casuarina woodland,

southwest shore

Pl. 10. Denis Island: phosphate cliffs at Muraille Bon Dieu.

PIL, dlls iDeraats) alsialewayels

Pi, 125 Wea Wesleiael?

+ ee

detail of phosphate

Typha swamp

+

—

cliffs at Muraille Bon Dieu

ATOLL RESEARCH BULLETIN
NO. 253

TOPOGRAPHIC AND FLORISTIC CHANGE, DRY
TORTUGAS, FLORIDA, 1904-1977

by D. R. Stoddart and F. R. Fosberg

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

July 1981

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Contents
Abstract
Introduction
Environment
Descriptions of the keys
Bird Key
Bush Key
East Key
Garden Key
Loggerhead Key
Long Key
Middle Key
North Key
Northeast Key
Sand Key
Southwest Key
Flora
Vegetation
Conclusion
Acknowledgements
Plants recorded from the Dry Tortugas Keys
References
Tables

Figures

abate

16.

UAE

18:

Loe

20%.

List of Tables
(pages 33-55)

Dimensions and area of Bird Key.
Plants recorded from Bird Key.
Dimensions and area of Bush Key.
Plants recorded from Bush Key.
Dimensions and area of East Key.
Plants recorded from East Key.
Dimensions and area of Garden Key.
Plants recorded from Garden Key.
Dimensions and area of Loggerhead Key.
Plants recorded from Loggerhead Key.
Dimensions and area of Long Key.
Plants recorded from Long Key.
Plants recorded from Middle Key.
Dimensions and area of Sand Key.

Plants recorded from Sand Key.

Distribution of native plants (excluding sea-grasses) on

Tortugas keys.

Dry

Numbers of species in different categories recorded from the Dry

Tortugas keys.

Summary of area and plant species numbers for Dry Tortugas keys.

'Extinction' of plant species of Dry Tortugas keys,

Species-area data for Alacran Reef.

1904-1916.

HO}

eee

niglst

List of Figures

(pages 56-66)

The Dry Tortugas Bank.
Bird Key in 1904 and 1915-16. (Millspaugh 1907, Bowman 1918).

Bush Key and Long Key in 1915-16, 1937 and 1977 (Bowman 1918,
Davis 1942, and Stoddart in 1977).

Bush and Long Keys in 1977.

Sand Key and East Key in 1904, 1915-16 and 1977 (Millspaugh 1907,
Bowman 1918, and Stoddart in 1977).

East Key in 1977.

Garden Key in 1904, 1911-13, 1915-16, and 1937 (Millspaugh 1907,
Vaughan 1918, Bowman 1918, and Davis 1942).

Loggerhead Key in 1904, 1915-16, 1937 and 1977 (Millspaugh 1907,
Bowman 1918, Davis 1942, and Stoddart in 1977).

Loggerhead Key in 1977.

Sand or Hospital Key in 1904, 1915-16, and 1977 (Millspaugh 1907,
Bowman 1918, and Stoddart in 1977).

Species-area relationships for the Dry Tortugas keys and other
coral islands in the Gulf of Mexico and Caribbean Sea. Sources:
Pedro Cays: Chapman 1944, Steers 1940, Steers et al. 1940,

Zans 1958, Fosberg unpublished observations in 1962; Morant Cays:
Chapman 1944, Asprey and Robbins 1953; Alacran: Fosberg 1962;
Glover's Reef: Stoddart and Fosberg, in press; Dry Tortugas: this
paper.

iv

Oye

ple

List of plates

(following page 66)

Bush Key (foreground) and Long Key seen from Fort Jefferson.
Note the tall Conocarpus woodland on Bush Key and the extensive
mangrove on Long Key. Compare this view with that in Davis
(1940), plate 12.

Bush Key: herb mat dominated by Sesuvium along the south coast.

Bush Key: Tournefortia and Opuntia scrub on the narrow eastern
Spies

Bush Key: tall Conocarpus woodland on the eroding north shore.
Bush Key: Avicennia woodland in the eastern bay.

East Key: Uniola paniculata and Iva imbricata on the eastern
ridge.

East Key: Uniola paniculata at the eroding northeast point.

East Key: open vegetation of Uniola, Sesuvium, and Cakile
(foreground) in the south-central part of the key.

Loggerhead Key: tall Casuarina woodland in the middle of the key,
seen from the anchorage on the east side.

Long Key: Rhizophora woodland on the west shore.

Sand Key: seen from the west end.

TOPOGRAPHIC AND FLORISTIC CHANGE, DRY
TORTUGAS, FLORIDA, 1904-1977

by D. R. Stoddart! and F. R. Fosberg?

ABSTRACT

Topographic and floristic surveys of the Dry Tortugas keys in
1904, 1915 and 1937 have been used in discussions of the changing
relationships between area and floristic diversity on small islands
over time, and of the processes of colonization and extinction. ie IS
shown that earlier topographic surveys are in general too unreliable
to be so used. A list of Dry Tortugas plants, including all published
records as well as new collections made in 1962 and 1977, is presented,

together with maps of the keys made in 1977. The total flora of about
130 species includes at least 35 native species, including 5 species
of sea-grasses and 4 species of mangroves. Introduced species are

largely confined to the two largest islands, and the floras of the
smaller keys are dominated by a small number of native species.

INTRODUCTION

The Dry Tortugas Bank (Figure 1) lies at 24°40' N, 82°50' W,
110 km west of Key West, Florida and 165 km north of the nearest point
of Cuba. It has a maximum NE-SW dimension of about 18 km, and is
5-8 km wide. As outlined by the 60 ft (18.3 m) isobath, the Bank
covers almost exactly 100 sq km. Reef shoals in the centre and
eastern part of the Bank rise to less than 6 ft (1.8 m) and have a
total area of 245 ha, or only 2.5 per cent of the total Bank area.
These shoals have historically supported eleven sand keys, of which
three disappeared between 1775 and 1875 and a further three during the
present century. The total land area of the eight keys existing in
1900 was about 50 ha; of this, two islands (Loggerhead and Garden Keys)
comprised some 70 per cent.

1. Department of Geography, Downing Place, Cambridge, England.
2. Smithsonian Institute, Washington, D.C. 20560, U.S.A.

Manuscript received August 1980 --Eds.

The reefs and keys of the Dry Tortugas have considerable
historical interest. Here Agassiz (1888) first mapped the distribution
of coral communities on any extensive reef tract (though his map,
showing extensive areas of the now absent Acropora palmata, was
perhaps somewhat conjectural); here too the presence of the Tortugas
Laboratory of the Carnegie Institution led to a long series of
research studies published between 1908 and 1942. The reefs have been
studied by Vaughan (1914), Brooks (1962) and Jindrich (1972), the latter
emphasising the role of hurricanes in controlling community succession
and sediment accumulation. More recently, Shinn et al. (1977) have
shown from shallow borings that the topographic highs on the Bank are
Holocene reef constructions at least 8 m thick which have accreted at
rates of 1.9-4.5 m/1000 yr.

The keys themselves have been the subject of a series of
topographic, geological and botanical studies. Detailed accounts of
individual islands are available at intervals over more than 70 years,
and this presents an opportunity to study changes in a way not
possible for most other reef islands in the world. Indeed, Millspaugh
(1907, 191), who published the results of O.E. Lansing's first surveys
in 1904, stated that "the principal value of the survey lies ... in
the historical record of the present flora". The main studies
available are as follows:

1904, March 19-22 O.E. Lansing, Jr., reported by
Millspaugh (1907)

TOMS awiulivae i ONG H.H.M. Bowman (1918)

1926, June W.R. Taylor (1926, 1928); algae
and sea-grasses

1931, June-July; G. Tandy, land plants; no
publication

1933, August and collections not seen

ASIEN AES IS XS} Davis (1940, 1942)

1962, August F.R. Fosberg

1977, May-June D.R. Stoddart

The most important of the earlier studies are those of Millspaugh,
Bowman and Davis. In addition a great deal of incidental topographic
and botanical information is contained in extensive studies of the

Dry Tortugas sooty tern colonies, notably by Stevenson (1938), Sprunt
(1948, 1950, 1962-63), Robertson (1964), and Dinsmore (1972).
Robertson in particular makes a major attempt to collate all previous
information, published and unpublished, on the topographic development
and nomenclature of the keys, and his results have been used in the
presen paper.

Given this data base, it is not surprising that MacArthur and
Wilson (1968), 51-55) used the Dry Tortugas as one of their two examples
(the other being Kapingamarangi Atoll) of the relationship between
floristic diversity and area on small islands. MacArthur and Wilson
used plant diversity levels in 1904 and 1916 (the Lansing and Bowman
surveys) to calculate extinction rates during this period, and related
these to island areas derived from Bowman's surveys.

Plant collections were made by the present authors on the Dry
Tortugas keys in 1962 and 1977, and the islands were mapped by pace-
and-compass methods in 1977. Comparison of the 1977 maps with earlier
maps shows beyond doubt that the maps published by Millspaugh are
thoroughly unreliable, and indeed none of his maps possesses any scale
atl all: Even worse, the scales of maps published by Bowman in 1918
are variable and misleading: the scales of each of his maps are given
only in captions, and it is clear that the printer has differentially
reduced the figures without a corresponding correction in the cited
scale. The island areas derived from Bowman's maps by MacArthur and
Wilson are thus wrong. Davis's (1942) maps, in contrast, are
meticulous, but unfortunately he maps only Bush, Garden, Loggerhead and
Long Keys.

In view of this situation, and especially of the theoretical
discussions already based on the Dry Tortugas data, this paper does

two things. First, we present maps of the keys in 1977, compare them
with previous maps, and attempt to provide reasonable estimates of
Size and area at the times of the successive surveys. In this section

the earlier work of Robertson (1964) is of outstanding importance.
Second, we list the plant collections made in 1962 and 1977 and collate
the previous records given by Millspaugh, Bowman and Davis. This
collation is based only on literature records; no herbarium material
from the earlier surveys has been examined; nor have we seen any lists

or collections from Tandy's surveys in 1931 and 1933. These data are
Ehengused tomprovide) revased illomi strc wists, form cach masland at) the
time of each survey. Finally, the usefulness of these revised data

for the kind of analysis carried out by MacArthur and Wilson is
discussed.

ENV ITRONMENT

Rainfall recording at Key West began in 1832; the long term annual
mean is 961 mm, with extremes of 1770 mm (1870) and 520 mm (1838).
There is a dry season from December to April, with an average of less
than 50 mm rainfall per month, a secondary rainfall peak of 105 mm in
June, and the main wet season from August to October. September is
the wettest month with 165 mm. Hurricanes are frequent between
August and December and may bring heavy rainfall as well as high winds
and seas; the highest individual monthly rainfall recorded is 533 mm
in November 1954. Mean daily temperature ranges from 21.3°C in
January to 29.1°C in August; cold spells of several days, with
temperatures exceptionally approaching O°C, may occur during 'northers'

in’ the! dry season’. Prevailing winds are easterly, and so are currents
(Vaughan 1935), so that the fauna and flora of the Keys are probably
mainly derived from the Florida Keys and mainland. The Dry Tortugas
are almost tideless: range at springs (MHHW to MLLW) is only 0.2 m.

DESCRIPTIONS OF THE KEYS
Bird Key

Bird Key, the Booby Key of Gauld (1790), was located 1.3 km
southwest of Garden Key; it disappeared in 1935. Audubon (1835,
263-269) visited it (and called it Bird Key) in May 1832, when it was
occupied by a party from Havana collecting tern and noddy eggs.

Robertson (1964) quotes a survey by Tatnall and Gednery in 1829
which gave an area of 4 acres 2 roods 20 poles, equal to 18,210 m’,
and an estimate by Scott in 1890 of about 8 acres, or 32,400 m?.
Millspaugh (1907) gave dimensions of 500 x 250 ft (= 153 x 76m),
derived from Lansing's map of 19 March 1904, and if this is used to
scale the map the resulting area is 6,440 m’. Bowman's (1918) map,
at its cited scale of 1:6340, yields an area of 47,695 m?; MacArthur
and Wilson (1968) derive an area of 13,935 m* from Bowman's account;
while correcting Bowman's scale to 1:2717 by reference to his text,
where he gives dimensions of 500 x 300 ft (= 152 x 92 m), yields an
area of 9,375 m?. Watson (1908), however, gave dimensions of
400 x 300 yds (= 366 x 274 m). Table 1 summarises dimension and area
estimates for the island, and Figure 2 gives the maps by Lansing and
Bowman at the same scale and orientation: it is clearly impossible to
estimate the size of the island with any confidence from these
conflicting data.

Wurdemann (1861) in 1857 described a scrub of Suriana 7-8 ft
(2.1-2.4 m) high, with some Opuntia. Lansing found a 'dense growth of
Suriana maritima', with a marginal zone of herbs and shrubs, especially
on the eastern side (Millspaugh 1907, 233). Watson (1908, 192) also
mentioned the Suriana cover, as well as Opuntia. Much of the Suriana
was killed by a hurricane on 15-17 October 1910, after which Bowman
(1918) described Euphorbia mesembrianthemifolia as dominant, with
extensive Portulaca and Opuntia. The vegetation was stripped in
another storm on 10-11 September 1919 (Robertson 1964, 8, 60-61), and
the key itself was destroyed by a hurricane variously dated as 1933,

1935. and: 1938). By 1934 it was no more than a sandbar. In January
1940 it was a sandbar 40 ft (12 m) long and 2 ft (0.6 m) high
(Robertson 1964, 9); it was not seen in 1977.

Table 2 lists the plant species recorded by Lansing on 19 March
1904 and by Bowman in 1915-1916. The total flora of 15 species
includes only one certain introduction, Cocos nucifera, which was
probably planted between the two surveys. Bartsch (1919, 470)
mentions "a few ornamental shrubs near the buildings", but there is no
record of their identity:

Apart from the noddies and sooty terns, Robertson (1964, 25)
records that pigs were taken to the island from Long Key in July 1865.

Bush Key (Plates 1-5)

Bush Key, immediately east of Garden Key, has had a varied
topographic history. It does not appear at all in Gauld's (1790)
survey of 1773-75, but according to Bartsch (1919) it was vegetated at
the time of Audubon's visit in 1832. Robertson (1964) quotes the
Tatnall and Gednery survey of 1829 as yielding an area of 5 acres
3 roods 22 poles, or 23,270 sq m. Robertson summarises the subsequent
history as follows: a sizeable island by the 1850s; hurricane damage
ca 1870; a sandbank in 1889; not mentioned as a Separate island in
1890; used as a source for dredged sand for fill at Garden Key in
1901-5. The island was stated to be awash in Millspaugh's (1907)
account of Lansing's 1904 survey.

Bowman (1918) prepared the first map in 1915-16 (Figure 3), ina
composite map of Bush and Long Keys, said to be on a scale of 1:6340.
Bush was given dimensions of 583 x 342 m, and was said to have shrubs
on it 12 years old. However, Bartsch (1919, 469) said in 1917 that all
vegetation had 'long since' been swept away, and that the island was
"barren'; it is, however, possible that he was really describing Long
Key rather than Bush. It is clearly difficult to relate Bowman's
map to the present topography of the key, or even to Davis's map of
1937-38; Davis (1942, 187) indeed states that Bowman's map is wrongly
oriented by 90°, but even reorientation does not make identification
easier.

Davis's map is the first reliable one, giving an area of
tS 9O0Or mer. Davis (1942, 188) shows three small ponds at the western
end of the key, said to have resulted from dredging in 1905, to have
been cut off from the sea since about 1919, and to have been infilled
during the next decade as the eastern spit gradually extended towards
Long Key. He noted (1940, 377) a ‘young swamp of Laguncularia' around
one of these ponds, together with recently established Avicennia and
Rhizophora. Many Rhizophora seedlings also drifted ashore here during
dispersal experiments in 1937-38 (Davis 1940, 373). In 1942 bombing
practice led to a fire 'that burned all vegetation' (Robertson 1964,
3}5))) 6 In 1946 Sprunt (1948, 6) found a 'remarkable spread' of
vegetation. There was an eastern pond with Laguncularia, Rhizophora
and some Suriana, and a western pond with Opuntia, Salicornia and
Tournefortia on its margins; Salicornia and Tournefortia covered
large areas of the key.

In 1962 Fosberg found a dense thicket of Laguncularia with a few
Rhizophora in the main depression; a second small pond covered with
Rhizophora 4-5 m tall; and much of the island occupied by Suriana
scrub 2 m tall and by Opuntia, with scattered Sesuvium, Sporobolus and
Uniola. Areas at the western end used by the sooty terms were almost
bare of vegetation. In 1977 the island was much narrower than xin 1937.
The location of Conocarpus and Laguncularia thicket indicates that the
narrowing has been by erosion along both north and south shores. The

Main change in the vegetation has been the development of Avicennia
woodland within the eastern spit. Dinsmore (1972, 131-132) gives an
accurate brief account of the vegetation.

Table 4 lists the plants recorded in 1915-16, 1937, 1962 and 1977.
Of the total of 28 species, 25 are native, in spite of the very large
number of introductions on the adjacent Garden Key. Casuarina, the
only certain introduced species,was first recorded in 1977, and could
have dispersed naturally from Garden Key.

East Key (Plates 5-8)

East Key was charted by Gauld (1790) in 1773-75. Tattnall and
Gednery in 1829 give an area of 12 acres or 48,600 m* (Robertson 1964).
The key was said to be eroded during the nineteenth century, but
Agassiz's (1888) chart shows the island as one of the largest in the
group, with a length of about 500 m and a pronounced NW-SE orientation.

Scott in 1890 gave an area of 18 acres or 72,800 m°’. Lansing mapped it
on 21 March 1904, when it was 'a mere sand bank about 280 x 50 feet

in area' (Millspaugh 1907, 224-225). Using these dimensions to scale
his map gives an area of 2170 m?; if, however, the measurements were

really in yards rather than feet the area would be 6500 m°’. Bowman's
(1918) map at its stated scale of 1:6340 yields an area of 115,610 m?;
MacArthur and Wilson (1968) estimate the area from Bowman's survey at
111,480 m?; but if Bowman's dimensions of 540 x 250 m are used to
rescale the map to 1:5512 the area becomes 87,220 m?. Other estimates
are given in Table 5. Unfortunately the island was not mapped by
Davis. Its area in 1977 was 27,145 m*, and its orientation had
altered to NE-SW. The sediments of the key have been studied by
O'Neill (1976). The variation in size alone of the maps of the
island shown in Figure 5 strongly suggests that gross errors have
entered the earlier records.

Robertson (1964, 12) quotes Holder's (1892) observation in 1860
that the vegetation comprised 'a dense stand of bay cedar bushes
[Suriana] and numerous mangroves'; records in 1875 mention it 'partly
covered with a growth of cedar', in 1889 as having 'a few bushes on it',
and in 1890 as 'covered in parts with stunted bushes'. According to
Lansing in 1904 "the vegetation consists principally of Cenchrus and
Euphorbia with a sprinkling of Uniola at the southern end; two
isolated patches of Sesuvium near the centre of the islet; anda few
other species scattered without definite association" (Millspaugh
1907, 225): Lansing records Scaevola, Iva and Tournefortia, but not
Suriana. In 1915-16 Bowman (1918) found the key "almost entirely
covered with vegetation", with "large well-grown bushes", but again
he does not record Suriana. Bartsch (1919, 469) describes "a dense
growth of Bermuda grass [meaning Uniola?] on the flattened, upper
portion, with a scattered growth of Scaevola bushes and other plants".
Davis (1942) mentions a thicket of Suriana on the highest ridge,
together with Ipomoea, Sesuvium and Cenchrus. In 1946 Sprunt (1948,
17) found that "Sea-oats (Uniola) or sea-lavender (Tournefortia) and
some bay cedar (Suriana) are the principal growths, and there is a

great deal of the goat's-foot vine (Ipomoea pes-caprae) running along

the beach". Robertson (1964, 13) in about 1960 found "sizeable bushes
of bay cedar [Suriana], sea lavender (Tournefortia gnaphalodes), and
Scaevola plumieri". In 1977 the island was dominated by open Uniola

grassland with Scaevola, Suriana, Iva, Tournefortia and herbs.

Table 6 shows the plants recorded from East Key by Lansing, Bowman,
Davis and Stoddart. All ten species are native. Suriana, not
recorded by Lansing and Bowman, was certainly present in the nineteenth
century, and Ipomoea pes-caprae, also not recorded by them, may have
simply been overlooked. The only mangrove records for East Key are
the Rhizophora seedlings which drifted there in 1937 and 1938 during
Davis's dispersal experiments (Davis 1940, 373).

Garden Key

Garden Key, the Bush Key of Gauld's (1790) survey of 1773-75, has
been enormously altered by the construction of Fort Jefferson, said to
be the largest brick construction in the western hemisphere, during
1846-1864. The original sand key, on which a lighthouse was built in
1825, was estimated at 7.5 acres (30,350 m*) by Tatnall and Gednery
in 1829, and at 8.8 acres (35,610 m?) by Bache in 1845 (Robertson
1964). The total area of the Fort, including the moat, is 64,400 m?,
and the area within the walls 34,000 m?. The Fort was abandoned in
1875; the island was used as a coaling station by the U.S. Navy in
1901-1905, when its areas was increased and modified by dredging in
the channel to the east and on the site of Bush Key; and it was again
abandoned in 1910 and once more in 1925. It was declared a National
Monument in 1935.

The island was mapped by Lansing on 22 March 1904 (Millspaugh
1907, 228-231), by Vaughan (1918), by Bowman (1918), and, in greatest

detail, by Davis (1942). Fort Jefferson itself provides a common
scale to all these maps. By calculation Lansing's map is at 1:3680;
Bowman's is said to be at 1:6340 but is actually at 1:4900; and
Davis's is at 1:4646. Comparison of the maps (Figure 7) shows that

Bowman's is clearly inaccurate (e.g. in the relative location of the
coaling sheds), and that as a result his map overestimates the total
area of the island by about 50 per cent.

Millspaugh (1907, 228-231) and Bowman (1918, 125-128) map the
wwegetation outside) the Fort and describe that: inside; \Davis) (1942,
184-187) gives the most detailed account and distribution map. The
island was not remapped in 1977. In addition to the main sources,
Bartsch (1919, 469-470) gives a brief note on the vegetation. Table 8
lists over 100 species of plants recorded from Garden Key. Of these
only 24 are native, and all of the native species were recorded by
Millspaugh and by Bowman.

Loggerhead Key (Plate 9)

Loggerhead or Turtle Key (of Gauld 1790) is the westernmost of the
group. A lighthouse 50 m high was built on it in 1856-1860. The
island was the site of the Tortugas Laboratory for more than thirty
years (only the foundations now remain); and it is permanently
inhabited. Maps are available for 1904, 1915-16, 1937-38 and 1977
(Figure 8). These show variability in length, which is undoubtedly
seasonal, and also a general decrease in width. Both types of change
are traced by relict beachrock distributions. Loggerhead Key is the
only island in the group to possess actively forming beachrock. This
was noted by Agassiz (1888, 67), but characteristically misinterpreted
as the equivalent of the last interglacial limestones of the Florida
Keys. More recent studies of the Loggerhead Key beachrocks are those
of Field (1919, 1920), Ginsburg (1953) and Multer (1971).

The vegetation has changed greatly over the last century.
Initially, ca 1840, according to Bowman (1918, 120), the island was
covered with ‘a large stand of old white buttonwood trees, Conocarpus
erectus L.' These were either cut down or burned by residents. By
the time of Lansing's visit in 1904 the centre of the island was
covered by a 2 m tall scrub of Suriana. Bowman (1918, 121) found
Suriana dominant, supplemented by an Opuntia association. Bartsch
(1919, 470) also briefly describes this vegetation. Today the island
is dominated by a tall woodland of Casuarina, originally introduced by
the director of the Tortugas Laboratory ca 1910. Casuarina seedlings
were growing all over the island by the time of Davis's survey in
1937-38, though Suriana was still dominant. It is also interesting
that many of the Rhizophora seedlings released up to 65 km east of the
Dry Tortugas by Davis in 1937 and 1938 arrived on the east beach at
Loggerhead Key (Davis 1940, 371-377).

Both Millspaugh (1904, 235) and Bowman (1918, 121) found the key
remarkably free from common weeds. Table 10 lists plants recorded
from the island. Of the total of 57 species, 19 are native.
Fifteen of the 24 definitely introduced species were recorded before
LOL.

Long Key (Plate 10)

Long Key, the Rocky Key of Gauld (1790), was first mapped by
Bowman (1918, 129): it was described as awash and without vegetation
by Lansing in 1904. The difficulties of interpreting Bowman's map
(Figure 3) have already been discussed in the account of Bush Key.
Bartsch (1919, 469) mentioned scattered Scaevola and grasses on the
southern end of the island, which otherwise was a simple boulder ridge.
Davis (1940, 381-382, plate 12; 1942, 189-190) was responsible for
planting mangrove seedlings: a few in 1937, 4100 in 1938, 575 in 1940.
2500 were alive in February 1942. Additional seedlings also drifted
ashore here during dispersal experiments (Davis 1940, 373). In 1940
also there were 25 Avicennia bushes, four of which were more than
60 cm tall. Other plants mentioned by Davis were the mangrove

associates Batis, Salicornia and Sesuvium. Since Davis's survey the
key has reduced in area by about 80 per cent.

By 1962 Fosberg found a considerable low scrub of Avicennia and
much Salicornia on the leeward side, with mats of Sesuvium, and small
numbers of Rhizophora up to 1m tall. In 1977 Stoddart found a
woodland of Rhizophora and Avicennia 4.5 m tall on the west side of the
beach ridge, with strand shrubs and herbs on the seaward side.

Table 12 documents the plant species recorded from Long Key.
Fifteen of the 16 species recorded are native; Casuarina is the only
introduced species, recorded as a seedling 50 cm tall.

Middle Key

Middle Key, the Bird Key of Gauld (1790), is the smallest of the

vegetated islands of the group. It lacked vegetation in 1875
(Robertson 1964). Agassiz's map (1888) shows it to be about 140 m
long. In March 1904 Lansing found it awash, with no vegetation.

Bowman (1918, 131) described it as an oval patch of sand 80 x 50 ft
(24 x 15 m), with an area of 194 m?, with clumps of dead Cakile.
Bowman's map, said to be at 1:6340, is actually at 1:520, and this
yields an area of 370 m?. Davis (1942) described it as 75 ft (23 m)
long in 1937, with Cakile, Iva and Euphorbia. In July 1938 it was
intertidal, and in 1940 and 1942 non-existent. Fosberg in 1962
described it as a crescent-shaped sand bar with no vegetation, and it
remained the same in 1977.

Table 13 lists the plant records from Middle Key: all three
recorded species are native.

North Key
This was charted by Gauld (1790) in 1773-75. It was a sandbar in
1860, and had disappeared by 1875 (Robertson 1964, 16). According to

Bartsch (1919, 492-493) it may have been the island of a few acres in
area named Booby Island by Audubon in 1832.

Northeast Key

This was charted as Sandy Key by Gauld (1790) in 1773-75. In the
1850s it possessed a tern colony, but had disappeared by 1875
(Robertson 1964, 16). It was not seen in 1977.

Sand Key (Plate 11)

Lansing mapped Sand Key (or Hospital Key) (the Middle Key of Gauld
1790) on 21 March 1904, when it was a small oval island (Figure 10) of
only 80 x 50 ft (24 x 15 m), though possibly these measurements should
be in yards (= 73 x 46 m): on Agassiz's (1888) chart the island is
about 160 m long. Bowman (1918) described it as measuring 90 x 45 ft
(27 x 14 m) and mapped it at a scale said to be 1:6340 but actually
1:460. The true area at both surveys, assuming the dimensions in feet

10

to be correct, was about 225 m? (Table 14). The flora was confined to
few scattered herbaceous species; there were no plants on it at all at
the time of Bartsch's (1919, 469) survey. In 1938 Davis gave
dimensions of 110 x 70 ft (34 x 21 m) but did not map it. Ttshad va
sparse vegetation, but Rhizophora seedlings which drifted there in
1937-38 failed to become established (Davis 1940, 373). It was
unvegetated in 1946 (Sprunt 1948). Fosberg noted 'a patch of bushes
and a patch of grass" in 1962. Robertson (1964, 14) described it as
"always a small, shifting sandbar with little vegetation ... no
permanent plant cover has been established". In 1977 it was much
larger than in the past, but again unvegetated (Figure 10).

A hospital was built on the key in the1870s, and became derelict
and ruined at an unknown data. Broken blocks of masonry were noted
on the key by Bowman in 1938, Davis in 1938, and again in 1977. Davis
also noted relict beachrock, which is extensive in shallow water south
of the key.

Table 15 lists plants recorded from the key: all eight species
are native.

Southwest Key

This was charted by Gauld (1790) in 1773-75; it had disappeared
by 1875 (Robertson 1964, 16).

FLORA

The flora comprises 35 native or probably native species (including

five species of sea-grasses and 4 species of mangroves); at least 73
introduced species; and 23 species of uncertain status; giving a
total of 131 species. This compares with 25 species (20 native) at

Alacran Reef, Gulf of Mexico; 16 species (15 native) at the Morant
Cays, 11 species (6 native) at the Pedro Cays, and 63 species at
Glover's Reef, all in the Caribbean Sea. It is striking that the
smaller Dry Tortugas islands, other than Garden and Loggerhead Keys,
are dominated by native species, and that a very small number of native
species occur on most of the islands. These include Cenchrus incertus,
Uniola paniculata, Cyperus planifolius, Sesuvium portulacastrum, Cakile
lanceolata, Suriana maritima, Euphorbia mesembrianthemifolia, Ipomoea
pes-caprae, Tournefortia gnaphalodes, Scaevola plumieri and Iva
imbricata, all present on at least five islands. The other two-thirds
of the native species are more patchily distributed (Table 16).

The native flora is characteristic of small Caribbean/West Indian
coral islands, but with some distinctive additions and several
omissions. The five native grasses include the tall and distinctive
sea-oats Uniola paniculata. This species extends along the coast of
Florida, round the Gulf of Mexico into Tabasco, through the Bahamas,
and along the north coast of Cuba (Yates 1966), but it is otherwise
absent from the Caribbean strand flora. The other grasses are

ial

widespread strand species. The single sedge (Cyperus planifolius) is
also a widespread reef-island species.

The shrubs are dominated by Suriana maritima (also extensive at
Alacran); Tournefortia gnaphalodes, a widespread Caribbean beach shrub;
Scaevola plumieri, common in the northeastern and eastern Caribbean;
and Iva imbricata, a variable plant of the tropical and subtropical
Atlantic coast of North America but not otherwise found on Caribbean
coral islands. Borrichia arborescens, which extends through the
Florida Keys to the Marquesas, and is also extensive in Cuba and the
Bahamas, as well as further south (Semple 1978), is curiously absent
from the Dry Tortugas.

The herbaceous flora includes common reef island species of
Portulaca, Sesuvium, Cakile, Euphorbia, and the vines Ipomoea macrantha,
Ipomoea pes-caprae and Canavalia rosea. Cassytha filiformis is notably
absent. On some islands with suitable substrate conditions there is a
Mangrove community (Conocarpus, Laguncularia, Rhizophora, Avicennia)
with associated Batis maritima, Atriplex pentandra, Salicornia bigelovii,
Alternanthera maritima and Philoxerus vermicularis, but the mangroves
are much less extensive than in the nearby Marquesas and may be a
recent development (Bowman 1918, 115, describes mangroves as ‘entirely
Waclksimnei) ir.

There is no indubitably native tree species,though the status ofCordia
sebestena, Bursera simaruba, Coccoloba uvifera and Thespesia populnea
is open to question. All are common Caribbean coral island species.

VEGETATION

Both Bowman (1918, 115) and Davis (1942) have described vegetation

units from the Dry Tortugas. Bowman distinguished a Uniola grass
community (with Cakile, Cenchrus, Sesuvium, Tournefortia, Ipomoea and
Scaevola); a Suriana scrub community (with Canavalia); an Opuntia
community (with Paspalum, Ipomoea macrantha, Ipomoea pes-caprae, and
Melanthera aspera); and a Euphorbia [Chamaesyce] community with Iva
imbricata and Cyperus planifolius. Davis's categories were broader

and described in terms of topographic units common to the Florida Keys
as a whole. They included:

(a) Strand-beach Associes (Sesuvium, Cakile, Ipomoea, Sporobolus,
Uniola, Euphorbia, Tournefortia) .

(b) Strand-dune Associes (Uniola, Euphorbia, Hymenocallis,
Tournefortia, Suriana).

(c) Strand-scrub Associes (dominated by Suriana, with Caesalpinia,
Solanum and Opuntia).

In addition he describes an Opuntia Associes, a Euphorbia [Chamaesyce]
Associes, an Agave Associes, and a Laguncularia Associes. He comments
on the absence of mangrove communities, except around the ponds on Bush
Key.

12

These categories remain useful for describing vegetation units on
the keys, but an additional unit of Casuarina woodland must be added
for the tall woodland which has grown up on Loggerhead Key since 1940.

CONCLUS ION

It will be apparent from this paper that the uncertainty over past
estimates of dimension and area of the Dry Tortugas keys is too great
for acceptable estimates to be derived from the surveys made in 1904
angio T 516). This results from a combination of surveying error,
printer's error, and natural change in the islands themselves. Only
Davis's surveys in 1937-38 of Bush, Garden, Loggerhead and Long Keys
meet required standards of reliability. Since there is also much
evidence of considerable topographic changes in the islands themselves,
it is not acceptable in analyses of area-diversity relationships to
standardise island area on the basis of any one survey, however
reliable it may be.

Second, we do not believe that the plant species records from the
earlier surveys can be made to bear inferences of colonisation or
extinction between the surveys, except for those islands such as Bird,
Middle and Sand Keys which underwent catastrophic destruction during
cyclones and were then recolonised. On the larger islands inadvertent
non-recording is at least as plausible an explanation as colonisation
or extinction. Indeed, apart from the cyclone-damaged islands, where
floristic change resulted from massive substrate disturbance, we are
impressed by the stability of the native floras over time. The fact
that on all of the islands the total number of native species recorded
has increased between 1904 and 1977 we interpret as simply the result
of collecting intensity rather than any true increase in the number of
native species present.

This is not to deny that processes of the kind envisaged by
MacArthur and Wilson (1968) may operate on small reef islands, but
rather to say that the data from the Dry Tortugas are not of sufficient
quality to support their theoretical speculations. Furthermore, we
feel it highly likely that more species remain to be recorded on the
keys, and that such new records should not themselves be inferred to be
new arrivals.

The data considered in this paper are summarised in tabular form
in Table 17 (numbers of all species, of native species, of introduced
species, and of species of uncertain status, recorded for each island
at the time of each survey) and in Table 18 (the area, total number of
species, and number of native species for each island at each survey).
Table 19 revises MacArthur and Wilson's (1968) Table 5, which sets out
data on presumed extinctions between 1904 and 1916. Finally, Figure 11
uses the 1977 island areas and the total recorded number of species for
each island (other than Garden and Loggerhead Keys) to give a visual
impression of the relationship between species number and island area
both for the Dry Tortugas and for other Gulf of Mexico and Caribbean

3}

groups of reef islands. In addition to a rather diffuse area control,
Figure 11 also shows that wet islands close to continental areas (such
as Glover's Reef) have higher species numbers than more remote and
drier islands such as the Pedro and Morant Cays, with the Dry Tortugas
in an intermediate position. These relationships have been analysed
further by Stoddart and Fosberg (in press).

The problems examined here are not, of course, unique to the Dry
Tortugas. It is worth recalling that Millspaugh (1899) was also
responsible for a series of early maps and species records for the
islands of Alacran Reef, Gulf of Mexico. These islands were remapped
by Folk (1967) in 1960, and the flora restudied by Bonet and Rzedowski
ClIG2) Mandmerosbexrg §(1962)) any 961i: Table 20 summarises the Alacran
data from these surveys. Island areas derived from Folk's maps vary
from 1.4 to 6 times the areas measured in 1899, the average increase
being about four times; Folk (1967, 415, 343) used these apparent
increases in a detailed commentary on stages of island evolution.
Similarly Fosberg (1962, 4) commented on the 'profound changes' which
had occurred since Millspaugh's visit. In view of the ambiguities
demonstrated in the Dry Tortugas data, it would perhaps be wise to be
sceptical of the Alacran — and indeed other early — records which
purport to indicate massive changes over short periods of time.

The temptation to use such data in support of general theory is
considerable, but we need to remember that such theory is no better
than the data by which it is supported.

ACKNOWLEDGEMENTS

We thank Dr W.B. Robertson, Jr., and Dr Gary E. Davis for making
our visits to the Dry Tortugas possible in 1962 and 1977.

14

PLANTS RECORDED FROM THE DRY TORTUGAS KEYS

This list derives mainly from the published records of Millspaugh
(1907), Bowman (1918) and Davis (1942), together with determinations
by F.R. Fosberg of collections made by the authors in 1962 and 1977.
Species marked N are judged to be native; those marked I to be
introduced; and those marked ? are of uncertain status.

PANDANACEAE

I Pandanus tectorius Park.
Garden Key: Fosberg sight record in 1962.

ZOSTERACEAE

N Halodule wrightii Aschers. (sensu lato)
Garden Key: Fosberg 43018 (US).
Bird Key: W.R. Taylor, 7 June 1926 (MICH) (den Hartog 1970, as
H. beaudettei (den Hartog) den Hartog).
Long Key: W.R. Taylor, 25 June 1926 (MICH) (den Hartog 1970, as
H. beaudettei (den Hartog) den Hartog).

N Syringodium filiforme Ktitz.
Garden Key: Fosberg 43013 (US).
Loggerhead Key: W.R. Taylor, 19 June 1926 (MICH) (den Hartog
1970)":

HYDROCHARITACEAE

N Halophila decipiens Ostenfeld
Dredged, 15 fathoms, H.H.M. Bowman, July 1915 (MICH) ;
10 fathoms, Tandy 1111 (BM), Tandy 1254 (BM); 18-20 fathoms,
Tandy 1570 (BM) (all citations from den Hartog 1970).

N Halophila engelmanni Aschers.
Dredged, 18-20 fathoms, Tandy 1569 (BM), Tandy 1599 (BM) (den
Hartog 1970).

N Thalassia testudinum Banks ex K6nig
Bird Key: Tandy 1068 (BM) (den Hartog 1970).
‘Long Bush Key': Tandy 1264 (BM) (den Hartog 1970).
GRAMINEAE

I Brachiaria sp.
Garden Key: Fosberg 43021 (US).

15
? Cenchrus echinatus L.
Garden Key: Millspaugh 1907; Davis 1942; Fosberg 43059 (US).
Loggerhead Key: Bowman i918; Stoddart 8056 (US).

? Cenchrus myosuroides H.B.K.
Loggerhead Key: Fosberg 43041 (US).

N Cenchrus incertus M.A. Curtis

Bird Key: Millspaugh 1907 (as C. tribuloides); Bowman 1918.
Bush Key: Bowman 1918.
East Key: Millspaugh 1907 (as C. tribuloides); Bowman 1918;

Davis 1942; Stoddart 8033 (US).

Garden Key: Millspaugh 1907 (as C. tribuloides); Fosberg 42983,
42990, 43060 (US).

Loggerhead Key: Bowman 1918; Fosberg 43045 (US).

I Cynodon dactylon (L.) Pers.
Garden Key: Bowman 1918 (as Capriola dactylon); Davis 1942;
Fosberg 42996 (US).

I Digitaria horizontalis Willd.
Garden Key: Fosberg 42981 (US).

? Digitaria sanguinalis (L.) Scop.
Garden Key: Millspaugh 1907 (as Syntherisma fimbriatum) ;
Bowman 1918 (as Syntherisma marginatum); Davis 1942;
Fosberg 42985 (US).

HE GATLOSELS Cll varns (sie) ake ie
Garden Key: Fosberg 43017 (US).

? Eragrostis prolifera (Sw.) Steud.
Garden Key: Fosberg 43019 (US); Stoddart 8065 (US).
Loggerhead Key: Stoddart 8052 (US).

I Eragrostis tenella (L.) Beauv. ex. R. & S.
Garden Key: Davis 1942.
Loggerhead Key: Fosberg 43032 (US).

N Eustachys petraea (Sw.) Desv.
Bush Key: Davis 1942 (as Chloris).
Garden Key: Millspaugh 1907; Davis 1942 (as Chloris);
Fosberg 43056 (US).
Loggerhead Key: Fosberg 43051 (US); Stoddart 8058 (US).

I Panicum maximum Jacq. }
Loggerhead Key: Fosberg 43047 (US).

2? Paspalum caespitosum Fleugge
Garden Key: Bowman 1918; Davis 1942.
Loggerhead Key: Bowman 1918.

16

N Paspalum distichum L.
Bird Key: Millspaugh 1907.
Garden Key: Millspaugh 1907.

? Sporobolus domingensis (Trin.) Kunth
Garden Key: Davis 1942.

? Sporobolus purpurascens (Sw.) Hamilt.
Garden Key: Millspaugh 1907.

N Sporobolus virginicus (L.) Kunth
Bush Key: Davis 1942; Fosberg 43000 (US); Stoddart 8004 (US).
Garden Key: Bowman 1918; Davis 1942.
Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Fosberg 43029 (US).
Long Key: Stoddart 8029 (US).

I Stenotaphrum secundatum (Walt.) O. Ktze.
Garden Key: Fosberg sight record in 1962.

N Uniola paniculata L.

Bird Key: Millspaugh 1907; Bowman 1918.

Bush Key: Bowman 1918; Davis 1942; Fosberg sight record in
1962;; Stoddart 8001 (US):.

East Key: Millspaugh 1907; Bowman 1918; Stoddart 8031 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942.

Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Fosberg 43038 (US); Stoddart 8048 (US).

Sand Key: Millspaugh 1907; Davis 1942.

CYPERACEAE

N Cyperus planifolius L.C. Rich.

Bird Key: Millspaugh 1907 (as C. brunneus).

Bush Key: Fosberg 42999 (US); Stoddart 8018 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942 (as
C. brunneus); Fosberg 42982, 43054, 43055 (US).

Loggerhead Key: Bowman 1918 (as C. brunneus); Fosberg 43033
(US); Stoddart, 8053 .(US).

Long Key: Bowman 1918 (as C. brunneus).

PALMAE

I Cocos nucifera L.
Bird Key: Bowman 1918.
Garden Key: Fosberg sight record in 1962; Stoddart sight
mecord iam 1977.
Loggerhead Key: Bowman 1918; Davis 1942; Stoddart sight
Gecord in 19777

I Phoenix canariensis Hort ex Chabaud.
Garden Key: Bowman 1918; Davis 1942.

187,

I Phoenix dactylifera L.
Garden Key: Bowman 1918; Davis 1942; Fosberg sight record
ag ih «

i -Priéchardia Spi
Garden Key: Fosberg sight record in 1962.

I Washingtonia filifera (Lindl.) Wendl.
Garden Key: Fosberg sight record in 1962.

COMMELINACEAE

I Rhoeo spathacea (Sw.) Stearn
Garden Key: Fosberg sight record in 1962.

LILIACEAE

I Agave decipiens Baker
Garden Key: Bowman 1918.

I Agave americana L.
Garden Key: Davis 1942,
Loggerhead Key: Davis 1942.

I Agave rigida Mill.
Loggerhead Key: Bowman 1918.

I Agave sp.
Loggerhead Key: Fosberg sight record in 1962; Stoddart sight
record in 1977.

I Aloe barbadensis Mill.
Garden Key: Fosberg sight record in 1962.
Loggerhead Key: Bowman 1918; Davis 1942 (both as Aloe).

I Asparagus sprengeri Regel
Loggerhead Key: Bowman 1918.
Without locality: Davis 1942 Table 7.

? Hymenocallis latifolia (Mill.) Roem.

Bird Key: Bowman 1918 (as H. caymanensis Herb.).

Garden Key: Millspaugh 1907 (as H. caribaea (L.) Herb.) ;
Bowman 1918 (as H. caymanensis Herb.); Davis 1942 (as
H. caymanensis Herb.).

Loggerhead Key: Millspaugh 1907 (as H. caribaea (L.) Herb.);
Bowman 1918 (as H. caymanensis Herb.); Davis 1942 (as H.
caymanensis Herb.); Fosberg 43050 (US); Stoddart sight
satevoroy tel aljel SL S)77/

I Hymenocallis littoralis (Jacq.) Salisb.
Garden Key: Fosberg sight record in 1962.

18

Yucca aloifolia L.
Loggerhead Key: Bowman 1918; Davis 1942.

MUSACEAE

Musa sapientum L.
Garden Key: Fosberg sight record in 1962.

CASUARINACEAE
Casuarina equisetifolia L.
Bush Key: Stoddart 8008 (US).
Garden Key: Davis 1942.
Loggerhead Key: Bowman 1918; Davis 1942;
Long Key: Fosberg 43005 (US).
MORACEAE

FIeus carica TG.
Without locality: Davis 1942 Table 7.

Ficus hispida L.f.
Loggerhead Key: Bowman 1918; Davis 1942.

Ficus palmata Forsk.?
Loggerhead Key: Fosberg 43048 (US).

POLYGONACEAE

Coccoloba uvifera L.

Stoddart 8050

(US).

Garden Key: Bowman 1918; Davis 1942; Fosberg sight record in

1962; Stoddart 8062 (US).

Loggerhead Key: Bowman 1918; Stoddart 8041

BATIDACEAE

Batis maritima L.
Bush Key: Stoddart sight record in 1977.
Long Key: Davis 1942.

(US).

19

CHENOPODIACEAE

Atriplex pentandra (Jacq.) Standl.

Bush Key: Bowman 1918 (as A. cristata); Stoddart 8019 (US).
Garden Key: Millspaugh 1907; Bowman 1918 (both as

A. cristata); Stoddart 8081? (US).
Long Key: Bowman 1918 (as A. cristata); Fosberg 43004 (US);

Stoddart scO Zia (US)i-
Without locality: Davis 1942 Table 7 (as A. cristata).

Salicornia bigelovii Torr.
Long Key: Davis 1942; Fosberg 43007 (US); Stoddart 8026 (US).

AMARANTHACEAE

Alternanthera maritima J. St. Hil.
Bush Key: Bowman 1918; Davis 1942.
Garden Key: Bowman 1918.
Long Key: Bowman 1918.

Alternanthera philoxeroides (Mart.) Griseb.
Bush Key: Stoddart 8009 (US).

Amaranthus viridis L.
Garden Key: Millspaugh 1907.

Amaranthus sp.
Without locality: Davis 1942 Table 7.

? Philoxerus vermicularis (L.) Beauv.

Bush Key: Fosberg 42975 (US).

Garden Key: Millspaugh 1907 (as Lithophila vermicularis).

Without locality: Davis 1942 Table 7 (as Lithophila
vermicularis).

NYCTAGINACEAE

Boerhavia coccinea Mill.
Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942
(all as B. viscosa); Fosberg 43061 (US).
Loggerhead Key: Bowman 1918; Davis 1942 (both as B. viscosa);
Stoddart 8055? (US); Fosberg 43035 (US).
Without locality: Davis 1942 Table 7.

Bougainvillea glabra Choisy
Garden Key: Fosberg sight record in 1962.

20

PORTULACACEAE

N Portulaca oleracea L.
Bird Key: Millspaugh 1907; Bowman 1918.
Garden Key: Millspaugh 1907; Bowman 1918; Fosberg 43008,
43009 (US); Stoddart 8073 (US).
Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942.

AIZOACEAE

N Sesuvium portulacastrum (L.) L.

Bird Key: Millspaugh 1907; Bowman 1918.

Bush Key: Bowman 1918; Davis 1942; Fosberg sight record in
1962; Stoddart 8011 (US).

East Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Stoddart 8032 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942.

Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Fosberg sight record in 1962.

Long Key: Bowman 1918; Davis 1942; Fosberg sight record in
1962; Stoddart 8029 (US).

Sand Key: Millspaugh 1907.

PAPAVERACEAE

I Argemone mexicana f. leiocarpa (Greene) G.B. Ownbey
Garden Key: Millspaugh 1907 (as Argemone leiocarpa).

N Cakile lanceolata (Willd.) O.E. Schulz

Bird Key: Millspaugh 1907 (as C. fusiformis); Bowman 1918.

Bush Key: Bowman 1918; Stoddart 8002 (US).

East Key: Millspaugh 1907 (as C. fusiformis); Bowman 1918;
Stoddart 8035° (US)

Garden Key: Millspaugh 1907 (as C. fusiformis); Bowman 1918;
Stoddart 8080 (US).

Loggerhead Key: Millspaugh 1907 (as C. fusiformis); Bowman 1918.

Middle Key: Bowman 1918; Davis 1942.

Sand Key: Bowman 1918.

CRUCIFERAE

I Lepidium virginicum L.
Garden Key: Bowman 1918; Fosberg 43025 (US).

LEGUMINOSAE

I Abrus precatorius L.
Garden Key: Fosberg sight record in 1962.

1 BL

Caesalpinia bonduc (L.) Roxb.
Garden Key: Bowman 1918 (as Guilandina crista).
Loggerhead Key: Millspaugh 1907 (as Caesalpinia crista) ;
Stoddart 8059 (US).

Caesalpinia pulcherrima (Ging ))s SK
Garden Key: Fosberg sight record in 1962.

? Caesalpinia sp.

Loggerhead Key: Fosberg 43042 (US).

Cajanus cajan (L.) Millsp.
Garden Key: Bowman 1918 (as Cajan cajan).
Without locality: Davis 1942 Table 7.

Canavalia rosea (Sw.) DC.

Garden Key: Millspaugh 1907 (as C. obtusifolia); Bowman 1918;
Davis 1942 (both as C. lineata); Fosberg 43057 (US);
Stoddart 8063 (US).

Loggerhead Key: Millspaugh 1907 (as C. obtusifolia); Bowman
1918; Davis 1942 (both as C. lineata); Fosberg sight
record in 1942.

Ceratonia siliqua L.
Garden Key: Fosberg 43064 (US, POM).

Crotalaria pallida Ait.
Garden Key: Stoddart 8078 (US).

Delonix regia (Boij.) Raf.
Garden Key: Fosberg sight record in 1962; Stoddart 8070 (US).

Desmanthus depressus H. & B.
Garden Key: Fosberg 43011 (US).

Desmodium incanum (Gmel.) Schinz
Garden Key: Fosberg 43067 (US).

Rhynchosia minimum (L.) DC.
Without locality: Davis 1942 Table 7 (as Dolicholus minimus) .

Rhynchosia parviflorus DC.
Garden Key: Bowman 1918 (as Dolicholus parviflorus) .

Sesbania sp.
Garden Key: Millspaugh 1907 (as S. sericea); Bowman 1918 (as
Glottidium vescarium); Davis 1942 (as S. macrocarpa).

Tamarindus indica L.
Garden Key: Bowman 1918; Davis 1942; Fosberg sight record in
1962.

22

ZYGOPHYLLACEAE

Tribulus cistoides L.
Loggerhead Key: Millspaugh 1907.

RUTACEAE

Citrus aurantiifolius (Christm.) Swingle
Garden Key: Fosberg sight record in 1962.

SURIANACEAE

Suriana maritima L.

Bird Key: Millspaugh 1907; Bowman 1918.

Bush Key: Bowman 1918; Davis 1942; Fosberg sight record in
1962; Stoddart 8020 (US).

East Key: Davis 1942; Stoddart 8037 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942;
StoddazeRsOvseWusir

Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Stoddart 8051 (US).

Long Key: Bowman 1918; Fosberg 43006 (US); Stoddart 8023
(US).

BURSERACEAE

Bursera simaruba (L.) Sarg.
Garden Key: Bowman 1918; Davis 1942 (both as Elaphrium
Simaruba); Fosberg sight record in 1962.
Loggerhead Key: Fosberg 43027 (US).

EUPHORBIACEAE

Breynia disticha (J.R. & G. Forst.)
Garden Key: Fosberg sight record in 1962.

Codiaeum variegatum (L.) Blume
Garden Key: Fosberg sight record in 1962.

Euphorbia adenoptera Bert.
Garden Key: Millspaugh 1907.

Euphorbia blodgettii Engelm. ex Hitchc.
Garden Key: Fosberg 42980, 42995 (US).

Euphorbia glomerifera (Millsp.) Wheeler
Garden Key: Bowman 1918 (as Chamaesyce hypericifolia) ;
Fosberg 42994 (US), 43062 (US, POM).

23

? Euphorbia heterophylla L.
Poinsettia pinetorum Small
Euphorbia havanensis Willd.

Bush Key: Stoddart 8005 (US).

Garden Key: Millspaugh 1907 (as E. havanensis); Davis 1942
(as Poinsettia cyathophora); Fosberg 42989 (US);
Stoddart 8075 (US).

Loggerhead Key: Bowman 1918 (as Poinsettia pinetorum); Davis

1942 (as Poinsettia cyathophora); Fosberg 43046 (US);
Stoddart 8054 (US).
Some of these seem intermediate between this species and
Euphorbia cyathophora Murr.

I Euphorbia hirta L.
Garden Key: Fosberg 43015 (US).
Loggerhead Key: Fosberg 43031 (US).

N Euphorbia mesembrianthemifolia Jacq.
Chamaesyca buxifolia in Bowman 1918.
Euphorbia buxifolia in Millspaugh 1907, Davis 1942.
Bird Key: Millspaugh 1907; Bowman 1918.
Bush Key: Bowman 1918; Davis 1942; Fosberg 42976 (US);
Stoddart 012) (US)r
East Key: Millspaugh 1907; Bowman 1918; Stoddart 8034 (US).
Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942.
Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Fosberg sight record in 1962; Stoddart 8045 (US).
Long Key: Bowman 1918; Fosberg sight record in 1962;
Stoddart 8022 (US).
Middle Key: Davis 1942.
Sand Key: Millspaugh 1907; Davis 1942.

I Euphorbia prostrata Ait.
Garden Key: Fosberg 42991 (US).
Loggerhead Key: Fosberg 43030 (US).

I Pedilanthus tithymaloides (L.) Poit.
Garden Key: Fosberg sight record in 1962.
Loggerhead Key: Bowman 1918.
Without locality: Davis 1942 Table 7 (as Tithymalus smallii
(Millsp.) Small).

I Phyllanthus amarus Schum. & Thonn.
Garden Key: Fosberg 43014 (US).

I Ricinus communis L.
Garden Key: Bowman 1918.

24
MALVACEAE
I Hibiscus rosa-sinensis L.
Garden Key: Fosberg sight record in 1962.

Loggerhead Key: Bowman 1918.

I Malvaviscus arborea Cav.
Garden Key: Fosberg sight record in 1962.

I Sida acuta Burm. f.
Garden Key: Millspaugh 1907; Bowman 1918 (both as Sida
carpinifolia); Fosberg 43016 (US).

I Sida procumbens Sw.

Garden Key: Millspaugh 1907 (as Sida diffusa); Bowman 1918;

Fosberg 43012 (US).

I Thespesia populnea (L.) Solander ex Correa
Garden Key: Bowman 1918; Davis 1942; Fosberg sight record
in 1962; Stoddart 8068 (US).

Loggerhead Key: Bowman 1918; Davis 1942; Stoddart 8042 (US).

STERCULIACEAE

I Waltheria indica L.
Garden Key: Fosberg 43068 (US).

CARICACEAE

I Carica papaya L.
Loggerhead Key: Millspaugh 1907 (as Papaya); Bowman 1918.

CACTACEAE

N Opuntia dillenii (Ker.) Haw.
Bird Key: Millspaugh 1907; Bowman 1918.
Bush Key: Davis 1942; Fosberg sight record in 1962;
Stoddart sight record in 1977.
Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942.
Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Stoddart sight record in 1977.

LYTHRACEAE

I Lawsonia inermis L.
Garden Key: Fosberg 43065 (US).

i

8

i

25
COMBRETACEAE

Conocarpus erectus L.
Bush Key: Davis 1940, 1942; Fosberg sight record in 1962;
Stoddart 8015 (US).
Garden Key: Bowman 1918; Davis 1942; Fosberg 42992, 42993
(US); (Steddart 8066, 80675 (US)

Laguncularia racemosa (L.) Gaertn. f.
Bush Key: Davis 1940, 1942 (as Laguncularia); Fosberg sight
wecond, in 1962) {Stoddares00m, S06. (US).
Long Key: Fosberg 43001 (US).

Terminalia catappa L.
Garden Key: Bowman 1918 (as Terminalia); Fosberg sight record
gy OOD.

MYRTACEAE

Psidium guajava L.
Garden Key: Bowman 1918; Davis 1942; Fosberg 43069 (US).

RHIZOPHORACEAE

Rhizophora mangle L.

Bush Key: Bowman 1918; Davis 1940, 1942; Fosberg 42977 (US);
Stoddart 8014 (US).

East Key: Davis 1940 (drift seedlings).

Garden Key: Bowman 1918; Davis 1940 (drift seedlings).

Loggerhead Key: Davis 1940 (drift seedlings).

Long Key: Bowman 1918; Davis 1940, 1942; Fosberg 43002 (US);
Stoddart 8027 (US).

Sand Key: Davis 1940 (drift seedlings).

ARALIACEAE

Polyscias guilfoylei (Cogn. & March.) Bailey
Garden Key: Fosberg sight record in 1962.

GENT ITANACEAE

Eustoma exaltatum (L.) G. Don
Garden Key: Fosberg 42979 (US).

APOCYNACEAE

Catharanthus roseus (L.) G. Don
Garden Key: Fosberg sight record in 1962.
Loggerhead Key: Bowman 1918 (as Vinca roseus); Fosberg
sight record ain 1962.

26

N

Nerium oleander L.
Garden Key: Bowman 1918; Davis 1942 (both as Oleander) ;
Fosberg sight record in 1962.
Loggerhead Key: Bowman 1918.

Ochrosia elliptica Labill.
Garden Key: Stoddart 8064 (US, POM).

Thevetia peruviana (Pers.) Schum.
Loggerhead Key: Bowman 1918 (as Cerbera thevetia).

ASCLEPIADACEAE

Cryptostegia grandiflora R. Br.
Garden Key: Fosberg sight record in 1962.

CONVOLVULACEAE

Ipomoea macrantha R. & S.

Bush Key: Stoddart 8017 (US).

Garden Key: Bowman 1918 (as Calonyction tuba) ;
Stoddart 8079 (US).

Loggerhead Key: Millspaugh 1907 (as Calonyction album) ;
Bowman 1918; Davis 1942 (both as Calonyction tuba) ;
Fosberg 43052 (US).

Long Key: Stoddart 8025 (US).

Ipomoea pes-caprae (L.) R. Br. subsp. brasiliensis (L.) v. Ooststr.

Bush Key: Bowman 1918; Davis 1942.

East Key: Davis 1942; Stoddart 8038 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Stoddart 8074 (US).

Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;

Fosberg 43044 (US); Stoddart 8043 (US).
Long Key: Fosberg 43003 (US).
Sand Key: Millspaugh 1907.

BORAGINACEAE
Cordia sebestena L.

Garden Key: Bowman 1918; Fosberg sight record in 1962;
Stoddart 8069 (US).

Loggerhead Key: Millspaugh 1907; Bowman 1918 (as Sebesten
sebestina); Fosberg sight record in 1962; Stoddart 8044

(US).

Heliotropium curassavicum L.
Garden Key: Millspaugh 1907; Fosberg 43066 (US).
Without locality: Davis 1942 Table 7.

Heliotropium angiospermum Murray
Garden Key: Fosberg 43024 (US).

N

27

Tournefortia gnaphalodes (L.) Kunth

Bird Key: Millspaugh 1907; Bowman 1918.

Bush Key: Bowman 1918; Davis 1942; Stoddart 8003 (US).

East Key: Millspaugh 1907; Bowman 1918; Stoddart 8040 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Stoddart 8066 (US).

Loggerhead Key: Millspaugh 1907; Bowman 1918; Davis 1942;
Fosberg 43043 (US); Stoddart 8049 (US).

Long Key: Bowman 1918; Stoddart 8024 (US).

Sand Key: Davis 1942.

VERBENACEAE

Avicennia germinans (L.) L.
Bush Key: Davis 1940; Fosberg 42998 (US); Stoddart 8013 (US).

Garden Key: Bowman 1918 (as A. nitida); Davis 1940, 1942;
Fosberg 43058 (US).
Long Key: Davis 1942 (as A. nitida); Fosberg sight record in

1962;7 “Stoddanrers0so0n (Us)

Lippia nodiflora (L.) Michx.
Garden Key: Bowman 1918 (as Phyla nodiflora); Fosberg 43010,
43020 (US).
Without locality: Davis 1942 Table 7.

Stachytarpheta jamaicensis (L.) Vahl
Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942 (all as
Valerianoides jamaicense); Fosberg 42997 (US);
Stoddart 8077) (US) =
Loggerhead Key: Stoddart 8046 (US).

LABIATAE
Salvia serotina L.
Loggerhead Key: Millspaugh 1907; Bowman 1918; Fosberg 43026
(US).
Without locality: Davis 1942 Table 7.

SOLANACEAE

? Solanum bahamense L.

Bush Key: Fosberg 42974 (US).
Loggerhead Key: Fosberg 43034 (US).

SCROPHULARIACEAE

Capraria biflora L.
Garden Key: Millspaugh 1907 (as C. saxifragifolia) ;
Fosberg 43023 (US).
Loggerhead Key: Millspaugh 1907; Bowman 1918; Fosberg 43028
(Us).
Without locality: Davis 1942 Table 7.

28
GOODEN IACEAE

N Scaevola plumieri (L.) Vahl
Bird Key: Millspaugh 1907; Bowman 1918.
Bush Key: Bowman 1918; Fosberg 42978 (US); Stoddart 8006 (US).
East Key: Millspaugh 1907; Bowman 1918; Stoddart 8039 (US).
Garden Key: Bowman 1918; Stoddart 8072 (US).
Loggerhead Key: Millspaugh 1907; Bowman 1918; Fosberg 43053

(US) i Stoddart SO4vaWus)..

Sand Key: Bowman 1918.

COMPOSITAE

2? Ambrosia hispida Pursh
Loggerhead Key: Fosberg 43039 (US).

I, Bidens albaca(l.). DC.
Garden Key: Millspaugh 1907; Bowman 1918 (both as

B. leucantha); Fosberg 42987 (US).
Loggerhead Key: Davis 1942 (as B. leucantha); Stoddart 8060
(USE

I Conyza canadensis (L.) Cronquist
Garden Key: Bowman 1918; Davis 1942 (both as Leptilon
canadense); Fosberg 42984 (US).
Loggerhead Key: Fosberg sight record in 1962.

I Gaillardia pulchella Foug.?
Garden Key: Fosberg 43022 (US).

N Iva imbricata Walt.

Bird Key: Bowman 1918.

Bush Key: Bowman 1918.

East Key: Millspaugh 1907; Bowman 1918; Stoddart 8036 (US).

Garden Key: Millspaugh 1907; Bowman 1918; Davis 1942.

Loggerhead Key: Millspaugh 1907; Fosberg 43040 (US);
Stoddart. 80525 (Us)

Middle Key: Davis 1942.

Sand Key: Millspaugh 1907; Bowman 1918.

N Melanthera aspera var. glabriuscula (O. Ktze.) Parks.

Bush Key: Stoddart 8010 (US).

Garden Key: Millspaugh 1907 (as M. nivea); Bowman 1918;
Davis 1942 (both as M. brevifolia); Fosberg 42986 (US);
Stoddart 8076 .(US).

Loggerhead Key: Millspaugh 1907 (as M. nivea); Bowman 1918 (as
M. brevifolia); Fosberg sight record in 1962;
Stoddart, 8061. (US).

I Sonchus oleraceus L.
Garden Key: Millspaugh 1907; Bowman 1918; Fosberg 42988 (US).
Without locality: Davis 1942 Table 7.

29

REFERENCES

Agassiz, A. 1888a. The Tortugas and Florida Reefs. Mem. Am. Acad.
MAES Coslg IWoblileys ihe MOGs sie

Agassiz, A. 1888b. The Florida Reefs. ElneioneSre Bp joo -DANA” alive
Three cruises of the United States Coast and Geodetic Steamer
'Blake' in the Gulf of Mexico, in the Caribbean Sea, and along
the Atlantic coast of the United States, from 1877 to 1880.
London: Sampson Low, Marston, Searle and Rivington. Soil 5 | Saale
DAO) jojoe

Asprey, G.F. and Robbins, R.G. 1953. The vegetation of Jamaica.
EeEol. Monogr. 23: 359-412.

Audubon, J.J. 1835. Ornithological biography, or an account of the
birds of the United States of America. Volume 3. Edinburgh:
A. and C. Black aesval OSG mpp\.

Bartsch, be i OmoN The bird rookeries of the Tortugas. Smithsonian
instEnwAnn sy eRepeG suekOr HL Dl/aes KAO I-50 0%

Bonet, F. and Rzedowski, J. 1962. La vegetacidén de las islas del
Arrecife Alacranes, Yucatan (México). An. Esc. Nac. Cienc.
Biol. Mex. i1(1-4): 15-50.

Bowman, H.H.M. 1918. Botanical ecology of the Dry Tortugas.
Papers Dept. mar. Biol. Carnegie Instn Wash. 12: 109-138.

Brooks, H.K. 1962. Reefs and bioclastic sediments of the Dry
Tortugas (abstract). Geol. Soc. Am. Spec. Pap. 73: 1-2.

Chapman, V.J. 1944. 1939 Cambridge Expedition to Jamaica. Welseig io
A study of the botanical processes concerned in the development
of the Jamaican shoreline. JimeGDrnniemSOc. Bot. 2: 407-447.

DENS» Wdlslo 5 wie5 ) LSHO- The ecology and geologic role of mangroves
in Florida. Papers Tortugas Lab. 32: 303-412.

Davis, J.H., Jr. 1942. The ecology of the vegetation and topography
of the sand keys of Florida. REVQeInS Wosmebiees Ieiag) Se) itso S)s)-
DenwHartkog,) C2 ) 970% The sea-grasses of the world. Verh. Kon. Ned.

Akad. Wet. afd. Nat., Tweede Reeks, 59(1): 1-275.

Daler, Yow | LYV7A- Sooty tern behavior. BUTT Wer Lanse. wMUS:| 9BLOL..
Seis MSs NACE).

England, G.A. 1928. Bird Key. Saturday Evening Post, 201: 14-15,
SSI; (shshq

30

Field, R.M. 1919. Investigations regarding the calcium carbonate
oozes at Tortugas, and the beach-rock at Loggerhead Key.
Carnegie Instn Wash. Ybk. 18: 197-198.

Biecld ReM. l9Z0" Origin of the beach rock (coquina) at Loggerhead
Key, Tortugas. BULE Geole Soc. tAms V3: BH215)2

FOLK) UR. 1967. Sand cays of Alacran Reef, Yucatan, Mexico:
morphology. Jie IGEOL Tai “41 2—43)7".

Fosberg, F.R. 1962. A brief survey of the cays of Arrecife Alacran,
a Mexican atoll. Atoll Ress Bulle, 93: W—25-

Gauld, G. 1790. An accurate chart of the Tortugas and Florida Kays
or Martyrs, surveyed by George Gauld, A.M. in the years 1773, 4,
& 25. London: W. Faden. 2nd edition, 1820.

Ginsburg, sven 195s Beachrock in south Florida. J. sedim. Petrol.
23) 185—92"
Holliders; aC ati S92 Along the Florida reef. New York:

Diy Appleton Lx, 2 ONDE

Jiandrichtm Ver =LOW2 Biogenic buildups and carbonate sedimentation,
Dry Tortugas reef complex, Florida. State University of New
York at Binghamton, Ph.D. thesis. 96 pp. (University Microfilms
International No. UM 72-24065).

ashley kaos su Oto. Notes on the nesting activities of the noddy
and sooty terns. Papers Tortugas Lab. 7: 61-83.

Millspaugh, C.F. 1907. Flora of the sand keys of Florida. Field

Columbian Mus’.. Publ) AV8 (Bots Ser. 92): 191-245.
Millspaugh (Ciro 1916. Vegetation of Alacran Reef. Field Mus.
Nats AVsSe. Bobs Ser... Cli 221—430.

Multern,;e He Ger sie. Holocene cementation of skeletal grains into
beachrock, Dry Tortugas, Florida. In iO..P. Bracker;) ted:
Carbonate cements (Baltimore: Johns Hopkins Press), 25-26.

O'Neitly cs LING: Sedimentology of East Key, Dry Tortugas (abstract).
Florida Scientist, 89 (suppl 2) 110).

Robertson, W.B., Jr. 1964. The terns of the Dry Tortugas. iS{OHET I)
FlacSt. Mus. Biol S"Sei. Sits "1-94".

SCOtE, WeE.D. 19047 The story of a bird lover. New York:
Macmillan. “xi, 372-pp.

Semple; J.C. 1978. A revision of the genus Borrichia Adans.
(Compositae). Ann. Missourl bot. Gdn. 65: | 681-693'.

Shul

Sham eAun) HUGSON) mWisHy, Holley, mR ab. and slicizi,)9Bin § OW < Lopographalc
control and accumulation rate of some Holocene coral reefs: south

Florida and Dry Tortugas. Proc. 3rd Int. Symp. Coral Reefs,
DAS B-Heo
Smell p Wolk 19S, Flora of the Florida Keys. New York. xii,
162 pp.
Sprunt, A. 1948. The tern colonies of the Dry Tortugas keys. Auk,
S58 “ole

Spiguinitay: Ave | 950): A list of the birds of the Dry Tortugas keys 1857-
LEO}. jaulonguela Wesigeulicn, 238 A9-G0, 73aKs5 UO it il

Spwunite A. | 1962-63 Birds of the Dry Tortugas. Florida Naturalist,
35) (IISA) 8° SS-CO, S235, UA oNsAe) Ske (eS) 8) AIA py DADS Sic

Steers, J.A. 1940. The coral cays of Jamaica. Geogrl J. 95:
30-42.

Steers, J.A., Chapman, V.J., Colman, J., and Lofthouse, J.A. 1940.
Sand cays and mangroves in Jamaica. Geogrildin 96) yoOb=s28.5

Stevenson, J.O. 1938. The tern colonies of Dry Tortugas. Bird-
Lore, 40: 305-309.

Stoddart, D.R. and Fosberg, F.R. in press. Species-area relationships
on small islands: floristic data from Belizean sand cays.

Smithsonian Publs mar. Biol.

Taylor, W.R. 1926. The marine flora of the Dry Tortugas. Rev.
ANG Ole 293. hisses

avo naWiekye 1) LOZ: The marine algae of Florida with special

reference to the Dry Tortugas. Papers Tortugas Lab. 25:
220)

Vaughan, T.W. 1914. The building of the Marquesas and Tortugas
atolls and a sketch of the geologic history of the Florida reef
tract. Papers Tortugas Lab. 5: 55-67.

Vaughan, T.W. 1918. The temperature of the Florida coral-reef
lealCiter. Papers Dept. Marabiol. Carnegie instn Wash. 9: 319-339.

Vaughan, T.W. 1935. Current measurements along the Florida coral

reef tract. Papers Tortugas Lab. 29: 129-141.

Watson, J.B. 1908. The behavior of noddy and sooty terns. Papers
Tortugas Lab. 2: 187-255.

32

Watson, J.B. and Lashley, K.S. 1915. An historical and experimental

study of homing. Papers Dept. mar. Biol. Carnegie Instn Wash.
ieee — COR
Yates, HO. 7966. Revision of grasses traditionally referred to

Uniola, I. Uniola and Leptochlodpsis. Southwestern Naturalist,
11: 3972-394.

Zansy- Visi L9S8r The Pedro Cays and Pedro Bank: report on the
survey of the cays 1955-57. Bull. geol. Surv. Dept. Jamaica, 3:
1-47.

Table 1.

Source

DIMENSIONS
Millspaugh 1907, 233
Watson 1908, 191

Bowman 1918

AREA
Tatnall and Gednery 1829

Seowe W047 2us
Millspaugh 1907

Watson and Lashley
LDL 5 SS

Lashley 1915, 61
Pearson in 1915
Ashe and Lowe in 1918
England 1928, 14

Bowman 1918

Source:

Dimensions and area of Bird Key

Original estimate

500 x 250 ft
400 x 300 yds
500 x 300 ft

4 ac 2 roods 20
poles

Cale ac
from map

ca 6,000 sq yds

less than 5 ac
8 ac

Camomac

less than 5 ac

from map at
given scale

from map by
MacArthur and
Wilson 1968

from map, corrected
scale

38

Metric equivalent

ILS} 576) hin
366 x 274 m
52) xa Dem

8721 Ome

ca 32,400 m?
6,440 m?
Can) OM me

less than 20,200 m?
32,400) m-

ca 24,300 m?

less than 20,200 m?
47,695 m?

13) pS) vine

Q-37S int

records in Robertson (1964) and papers cited

34

Table 2. Plants recorded from Bird Key
Status 1904 sens)
Cenchrus incertus N x x
Paspalum distichum N x -
Uniola paniculata N x x
Cyperus planifolius N x -
Cocos nucifera I = xe
Hymenocallis latifolia @ = x
Portulaca oleracea N xX x
Sesuvium portulacastrum N x x
Cakile lanceolata N x x
Suriana maritima N x x
Euphorbia mesembrianthemifolia N x x
Opuntia dillenii N x x
Tournefortia gnaphalodes N xe x
Scaevola plumieri N x x
Iva imbricata N - x
Total 1904 1915

Number of species 15 2 13
Native species eS PZ 11
Introduced species 1 O 1
Uncertain status 1 O 1

N: native I: introduced ?: uncertain status

Table 3. Dimensions and area of Bush Key

Source Original estimate

DIMENSIONS

Bowman 1918 from map

Davis 1942 from map

Siwoclsleusic ali LDV from map
AREA

Tatnall and Gednery 1829 Siac 1/3) roods’ 22) polles

Davis 1942 i) AG

Sprunt 1948 110,000 sq yds

Robertson 1964, 12 2ZOVac

Stoddart in 1977 from map

Source: records in Robertson (1964) and papers cited

35

Metric equivalent

583 x 342 m | |
976 x 333m

SeOlinxs 2 Ofam

234.20 Om me
157,90 0lams
91,960 m?
80,900 m?

81,330 m?

36

Table 4. Plants recorded from Bush Key
Status
Cenchrus incertus N
Eustachys petraea N
Sporobolus virginicus N
Uniola paniculata N
Cyperus planifolius N
Casuarina equisetifolia a
Batis maritima N
Atriplex pentendra N
Alternanthera maritima N
Alternanthera philoxeroides @
Philoxerus vermicularis N?
Sesuvium portulacastrum N
Cakile lanceolata N
Suriana maritima N
Euphorbia heterophylla ?
Euphorbia mesembrianthemifolia N
Opuntia dillenii N
Conocarpus erectus N
Laguncularia racemosa N
Rhizophora mangle N
Ipomoea macrantha N
Ipomoea pes-caprae N
Tournefortia gnaphalodes N
Avicennia germinans N
Solanum bahamense N?
Scaevola plumieri N
Iva imbricata N
Melanthera aspera N
Total
Number of species 28
Native species 25
Introduced species 1
Uncertain status 2
N: Native I: introduced

TOUS

*

~ mM I

ESTES)

13
13
0
0

OST,

L957

13
13
0
0

ous

1962 LO
x
x x
x x
= x
= x
= x
= x
>3€ —
x x
= x
x x
= x
x x
x x
x x
x x
x x
= x
= x
x x
>34 —
x x
= x
1962 LOTT.
14 Pal
14 18
0 if
0 2

uncertain status

Table 5.

Source

DIMENSIONS
Agassiz 1888
Millspaugh 1907

Bowman 1918
Davis 1942
Stoddart in 1977

AREA
Tatnall and Gednery 1829
Scott 1904 (in 1890)
Millspaugh 1907

Bowman 1918

Sprunt 1948
Stoddart in 1977

Source:

Dimensions and area of East Key

Original estimate

from chart
ASO) $2 50) sete

1/3 mile long,
1/6 mile broad

LAO) >< OO see

from map

V2 ele
18 ac

from map
(dimensions in ft)

from map
(dimensions in yds)

from map at
given scale

from map estimated
by MacArthur and
Wilson 1968

from map, corrected
scale

85,000 sq yds

from map

37

Metric equivalent

500 m long
85 xaiSem
ca 540 x 250m

366 x 188 m

345 x 103 m

48,600 m?
72,800 m?
Zw] Omme

6,500 m?

11,5) 6\Olme

111,480 m?

87,220 m?

fly 07 Omens
DA Nas juke

records in Robertson (1964) and papers cited

38

Table 6. Plants recorded from East Key
Status
Cenchrus incertus N
Uniola paniculata N
Sesuvium portulacastrum N
Cakile lanceolata N
Suriana maritima N
Euphorbia mesembrianthemifolia N
Ipomoea pes-caprae N
Tournefortia gnaphalodes N
Scaevola plumieri N
Iva imbricata N
Total
Number of species 10
Native species 10
Introduced species O
Uncertain status O
N: native I: introduced

1904

1904

8

8
0
0

1915 1937 LO

TOS 1937 LOW

aus

8
8
@)
0)

4 10
| 10
0 0
0 0)

uncertain status

This tables does not include the strand records of Rhizophora mangle
resulting from Davis's (1940) drift experiments in 1937 and 1938.

39

Table 7. Dimensions and area of Garden Kay

Source Original estimate Metric equivalent
DIMENSIONS
Millspaugh 1907 from map iS) Ss SLO Tn
Vaughan 1918 from map Dae xens 9 im
Bowman 1918 from map 120 e73,60) sm
Davis 1942 from map 590 x 320 m
AREA
Tatnall and Gednery 1829 Uo. Ge 30/7350) m2
Bache in 1845 8.28 ac 35,610 m?
Millspaugh 1907 from map 92,880 m?
Vaughan 1918 from map *96,550 m?
Bowman 1918 from map 150,540 m?
Davis 1942 from map 112,420 m?

Source: records in Robertson (1964) and papers cited

*with the separate sandbar to the northwest the area is 101,200 m?

40

Table 8. Plants recorded from Garden Kay

Status 1904 1915 1937 1962 1977
Pandanus tectorius
Brachiaria sp.
Cenchrus echinatus
Cenchrus incertus

A wHH
™
|
™

Cynodon dactylon
Digitaria horizontalis
Digitaria sanguinalis
Eragrostis ciliaris

eh OO) Lah teal
|
|
!

Eragrostis prolifera
Eragrostis tenella
Eustachys petraea
Paspalum caespitosum

VAwHs
™
I
a mM OM
*
I

Paspalum distichum
Sporobolus domingensis
Sporobolus purpurascens
Sporobolus virginicus

HA) Fosyycend) 14
|
I
x
|
|

Stenotaphrum secundatum
Uniola paniculata
Cyperus planifolius
Cocos nucifera

H2A2AH
~
x x
~

l
|

!
!
|
*
*%

Phoenix canariensis
Phoenix dactylifera
Pritchardia sp.
Washingtonia filifera

HH HH
| |
I *
| *

x mM I
! !

Rhoeo spathacea
Agave decipiens
Agave americana
Aloe barbadensis

betes | tle tea|

Hymenocallis latifolia

Hymenocallis littoralis
Musa Sapientum
Casuarina equisetifolia

FAs)
I
|
|
~
|

Coccoloba uvifera ?
Atriplex pentandra N
Alternanthera maritima N -
Amaranthus viridis I

*
mx x

!

|

*%

Philoxerus vermicularis N? x - — = =
Boerhavia coccinea
Bougainvillea glabra
Portulaca oleracea

4H ev
|
|
|
x x
|

41

Table 8 cont.

Sees WSO ley MOST sXeo NST 7)

Sesuvium portulacastrum N Xx Xx xX > =
Argemone mexicana I x - - - -
Cakile lanceolata N x x = = x
Lepidium virginicum i - x - 5K -
Abrus precatorius I - - - x -
Caesalpinia bonduc g = x = = =
Caesalpinia pulcherrima iL - - - x -
Cajanus cajan if = x - - -
Canavalia rosea N x x xX x x
Ceratonia siliqua if = = = x =
Crotalaria pallida iL - = - ~ x
Delonix regia ag - = = x x
Desmanthus depressus 2 = - - X _
Desmodium incanum I - = - x -
Rhynchosia parviflorus I - x - - -
Sesbania sp. I x x X - -
Tamarindus indica I = x x x -
Citrus aurantiifolius if - - - x -
Suriana maritima N x x x - x
Bursera simaruba B = x x x -
Breynia disticha I - - - x -
Codiaeum variegatum i - - - x -
Euphorbia adenoptera @ x - - - -
Euphorbia blodgettii @ = = = x =
Euphorbia glomerifera I - x - x -
Euphorbia heterophylla ? x - x x x
Euphorbia hirta I - - - x -
Euphorbia mesembrianthemifolia N x x x - -
Euphorbia prostrata I - - - x -
Pedilanthus tithymaloides I = = - x =
Phyllanthus amarus i = = - X =
Ricinus communis ai - x = = =
Hibiscus rosa-sinensis i - - - x -
Malvaviscus arborea I = = - x -
Sida acuta if x x = x -
Sida procumbens I x x - x -
Thespesia populnea I - x x x x
Waltheria indica I = - - -
Opuntia dillenii N x x x - -
Lawsonia inermis I - - - x -

42

Table 8 cont.

Status ‘1904 41905 S937. 1962) 1977
Conocarpus erectus
Terminalia catappa
Psidium guajava
Rhizophora mangle

ZAHHY

Polyscias guilfoylei
Eustoma exaltatum
Catharanthus roseus
Nerium oleander

HAH H
!
|
|

a

Ochrosia elliptica
Cryptostegia grandiflora
Ipomoea macrantha
Ipomoea pes-caprae

Z22wHH
|
*
I
I
x

Cordia sebestena
Heliotropium curassavicum
Heliotropium angiosperumum
Tournefortia gnaphalodes

ZAwHAH
x
|
|
mm OM OM
I

Avicennia germinans

Lippia nodiflora
Stachytarpheta jamaicensis
Capraria biflora

NOH &
|

wm Mm OM
|

Scaevola plumieri N x
Bidens alba I x > -
I x
EE

|
l
*

Conyza canadensis
Gaillardia pulchella

*
~ Mm mM
|

Iva imbricata N x x x ~ -
Melanthera nivea N
Sonchus oleraceus IL x x - x -

*
*
~
*%
*

Total. 1904 1915 “19387 “962"saSMa

Number of species 107 35 5 il 36 del: Pay
Native species 26 19 21 14 2) 11
Introduced species 64 8 22 13 5 7
Uncertain status 17 8 8 S) 11 3

N: native I: introduced ?: uncertain status

Table 9.

Source:

Source

DIMENSIONS

Agassiz 1888
Millspaugh 1907
Bowman 1918
Davis 1942
Stoddart in 1977

AREA

Tatnall and Gednery 1829
Millspaugh 1907

Bowman 1918

Davis 1942
Robertson 1964, 14
Stoddart in 1977

Dimensions and area of Loggerhead Key

Original estimate

from chart

3/4 x 1/8 mile
from map

4500 x 600 ft

from map

ca 30 ac
from map

from map at
given scale

from map estimated
by MacArthur and
Wilson 1968

from map, corrected
scale

from map
ca 30 ac

from map

records in Robertson (1964) and papers cited

43

Metric equivalent

1140 x 190
1207x7200 om
GSS ro) Ssh in)
1380 x 208 m
1280x243) m

B

1217400) m2
242,, 300m:
473,840 m?

19577090)pme

278,665 m?

209,920 m?
Ca e214. 00) im?
22 i, hOOmme

44

Table 10. Plants recorded from Loggerhead Key

Statuses 1904" tolsy hose TI62e muSra

Cenchrus echinatus ? = x - - x
Cenchrus myosoroides g = = = x -
Cenchrus incertus N - x = OK =
Eragrostis prolifera 2 = = = - x
Eragrostis tenella I - = = x =
Eustachys petraea N = = = x x
Panicum maximum I = = - x =
Paspalum caespitosum @ = x = - -
Sporobolus virginicus N x x x x =
Uniola paniculata N x x x x x
Cyperus planifolius N - x ~ x x
Cocos nucifera aL ~ x x - x
Agave americana I = - x - -
Agave rigida I = x - - -
Aloe barbadensis I = x x - -
Asparagus sprengeri it = K = = -
Hymenocallis latifolia a x x x x x
Yucca aloifolia I = x x - -
Casuarina equisetifolia I - x x - x
Ficus hispida a = x x ~ -
Ficus palmata i = = = x -
Coccoloba uvifera B = x - - x
Boerhavia coccinea ? = x Ds x x
Portulaca oleracea N x x x - -
Sesuvium portulacastrum N x x x > =
Cakile lanceolata N x x - - -
Caesalpinia bonduc 2 x = - x x
Canavalia rosea N xX x x x -
Tribulus cistoides G x - - - -
Suriana maritima N x br x - x
Bursera simaruba ? - - - x -
Euphorbia heterophylla 2 - x x x x
Euphorbia hirta i = - - x -
Euphorbia mesembrianthemifolia N x x x x Be
Euphorbia prostrata I = = = x =
Pedilanthus tithymaloides at = x = = =

Table 10 cont.

Hibiscus rosa-sinensis
Thespesia populnea
Carica papaya

Opuntia dillenii

Catharanthus roseus
Nerium oleander
Thevetia peruviana
Ipomoea macrantha

Ipomoea pes-caprae

Cordia sebestena
Tournefortia gnaphalodes
Stachytarpheta jamaicensis

Salvia serotina
Solanum bahamense
Capraria biflora
Scaevola plumieri

Ambrosia hispida
Bidens alba
Conyza canadensis
Iva imbricata

Melanthera aspera

Number of species
Native species
Introduced species

Uncertain status

N: native

IL 3

Status

ZwWHH sy

Total
57

introduced

1904

x mM

1904

SS

TOUS

45

LOST MYS2 syd

x = x
x = x
_— xX —
x x =
x x x
= x x
x x x
- - x
— >< —
_ 5S _
— xX —
= x x
_ x _—
x = x
— x —
= x x
= x x

LOST Uo MOY)

uncertain status

This table does not include the strand records of Rhizophora mangle
resulting from Davis's (1940) drift experiments in 1937 and 1938.

46

Table 11. Dimensions and area of Long Key
Source Original estimate Metric equivalent

DIMENSIONS

Davis 1942 from map 5 2exale Oem

Stoddart in 1977 from map 224 x 73m
AREA

Davis 1942 from map 47,300 m?

Stoddart in 1977 from map 8,120 m?

Source: papers cited

Table 12. Plants recorded from Long Key

Sporobolus virginicus
Cyperus planifolius
Casuarina equisetifolia
Batis maritima

Atriplex pentandra
Salicornia bigelovii
Alternanthera maritima
Sesuvium portulacastrum

Suriana maritima

Euphorbia mesembrianthemifolia
Laguncularia racemosa
Rhizophora mangle

Ipomoea macrantha

Ipomoea pes-caprae

Tournefortia gnaphalodes
Avicennia germinans

Number of species
Native species
Introduced species

Uncertain status

N: native T:

Status 1915

Sy

47

N a
N Xx
I =
N =
N x
N =
N x
N x
N x
N x
N =
N Xx
N =
N =
N x
N =

Total 1915

16 8

iS) 8

1 O

0) 0)
introduced

HOB

5)

LIGZ OT,
= x

x _—
x x

x x

x x

x x

x x

x —
x 8

x
>< —
= x

x x

LIGZ LO,
10 10
9 10
1 @)
0) 0

uncertain status

48

Table 13. Plants recorded from Middle Key

Status OWES L987
Cakile lanceolata N x x
Euphorbia mesembrianthemifolia N = x
Iva imbricata N = 2g

Total 1945 1937
Number of species 3 i 3
Native species 3 1 3
Introduced species O @) @)
Uncertain status @) O O

N: native I: introduced ?: uncertain status

Table 14. Dimensions and area of Sand Key

Source
DIMENSIONS

Agassiz 1888
Millspaugh 1907

Bowman 1918
Davis 1942

Stoddart in 1977

AREA
Millspaugh 1907

Bowman 1918

Stoddart in 1977

Source: papers cited

Original estimate

from chart
SOR se Ol hit
from) Chast
TWO x 7/05 Ee

from map

from map

from map at
given scale

from map estimated
by MacArthur and
Wilson 1968

from map, corrected
scale

from map

49

Metric equivalent

160 m long
DAS x voy
2x
34 x 21 Mm

113 x 425m

228m

417733 m*

3,344 m?

DAD vie

B)75 2) m2

50

Table 15. Plants recorded from Sand Key

Status 1904 1915 i937

Uniola paniculata N x = x
Sesuvium portulacastrum N x - -
Cakile lanceolata N = x -
Euphorbia mesembrianthemifolia N x = x
Ipomoea pes-caprae N x = =
Tournefortia gnaphalodes N = = x
Scaevola plumieri N = x -
Iva imbricata N x x -

Total 1904 1915 193i

Number of species 8 5 3 3
Native species 8 5 3 3
Introduced species O O O 6)
Uncertain status O O O 6)
N: native I: introduced ?: uncertain status

This tables does not include the strand records of Rhizophora mangle
resulting from Davis's (1940) drift experiments in 1937 and 1938.

Syl

Table 16. Distribution of native plants (excluding sea-grasses) on Dry
Tortugas keys

Bird Bush East Garden Loggerhead Long Middle Sand

Cenchrus incertus x fy x x x x = = =
Eustachys petraea as x = x x = = =
Paspalum distichum x = = x = = = =
Sporobolus virginicus —- x = x x x - -
Uniola paniculata x OF 5c ore x - - Sx
Cyperus planifolius x x - x x x - -
Batis maritima = xX = = = x - -
Atriplex pentandra = x = x - x - -
Salicornia bigelovii = = = = = x - -
Alternanthera maritima — x ca x = x = =
Philoxerus ry x a x = = = =
vermicularis
Portulaca oleracea x = = x x = = =
Sesuvium x me Si x x x - x
portulacastrum
Cakile lanceolata x x x x x - x x
Canavalia rosea = = - x x - - -
Suriana maritima x x x x x x - -
Euphorbia x x x x x x x x
mesembrianthemifolia
Opuntia dillenii x 5 - x x - - -
Conocarpus erectus - x = x = - - -
Laguncularia racemosa - x = = = x - -
Rhizophora mangle = x = x = x - -
Ipomoea macrantha = x - x x x = =
Ipomoea pes-caprae = x x x x x = x
Heliotropium - - = aX - - ~ -
curassavicum
Tournefortia x x x 3K x x - x
gnaphalodes
Avicennia germinans = x = x - x - =
Solanum behamense = x - = x - - -
Scaevola plumieri x x x x x = = x
Iva imbricata x x x x x - x x

Melanthera aspera = x - x x - = a

52

Table 17. Numbers of species in different categories recorded from the
Dry Tortugas keys

IIOAP stole W987 1962 1977 Total

TOTAL NUMBER OF SPECIES

Bird 2 13 = = = 15
Bush 1 13 13 14 Mlk 28
East 8 8 4 ) 10 10
Garden 35 51 36 71 24 107
Loggerhead 22 38 22 32 24 56
Long = 8 5 10 10 16
Middle - 1 3 = = 3
Sand 5 3 3 = = 8

NUMBER OF NATIVE SPECIES

Bird 12 ipl = = - iL}
Bush 1 3 13 14 18 25
East 8 8 4 0) 10 10
Garden ig) 21 14 @) iM 26
Loggerhead 15 16 Abt iS) al 19
Long - 8 5 9 10 115)
Middle - 1 3 - = 3
Sand 5 8 3 = = 8

NUMBER OF INTRODUCED SPECIES

Bird O 1 - - - 1
Bush 0 6) 0 O 1 1
East ¢) O O O O 0)
Garden 8 22 ih} Syl 7 64
Loggerhead 2 14 8 8 6 23
Long - O 0) 1 O 1
Middle - O O - - O
Sand O O O - - 0)
NUMBER OF SPECIES OF UNCERTAIN STATUS

Bird O 1 - - - 1
Bush O 0) O ¢) 2 2
East O O O O O 0)
Garden 8 8 9 iat 3 A
Loggerhead 5 8 3 g 7 14
Long = O O O @) O
Middle - O O - - O
Sand 0 @) O = - )

Table 18.
keys

Bird

Bush

East

Garden

Loggerhead

Long

Middle

Sand

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

Area sq m
Total no. species
No. native species

*area considered unreliable

1904 LOS iL S)8)7/

6,440 9,375 =

12 13 =
12 11 =
= = Is) 5 SO
3 3) 14
3 LS} 14
67000877, 220% -
8 8 4
8 8 4

92,880 150,540* 112,420

32) Si 36
WY) 21 14
242,300 278, 700* 209,900
22 39 22
15 16 fe
= = 47,300
0 8 5
) 8 5

@) 194 =
0) 1 3
@) 1 3

228* DDO -

1977

Sit 350:
21
18

27,145
10
10

BA
il
227,700

24
lp:

Sp i

538

Summary of area and plant species numbers for Dry Tortugas

Total

107

16
15

54

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tall Conocarpus woodland on the eroding north shore.

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Pl. 7. East Key: Untola pantculata at the eroding northeast point.

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shore.

ATOLL RESEARCH BULLETIN
NO. 254

NATURAL HISTORY OF RAINE ISLAND, GREAT
BARRIER REEF

by D. R. Stoddart, P. E. Gibbs, and D. Hopley

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

July 1981

CONTENTS
Introduction
General description
Geology and geomorphology

Topography

Lithology

Stratigraphy

Sediments

Interpretation and subsequent observations
Phosphate deposits

Water supply
Vegetation and flora
Invertebrates

Marine Turtles

Birds

Ducula spilorrhoa spilorrhoa
Rallus philippensis yorki
Gallinula porphyrio melanotus
Puffinus pacificus

Pterodroma arminjoniana heraldica
Fregata minor

Fregata ariel

Sula leucogaster plotus

Sula sula rubripes

Sula dactylatra personata
Pelecanus conspicillatus
Phaethon rubricauda

Sterna bergii

Sterna fuscata nubilosa
Sterna anaethetus anaethetus
Anous stolidus pileatus

Anous minutus minutus

Larus novaehollandiae forsteri
Arenaria interpres interpres
Pluvialis squatarola
Pluvialis dominica fulva
Numenius phaeopus variegatus
Limosa lapponica baueri
Calidris ruficollis

Calidris acuminata

Egretta alba modesta

Egretta sacra

CO oO @O WO x OV

alia

Nycticorax caledonicus hilli
Merops ornatus

Hirundo nigricans nigricans
Myiagra rubecula rubecula
Aplonis metallica metallica

Disturbance by Man
Construction of the Beacon
Wreck of the Enchantress
Beche-de-mer Fishery
Phosphate mining

Acknowledgements

References

32
32
32
32
32

33
34
36
37/
S)7/
38

39

Or

List of Figures

The northern Great Barrier Reef, showing the location
of Raine Island.

Raine Island and its reef.

Monthly distribution of rainfall at Willis Island,
NOZTSOW (datvaytronataylorm 97S)

Raine Island in 1844, from Jukes (1847), vol. I,
De SSE5

ProOrilewor Raine: island, Leomigukes! (i847) vol. i,
4 IIe

Map of Raine Island. The areas in the central guano

flat are the seabird survey areas detailed in Table 2.

Topographic profile of Raine Island.

Sediment samples from Raine Island.

Histogram of curved carapace length of turtles measured

on the nest, night of 3 November 1973.

Section and elevation of the Raine Island Beacon, 1844,

after Bateson (1972), p. 201.

List of Tables
Visitors to Raine Island.

Sula leucogaster and Sula dactylatra in sample areas
of the central guano flat, midday, 3 November 1973.

aLgtal

iv

TS

List of Plates
(grouped at the end of the paper)
Raine Island in 1844, from Jukes (1847).
The central guano flat from the west end of the island, looking
towards the Beacon. The ridge in the foreground is the site of

the old guano railway.

Eastern end of the central guano flat, with Brown and Masked
Gannets.

Mounds of rubble and stones in the central guano flat.
Massive beachrock on the northeast shore.

Grooved and furrowed beachrock on the northeast shore.
Beachrock now distant from the beach on the north shore.

Phosphatic beachrock unconformably overlying old eroded beachrock
at the east end of the island.

Cliffs in phosphate rock at the southeast end of the island.
Irregular lower surface of the phosphate rock, forming caves, on
the south side of the island. Note the nesting Tropicbird
beneath the overhang.

Detail of the columnar structure of the phosphate rock.

Detached remnants of phosphate rock, south side of the island.
Western sand beach, showing early morning turtle tracks.

Turtle nests on the beach crest at the west end of the island.
Green Turtle on the northeast beachrock.

Turtle remains in Lepturus grassland of the high ridge.

Shearwater burrows in fine guano at the west end of the central
guano flat.

Mound in the central guano flat with juvenile Lesser Frigate-birds.

19-21. Juvenile Lesser Frigate-birds.

Papo

236

Brown Gannets lining the beachrock at the east end of the island.

Brown Gannets on beachrock on the northeast shore.

24. Brown Gannets on the nest in the central guano area. Note
the twigs outlining the nest.

25\. Brown Gannets nesting in the Lepturus grassland of the high
ridge. No twigs surround the nest in the vegetated areas.
26-27. Masked Gannets on the nest in the central guano area. The nest

is simply a depression in the sand.
28. Hatchling Masked Gannet in the central guano area.

29. Hatchling Masked Gannet with regurgitated flying fish Cypsilurus
melanocercus in a nest in the central guano area.

30-31. Red-footed Gannets on nests built on low Abutilon shrubs on the
high ridge at the west end of the island.

32. Fledgling Red-footed Gannet on the nest at the west end of the
island.

33. Juvenile Red-tailed Tropic-bird under the phosphate cliffs at the
east end of the island.

34. The Beacon and grave at the east end of the island, seen from the
south in November 1973.

35. The Beacon as erected in 1844, from Nautical Magazine, vol. 14
(USIS)p Ao BAST s

36. The Beacon in November 1973. The dark area on the lower wall is
that repaired by H.M.A.S. Gascoyne in 1961.

37. Inscriptions inside the walls of the Beacon.

38. The grave of Annie Eliza Ellis.

- =
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Fig. 1. The northern Great Barrier Reef, showing: the location of
Raine Island.

NATURAL HISTORY OF RAINE ISLAND, GREAT
BARRIER REEF

by D. R. Stoddart', P. E. Gibbs”, and D. Hopley?®

INTRODUCTION

Raine Island, on the northern Great Barrier Reef, is a small sand
cay 850 m long and up to 430 m wide. It was the subject of now
classic descriptions by Joseph Beete Jukes and John MacGillivray during
the voyage of H.M.S. Fly in 1843-45, when a substantial stone Beacon
was built there. Several other scientific parties, including that of
H.M.S. Challenger, have visited the island, but no general account
exists of it. This is the more surprising, since it is the most
important breeding station for tropical seabirds in Australia, and has
recently been shown to be one of the world's largest nesting sites for
the Green Turtle.

Much of Raine was originally covered with guano and lightly
cemented phosphate rock, which formed the basis of a brief but
considerable mining industry in 1890-92. Raine is, in fact,
interesting not only in itself but also as a representative of a class
of small, semi-arid guano and phosphate islands in the reef seas.
Among those recently studied are several in the Amirantes in the
western Indian Ocean; Laysan in the Leeward Hawaiian Islands; and
several of the smaller Phoenix Islands in the central Pacific. Raine
is the smallest of all of these: it is half the size of McKean and
Birnie Islands in the Phoenix group, and half the size of the Marie-

Louise in the Amirantes. Bird Island in the Seychelles is three
times as big as Raine, and most of the other phosphate islands are
much larger. But in spite of its small dimensions, Raine is of

sufficient interest to merit bringing all the available information on
it together, together with new observations from the work of the Royal
Society and Universities of Queensland Expedition to the northern Great
Barrier Reef in 1973.

Ihe Department of Geography, Downing Place, Cambridge, England.

Ao Marine Biological Association of the U.K., The Laboratory, Citadel
Hill, Plymouth, England.

3 Department of Geography, James Cook University of North Queensland,
Townsville, Australia.

Manuscript received August 1980 -- Eds.

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GENERAL DESCRIPTION

Raine Island is located at the leeward end of an oval patch reef
3.5 km long and 0.75 km wide, with an area of 210 ha, north of the
Great Detached Reef of the. northern Great Barrier Reef system, and
100 km ENE of Cape Grenville, the nearest point on the Queensland
mainland (Figures 1 and 2). Its co-ordinates are 11°36'S and
144°0O1'E. Jukes (1847) described the island as 1000 yds (915 m) long,
500 ft (150 m) wide, and up to 20 ft (6 m) high above high water level.
The central part of the island he described as covered with a cap of
calcareous sandstone, terminating on its seaward sides in a cliff
4-5 ft (1.2-1.5 m) high. The Raine Island Reef has been termed a
"Small coral atoll" (Tizard et al. 1885, II, 530), but in the absence
of a lagoon this usage seems inappropriate.

The first full descriptions of the island are those of Jukes
(1847) and MacGillivray (1846), based on observations made during the
time that a beacon was being built on the eastern end of the island by
Capt F.P. Blackwood and H.M.S. Fly in 1844. The island was
subsequently occupied during 1890-92, when tens of thousands of tons of
phosphate were dug and exported. Twenty years later, at the time of
W. Macgillivray's visits, seabirds were abundant, and the island is
still "probably the most important breeding station for tropical sea-
birds in Australian seas" (Warham 1961, 77). It is also a major
nesting site for Green Turtles, though this has attracted less attention
in the scientific literature.

No weather records have ever been kept at Raine. During April
to November the Southeast Trades blow constantly; during December to
March the Trades retreat to the south and are replaced by calms and
northerlies. Most rainfall occurs during this second period. Mean
annual rainfall has been estimated by A.T. Bath (in Warham 1961) as
1000 mm (40 inches); this is very close to the mean for Willis Island
in the Coral Sea (16°18'S, 149°59'E), where the mean annual rainfall
1921-1971 was 1098 mm (Figure 3). Reiwvab wel ey 4Oreroinralhvat Nil ans,
and probably also at Raine, is low; annual totals at Willis over the
period of record range from a minimum of 240 mm (in 1966) to a maximum
of 2024 mm (in 1959). Overnight dew is probably an important source of
moisture on these islands.

Tidal range has been estimated as 10-12 ft (3.0-3.7 m) (Blackwood
1844a, 540; Fairbridge 1950, 352). Mean tidal range at springs for
islands closer to the mainland between Cape Grenville and Cape Melville,
however, ranges from 5.2 to 6.8 ft (1.6-2.1 m), increasing northwards
to 8.4 ft (2.6 m) near Cape York.

GEOLOGY AND GEOMORPHOLOGY

Topography

MacGillivray (1846, 1474) described the following main features of
the island:

(a) a steep beach, 20 yards (18 m) or more in width.

(b) a "more or less continuous, low, wall-like border of coral
rock, its faces much decomposed by the weather, and hollowed
out in a Singular manner. ... This rock, which ranges from
a conglomerate to a coarse sandstone, is in general soft and
crumbles readily. ... This bed of sandstone is not more than
a few feet in thickness, and exhibits proof of its recent
formation, by containing shells and fragments of coral,
specifically identical with living inhabitants of the reef,
and occasionally eggs of turtle, apparently as if, while
in situ in the loose sand of what was then a mere sand-bank,
some chemical agency had converted the whole to a bed of
stone". According to MacGillivray the edge of the cliff
reached a height of 24 ft (7.3 m) above sea level.

(c) a lower central area with a surface cover of "rich black
mould".

Jukes (1847, I, 339) presented a schematic section through these
features.

Figures 4 and 5 give Jukes's map and profile of Raine Island as
surveyed in 1844. Figure 6 is a map of Raine surveyed by pacing and
compass traverse in November 1973, and Figure 7 is a profile across the
centre of the island surveyed at that time. The datum is inferred from
the measurement of a still water level at a given time, and comparison
with tide predictions for Sir Charles Hardy Islands, the nearest place
for which predictions are available; the Standard Port is Cairns
(Department of Harbours and Marine 1973). These data show the
following topographic features:

(a) the outer beach, generally 20 m wide and 5 m high, with
intermittent beachrock;

(b) the unvegetated beach crest, horizontal or falling slightly
landwards, 20-25 m wide;

(c) a low area of tussock grassland, varying in width from 10 to
80 m but averaging 30-40 m, with average elevation 4m ora
little less;

(ad) a cliff of cemented sandstone, of intricate outline in detail
and with some isolated islands of sandstone in the tussock
grassland, undercut and cavernous, generally 1-1.5 m high,
with the outer edge reaching a uniform height of 6 m;

from

(e) a high ridge of uncemented sand, with grass and herb cover,
widest in the south (average 100 m but varying from 75 to
110 m) and narrower in the north (average 35 m, varying from
25 to 75 m), with a maximum elevation of about 9 m;

(f£) a central, unvegetated flat, with a superficially-cemented
surface of white guano, with an elevation of 6 m accordant
with the height of the peripheral cliff edges.

Areas of the different topographic units derived by planimetry

the map in Figure 5 are as follows:

outer beach slope 36 V2
unvegetated beach crest 3.23
tussock grass depression 6.75

high ridge crest )55x3}
central guano flat 4.04
total area Dee

ha
ha
ha
ha
ha

ha

These figures do not include the area of intertidal beachrock, which
covers another 0.68 ha.

Lithology

oolite in texture and appearance.

The nature of the "coral rock" was first studied by Jukes, both at
outcrop on his first visit in 1843 (Jukes 1847, I, 128) and in quarries
opened during the erection of the Beacon in 1844 (1847, I, 337-340).

He found that "the stone was made up of small round grains, some of them
apparently rolled bits of coral and shell, but many of them evidently
concretionary, having concentric coats

not unlike some varieties of

It contained larger fragments of

coral and shells, and some pebbles of pumice, and it yielded
occasionally a fine sand that was not calcareous, and which was

probably derived from the pumice.

Some parts of it made a very fair

building stone, but it got softer below, till it passed downwards into a
coarser coral sand, unconsolidated and falling to pieces on being
touched" (1847, I, 127-128). Excavations for the quarries revealed
drusy cavities with gypsum, many recent shells, and one or two nests of
turtle eggs (1847, I, 128, 340), as well as pumice pebbles (1847, I,

SBi/) i.

round the shore.
dip of seven degrees.

In 1874 Moseley (1879, 299-300) observed that the rock was evenly
bedded, the bedding being horizontal in the centre and towards the sea

On the northeast side near the Beacon he quotes a
Moseley also contrasts the white sand, consisting

of shells, corals and Foraminifera, of the beaches, with the areas above
high water mark, which were redder and had a consolidated crust (1879,

300).

es

Stratigraphy

Two holes were dug during the Fly expedition in the centre of the
island. One dug by Jukes to a depth of 5 ft 1 in (1.5 m) showed the
following section:

O - 6 inches ( 0 - 15 cm) "Good black vegetable mould"

6 - 9 inches (15 - 23 cm) "Stone, brown mottled with white,
hard and coarse grained"

9 = 25 inches’ (23°-"63"%em) “Rich moist’ bilack ‘soil’; dikesbog—
earth"

25 - 61 inches (63-155 cm) "Stone of a light brown colour,
rather soft, but tough, and
yielding slowly to the pickaxe"

(Gukes* T8427) ee 1 2ap)is

MacGillivray's pit the following year reached a depth of 16 ft
(4.9 m), with the following section:

0 - 3 inches (0 Sate. hi) "Fine coral sandstone"

Ww
!

V2 inches, (756) — 30/25 cm) "Moist pulverulent black earth,
resembling peat, but without
any trace of woody fibres"

a
|

13 feet (30.5 cm - 3.96 m) "Successive deposits, varying
from coarse coral conglomerate
to unconcreted calcareous sand
mixed with a few small
scattered fragments of coral
and shells"

1b)

14 feet (3.96 - 4.27 m) "Masses and large fragments
of coral (of species now alive
on the reef), bleached and
water-worn, loosely inbedded

in coarse sand"; inflow of
seawater took place at this
depth.

(MacGillivray 1846, 1474).
Sediments

Sediment samples were taken from the beach slope, beach berm and
high ridge on the north, west and south sides of the island; these
were mechanically analysed and components identified in the fraction
coarser than 250 microns. Components were classed as coral and

coralline algae, molluscs, platey Foraminifera (mainly Marginopora),
spherical Foraminifera (including Baculogypsinoides), and Halimeda.

All the samples are remarkably homogeneous. They lie in the size range
of coarse sands (0.4 - 0.9%); are well sorted (0.3 - 0.5¢); have near
symmetrical distributions; and are mainly platykurtic. Two
components dominate, molluscan fragments and spheroidal Foraminifera.
Only on the leeward side do coral and coralline algal fragments
contribute significantly to the coarse fraction. Molluscan fragments
are more common on the beach and spherical Foraminifera on the berm

and ridge, presumably because of the greater mobility of the latter.
Small discrete lenses of platey Foraminifera, Halimeda and pumice were
noted at the time of collection. Figure 8 gives sample curves for a
beach (curve 1) and high ridge (curve 2) sample.

Two samples were also taken from the centre of the island, one
(curve 3 in Figure 8) from the central depression, and one (curve 4)
from its surrounding ridge. The central depression sample has a mean
size of 1.0¢, is moderately sorted, symmetrical and leptokurtic; it
differs from the beach and berm sediments in its fine tail (coarser
than 2.0¢). The north ridge sample is much finer (mean size 2.3¢),
is poorly sorted (1.25¢), symmetrical and platykurtic. Its
distribution differs markedly from that of the beach sands.

Interpretation and subsequent observations

Jukes (1847, I, 339-340) believed that the geology of the island
could be explained by (1) the formation of a sand cay at the leeward
end of the reef; (2) the formation of a crust by solution and
redeposition of calcium carbonate above high water mark: (3) the
cessation of accumulation and the cliffing of the lithified material;
and (4) the later resumption of sand accumulation to form the present
peripheral sand ridge. The age of the whole island he believed was
indicated by the thickness of up to 2 ft of "vegetable mould" in the
central depression. Moseley in 1874 compared the rocks of Raine with
the "calcareous sand rock" or aeolianite of Bermuda, except that the
former was more evenly bedded. Jukes's interpretation is largely
confirmed by recent observations, though following the large-scale
guano digging of 1890-92 it is no longer possible to be sure that
surface features are not of human origin, at least in the central
depression.

Many later interpretations, often involving sea-level changes,
are less closely related to the facts of lithology and stratigraphy
than was Jukes's. Rattray (1869, 303) described Raine as "consisting
of hard compact brecciated coral conglomerate, with a shelving beach of
coarse coralline and shelly sand, and a scanty superstratum composed of
the coral debris sparingly mixed with vegetable matter, and a thin
layer of guano". He suggested that the present island had been formed
beneath the sea and had then undergone emergence, and he termed the
island an "extinct" coral reef. Neither Jukes nor MacGillivray had
described a coral conglomerate or any raised reef. Several subsequent
workers accepted Rattray's inference of emergence, however, from
Agassiz (1898, 124) onwards. Fairbridge (1950, 352) states that Raine

|
|
|

10

is "in part at least, an emerged beach rock. With a tidal range of
10-12 feet and an extra 2-3 feet to include mean swash, an emergence of
about 10-11 feet is indicated. Rattray (1869) also mentioned a raised
coral breccia here. A definitely emerged coral reef occurs some 12
miles southwest of Raine Island (Jukes 1847). he *is2—3' miles tlong
and 0.25 miles wide, with Porites colonies in the position of growth
NO ee feet above the present reef growth level". Fairbridge ina
later paper (1967, 403) called Raine "the only emerged reef island on
the whole outer barrier; Jukes observed that it consists of a
calcareous aeolianite, with an emerged beachrock terrace here ascribed
to the 3 m mid-Holocene stage. The aeolianite, by analogy with other
offshore occurrences in Australia ... is probably of late glacial
(Wtirm) age, when sea level was low enough to permit wide beaches to
form and dune accumulations to develop". According to Bennett (1971,
37), Raine "consists of older, solidified reef material formed at a
period when the sea level was higher", and the stone used for the
Beacon was "quarried out of the solid reef".

These conclusions evidently go substantially beyond the facts so
well recorded by Jukes and MacGillivray in 1843-44, and are not supported
by evidence collected then or observable now. The 3 m mid-Holocene
"emerged beachrock terrace" of Fairbridge is not beachrock, is not
emerged (in the sense of being now at a higher level with respect to
sea level than when it was formed), and does not stand at 3 m. The
cliff edges stand at 6 m and are accordant with the central guano flat.
All the beachrock at Raine Island is at low intertidal levels, and there
is no evidence that any beachrock there is raised: a radiocarbon date
(ANU-1591) on a Tridacna valve from beachrock at spring-tide swash level
is t80's 65 4yr ‘BaP. (Polach et al. 1978, 151): There is no evidence
that the main cemented horizon which outcrops in the cliffs is an
aeolianite (cross-bedding and similar sedimentary structures seem to be
completely lacking), and all the evidence indicates that it isa
superficial cementation phenomenon, decreasing with depth, in the way

that Jukes envisaged. It is a cay sandstone in which the cement is
partly phosphatic, and cementation probably represents a continuing
process rather than a discrete event. The "raised reef" 12 miles

southwest of Raine Island, described by Jukes and to which Fairbridge
refers, was also visited in 1973: it is not a raised reef and there
are no Porites colonies in the position of growth: it is simply a
deposit of large storm boulders near the reef edge, all now much eroded,
and having no significance with respect to changes in sea level.

Phosphate deposits

No detailed account of the Raine Island phosphates, mined in the
central area in 1890-92, has been published, and no analyses of the
material are known. According to Saville-Kent (1893, 120), "the
deposits occur under three distinct conditions: Firstly, in layers some
fifteen inches thick, immediately beneath the upper crust of coral
conglomerate, which constitutes the encircling plateau described by
Mr Jukes; secondly, in pothole-like hollows in the same location; and
thirdly, in trench-like depressions in the central black earth basin".

ial

The cliffs probably retain much of their original appearance,
since the mining was carried out in the central flat; they closely
resemble comparable cliffs on other islands with phosphatic cementation,
such as Denis Island, Seychelles (Fryer 1910, 18-19, Plate 2). The
cementation is greatest at higher levels and diminishes with depth;
the lower surface of the cemented horizon is irregular, and forms a
series of downward-directed lobes and pillars. These probably indicate
differential permeability in the originally uncemented sands. Weer
Scoffin has examined specimens from the upper part of the cliffs as
part of a wider study of coral island phosphates. The grains in the
sands are of variable origin: Foraminifera 40%, Halimeda 30%, coral 10%,
mollusc 10%, unknown 10%. Phosphatic cement occurs as a wavy thin film
with laminated structure coating the individual grains; the grains
themselves have been altered centripetally for distances of a few

microns from the grain margins. The phosphate mineral is
hydroxylapatite; the other main mineral present is aragonite, with a
trace of calcite. The presence of aragonite suggests a younge age for
the material. Chemical composition is 31.0% P205, 54.0% Cao.

WATER SUPPLY

There is no standing surface water on Raine, except possibly for

a few days immediately after storms. During the Fly expedition water
was imported from Sir Charles Hardy's Islands for the men building the
Beacon. Jukes (1847, II, 266) says that "one or two wells were sunk

in the island, but no fresh water was procured; although in one of the
wells, at a depth of 16 feet, the water was only brackish, and could
be used to slack the lime, although very unpalatable to the taste".
Arundel in 1890 had to construct a seawater distillation plant for his
work force. Warham (1961, 78-79) also states that there is no fresh
water and that "various attempts to strike it by digging in the past
have failed".

A carved inscription on the inside of the walls of the Beacon,
however, apparently of mid-nineteenth century date (it is mentioned in
Arundel's diary in 1890), states that fresh water can be obtained at a
depth of 7 feet.

VEGETATION AND FLORA

The vegetation of Raine Island consists of low shrubs, herbs and
grasses; there are no trees. To Blackwood (1844a, 539) it was the
"quantity of coarse green vegetation on it" which distinguished Raine
from most of the other small sand cays in this northern reef area;
he was clearly thinking of unvegetated sand cays in the vicinity, such
as those on Ashmore Reef.

Jukes (1847, I, 127) described "a low shrubby vegetation, partly
of reed-like or umbelliferous plants, and partly with a close green
carpet of a plant with succulent leaves and stem, which we subsequently
found was good to eat, and so went with us by the name of ‘spinach'"

————

12

(this was probably Portulaca). MacGillivray (1846, 1475) gave a
fuller account, though unfortunately unsupported by specimens: "Of

the Botany of the islet I can give but a very meagre account, for a
collection of about twenty species, found by me, unfortunately went to
decay for want of a proper place of storage on board ship. Several of
these species I have elsewhere observed on the main land of New Holland;
among others, a long, trailing, woody plant with conspicuous yellow
blossoms [probably Tribulus cistoides], and a large white-flowered
convolvulus [probably Ipomoea macrantha]. Two species, very abundant,
on the island, were used by us as vegetables, one of them under the
name of spinach [Portulaca], for which it was considered a very fair
substitute, and as such served out to the ship's company. Most of the
plants of the island are more or less succulent; there is but one
shrub, a slender Acacia, five or six feet high, with small yellow
flowers [probably Sesbania]". The surrounding sand area had "a few
scattered tufts of grass and other herbage" (MacGillivray 1846, 1474).
A variety of cultivated plants was introduced during the Fly expedition
and during the visit of the Heroine in 1845; they are described later.

The Challenger Expedition in 1874 recorded 11 species of flowering

plants, including two grasses. They noted fungi on dung, but no
mosses, ferns or lichens. There was also no trace of the Fly vegetable
gardens or of other introductions. No species list appears to have

been published from these collections (Moseley 1879).

In 1910 W. Macgillivray (1910, 224) found "a coarse grass, a kind
of pig-face weed, and a low perennial shrub of horizontal growth, not
more than 1 to 13 feet from the ground anywhere, and bearing grey-green
leaves and a yellow flower". These were probably Lepturus repens,
Portulaca sp., and Tribulus cistoides, respectively.

In February 1959, after good rains, Warham (1961, 78) found only
Six species of flowering plants, "the principal one being a kind of
mallow (Abutilon graveolus or indica) which was flowering, a spinach-
like amaranth (Amaranthus viridis) and a low bushy plant Tribulus
cistoides. This bore orange flowers and is probably the ‘'acacia' of
the early visitors of the island [Abutilon and Tribulus appear confused
in; this account] . Two grasses also flourished: Eleusine indica, an
introduced species, and Lepturus repens, a native plant. This grew
quite thickly on the dunes on the south-east side of the island. A
low succulent also grew around the edges of the central depression
[probably Portulaca sp.]. None of these plants was more than two feet
folie faye

The only published list of Raine Island plants appears to be that

of Hindwood et al. (1963, 44), based on collections made in November
1961 and lodged in the Queensland Herbarium, Brisbane, and the
Division of Plant industry, C.SiLok.On,. Canberra. Six species are

recorded, and it is likely from Warham's records two years earlier
that others were missed.

13

Plants were collected in 1973 and were identified by Miss S.
Reynolds at the Queensland Herbarium, Brisbane. The following list
includes the plants cited by Hindwood et al. (1963).

Gramineae ;
Eleusine indica (L.) Gaertn.
Stoddart 5061. Also cited by Warham (1961, 78).
Lepturus repens R.Br. cf. var. repens
Stoddart 5051, 5059. Also cited by Warham (1961, 78)
and Hindwood et al. (1963, 44).

Nyctaginaceae
Boerhavia sp.
Stoddart 5054 (specimen lost). Hindwood et al. (1963, 44)
also cite Boerhavia tetrandra Forst., but the specimen
collected in 1973 is not this species.

Amaranthaceae
Achyranthes aspera L.
Stoddart 5053. Cited by Hindwood et al. (1963, 44).
Amaranthus viridis L.
Stoddart 5057, 5058? Warham (1961, 78) also cites
Amaranthus viridis.

Portulacaceae
Portulaca oleracea L. (sensu lato)
Stoddart 5056. Possibly the plant mentioned by
W. Macgillivray (1910, 224) and Warham (1961, 78).

Cruciferae
Coronopus integrifolius Spreng.
Stoddart 5060

Leguminosae
Sesbania cf. aculeata Poir.
Stoddart 5050. Sesbania aculeata is cited by

Hindwood et al. (1963, 44).

Zygophyllaceae
Tribulus cistoides L.
Stoddant 5055% Cited by Warham (1961, 78) and
Hindwood et al. (1963, 44); possibly the plant
mentioned by MacGillivray (1846, 1475).

Malvaceae
Abutilon asiaticum var. australiense (Noch. ex Britt.) Fosb.
Stoddart 5052. Cited as A. indicum (L.) Sweet by
Hindwood et al. (1963, 44); probably the plants
mentioned by W. Macgillivray (1910, 224) and Warham
(CSS a 5 78))) 6

14

Convolvulaceae
Ipomoea macrantha R. and S.
Not collected in 1973 or by Hindwood et al. (1963).
Possibly the plant mentioned by MacGillivray (1846,
EATS

INVERTEBRATES

Most of the collections of invertebrates made at and near Raine
Island have been of marine animals during the Challenger Expedition.
These are listed by Murray (1895, 682-688); they comprise mainly
species of Ostracoda, Mollusca and Foraminifera, with only a few land
or shore animals.

The most conspicuous, indeed the only, semi-terrestrial crustacean
is the ghost crab Ocypode. Moseley collected a single male specimen of
O. ceratophthalma (Pallas), recorded by Miers (1886, 238-239). itoy lS) 7/3}
the population was predominantly O. ceratophthalma with a few O.
cordimana Desmarest, the two species being represented in a sample of
60 andividuals “invalratzovo£ (6.5) 3 1. Coenobitid hermit crabs,
elsewhere common on guano islands in the central Pacific and Indian
Oceans and also in the Caribbean, appear to be completely absent on
Raine and other Great Barrier Reef islands. Mole-crabs of the genus
Hippa inhabit the coarse sands of the lower beach levels; although
H. pacifica (Dana) was the only species collected in 1973, it seems
likely that H. celaeno (de Man), a commoner species on other reef
islands in the region (see Gibbs 1978), is present also.

Among insects, MacGillivray (1846) noted a large scarab, Coleoptera
including Hister, Necrobia cf. ruficollis, and Dermestes murinus;
Pimelia; and ixodid ticks. Moseley (1879, 302) found an earwig
Forficula under stones and a very common locust Acridium. No further
invertebrate collections appear to have been made.

MARINE TURTLES

The Green Turtle nesting season at Raine Island is concentrated
from October to December; visitors outside these times have usually
made few turtle observations. Parsons (1962) in his standard work on
the Green Turtle makes no mention of Raine Island, and while Bustard
(1972) includes Raine Island in a map as an important nesting site he
makes no mention of it in his text. Yet Raine is certainly one of the
largest Green Turtle nesting sites in the world, and could be the
largest remaining still undisturbed by man.

Jukes's first visit in July 1843 produced few records. "There
were a few turtle tracks on the beach, but we did not succeed in taking
any, though many dead ones were scattered about the island", especially
at the foot of the cliff (Jukes 1847, I; 130): During the 1844 visit,
MacGillivray (1846, 1479) found that "during the months of June, July
and August, the turtle occurred at irregular intervals, generally

5)

singly, but in the beginning of September they became more numerous".
In the following year a party from the Bramble found the cliffs and
quarries "full of the remains [of turtles] ... who had fallen over on
their backs and perished miserably". They took 14 turtle by turning
them on the beaches at night, some of them being of 250-300 lbs weight
(roughly 110-140 kg). Sweatman (MS, 94-95) gives a graphic account of
taking turtle during this visit. Mackenzie (1845, 494), calling with
the Heroine on 25 January 1845, "obtained fourteen large turtles, each
averaging four cwt [ca 200 kg]". In August 1874 the Challenger found
only the carapaces of numerous dead turtles (Moseley 1879, 302).

Conversely, Macgillivray (1910, 224), on 30 October 1910, found
"great numbers of turtles ... on the beach and in the shallow water
round the boat". On 4 December 1913 he found that "about thirty
turtles were crawling up the beach near where we landed, and the
shallow water contained hundreds of them", coming ashore at night to
nest (Macgillivray 1917, 67).

During the Fly visit, MacGillivray (1846, 1478-1479) noted that
all the turtle seen were Green, Chelonia mydas, and that they laid
clutches of at least fifty eggs. He found that the hatching young
suffered predation by frigate birds and sooty terns, and sixty years
later Macgillivray (1917, 84) recorded predation of eggs by Crested
Tern Sterna bergii and Silver Gull Larus novae-hollandiae.

Green Turtle were nesting during the visit to Raine Island in
LOWS Tracks were counted round the island each morning. Measured
at the foot of the beach, the perimeter of the island occupied by sand
(including the low and discontinuous narrow beachrock of the northern
coast) totals 1600 m, and that occupied by beachrock 500 m (76 and 24%
respectively). Numbers counted on the beach only were as follows:

Tracks Animals

Night of 31 October 324 162
1 November 245 WAS)
2 November 230 IIS)
3 November 270 13 )5

During the night of 3 November, animals were also counted on the nest
both on sandy beaches and also inland from beachrock shores. The
ratio of tracks:animals on the beaches can then be used to infer the
number of tracks which might be expected on the beachrock shores, to
give an estimate of the total number coming ashore for the whole island
on that night. The results are:

Beach Beachrock Whole island

(a) Animals counted on nest 87 38 125
(b) Numbers inferred from tracks SYS)
Ratio (b): (a) 155

Inferred total numbers BIS) 59 194

16

In subsequent surveys, Birtles (1978) estimated the numbers of nesting
turtles at 11,800 in 1974, 50 in 1975, 1000 maximum in 1976, and

50-100 in 1977. Kowarsky (1978) also made ground and aerial
observations in January 1976, but although tracks were numerous he
could not make any estimate of numbers. For comparison, at Ile Europa,
Mozambique Channel, sometimes stated to be the largest Green Turtle
colony in the world, Hughes (1974) estimated 5,000 emergences a year,
and Servan (1976) 1300 a year.

The prime nesting habitat on Raine is the unvegetated sand areas
of the beach crest, though a considerable number of turtles do nest in
the outer Lepturus tussock grassland between the beach and the cliffs.
Planimetric measurements on Figure 5 indicate a total area of
unvegetated beach crest of 32,300 sq m. A turtle 1 m long requires an
area of not less than 1.54 sq m for its nesting depression. Hence
there are approximately 21,000 potential non-overlapping nest sites in
the beach crest area. Two hundred turtles nesting in one night would
have 161.5 sq m each in which to choose a nesting site. Each site
would have roughly 1 chance in 100 of being selected. These
calculations give some idea of the possible potential size of the Raine
Island colony. Elsewhere on the Great Barrier Reef, on vegetated
islands, the size of nesting area available is a limiting factor in the
number of successful nestings (Bustard and Tognetti 1969): at Raine
Island this is unlikely to be so. Kowarsky (1978) quotes a clutch size
at Raine of 105.8 + 22.0 eggs (n = 6).

The curved carapace length of females on the nest was also
measured on the night of 3 November, and Figure 9 presents a histogram
of the results. 124 animals were measured. The mean length was
109 cm, standard deviation 5.29 cm; the smallest measured was 90 cm
long, and the largest 122 cm. Bustard (1972, 138-141) quotes average
figures of 107 cm curved length for Great Barrier Reef Green Turtles,
and ranges of 89-127 cm. Curved length can be converted to straight
length using the formula derived for Aldabra turtles by Frazier (1971,
390), and the calculated straight lengths then used to compare
populations in different parts of the world. The Raine Island
turtles are very similar in size to those of Aldabra, markedly bigger
than those of Ceylon, Yemen and Sarawak, and markedly smaller than those
of Guyana, Ascension Island and Surinam (data listed by Frazier 1971,
380, and Servan 1976, 424). In addition to the 1973 data, Kowarsky
(1978) gives mean straight length of 100.2 + 5.5 and mean curved length
108236 +°5.1- cm’ for four individuals at Raine Tsilland*

While measuring the turtles in 1973 it was observed that animals
differed markedly in degree of curvature of carapace. In some the
curvature was so shallow that it is possible they belong to Chelonia
depressa Garman, the Flatback Turtle (Bustard 1972, 74-88) rather than
to Chelonia mydas L.

According to Limpus (1978), tagged turtles from Raine Island have
been recovered from south of Cooktown, Queensland, through the Torres
Strait and southern Papua New Guinea to Aru Island, Indonesia.

1L7/

In addition to the Green Turtle, Boulenger (1889) also records the
Hawksbill Eretmochelys imbricata (as Chelone imbricata) from Raine,
but no subsequent records are known.

BIRDS

Raine Island is a major breeding station for seabirds, and
probably the most important in terms of numbers of species on the Great
Barrier Reef. It has large breeding colonies of Brown and Masked
Gannets, Noddies, Sooty Terns, and Wedge-tailed Shearwaters. ite) aS
also a breeding site for the Red-tailed Tropic-bird and the Lesser
Frigate-bird, otherwise rare on the Great Barrier Reef. There are no
comparable seabird-breeding stations (except for colonies of Noddies
and Sooty Terns) between Raine and the Capricorn and Bunker Islands at
the southern end of the Reef. There is also a resident land bird, the
Banded Landrail, many shorebirds, and a number of migrants and vagrants
recorded.

The first ornithological observations were those of Jukes and
MacGillivray in 1843-44 during the visits of the Fly. Moseley made
observations and collections during the Challenger visit in August 1874.
The first extensive records are those of MacGillivray and M'Lennan in
MOMOP Ot ancds LOS The most detailed account of Raine Island birds
is that of Warham (1961), following his visit in February 1959.
Warham's list is used as the basis for the present account, though
nomenclature follows Storr (1973); we are most grateful to C.W. Benson
for his assistance with and comments on this list. Following Warham,
K.A. Hindwood and others also visited Raine during a survey of seabirds
of the Coral Sea in November 1961; their records are also incorporated
here.

In addition to the birds listed here, Ellis (1937, 179) also
records "several long-legged and long-billed ibises", "three common
crows", and "several diminutive hawks". MacGillivray (1846) also
records several species of uncertain synonymy, including Charadrius
virginianus, Thalasseus strenuus, Callocalia arborea, and Porzana.

Ducula spilorrhoa spilorrhoa (Gray) Torres Strait Pigeon

One migrant bird is recorded (as Myristicivora spilorrhoa) by
PLES (193865 178):

Rallus philippensis yorki (Mathews) Banded Landrail

Recorded as Rallus philippensis? by MacGillivray (1846, 1476-77) in

1844, and subsequently figured by Gould (1848, VI, 76). It was "very
abundant all over the island", under vegetation, beneath overhangs, and
in shearwater burrows. MacGillivray found it very wary, though it ran

and rarely flew. He caught "great numbers" (up to 45 in a day) with
his dog.

18

In 1874 Moseley collected two males and one female (Forbes 1878).
Under the name Rallus pectoralis Moseley described it as tame, and
easily knocked down with sticks or caught by hand. In August there
were full-fledged young (Moseley 1879, 301). On the basis of this
record Kikkawa (1976) lists R. pectoralis Lewin Water Rail as a separate
species (but see discussion below).

During 1890-92 there were initially large numbers, but "a welcome

addition to the table they were. Their numbers were materially
lessened during our occupation of the island, but some were left to
carry on the restocking" (Ellis 1937, 177-178). Macgillivray in

October 1910 simply records "numbers" running over the island (1910,
226).

There are no further records until Warham's visit in February 1959.

It was then "very numerous and ... often seen in the daytime skulking
among the herbage or running about beneath the caves and cavities under
the iciaitis At night they spread out all over the island and were
often seen on the dunes where they fed on turtle eggs exposed by the
subsequent diggings of other turtles. -.». Two nests were discovered

.. their eggs being hidden beneath the thick herbage" (Warham 1961,
83)°2 Warham also noted that the rails could be approached closely at
night. The description agrees with the situation in 1973, except that
dead birds were numerous on the ground, both on the high ridge and
beneath the cliffs. Several of these dead birds were collected.
Two males were also collected by Hindwood et al. (1963, 38) in
November 1961; one is in the Western Australian Museum (no. A8755)
andthe other iin’ the €7S-s1.R:0. collection (no: CSIRO’301):

Storr (1973) states that R. p. yorki, a small, dark race, is very
common in the Torres Straits islands and on low islands of the
Queensland coast from Pandora Cay to Lady Musgrave Island (in 1973 we
observed rails on Hope Islands, Turtle Islands, Three Isles, and
Pelican Island). A second race, R. p. pectoralis, is of uncertain
status but evidently rare in Queensland. The pectoral band in yorki
is darker (chestnut cinnamon rather than pale cinnamon) and narrower
(6-10 mm wide rather than 14-30 mm). C.J.0O. Harrison has kindly re-
examined specimens from Raine Island in the British Museum (Natural
History), together with those taken as dead birds in 1973. The three
Challenger specimens which included a young bird, suggesting recent
breeding, are of the southern race, pectoralis. S.A. Parker has
examined Raine Island specimens in Australian museums, and found them
to be yorki, and the specimens taken in 1973 were also of this form.
Harrison comments: "It seems probable from the evidence that both
Australian races are migratory to some degree, and that both could
potentially occur in Raine Island in passage. There are other
references in literature to the species breeding on the island but no
subspecific recognition of them. There is no clear indication of the
status or origin of the birds which do or did (intermittently?) breed
on the island. Dead bodies might be those of failed migrants."

19

Gallinula porphyrio melanotus (Temminck) Eastern Swamp-hen

This common Torres Strait migrant is named (as Porphyrio melanotus)

by Warham (1961) from an observation by Ellis (1936). Ellis refers to
"some game-birds with dark blue plumage, a patch of white below the
tail, strong red beak, and long red legs and feet ... known as

pukekos in New Zealand", which were seen on "several occasions"
GalMlbisc rs 936), 79): =

Diomedea chrysostoma Forster Grey-headed Albatross

This is listed for Raine Island by Kikkawa (1976) on the basis of
a reference by Hull (1925). Hull saw a bird near Lizard Island in
June 1924 which "answered fairly well to the description of Diomedea
chrysostoma, except for the neck shading. The mate ...a Filipino,
said he had seen a similar bird at Raine Islet ..." (Hull ISAS Ao AUS)
This is clearly an uncertain record.

Puffinus pacificus (Gmelin) Wedge-tailed Shearwater

In May 1844 MacGillivray (1846, 1478) noted (as Puffinus ------ ?)
"a small colony of these birds among some rank herbage which concealed
the entrances to their burrows, in which they were easily caught ...
it is possible that they are, in a great measure, nocturnal" as at
Heron Island.

Under the name Puffinus sphenurus, Macgillivray and M'Lennan made
several observations during 1910-1913. In October 1910: "At the
north-western end of the island many burrows of a Petrel are found.

We dug out several to a depth of 4 or 5 feet, and found them to contain
either one or a pair of birds (Puffinus sphenurus) . There are no eggs.
It seems that the birds are only cleaning out their burrows and will
probably lay in a month's time" (Macgillivray 1910, 226). In July
1911 the shearwater was again "in its burrows". M'Lennan "obtained
several skins, and noted that each burrow contained two birds. He
visited the burrows after dark, and waited for some time, and saw only
one bird leave, but could hear mournful calling in all directions. A
male and female in each burrow" (Macgillivray 1914, 141). In December
1913 Macgillivray (1917, 83-84) noted "many burrows", some 6 ft long,
in the centre of the island, most with one egg and some with downy
young.

In February 1959 Warham (1961, 80, 83) found "some hundreds of
burrows", with "large young in down, some with feathers 1.25 inches
long on the wings". In November 1961 Hindwood et al. (1963, 38) noted
breeding, without further detail. In 1973 inhabited burrows were
concentrated on the high ridge at the western end of the island.

This colony of shearwaters is the only one on the Great Barrier
Reef north of the Bunker and Capricorn Island (Lavery and Grimes 1971;
Serventy et al. 1971, 124). At the Heron Island colony, birds are
present from October to April (Moulton 1961). In 1960 the first birds

20

arrived at Heron Island on 8 October, and the first eggs were laid on
15 December. Each day the birds left the island at 5 a.m. and
returned at 7.45 p.m. (Gross, Moulton and Huntington 1963).

Pterodroma arminjoniana heraldica (Salvin) Trinidade Petrel

Recorded for the first time in Australia and photographed on
22 February 1959 by Warham (1959, 1961: Serventy et al.1971). Warham
also found an egg which may belong to this bird.

Fregata minor (Gmelin) Greater Frigate-bird

There are several early references to this species on Raine Island,
possibilyiin, erronifor.thatarilels Storr (1973, 8) states that minor
apparently breeds in Torres Strait, while ariel breeds on Rocky Island
(Gulf of Carpentaria), Raine Island, and ‘Gillett Cay', Swains Reefs,
in the southernmost sector of the Great Barrier Reef, April to August.
Bennett (1971, 60) figures juvenile birds on the nest at the latter
location. Serventy et al. (1971, 155) state that minor is not known
to breed at Raine Island. It is normally a tree-nesting species, and
thus might not be expected there, though small numbers are found on
treeless islands in the Phoenix Group, Central Pacific.

F. minor is recorded from Raine Island as F. aquila by
MacGillivray (1846), quoting Lt J.M.R. Ince. Specimens were said to
have been sent to Gould, but they are not mentioned by Gould (1865).

F. minor is also mentioned during the Challenger visit by Tizard et al.
(1885), and one male and four females were so identified by Sclater and
Salvin (1878, 650) as collected by the Challenger on 31 August 1874.
These specimens were, however, cited as F. ariel by North (1912, 353),
and they were also so listed by Ogilvie-Grant (1898, 449). Two of

the specimens, in the British Museum (Natural History), have been found
and re-examined by C.W. Benson, who confirms them to be ariel and not
minor (see next species).

Warham (1961, 83) in 1959 saw 4-5 birds identified as F. minor,

all apparently adult females. It was possible that an area of 40

empty nests might belong to this species, since these were on the ground
and all the ariel nests he saw were on bushes. There is, however, no
breeding record of F. minor yet for Australia. Lavery and Grimes

(1971) cite Raine as a possible breeding site for this species, but its
status there remains to be established.

Fregata ariel Lesser Frigate-bird

Raine Island is one of only two Great Barrier Reef breeding
localities for this species (Lavery and Grimes 1971). In July 1843
Jukes (1847, 2, 129) found “young of all ages”. In May 1844,
MacGillivray (1846, 1478), under the name Atagen ariel, reported
"small colonies of about a dozen individuals. Its nest is formed of
small dry twigs, raised about a foot from the ground, or sometimes
placed upon a tuft of herbage, a foot in diameter"; he noted one egg.

Cad

Ince (in Gould 1848, VII, 72) described it "breeding in colonies at its
S.W. corner; the nest being composed of a few small sticks collected
from the shrubs and herbaceous plants which alone clothe the island,
and placed either on the ground or on the plants a few inches above it.
The eggs ... generally one, but occasionally two". Ince found stages
from fresh eggs to 2-3 week hatchlings. Stomach contents comprised
young turtles, squid and crabs.

In August 1874 the Challenger found 30-40 nests with young well
advanced (mostly in the air) but no eggs. All the nests were in a
small patch near the cleared area, and consisted of compact platforms
of twigs and grass 8 inches in diameter raised on bushes above the
ground surface (Moseley 1879, 301-302: Sclater and Salvin 1878). The
identity of these birds has been discussed under the previous species.
Two specimens were re-examined by C.W. Benson at Tring:

No. 80.11.18.119: adult male, no date
No. 80.11.18.442: young bird still partly in down, no date.

Both are F. ariel, as listed by Ogilvie-Grant (1898, 449).

iin Ocreoloere O10) Macepsiiikivsceyy, (UG), ZAR ionaeey WS)A > S55) se oybuovel
50 feathered young on the east side of the island near the centre,
presumably from eggs laid in May-June. The nests were flat platforms
of sticks and grass 4-6 inches above the ground. In July 1911 M'Lennan
found 150 nests: "Several colonies of Frigate-birds were seen near the
beacon ... They were eight in number, of from three to thirty nests.
I counted 150 nests altogether, several of which contained one egg
each; two of these were on the point of hatching. The rest of the
nests contained one young bird each, in all stages of plumage, from a
couple of days old to birds ready to fly" (Macgillivray 1914, 148).
In December 1913 nesting was over; fledged young were all over the
island and soaring in the air. Macguililivray Clg, Veil—182) stook and
described one male and one female specimen.

Warham in February 1959 estimated the population at 2000, many
possibly from elsewhere, and counted about 150 nests. There were no
eggs. The young were all able to fly though three or four still spent
all day on the nest. 80 per cent of the 500 birds in the air during
the day were juveniles. Each bird on the ground was perched on a
dwarf shrub of Abutilon or Tribulus. The frigates were seen harrying
Red-footed and Masked Gannets, and also fishing. Warham (1961, 80, 84;
Serventy et al. 1971, 156) also gives notes on behaviour. Hindwood
et al. (1963, 39) confirmed the breeding and collected one female in
November 1961 (Queensland Museum No. 0.9135).

In November 1973 adult males and females were numerous both perched
on the parapet of the Beacon and soaring in the air at the east end of
the island. There were two concentrations of juveniles not yet flying:
one of about 120 birds on a mound in the central guano flat (Figure 5),
and another of about 65 juveniles immediately to the north, on the
Lepturus tussocks between the cliff and the beach crest. Ogilvie

22.

(1975) estimated the population at 300-500. For notes on diet, see
under Sula leucogaster.

Sula leucogaster plotus (Forster) Brown Gannet

There are several large colonies of Brown Gannets on the Great
Barrier Reef islands, including those on Ashmore Banks and Pandora Cay
near Raine Island, and on Raine Island itself (Lavery and Grimes 1971,
Serventy et al. 1971, 172).

In the 1840s, however, numbers were apparently small. Jukes
(1847, I, 129) in July 1843 found a few with young, forming a flock
nesting separately from the Masked Gannet. In May 1844, under the
name Sula fiber, MacGillivray noted: "The well-known brown booby
breeds upon Raine's Islet, but in small numbers, as I found its nest
there only once" (MacGillivray 1846, 1478). Specimens, presumably
MacGillivray's, were reported as Sula fusca by Gould (1848, VII, 78),
and there is a specimen (10/Sul/2/C/6) marked 'Blackwood Collection'
in the University Museum of Zoology, Cambridge (C.W. Benson, pers.
comm.).Moseley collected a male in August 1874 (Sclater and Salvin 1878,
651) and described "a slight nest of green twigs and grass on the
ground" (Moseley 1879, 301), but gives no indication of abundance.

By the time of Macgillivray's first visit on 30 October 1910,
however, ninety per cent of the nesting birds consisted of this species.
In July 1911 M'Lennan found them all over the island, and Macgillivray
described them "in thousands all over the place" (1917, 181). At the
time of his October visit there were eggs, naked young, young in down,
and young fledged. Most clutches were of two eggs, variable in size
and shape. MacGillivray noted that in these cases only one survived,
probably because of the initial difference in size of the hatchlings.
The nest was simply a depression in the sand, 3-4 inches deep and 8-12
inches in diameter, with a few sticks surrounding it (Macgillivray
MO TOF an 2i24)) In July 1911 there were several nests with one egg, eight
with two, but no hatched young, and nesting had clearly only just
begun (Macgillivray 1914, 148). In December 1913, nesting was finished,
very few eggs were left, and there were young birds everywhere
(Macgillivray 1917, 181). Further observations by Macgillivray, with
a photograph, are given by North (1912, 351-353).

In February 1959 it was again the most plentiful bird: Warham
(1961, 80) estimated a total population of 7,000-9,000. He saw only
two sets of eggs; the young were mostly in down, feathered or flying,
but still fed by their parents. In addition to birds on the nest
there were about 2,000 birds roosting on the beaches in lines facing
the sea. Warham notes various aspects of behaviour and draws attention
to the difference in call between male (hissing) and female (a deeper
call).

In November 1961 Hindwood et al. (1963, 21; Hindwood 1964, 309)
estimated there were at least 2,000 pairs. The nests contained well-
incubated eggs or very small nestlings, and there were a few large downy

23

young. Most of the birds were in the central hare area. The
differential survival of larger nestlings in a clutch of two was again
noted.

The Brown Gannet was the most abundant nesting seabird in November
LQYSe Most of the birds were on the central flats. The nest was a
slight depression, often broken through the surface crust, surrounded
by twigs. Breeding was in the stage described by Hindwood in November
1961, except that few nests had young. In a count of 100 nests, 45
contained a single egg and 55 two eggs. Four contained hatchlings.
Counts were made of birds on the ground at midday on 3 November in five
sample areas of the central guano flats (Figure 5). These areas
totalled 2370 sq m, or 5.9 per cent of the area of the flats. 162
birds were present, giving a mean density in the sample areas of 6.8
per 100 m?. Extrapolating these figures to the whole of the central
flats gives a total number of birds of 2764. This figure does not
include adult birds absent while feeding, birds in the air as a result
of our own presence, or birds in the high-ridge grasslands surrounding

the flats. Details of the counts are given in Table 2. Data given
by Nelson (1970) suggest a mean nesting density of 15 birds (7.5 pairs)
per 100 m?, with some colonies very much more crowded. The largest

colony recorded by Nelson is of 8,000-10,000 birds; our figures suggest
a total population half this size, and probably rather more than the
2,000 pairs estimated by Hindwood in 1961. Since our visit P. Ogilvie
(1975) has estimated the population at 2,000-3,000.

Collections were made of regurgitations produced by adults of the
gannets Sula dactylatra and S. leucogaster and the young of the lesser
frigate Fregata ariel. The diet of the gannets appeared to be chiefly
fish, with the exocoetid Cypsilurus melanocercus (Ogilvy) forming the
predominant element, but a few small individuals (mantle length less
than 10 cm) of the ommastrephid squid Symplectoteuthis oulaniensis
(Lesson)? were noted. Similar small-sized specimens of this squid
formed the bulk of diet of the young frigate birds. The importance
of both the Exocoetidae and the Ommastrephidae in diets of birds in
the Pacific has been discussed by Ashmole and Ashmole (1967).

1. The nomenclature of the forms of the 'species' Symplectoteuthis
oulaniensis remains to be elucidated. The specimens taken on Raine all
lack light organs (yellow patches) on the dorsal surface (see Clarke
1965) and thus correspond to the species A of Ashmole and Ashmole
(1967).

24

Table 2. Sula leucogaster and Sula dactylatra in sample areas of the
central guano flat, midday, 3 November 1973.

a ee eae ee
Sample area Area, sqm Sula leucogaster Sula dactylatra

Number Density/100 m? Number Density/100 m?

-_—_e_eee—_———e

A 570 40 7.018 16 2.807
B 550 32 5818 9 1636
C 460 32 62,957 5 1.087
D 390 28 Pe) 8 2.051
E 400 30 7.500 3 0.750
Total 2370 162 6.835 41 1a 730,

S——.:.:.— eeeeaeeaaaa=a=cc oa eee SS EE

The sample areas cover 5.86% of the 40,440 sq m of the central guano
flats. Extrapolation from the mean measured densities gives total
numbers of 2764 Sula leucogaster and 699 Sula dactylatra.

Sula sula rubripes Gould Red-footed Gannet
Under the heading "Sula ------ 2?" MacGillivray (1846, 1478)

described "a small species of gannet, which we named provisionally the

"white booby'. Its change of plumage are remarkable and puzzling.

Early in June, this species, the young having been for some time able to
fly, forsook the island during the day, returning at night to roost in
a large body of several hundred". Gould (1848, VII, 79) reported this
species as Sula piscator, breeding in "great numbers", and quotes
MacGillivray's estimate of a population of "several hundreds". He
also notes the variable colour. MacGillivray's specimen was later
described as Sula nicolli by Grant and Mackworth-Praed (1933),
distinguished by red feet, ash-brown upper parts, white rump and tail,
but Murphy (1936) considered it to be only a colour phase of Sula sula,
and Warham (1961) agrees. Moseley collected a female in August 1874,
and this was reported as S. piscator in Sclater and Salvin (1878, 652).
Moseley (1879, 301) described "one or two of its nests made in the
bushes, like those of the noddies, raised six inches from the ground".

More information comes from the visits by Macgillivray and M'Lennan.
In October 1910 Macgillivray (1910, 225) found "nests in groups in
different parts of the island. All of the nests are placed on the
horizontal shrubby growth, and are a clear foot or more from the ground.
The nest consists of a substantial interwoven platform of sticks, about
8 to 12 inches in diameter, depressed to about an inch in the centre for
the reception of the single egg". The young were then fully fledged
and many roosting. In July 1911 M'Lennan found "great numbers"

25

building nests, several nests with one egg each, and a few with young
(Macgillivray 1914, 148). In December 1913 a few stragglers were

still nesting. A few nests had one egg, more had young, but the rest
had left the nest. Macgillivray believed that the colour changes were
age colour phases (1917, 180-181). Other observations by Macgillivray,
with a photograph taken in October 1910, are given by North (1912, 348-
349).

Warham (1961, 80) estimated the population as 300 in February
1959). Nesting was then over; he saw one egg and two pairs with downy
young, and inferred that the breeding season lasted from June to
January. The nests were built 2 ft from the ground on what he termed
Tribulus bushes. He also found large roosts along the south side of
the island, mostly in low shrubby bushes six inches from the ground or
in thick grass. Most of the day is spent at sea, and the birds appear
during the late afternoon. Warham also describes the colour phases.
Hindwood et al. (1963, 38, 41) collected two females (C.S.I.R.O.
numbers 299, 300).

In November 1973 there were two areas with Red-footed Gannets.
One on the high ridge at the northwest end of the island, with about
50 birds during the day, consisted of nests on low Abutilon bushes
1-2 feet above the ground. Many of the nests had young in down.
Another area on the high southern ridge comprised about 100 roosting
birds during the day (Figure 5). Ogilvie (1975) estimated the
population at 150.

According to data given by Lavery and Grimes (1971) and Serventy
ebval.. (1971, 173-5). this small colony on Raine Island is) the only
breeding station for the Red-footed Gannet on the Great Barrier Reef.

Sula dactylatra personata Gould Masked Gannet

Jukes (1847, I, 128) found a few with young in July 1843.
MacGillivray (1846, 1478) reported it, as Sula personata, as "never
very numerous" and said it left the island entirely during the day.
Gould (1848, VII, 77), however, quoting Lt J.M.R. Ince, referred
(again under S. personata) to "considerable numbers". Moseley gives
no indication of abundance in 1874, but collected two females (Sula
cyanops in Sclater and Salvin 1878, 652); he described the nest as "a
circular hole in the earth, about 13 inch deep" (Moseley 1879, 301).

In October 1910 Macgillivray found them nesting "promiscuously all
over the islet" (Macgillivray 1910, 225), with fresh eggs, hatching
eggs and young (Macgillivray 1917, 180). He reported that the nests
"differ in no wise" from those of the Brown Gannet, except of course
~that there are no surrounding twigs. On 10-15 July 1911 M'Lennan found
a few nesting sites but no eggs; one egg was laid during his visit
(Macgillivray 1914, 148). In December 1913 nesting was almost over
and only one nest had eggs (S. cyanops in Macgillivray 1917, 180).
Further information is given from Macgillivray's 1910 visit by North
(1912, 344-345), including a photograph. The nest depressions were

26

said to be 3-4 inches deep and 8-12 inches in diameter. Two-thirds
of the clutches were of two eggs.

Warham (1961, 80-82) estimated the population in February 1959 as
400-500. The birds were found only in the central depression. Many
of the nests had eggs or small young, and there were also downy chicks
and flying juveniles. He inferred a breeding season from July to
March. In contrast to the Brown Gannet, very few Masked Gannets
roosted on the beaches. As with the Brown Gannet there is a sexual
difference in call, the male whistling and the female more raucous.
The nest is simply a depression in the sand, with no twigs. He noted
differential survival of the larger bird in clutches of two. During
the day many birds were at sea, and the number on the island increased
during the night. Hindwood et al. (1963, 34) estimated the population
at about 1000 pairs in November 1961.

The Masked Gannet was numerous on the central flats in November
1973, though less abundant than the Brown. Counts of birds on the
ground were made at midday on 3 November in the five sample areas shown
in Figure 5; details are given in Table 2. There were 41 birds in
the 2370 sq m sampled, giving a mean density of 1.73 per 100 sq m.
Extrapolation at this density gives a total number of 699 birds for the
whole of the central flats, or about one quarter the number of Brown
Gannets present. This figure excludes birds at sea or temporarily in
the air, or in areas other than the central flats. The total is
consistent with the estimate of Hindwood et al. (1963), also in
November, and with Ogilvie's (1975) subsequent estimate of 800-1000
bindist The mean density is close to that reported for Masked Gannet
colonies at Kure and the Galapagos Islands by Nelson (1970), and the
colony is large in terms of the size ranges Nelson cites. Of 100
nests counted in 1973, 47 contained one egg or young and 53 two eggs
or young. 35 per cent of the nests contained young. A marked size
disparity was noted in clutches of two. For notes on diet see under
S. leucogaster.

According to Lavery and Grimes (1971) and Serventy et al. (1971,
175-177) Raine Island is the main breeding station of this species on
the Great Barrier Reef, with a further large colony at nearby Pandora
Cay and another in the far south on Masthead Island. The colony is
comparable in size with the colonies on Phoenix, Enderbury and McKean
Islands in the Phoenix Group.

Pelecanus conspicillatus Temminck Australian Pelican

Ellis (1937, 178) saw one bird during 1890-92. The nearest
breeding station is on Pelican Island, 255 km to the south (personal
observation, 1973); Serventy et al. (1971, 183) also record a breeding
colony near Thursday Island, Torres Strait, 225 km to the north.

2H)

Phaethon rubricauda Boddaert ‘Red-tailed Tropic-bird

This species is generally absent from the South Pacific, and Raine
Island as its only known breeding station in eastern Australia, other
than on sand cays in the Coral Sea (Serventy et al. 1971, 160).

MacGillivray (1846, 1477-1478) recorded it as Phaeton phaenicurus
(c£. Gould 1848, VII, 73). He caught about a dozen beneath the low
cliffs; there were no nests, but two were sitting on eggs. Two or
more were always hovering over the island. There are two specimens
(10/Phae/1/C/9, 18) collected in September 1844 in the University
Museum of Zoology, Cambridge (C.W. Benson,pers. comm.). In 1874
Moseley collected a young female — "the only one seen on the island".
This was reported as P. aethereus (i.e. the Red-billed Tropic-bird, not
the Red-tailed) by Sclater and Salvin (1878, 651), and Lord George
Campbell also stated that "we saw no red-tailed ones" (Campbell 1876,
162). Ogilvie-Grant (1898, 453), however, lists the Moseley specimen
as P. rubricauda. C.W. Benson has re-examined the specimen and
comments that "this identification seems correct. AVY) Joalagel sis} aepuliby,
feathered but not fully grown. It lacks the long tail-feathers of
adults, and the bill is also smaller (it measured, from base of skull,
66 mm) and black rather than red in colour".

Macgillivray (1910, 227; 1914, 148) saw none in October 1910.
M'Lennan, however, in July 1911, found a total of seven nests, three
having one egg and the rest with young. These included three nests on
the 9th (one with 1 egg and two with 1 young), two nests on the 10th
(one with 1 egg, one with 1 young), and one nest on the 15th (with 1
egg) (Macgillivray 1914, 148-149). Macgillivray (1917, 182-183) found
them still nesting in December 1913, all except one with young. In
February 1959 Warham again found seven pairs nesting, six with large
feathered young, and the other with an egg. Up to twelve birds were
seen in the air during the afternoons. Hindwood, etal, (1963 027;
Hindwood 1964, 309) found six pairs in November 1961. Two females
were collected (Queensland Museum No. 0.9133: C.S.1I.R.O. no. 298).
Several nests were seen in 1973, either in hollows beneath the cliffs

or beneath large beachrock slabs. No eggs were found, but large
feathered young were present. Two or three birds were generally in
the air over the island. Ogilvie (1975) found 23 nests, some with

eggs, and gave the population as more than 25.
Storr (1973) lists only this species as present on Raine Island.
Hydroprogne caspia (Pallas) Caspian Tern

Recorded by MacGillivray (1846), and listed as a breeding species
by Serventy et al. (1971, 209).

Sterna bergii Lichtenstein Crested Tern

This species was not mentioned by either Jukes or MacGillivray in
1843-44, It may be the species described by Gould (1848, VII, 23) as

28

Thalasseus pelecanoides, the Torres Straits Tern, on which he quotes
MacGillivray's observation of "three small parties upon a low ridge on
one side of the island", noting that it bred in June.

Otherwise the first record is that by M'Lennan on 9 July 1911, when
he observed a small colony in the centre of the island (Macgillivray
MORIA ew Ashi Macgillivray himself found two small colonies in the
centre in December 1913. "The birds themselves were very numerous ...
they were frequently seen in the early morning carrying off turtles'
eggs from the sandy shore when these had been unearthed by the nesting
reptiles" (Macgillivray 1917, 84). A few were seen by Warham (1961,
87) in February 1959, but were not breeding; and none were seen by
Hindwood et al. (1963, 38) in November 1961.

Lavery and Jones (1971) map large colonies of this species on
Michaelmas and Upolu Cays, near Cairns, and on the Barnard Islands;

Serventy et al. (1971, 219) record colonies in the Bunker and Capricorn
Groups.
Sterna fuscata nubilosa Sparrman Sooty Tern

The first record at Raine is that by Jukes in July 1843, when he
found young of all ages (Jukes 1847, I, 129). In May 1844 MacGillivray
(1846, 1476-77), under the name Thalassipora fuliginosa, found it
breeding in large colonies, on the ground, and said that many eggs were
taken. Gould (1848, VII, 33) refers to it as Onychoprion panaya.
Moseley in August 1874 found it breeding in grass around the shore;
Tizard et al. (1885) describe it as "exceedingly abundant" on "flat
ground above the shoreline covered with grass". "The stretches of
flat ground above the shore were absolutely full of the brown fledged
young of this bird: Eggs were already very scarce" (Moseley 1879,

SOP )F Saunders (1878) records Challenger material.

Macgillivray (1910) recorded Sooty Terns in October 1910, and in
July 1911 M'Lennan found them (reported as Onychoprion fuliginosa)
"nesting in tens of thousands all over the island" (Macgillivray 1914,
142). In December 1913, Macgillivray (1917, 85) found them coming
in from the sea at about 5 p.m. "in immense numbers", circling
without landing, then leaving again at dawn, presumably as a preliminary
to breeding. Sooty Terns are also recorded for Raine by Alexander
(i925)s:

In February 1959 Warham (1961, 87-88) found "vast numbers" on
the dunes, especially in the south. The birds were never seen on the
ground in 1959, though their numbers increased at night. Warham
stated that this species breeds between April and November, and is
present but does not alight between December and March. Hindwood
et al. (1963, 38) found in present in November 1961. In November
1973 Sooty Terns were abundant on the ground in Lepturus grassland on
the high ridge, especially towards the west end, during the hours of
darkness, but they were absent during the day.

29

According to Lavery and Grimes (1971) and Serventy et al. (1971,
227), Raine Island is the major breeding site of Sooty.Terns on the
Great Barrier Reef, though there are several other large colonies
north of Michaelmas Cay, a distribution very similar to that of the
Noddy Anous stolidus.

Sterna anaethetus anaethetus Scopoli Bridled Tern
Reported by Macgillivray (1910) and Macgillivray (1917, 85), who

recorded this species (as Onychoprion anaestheta) nesting in "great
numbers" in crevices and on piles of rock in the centre of the island

in December 1913. Warham (1961, 88) found very few in February 1959
and thought they had already left the island. They are said to breed
under the cliffs. Lavery and Grimes (1971) state that Raine is the

major breeding site for this species on the Great Barrier Reef, but
there are many lesser sites along the Reef (Serventy et al. 1971, 229).

Anous stolidus pileatus (Scopoli) Noddy

Jukes (1847, I, 129) found young of all ages in July 1843. aTEyal
May 1844 MacGillivray (1846, 1476-77) reported "myriads", building
nests of small twigs and coral fragments. In August 1874 Moseley
(1879, 300) found them breeding in grass around the shore, building "a
rude nest of twigs and grasses amongstthe low bushes, but often also ...
on the ground. There were plenty of eggs of this bird"; specimens
were taken (Saunders 1878).

In October 1910 Macgillivray (1910, 226) found "noddies ... in
great numbers, but ... not yet commenced to nest". In July 1911 they
were "nesting all over Raine Island" (M'Lennan in Macgillivray 1914,
142). In December 1913 they were "in great numbers, lining the
seashore of the island or in small colonies all over the island, mostly
composed of fully-fledged young and their parents. Many more were
continually over the sea ... and towards nightfall many morecame in to
roost on the island" (Macgillivray 1917, 85).

Warham estimated the population at 2000 in February 1959. Birds
were present during the day, congregating on the northern beach during
the afternoon, and many coming in from the sea at dusk. There were |
dense flocks on the central flats during the night. Warham thought
they were collecting prior to nesting (Warham 1961, 88). Hindwood
et al. (1963, 38) found them present in November 1961, and Ogilvie
(1975) found some 400 birds.

Lavery and Grimes (1971) and Serventy et al. (1971, 232-233) cite
Raine Island as a major breeding site for this species, together with
several other islands on the northern Great Barrier Reef; this
distribution is similar to that of the Sooty Tern.

30

Anous minutus minutus Boie White-capped Noddy

This species is recorded as Anous leucocapillus, White-capped Tern,
by Gould (1848, VII, 36), who quotes Lt Ince's observation that it is
"very abundant" at Raine Island; this observation probably refers to
the Common Noddy. There is no further record of this species at
Raine until Warham's visit in February 1959. He (1961, 88) found two
or three, and since this species nests in trees, he did not believe it
bred on Raine. Hindwood et al. (1963, 38, 42) collected a male in
November 1961 (Western Australian Museum No. A8768).

All the large colonies of this species on the Great Barrier Reef
are in the Bunker and Capricorn Islands in the far south (Lavery and
Grimes 1971; Serventy et al. 1971, 235-237). Its status on Raine
Island remains uncertain.

Larus novaehollandiae forsteri (Mathews) Silver Gull

This may be the species recorded as Xema Jamesonii in 1844 by
MacGillivray (1846, 1477), who noted "only a few pairs". Moseley
(1879, 300) collected it in August 1874 (Saunders 1878), but said it
did not breed. In December 1913 Macgillivray (1917, 86) found it "in
fair numbers ... constantly to be seen early in the morning on the
sandy strip above high tide mark on the look-out for turtle eggs that
had been scooped out by these creatures during the night". In
February 1959 Warham (1961, 88) found small numbers and no nests, and
noted that there was no breeding record from Raine; he estimated the
population to be 25. Hindwood et al. (1963, 38: Hindwood 1964, 310)
found "a few birds", and Ogilvie (1975) 3-4. This species is very
widely distributed along the Great Barrier Reef (Lavery and Grimes
1971; Serventy et al. 1971, 196), but is unimportant at Raine Island.

Arenaria interpres interpres (Linnaeus) Turnstone

This species was recorded from Raine Island in 1844 as Strepsilas
australis by MacGillivray (1846) and as Strepsilas interpres by Gould
(USASi Vai es9) i. Moseley (1879, 300) found flocks on the shore in
August 1874, but no nests; one male and two females were taken and
reported as S. interpres by Forbes (1878). Macgillivray (1971, 86)
found several small flocks on the shore in December 1913. Warham
(1961, 88) found small flocks everywhere except in the vegetated areas
in February 1959. It was noted as present by Hindwood et al. (1963,
38) in November 1961.

Pluvialis squatarola (Linnaeus) Grey Plover
The only record is that of a female collected by Macgillivray on

10 December 1913 and reported as Squatarola helvetica (Macgillivray
L917 5), 86)y

Shi

Pluvialis dominica fulva (Gmelin) Eastern Golden Plover

This species was "many times noted in all parts of Raine Island"
by Macgillivray (1914, 86) in December 1913. Warham (1961, 88) found
20 present on the beach, dunes and central flat in February 1959.
Hindwood et al. (1963, 38) report it not present in November 1961, but
in the same paper record a specimen collected there (Hindwood et al.
1963, 42), a male now in the Western Australian Museum (No. A.8754).

Numenius phaeopus variegatus (Scopoli) Whimbrel

The only record is of a single bird seen by Warham (1961, 89) in
February 1959. It was not seen by Hindwood et al. (1963, 38) in
November 1961.

Limosa lapponica baueri Naumann Bar-tailed Godwit

The only record is a sighting by Hindwood et al. (1963, 32) on
11 November 1961.

Calidris canutus (Linnaeus) Lesser Knot

Amiet (1957, 252) mentions an earlier sighting of this species "in
the vicinity of Raine Island". Kikkawa (1976, 319) lists the species
for Raine from this record.

Calidris ruficollis (Pallas) Red-necked Stint

This is probably the species recorded from Raine as Schoeniclus \
albescens by Gould (1848, VI, 31) and as Actodromas australis in Gould
(1865, II, 257) (for synonymy see Sharpe 1896, 545). Four were seen
feeding along the shore by Hindwood et al. (1963) in November 1961, and
reported as Erolia ruficollis. One male and one female were collected
andwarer an jhe Gaser RO. colilection! (Nos, 302)7,) 303). |

Calidris acuminatus (Horsfield) Sharp-tailed Sandpiper

Macgillivray (1917, 87) (under Pisobia acuminata) found it "common"
in December 1913. It was not present during Warham's visit in February
1959 (Warham 1961, 89).

Egretta alba modesta (Gray) White Egret

Warham (1961, 89) saw one bird on the south ridge on 13 February
11959):

Egretta sacra (Gmelin) Reef Heron

This was recorded as "Erodias, two species" by J. MacGillivray
(1846, 1477) in 1844. Both phases were present: "Some white and blue
herons frequented the reef, and probably are permanent residents,
judging from some deserted nests and fragments of eggs which I saw".
Gould (1848, VI, 60) cites it as Herodias jugularis Blue Reef Heron,

32

and notes that it "breeds among the recesses of the rocks". William
Macgillivray (1914, 145) saw one bird in 1911. It was not seen by
either Warham (1961, 89) in February 1959, or by Hindwood (1964, 311)
in November 1961, or by ourselves in November 1973.

Nycticorax caledonicus hilli Mathews Nankeen Night Heron

The first record of this distinctive species is M'Lennan's record
of "many" on 15 July 1911 (Macgillivray 1914, 146); he found no nests
but two young birds came into his camp. Macgillivray found one egg on
9 December 1913. Attenborough recorded one or two juveniles, unable
to nEllys pein ouiliys NO Sii By contrast Warham (1961, 80, 89) found "well
over a thousand" in February 1959, with several hundred clutches of
eggs on ridges, under overhangs, and on piles of rocks on the central
flat; there were few eggs. Warham considered that this species
occupied the niche on Raine that elsewhere was occupied by the Reef
Heron. But in November 1961 Hindwood et al. (1963, 33) found only
"about 50 birds" and "no signs that they were breeding". In November
1973 there was one large flock of about 150 birds, generally on the
beach at the east end of the island. Ogilvie (1975) later found more
than 2000 birds.

Merops ornatus Latham Australian Bee-eater
First recorded as a visitor by Ellis (1937, 178), who saw "quite a
number" during 1890-92. Ten were seen in two groups on 11 November
1961 by Hindwood et al. (1963, 7).
Hirundo nigricans nigricans Vieillot Tree Martin
First recorded by John MacGillivray in 1844. A few birds were
seen by Warham (1961, 89) in February 1959 and recordedas Hylochelidon
nigricans. It is recorded as Petrochelidon nigricans (Vieillot) by
Kikkawa (1976, 332).

Monarcha melanopsis (Vieillot) Black-faced Flycatcher

This is cited by Macgillivray (1918) and recorded by Kikkawa
(L976 nS S38)

Myliagra rubecula rubecula (Latham) Leaden Flycatcher
According to Warham (1961) this is the species cited as Rhipidura

by John MacGillivray in 1844. One was seen by W. Macgillivray (1918,

198) on 10 December 1913, and two the following day, all in the Beacon.

Aplonis metallica metallica (Temminck) Shining Starling

One bird was seen on the Beacon by W. Macgillivray on 10 December
LOA 35.

33

DISTURBANCE BY MAN

When the Fly arrived at Raine there was no trace of native
occupation (Jukes 1847, I, 130), and there is no reason to suppose that
the island had ever been occupied before this time. During the
nineteenth century, however, considerable disturbance took place, first
during the building of the Beacon in 1844, most severely during the
guano digging period before 1892, and to an unknown extent during
visits by beche-de-mer fishermen in the 1870s and 1880s. Accessible
food resources such as nesting turtles and their eggs, and ground-
nesting seabirds and their eggs, must have undergone severe if
spasmodic interference at these times. In addition to this direct
predation, some of these visitors introduced plants, mainly vegetables
which they attempted to cultivate, and on at least one occasion goats
were liberated too.

During the Fly's first visit, Jukes dined "upon young boobies and
frigate-birds and terns' eggs — the latter were excellent, and the
former very good, especially when cooked with a little curry powder"
(LB47 5 stp LZ9SNSO)) Ss In 1844 the island was occupied for four months
during the building of the Beacon. Jukes estimated in June that 3000
young birds (Noddies and Sooty Terns) and about 17,000 eggs had been
eaten by the shore party. Later, when a small party landed from the
Bramble in April-May 1845, Sweatman (MS, 93) records that 36 dozen
eggs a day were consumed by three men: "add to this turtle soup, eels,
mutton birds (which Clark cooked magnificently) fish, spinach and
damper ..." As is still the practice in egg collecting, Sweatman states
that "Our first act on landing was, of course, to break every egg on
the island so that all we collected afterwards were sure to be fresh,
and so abundantly did the birds lay that besides supplying our own
wants at the rate above stated we lined in three large casks for the
use of the ship". Gardens were also established in the centre of the
island by the Fly party, and coconuts, pumpkins, maize and other plants
introduced and cultivated (Jukes 1847, II, 267).

In January 1845 (Sweatman's manuscript says February), during the
Heroine's visit, Mackenzie (1845, 494) planted "cocoa-nut and various
other seeds, hoping they might be a benefit to some unfortunates
hereafter"; four months later the coconuts were reported "growing
very fast" (Anon. 1846, 549). Sweatman (MS, 91) then described them
as four feet tall but choked with weeds which had already obliterated
the paths and gardens. The Bramble party cleared the weeds from round
the young trees, built fences around them, and planted four more brought
from the Murray Islands (Sweatman MS, 96), but in spite of this these
introductions did not long survive. When the Challenger called in
1874, Moseley (1879, 300) planted pumpkins, tomatoes, water melons,

- Cape Gooseberries, and Capsicum, but no later visitor mentions any of
these surviving either.

Goats were introduced at an early stage, according to Lack (1953,
41) by the Fly, though there seems to be no comtemporary evidence that
this was so. In January 1845, however, the Heroine "left a male and
female goat, with two bags of rice" (Mackenzie 1845, 494). Sweatman

34

(MS, 92) in April found them "frisking about the island in good
condition but very wild and shy", and by August there were three young

(Anon. 1846, 549). There is a mention of "numbers" of goats being
present at the time of the wreck of the Enchantress in July 1850, but
otherwise no later record. The goats were doubtless eaten during the

beche-de-mer and guano digging periods, if they had not died out before,
and they were themselves probably responsible for the disappearance of
the Fly's vegetable gardens.

Early visitors continued to obtain food supplies from the seabirds
and turtles. The Heroine in 1845 "obtained fourteen large turtles ...
also an immense number of eggs, and the crew killed birds out of
number" (Mackenzie 1845, 494). The Bramble party also took fourteen
turtle by turning them on the beaches (Sweatman MS, 94-95). Buchanan
(1874, 127-128) records that the Challenger only left "after a
sufficient number of birds had been killed". Similar incidents,
though not recorded in the nautical or scientific literature, must have
been numerous throughout the nineteenth century.

Construction of the Beacon

A series of shipwrecks on the Great Barrier Reef near Great
Detached Reef (including those of the Charles Eaton in 1834, the
Ferguson in 1840, and the Martha Ridgway in 1841) led to a proposal
by Capt Blackwood? that a Beacon be erected on Raine Island to guide
ships using the Outer Passage route through the Barrier Reef. The
Beacon was originally to be triangular in plan and 60 ft high, but as
announced by Blackwood (1844b) it was proposed to build a tower 50 ft
high, 25 ft in diameter at the base, and 16 ft in diameter at the top,
and to make it more conspicuous by painting it with black and white
bands each one third the height of the building.

Blackwood fitted out for this expedition in Sydney, obtaining
"twenty picked convicts, chiefly masons and quarry men", and taking on
stores and prefabricated wooden huts. The captain had to spend
£180.14.5d on mason's tools since both the Government and the Engineers
Department refused to supply them. The Fly, with the Bramble and the
Prince George, sailed on 27 March 1844, and commenced landing on Raine
on 27 May. Jukes wrote to his sister Amelia on 26 May: "Our lumber
consists of twenty convicts, and an immense quantity of plank timber,
houses, barrows, jumpers, pickaxes, spades, and all manner of building,
digging, and blasting materials. All these will have to be carried in
boats through a heavy sea six miles, as the ship cannot anchor nearer

1. This account is based on the Captain's letters from H.M.S. Fly,
Blackwood's Remark Books, and the engineering drawings, ink sketches

and watercolour drawings in Sketches and views, vol. 6B, in the archives
of the Hydrographic Department, Royal Navy, as well as on published
accounts. We are grateful to the Hydrographer, Admiral D.W. Haslam,
for access to the records.

85

Raine's Islet than that. I expect this will take us two or three
weeks, as twenty of our crew are to be landed, and provisions and water
for forty men for three months are no trifling matter. The bread alone
will be 2500 lbs" (Jukes 1871, 224). On 29 May he wrote again, after
his first landing, to report that he had "got on board just now wet
through, and so tired I can hardly keep my eyes open. ... We have
terribly hard, heavy, disagreeable work landing things on this little
desolate islet, but hope finally to accomplish our object in building

a beacon pretty well" (Jukes 1871, 225).

The huts and tents were erected, and a quarry was opened at the
east end of the island. Jukes reported to his aunt on 21 June that
"Good building stone is procured in abundance, wooden houses and tents
are erected,lime is made, the foundation stone laid, and the first
course of masonry nearly complete. lLieutenant Ince, with a party of
forty men, occupies the island and superintends the work" (Jukes 1871,
DNS) Lime was made on the island by burning Tridacna and Hippopus
Shells. Water, however, had to be brought from Sir Charles Hardy's
Islands, and timbers were obtained from the wreck of the Martha
Ridgway 40 km to the south. After his first period of investigations
on Raine, Jukes found the routine tedious: he told Amelia that "This
period of inaction is becoming most oppressive to me. I would welcome
any danger or any hardship even that would break the monotony. I am
accordingly as dull, heavy, stupid, and spiritless as it is possible"
Gukes; 1871, 231).

But the shore party under Ince had the Beacon finished by mid- .
September. It had been designed by the Fly's carpenter, Stephen
Moore, — 'without any exception the best Ship's Carpenter I ever met',
Blackwood called him — and was described by Jukes as a circular tower
of stone, 40 ft high, 30 ft in diameter at the base, and with walls
5 ft thick, "divided into three stories, each of which was partially
floored, and made accessible bya ladder. It was roofed at the top by
a dome-shaped frame of wood, covered by painted canvas. Its summit
was thus raised 70 feet above low water mark. A large tank taken
from the Martha Ridgway was placed at the side, into which a series
of spouts were led from the roof, so that it would shortly be filled
with rainwater" (Jukes 1847, I, 267). The drawings for the Beacon,
in the Royal Navy's Hydrographic Department archives (Figure 10),
indicate that the foundation stone was 11 ft above low water mark, the
height to the top of the stone balustrade was 45 ft and to the top of
the canopy 63 ft 2 inches, giving an elevation overall from low water
mark of 74 ft 2 inches. It was painted in black and white vertical
stripes and was visible from thirteen miles. Blackwood formally
reported on the work by letter to the Admiralty from Surabaya on
26 October 1844.

Four months after it was completed, the Beacon was inspected by
the Heroine: "At daylight we went on shore to examine the beacon,
when we likewise found two wooden built houses and an oven. The
beacon is sixty feet in diameter, and about fifty feet high, and its
walls at the base three feet in thickness, painted red on the south-
east side, and the rest black, with a white cupola and black ball,

36

having a spout which runs off it into a deep tank, which we found full
of water. Altogether it is a substantial building, and well contrived
thoughout" (Mackenzie 1845, 494). Later the same year the same ship
"sent a boat ashore to leave a letter in the Post-Office" (since Raine
at this time, as Sweatman (MS, 96) also tells, performed the same kind
of function as the better known Booby Island in Torres Straits at a
later date), and found that "the tank was full of water, and the beacon
seemed to stand the weather very well" (Anon. 1846, 549). The Beacon
itself was visible from 8-9 miles from the deck of a ship and from
12-13 miles from the masthead (Horsburgh 1855, 883-884; Horsburgh 1852,
L, /98=800;" Findlay” 18847.971))-

When examined by Capt Denham in 1860 the dome had fallen, reducing
the height to 40 ft (Findlay 1884, 971), and the beams and floors had
gone and the iron tank disintegrated by the time of the Challenger's
visit in 1874 (Tizard et al. 1885; Swire 1938, II, 47-50). The ship's
plumber was sent ashore to cut the Captain's name and that of the ship
in the stone inside the tower (Swire 1938, II, 50), the whole surface
now being covered by overlapping inscriptions by casual visitors.

In 1959 Warham found that part of the base of the tower was
starting to collapse (Warham 1963, 5, photograph), but this damage was
repaired with bags of cement by H.M.A.S. Gascoyne in 1961. The Beacon
is of some historical importance, since it is the oldest stone structure
built by Europeans in North Queensland.

Wreck of the Enchantress

Lack (1953, 41) writes: "Five years later [after the Beacon was
built], Captain Anson of the brig Enchantress, Sydney to Sourabaya,
managed to put his ship ashore on the northern side of the Raine
Entrance despite the warning of the tower. A young passenger named
Buchan wrote a racy account of the incident to his relatives in England.
He related how they landed on the island, found numbers of the much
enduring goats, and quantities of sea birds' eggs on which they feasted
sumptuously". The crew rapidly became drunk and were taken off by a
second vessel, the Lady Margaret, only with difficulty.

Efforts to obtain details of this wreck and to trace the narrative
account mentioned by Lack have not been altogether successful. We are
indebted to Mr. H.E. Maude who has shown that the captain's name was
I'Anson, not Anson, and the passenger was Mr. B. Buchanan, not Buchan.
Maude notes that the Shipping Gazette and Sydney General Trade List,
vol. 7, nos 330 (13 July 1850), p. 186, records the departure of the
Enchantress in ballast from Sydney. The same journal, vol. 7, no. 351,
p. 322, reports its loss on the Raine reef on 24 July 1850, and prints
a letter from Buchanan giving the details to the owners of the ship.
The wreck occurred in the afternoon, and the ship's company escaped
and took shelter in the Beacon. The Enchantress was travelling with
the Lady Margaret, and everyone was taken off in small boats and
transferred to this ship by dusk on the following day. The details
given in Lack's published account do not derive from this letter by
Buchanan, and all efforts to trace the "racy account" Lack refers to

37/

have failed. The main interest of it lies in the references to goats
and to predation of seabird eggs which it evidently contains. There
is also an account of the wreck in the Sydney Morning Herald for 3 and
4 December 1850.

Beche-de-mer Fishery

During the 1880s Cooktown was the centre of the Queensland beche-
de-mer fishery, when holothurians were collected from shallow reef
flats, dried, and exported to China (Ward 1972). This trade had been
carried out at Raine Island at least since the early 1870s. Ellis
(1937, 71-72) recounts that about 1873 the fishery there was carried
out by two white and two or three natives (Chinese, kanakas and

aboriginals). The beche-de-mer were boiled, sun-dried and smoked on
the island, though in the absence of fuel this can hardly have been on
a very large scale. According to Ellis the aboriginals on one

occasion murdered one of the whites, one Chinese and two kanakas,
before fleeing from Raine; the bodies were buried by the surviving
white man near the centre of the island.

Phosphate mining

When phosphate was first discovered in commercial quantities on
Raine Island is not known. It was first mentioned when Raine, with
other islands, was included in a lease taken out in Hobart in 1862,
for seven years, for the purpose of guano digging (Crowther 1939).
There is no evidence that any digging under this lease took place at
Raine, although the Arundel papers contain an undated newspaper
cutting stating that "The Griffin, under the command of German Harry,
cleared yesterday for Rain Island under charter to a Southernspeculator,
who holds the right to take guano from that locality". Most of the
guano digging activity at that time was in the more accessible Bunker
and Capricorn Islands.

The J.T. Arundel Company, which had been digging on the central
equatorial Pacific islands during the 1880s, transferred its operations
from Baker to Raine in 1890. John Arundel records in his diary that
before starting work he consulted the master of the Jennie Scott, One-
eyed Robinson, who had worked on Raine in 1879 (though on what is not
known), about sources of water there and on neighbouring islands.
One-eyed Robinson told Arundel of the murder of twelve men by natives
on the island during the beche-de-mer fishery. Arundel went to Raine
with the Griffin and the Maile in August 1890 to organise the digging.
His diaries show that he was on the island from 20 August to 6 September
1890 and again from 13 February to 15 February 1891. The digging
operations were conducted by Albert Ellis, who also arrived in 1890
(though Saville-Kent (1893) suggests that extraction began as early as
1882). There was a staff of 9-10 Europeans, and one hundred Asian
(mainly Chinese) labourers; Arundel, a religious man, shipped Chinese
Bibles to the island for their use.

38

A tramway was built from the workings to a new jetty, anda
"locomotive" imported. A storage shed was built to house the guano
between shipments. Export was by sailing ships of 1000-1500 tons,
direct to Europe, with the exception of two which went direct to
Melbourne. The guano was packed in 60 lb sacks for ease of handling.
Water supply for so large a labour force was a major problem, anda
water condenser was built for this purpose. In 1892 the digging ended
and the equipment was dismantled and removed (Ellis 1937, 62-73, 98).
It is estimated that "tens of thousands of tons" of guano were exported
during this brief period (Hutchinson 1950, 256).

Sir Albert Ellis's mother died on Raine Island during this period,
and is buried in a substantial grave with a marble stone adjacent to
the Beacon. The inscription on the stone ends with a homily which seems
especially appropriate in so remote a location:

In loving memory
of
Annie Eliza
wife of
George C. Ellis
entered into rest
June 29th. 1891
aged 52 years
Her last words were
Father! not my will
But thine be done.
My i= iGod!i=xo£ =yhove
Reader!
Be ye also ready

ACKNOWLEDGEMENTS

This report results from a visit to Raine Island made during
Phase III of the Royal Society and Universities of Queensland Expedition
to the Great Barrier Reef in 1973. We thank the sponsors of the
expedition, and especially James Cook University of North Queensland,
which made its research vessel James Kirby available for this visit.

We are particularly grateful to C.W. Benson, Department of
Zoology, Cambridge University, for advice and comments on the birds of
Raine Island and for re-examining Raine Island specimens in the British
Museum (Natural History)and in the University Museum of Zoology, Cambridge;
and to C.J.O. Harrison, S.A. Parker, and H.J. Lavery for other comments
on birds. We are grateful to Dr J. Kikkawa for making available
unpublished material on his surveys of birds on the Great Barrier Reef,
and for sending data collected by P. Ogilvie (1975), which Mr. Ogilvie
has allowed to be included. Dr M.R. Clarke and Dr A. Wheeler

determined squid and fish respectively from seabird regurgitate. For
determinations of Crustacea we thank Dr R.W. Ingle, British Museum
(Natural History). Mr E.I. Butler, Marine Biological Association,

Plymouth, kindly carried out phosphate analyses of soil samples.

39

Professor H.E. Maude of Canberra helped greatly with the enquiry
about the Enchantress, and Dr Harold Fox pursued various lines of
enquiry in Cambridge and London. Dr J. Allen of Canberra kindly made
available the relevant portion of Sweatman's journal, through Dr Roger
McLean, and has allowed quotations to appear here. Mr R.A. Langdon of
the Pacific Manuscripts Bureau, Canberra, made available copies of the
Raine Island sections of J.T. Arundel's journals. Admiral Haslam
allowed access to papers of the Fly survey in Hydrographic Department
archives. Dr S. Everist, Director, The Queensland Herbarium,
Brisbane, arranged for Miss S. Reynolds to identify the plants
collected in 1973, and this material was also examined by Dr F.R.
Fosberg. Dr T.P. Scoffin, Grant Institute of Geology, Edinburgh
University, examined rock samples.

Mr M. Young and Mr R. Coe of the Department of Geography,
Cambridge University, prepared the maps and the photographs respectively.
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43

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44

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Island Sandbank, North Queensland. Emu’, “612 = 77-93.

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LOvC1s) se 2—9

Whitley, G.Ps. Loser John Gould's associates. Emu, 38: | 141-1672

Wildy credit 187 8x At anchor: a narrative of experiences afloat and

ashore during the voyage of H.M.S. "Challenger" from 1872 to 1876.
London: Marcus Ward and Co. 196 pp.

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b. Little cliff at edge of stone.
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10 feet.

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2015.

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Raine Island in 1844, from Jukes (1847).

The central guano flat from the est end of the island, look-
ing towards the Beacon. The ridge in the foreground is the
SiGevOn the olld euano) raqiway .

yemeae lata

Pl. 3. Eastern end of the central guano flat, with Brown and Masked
Gannets.

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Pl. 4. Mounds of rubble and stones in the central guano flat.

Pl. 5. Massive beachrock on the northeast shore.

i

on the northeast shore.

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beachrock at the east end of the island.

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Pl. 10. Irregular lower surface of the phosphate rock, forming caves,
on the south side of the island. Note the nesting Tropic-—
bird beneath the overhang.

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4
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island.

on the beach crest at the west end of the island

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rtle remains in Lepturus gr

Pl. 17. Shearwater burrows in fine guano at the west end of the
central guano flat.

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birds.

Pl. 19-20. Juvenile Lesser Frigate-birds.

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Pl. 21. Juvenile Lesser Frigate-birds.

Pl. 22. Brown Gannets lining the beachrock at the east end of the
island.

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Pl. 23. Brown Gannets on beachrock on the northeast shore.

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Note the twigs outling the nest.

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Pl. 25. Brown Gannets nesting in the Lepturus grassland of the
high ridge. No twigs surround the nest in the vegetated
areas.

Pl. 26. Masked Gannets on the nest in the central guano area.
The nest is simply a depression in the sand.

Pao 20.

Masked Gannets on the nest in the central guano area.
nest is simply a depression in the sand.

Hatchling Masked Gannet in the central guano area.

et

29.

Pl. 30. Red-footed Gannets on nests built on low Abuttlon shrubs on
the high ridge at the west end of the island.

ae oe oe
Fé Dy) a ne i »

Pl. 31. Red-footed Gannets on nests built on low Abuttlon shrubs on
the high ridge at the west end of the island.

- Fledgling Red-footed Gannet on the nest at the west end of
thé: island.

as Qo MA yee We rae
Juvenile Red-tailed Tropic-bird under the phosphate cliffs
at the east end of the island.

IPL 3336

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x

Pl. 34. The Beacon and grave at the east end of the island, seen
from the south in November 1973.

' Sh. Zagee Ol (Sig QE) eines qi “T96T UT SuAOCDSeD “S°Y°W°H Aq poatedeaz jeua ST TTeM
‘aurenboy ]oo1qnby Wors “HHL UT pejJosde se Uodweg ey, “GE *Tq ZAMOT SY} UO PoTe yTeP SUL “E61 FSQUBAON UT UCORSg SUL °9E “Td

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Pl. Sf. Inscriptions inside the walls of the Beacon.

of Annie Eliza Elias.

ATOLL RESEARCH BULLETIN
NO. 255

SHORT ORIGINAL ARTICLES

by various authors

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

July 1981

BD EOP Ras oN One r

In line with our policy of not issuing short articles as sepa-
rate numbers with their own title pages, the following articles are
offered as parts of a single number. They have accumulated over a
considerable period since the last Island News and Comments appeared.
They are grouped roughly according to their geographic and subject
interest.

Contents

An early report of the flora and fauna of the Aldabra Page
group, by E. P. Diamond il!

Geochemistry and mineralogy of carbonate rock samples
from Aldabra Atoll, Indian Ocean, by Stephen T. Trudgill 11

History of goats in the Aldabra Archipelago, by D. R.
Stoddart CB

Abbott's booby on Assumption, by D. R. Stoddart Pak

Penetration of host plant tissues by the stylets of the
ecoccoid Icerya seychellarum (Coccoidea: Margaroidea)
on Aldabra Atoll, by S. Blackmore 3355

Meteorological data from Ulul Island, Namonuito Atoll
by John Byron Thomas and Mary Durand Thomas 39

Marine benthic algae of Kayangel Atoll, Palau, by

Roy T. Tsuda 43
Qualitative assessment of asteroids, echinoids and

holothurians in Yap Lagoon, by Deborah A. Grosenbaugh h9

Acanthaster in the cultures of high islands, by
Charles Birkeland Dp)

Acanthaster as.a recurring phenomenon in Samoan history
by John M. Flanigan and Austin E. Lamberts 59

Distribution and abundance of the Crown-of-Thorns Starfish
(Acanthaster planet) around Tongatapu Island, Tonga,
by M. P. Francis 63

Rats as avian predators: discussion, by W. R. P. Bourne 69

Layard's bird hunting visit to Tromelin or Sandy Island in
December 1856, by R. K. Brooke 3

A submersible, rechargeable, electric drill, by W. H.
Easton 83

Artificial reefs in Discovery Bay, Jamaica, by
Michael J. Risk 91

Distribution of the decapods Brachyura and Anomura
(excluding Paguridea) of the Cryptofauna in the
reefs near Tulear, by Mireille Peyrot-Clausade 101

AN EARLY REPORT OF THE FLORA AND FAUNA OF THE ALDABRA GROUP

by E.P. Diamond!

Introduction

Among the records in the National Archives of Mauritius at Port
Louis is a copy of the earliest known detailed report on the flora and
EaunasOr) the Aldabra group, made by Sergt: F. Rivers o£ the Seychelles
Constabulary. It was sent by the Civil Commissioner for the Seychelles
to his superior, the Governor of Mauritiuson December 30, 1878. The
original was forwarded to London and a copy kept in the Seychelles
National Archives and another in the Mauritius Archives. A third copy
is to be found in the Glinther Papers in the British Museum (Natural
History) although I have not seen it. A later letter in the Mauritius
Archives suggests that another, more detailed report by Rivers may have
been made and sent to the Royal Society sometime in February 1879,
though I have been unable to trace it. The importance of this report
to the study of the changes on the Aldabra group lies in its early date
and the apparently knowledgeable and careful observation of the
reporter.

The report is dated 11 December 1878 and is signed by F. Rivers
who is said, in a covering letter, to be of Canadian extraction and to
have been a seaman before settling in Seychelles. The report is
English with creole names given for most of the plants and animals.

I have written the English common name, if there is one, in parentheses
after the creole one and, with the first reference of each species, have
included the scientific name as well. There may be some confusion in
the use of creole names, which will be discussed in the conclusion.
Detailed information about anchorages and geological formation of the
islands will not be included in this paper but can be found in the
report itself.

Astove

Rivers arrived on Astove on 10 October 1878. He reported that he
could find no traces of visits by fishermen and whalers, though there
was a well in the southeast which had been dug by a shipwrecked crew
"some years ago'.

lc/o Department of Zoology, University of Nairobi, Nairobi, Kenya
AtolleRes..( Bull) Noss255% N=10,- 198i):

2

Coconuts and Other Vegetation

Rivers reported that the island had no trees except a few stunted
‘mangliers' (mangroves of any species) and was covered with bush. He
mentioned two individual coconut trees, planted in recent years, and
implied that few if any others had yet been planted.

Fish

Rivers reported that fish ‘are not found in quantity', but that
the most common were; 'varvaras' (Lutjanus bohar), ‘capitaines rouges'
(Lethrinus kallopterus or L. nebulosus), 'carangues' (Alectis indicus or

Caranx sp.) and 'vielles' (Plectroponus maculatus or a serranid).
[These and all other translations of creole fish names are taken from
J.L.B. Smith and Mary Smith Fishes of the Seychelles, 1969. ]}

Turtles

In Seychelles creole, a green turtle is a 'torti' while a Hawksbill
turtle is called a 'caret". When translating 'torti' into English,
Seychellois often use only the word 'turtle' without the 'green'. Thus

when Rivers uses the word turtle alone he means only green turtles,
which is made clear by the context in the full report.

'Turtles' (green turtle, Chelonia mydas) were said to be very
abundant, although it was not the breeding season and Rivers estimated
that his ship, probably a small schooner, could take a complete cargo
of turtles in three days. He reported finding 12 young turtles in the
belly of a 'varvara'. Frigate birds (Fregata sp.) and ‘aigrette'
(egrets) were also said to prey on young turtles but no evidence was
given.

Birds

Aquatic birds were said to be few with frigates and 'aigrette'
present in small numbers. "Corbijeaux' (whimbrel, Numenius phaeopus)
and 'allouettes' (small waders and shore birds) were said to be
plentiful. Of the land birds, a 'ralle' (rail, probably Dryolimnus
Cuvieri) was mentioned, also the 'crow of Madagascar' (pied crow,
Corvus albus) a small species of 'colibri' (sunbird, probably Necterina
sovimanga) and a 'cardinal' (Foudia sp. with red plumage). A 'merle'
(bulbul, Hypsipetes sp.) and a ‘pigeon hollandais' (Alectroenas sp.)
were said to be present and 'smaller and paler than those found at
Seychelles'.

Other animals
Land crabs were said to be plentiful and 'cypaye' (coconut crab,
Birgus latro) was mentioned by name.

Discussion
Trees and other vegetation
Rivers describes the island as covered in bush with a few stunted

mangroves, which coincides with the description of Bayne et al. (1970b)
generally, although a large number of planted coconuts had changed the
appearance of the island by 1968. It is interesting to note, however,

that Rivers did not mention Casuarina woodland described by Bayne et al.
on the western rim of the island. Fosberg and Renvoize (1970) record a

report of its presence in 1919, suggesting that these Casuarina were
introduced between 1878 and 1919.

Marine Fauna
Bayne et al. do not mention fish but agree with Rivers that Astove

is a major breeding ground for green turtles. The presence of young
turtles in a fish's stomach implies that there had been laying two to
4; months before (late July or early August). Hawksbill turtles

(Eretmochelys imbricata) are not mentioned by Rivers and are said to
be rare by Bayne et al.

Land fauna other than birds
Giant tortoises (Geochelone gigantea) reported by Rothschild on

Astove in 1915 were not recorded by Rivers in 1878. This is perhaps a
reflection of the low numbers of the population at that date, seen also
on Aldabra (see Aldabra section). Both Rivers and Bayne et al. agree

on the abundance of coconut crabs.

Birds

Bayne et al. agree with Rivers report of the small number of
seabirds on Astove. They do not mention the frigates but these may
have been wanderers from Cosmoledo or Aldabra. It is with his account
of landbirds that Rivers differs markedly from all later reporters
(summarised by Benson 1970b). The rail, pied crow and small sunbird
are consistent with other reports but the 'merle'", the 'pigeon
hollandais’' and the cardinal are not. The word 'merle' is used in
Seychelles to mean a bulbul. It might possibly be applied toa
medium sized thrush-like bird of another genus but there is no bird
known from Astove to fit this description. Moreover Rivers was
apparently a careful observer familiar with Seychelles species. The
"pigeon hollandais' is so called after the Dutch flag with distinct
bands of red, white and blue. It is even less likely to be mistaken
for any other species. As another Alectroenas appears to have become
extinct at an early date on Farquahar (Stoddart and Benson 1970), it is
possible that the same thing occurred on Astove, between 1878 and
Dupont's visit in 1907. It is however to note that Dupont reported
seeing a Streptopelia picturata in 1907 which no other observer has ever
recorded. The 'cardinal' which, as its name implies, must have red
plumage at some stage of its life cycle is the Seychelles creole name
for Foudia madagascariensis. Rivers' cardinal must therefore have
been a Foudia sp. (see under Aldabra discussion).

Cosmoledo

On his arrival in the Cosmoledo group Rivers reported that it was
frequently visited by fishermen and whalers as indicated by turtle
“remains on the beaches. A hut and 'turtle park' (enclosure in the sea
for keeping turtles fresh before shipment) on Menai Island had been
recently burnt and destroyed. A hut on Wizard Island which had been
built from the wreckage of the Merry Monarch had also been burnt. There
were also signs of the felling of mangroves.

4

Trees and other vegetation

Rivers reported that some parts of the island would be suitable
for growing various palms. What he describes as the 'remnants of an
old grove of coconut trees' existed on Menai and those still standing
were bearing fruit abundantly. He believed palms took longer to mature
on Cosmoledo than on other islands because of the 'great sterility of
the islands'. Menai was described as ‘almost covered' with mangroves,
some of about 40 or 50 feet in height and 2 or 3 feet in diameter.

Fish

Fish were abundant and he estimated that 8 men could salt or dry
8000 lbs of fish a month. In Rivers" opinion, the capture of this
quantity would not exhaust the supply as no small fish were taken.
The most common species were 'captaines rouges', 'varvaras', 'carangues'
"'vielles', 'mullets! (Mulloidichthyus flavilineatus, a Mugilidae or
Polyremus kuru, 'croissants' (specific name unknown), 'chirugiens'
(surgeon fish, Acanthuridae), 'lions' (Holocentridae) and 'raies' of
different kinds (rays, Stoasodon marinari, Dasyatis uarnak, Taenuira
melanospila or Torpedo fuscomaculata).

Turtles and Crustacea

Both turtles (green turtles) and 'caret' (hawksbill turtles) were
abundant though the late October, early November of Rivers' visit was

"not yet the season'. Crabs were said to abound in the marshes.
Birds

Birds of all kinds were plentiful. "Fregates' (frigates), booby
‘of three different species' (Sula spp.) 'paille en queue' (tropic-

birds, (Phaethon lepturus or P. rubricauda), 'cordonniers' (lesser
noddies, Anous tenuirostris), 'fouquets' (wedgetailed shearwaters,
Puffinus pacificus), 'goelettes' (sooty tern, Sterna fuscata), and
"fanchins' (bridled terns, Sterna anaethetus) were listed. Rivers
says that the same land birds were found on Cosmoledo as on Astove,
but in greater numbers, and that there was also a 'tourtorelle rouge'
(turtle dove, Streptopelia picturata).

Other animals

Rivers reported a few goats on Menai but suggested that they did
not do well there, as many skeletons were found in the bushes. He
suggested that the reason may have been the scarcity of rain on the
island on the year of his visit.

Discussion
Coconuts and other vegetation
The present vegetation of the group as described in Bayne et al.
(1970a) is more varied than Rivers brief description suggests, no doubt
partly as a result of introductions since 1878. However, it is the
plants already present in his time which are most interesting. The
"old grove of coconuts' of 1878 was reported in 1822 by Capt. Moresby
(cited: in Bayne: et. az -1970a). Moresby also reported trees resembling
Casuarinas which Rivers did not mention, though he records them on other
islands.

Land fauna

Rivers did not report seeing giant tortoises, but the reasons may
have been the same as on Astove (see Astove discussion) . The goats
which Rivers thought were doing poorly still remained, though in small
numbers, in 1901 and had gone from Menai by 1968, though they were still
on North-east Island in 1961 (Bayne et al. 1970a).

Sea birds

Rivers reported 3 different species of sulids as did Bayne et al.
(LY) FO)» stigy ILQSS)e One species was undoubtedly Sula sula. Hteyn 16S
however very easy to mistake immature white boobies (Sula dactylatra)
for mature brown boobies (Sula leucogaster) . The only evidence for
the presence of Sula leucogaster is a skin in the National museum of
Kenya in Nairobi. This skin has been misidentified and in fact is
Sula dactylatra (A.W. Diamond pers. comm.) . It is, of course, remotely
possible that Rivers knew his boobies well and did in fact see a third
species, perhaps Sula abbotti known from Assumption at about that date,
collection by Abbott in 1893 (Stoddart, Benson and Peake 1970).

Rivers reported only 'cordonnier' or lesser noddies (Anous
tenuirostris) while Bayne et al. saw only common noddy (A. stolidus)
which in creole is called 'maqua'. It is possible that Rivers missed
seeing common noddies which nest only on small islets around the lagoon
(A.W. Diamond pers. comm.), but the fact that Bayne et al. (1970a) saw
no lesser noddies, suggests that he may have mistaken the two very
similar species. However, Since neither party was on the island for
more than a few days, the two species might both occur, possibly at
different seasons.

The fact that wedgetailed shearwaters reported by Rivers were not
seen by or reported to Bayne et al. (1970a) suggests that they may have

become extinct since 1878. The young birds and eggs are commonly eaten
by Seychellois fishermen who would therefore be aware if they were still
extant; the same habit would account for their extinction. The
extinction may also be the result of the rats, which were present by
Igo (clted by Bayne et al\.), 1 notyearlier.:

Land birds

Rivers records the land birds as the same as Astove i.e. a rail,
pied crow, sunbird, bulbul, cardinal and 'Pigeon hollandais', as well
as a turtle dove. The first three and the turtle dove were also
reported by Dupont in 1907 (Benson 1970a) . The pied crow and the
sunbird still exist andBenson suggests that futher investigation of
remoter parts of the atoll may reveal a relict populations of the rail
and turtle dove. For the possibilities of the extinction of the
bulbul 'cardinal' and 'pigeon hollandais', see the discussions on
‘“Aldabra and Astove.

Assumption Island
Rivers next went to Assumption where the extensive mining for

guano had not begun. He said, in fact, that there was nothing to
indicate that it had even been visited recently.

6

Coconuts and other vegetation

The island was covered with bushes of 'bois amanthe' (Pemphis
acidula) and other shrubs on which were found a small liane used for
dying, called 'orseille' which was common in Madagascar. There were
also a few stunted ‘affouche' (Ficus sp.) and 'bois de natte' as well
as mangroves. Rivers mentioned one coconut tree as marking the landing,
but gave no idea how many others there might have been. The “atrouche!
was apparently the preferred food of the goats on the island who would
eat even its bark.

Fish and turtles
Rivers reported that fish were found on the reefs but not in

quantity, the most common being 'carangues' and 'varavaras'. The
green turtles were said to be numerous and also 'better and fatter'
than those of other islands. Hawksbills, however, were not
numerous.
Birds

Aquatic birds were found, but not in large numbers and no names
were given. Of the land birds, Rivers said 'pigeon hollandais' is not

found but the rest were ‘like the other islands'.

Other animals

Goats had been left on the island by passing ships and Rivers
thought that there were 500 to 600 of them. He described them as
being of (2-0r 3" different species*allin "fine condition ™:

Discussion
It seems likely that Rivers spent only a short time at Assumption,
for he describes it in less detail than the other islands. By the time

of later reports, mining had started or was about to start and
subsequently destroyed the natural vegetation to a great extent.

Fish and turtles
The green turtle population which Rivers described as numerous has
been reduced drastically on Assumption (Stoddart, Benson and Peake 1970).

Birds

Rivers describes the island birds as being the same as the other
islands i.e. Cosmoledo and Astove, except for the absence of the
"pigeon hollandais'. Since he had previously said that there were
turtle doves on Cosmoledo and not on Astove, his remark is ambiguous.
The fact* that Nicoll in ‘1906.and Fryer in-1908 ‘both ‘recorded the. presence
o£ the turtle dove (Stoddart’et al. 1970) suggests that ~he intended® to
inelude it on this Jest. It is clear that he meant to include the
sunbird, pied crow, 'cardinal', rail and bulbul. The sunbird still
existed on Assumption in 1968 and probably the pied crow as well
(Stoddart et al. 1970). A rail (Dryolimnas abbotti) became extinct
between 1908 and 1937 (Stoddart et al. 1970). As in the case of Astove
and Cosmoledo, neither the bulbul nor the 'cardinal' were recorded by
any other visitors to the island. This omission is more striking on
Assumption because Abbott collected there, though not in the other
islands, in 1893, and yet failed to find either species. However, both

species are susceptible to rats which may have been the cause of their
extinction. It is unfortunate that Rivers did not list the seabirds
in detail, so that we cannot know whether he saw Sula abbotti which
Abbott collected 15 years later.

Aldabra

Rivers appears to have spent more time on Aldabra than on the other
islands. He gave details of place names and geography of the island
and reported that it was frequently visited. Although he speaks of
"fishermen that have been there for a long time fishing', he makes no
suggestion that there was a permanent settlement.

Coconuts and other vegetation

Coconuts and maize were grown on the small islands in the lagoon.
Michel was said to have about 20 grown coconut trees while La Poste on
Ile Picard was distinguished by the presence of a coconut tree. Two
or three stunted takamaka trees (Calophyllum inophyllum) were the
landmark for the well at Takamaka. Mangroves were growing on all the
islands with the largest being 4 to 5 feet in diameter. The trees
were, according to Rivers, commonly 2 to 3 feet in diameter and 40 to
50 feet high. Mangrove timber was said to be good for building because
it was straight and long lasting. Cedar Island had some 'filao'
(Casuarina) while 'vacoa Maron' (Pandanus sp.) occurred on all the
islands in quantity. The 'bois tanguin' of Madagascar (Euphorbia ?
abbotti?) was present as well as some 'bois de natte' and ‘affouche',
though Rivers did not specify where these trees occurred.

Fish, turtles and other marine fauna
Rivers reported that the fish were of the same species and in the

same numbers as they were on Cosmoledo. As well as these species there
were 'Licorne' (unicorn fish, Axinurus thynnoides) similar to the one at
Mauritius. Green turtles were as abundant as on Cosmoledo, but

hawksbill was far more numerous and ‘generally of better quality'. The

reefs were said to be full of shells, with 'pearl oysters' in quantity,
but Rivers was prevented from looking for them by bad weather.

Birds

Rivers lists 'serins' (see discussion), 'cardinal' (Foudia
eminentissima or F. madagascariensis,see discussion), pigeon hollandais,
hawks (Falco sp. probably F. newtoni), crows (Corvus albus), sunbirds,
turtle doves, rails, 'toulouse' (coucal, Centropus toulou), bulbuls
(Hypsipetes madagascariensis), '‘corbijeaux of two species, 'one white
and almost as large as a goose' [sacred ibis, Threskiornis aethiopica]',
the other the same as in Seychelles, (whimbrel), flamingoes
(Phoenicopterus ruber or P. minor), ‘veuve' and 'pie' (see discussion).
’Rivers reported that there were ‘almost all kinds of aquatic birds in
quantity, but lists only 'allouettes' (small shorebirds and waders) and
"cavalier' (crab plover Dromas ardeola).

Land animals
Rivers reported that 'by all appearances there is [sic] plenty of
tortoises on these islands'. However, though he saw 'traces' of them,

8

he never saw a tortoise, nor did he, or any of the fishermen he spoke
to, ever see a dead carcase or shell, though the fishermen sometimes
captured the animals for food. Rivers said that the largest tortoises
were from Ile Picard but that they were commonly caught on the plains
at Cinq Cases. He felt that, as it was the dry season when he was
there, the tortoises were 'hidden in holes' or shady places. When it
rained, they were said to come out and be more easily caught. On
Grand Ile 'some years ago' pigs were released which destroyed many
young tortoises, but the pigs were believed to have died out because

they were all males. There were numbers of 'flying foxes' (Fruit bats,
Pteropus seychellensis), and land crabs 'of every description' were
abundant. A few goats were found on Ile Picard and had been let loose
on other islands. There were also 'large locusts of the Madagascar
species'.

Discussion

Coconuts and other vegetation

It is apparent that on Aldabra in Rivers' day, as on other islands,
coconut trees were not so widespread as they are today. If, as Fosberg
(1971) suggests Casuarina is an introduced plant on the atoll, it was
present by 1878. Takamaka trees are still mainly restricted to the
area called after them. The largest mangroves described by Rivers are
larger than any now present on the atoll, but Rivers may have been
inaccurate in his estimation of proportions as his diameter seems
excessive (A.W. Diamond, pers. comm.).

Birds

Rivers reported 'cardinal' and 'serin' for Aldabra, but 'cardinal'
only for Astove, Cosmoledo and Assumption. "Cardinal' is present
Seychelles creole for the introduced Foudia madagascariensis. By
implication, it must be used for a bird which has red plumage at some
stage. It is never, for example, used for F. seychellensis, which is
never red. "Serin' on the other hand is used today for females of both
species of Foudia. However, when white-eyes were still common in
Seychelles, it was used at different times for Zosterops modesta and
Z. mayottensis. Rivers probably knew one of the Seychelles white-eyes.
It seems strange therefore, that he did not report white-eyes on Astove
or Cosmoledo where they were recorded after 1878 (Benson 1970a, 1970b),
especially since he did record 'cardinal' on these islands as well as on
Assumption. No other observer has recorded Foudia from these islands
and it might be suggested that Rivers mistook white-eyes for female
Foudia. This mistake is however, most unlikely, since, if he did not
know a white-eye he would have called a female Foudia a 'serin'.
Therefore, the birds on Astove, Cosmoledo, and Assumption must have been
Foudia with red plumage. Foudia eminentissima on Aldabra is in breeding
dress in November and December when Rivers was there and it is likely
that Foudia on other islands in the neighbourhood would have similar
breeding seasons. (September — March, according to Benson and Penny,
Ode) es The fody on Astove, Cosmoledo and Assumption may have been
Foudia eminentissima a subspecies of which appears on the Comores and
in Madagascar as well as on Aldabra, (Benson 1967). The reason for
its extinction on Astove and Cosmoledo is not known but it is reported
on Aldabra to be very susceptible to rats (Rattus rattus, Frith 1976),

9

as is Foudia seychellensis (Diamond and Feare, in press).

Rivers mentioned a bird like a 'veuve' which is in Seychelles
creole, a paradise flycatcher (Tersiphone corvina). However, he notes
that it is not the same species, as both male and female have long
tails and are black. This bird is clearly the Aldabra drongo
(DicruruS aldabranus) . Rivers also reported hearing a 'pie' which |
he did not see, though he implied that fishermen had done so. 'Pie'

or 'pie chanteuse' is creole for the magpie robin (Copsychus seychellarum),
the 'pie' implying a black and white colouring and the 'chanteuse' a
song. Although the fishermen probably both saw and heard this bird,
Rivers only heard it. It need not therefore have been a magpie robin,
which is, at present, restricted to Seychelles, but there seems no
other likely candidate either resident or migrant. It does not sound
like the Aldabra warbler at all. (Diamond, pers. comm.). As the
magpie robin was introduced to Alphonse at a later date, it is possible
that Rivers heard a bird that had been introduced to Aldabra, but this
is unlikely. All the other birds listed by Rivers still occur on
Aldabra.

Land animals
It is obvious from Rivers description that the tortoise population
of the islands was much lower in 1878 than at present (see summary in
Stoddart 1971). Rivers report merely confirms the findings of other
reporters that population levels were very low at the end of the
nineteenth century. Iie WbISe, lS) WMOTESIC, qelolelis, score SGeugilyy Walenlicoress pt ereehyisl
was much more difficult than today, with no outboard motors and well
cut paths. Tortoises in remoter areas would not have been found because |
these areas were not explored. Rivers record of goats appears to be the
earliest. It is however, possible that goats as well as pigs have
introduced and extirpatedmore than once. (Stoddart ioy7l)

Fish and other marine fauna

Though Aldabra is still a major breeding ground of green turtles,
it is impossible to compare present day population levels with Rivers
unquantified remarks. It is certain however that the population has
declined on Aldabra, although the hawksbill population may not have
done so. (J. Frazier 1971.) The hawksbill shell of "better quality'
was the famous Aldabra 'blonde'" shell. The pearl oyster and the
unicorn fish still occur.

References

Bayne, C.J. Cogan, B.H., Diamond, A.W., Frazier, J., Grubb, P.,
HUESOn) mA pe LOOGe), eMei DiajotOddarta, U Dake) andutaylor, wi. Di
1970a. Geography and ecology of Cosmoledo Atoll. ACOIIREST
Buby 3637-56"

Bayne, €.J., Cogan, B.H., Diamond, A.W., Frazier, J., Grubb, P.,
Hutson, A., Poore, M.E.D., Stoddart, D.R., and Taylor, J.D.
1970b. Geography and ecology of Astove. Atoll Res. Bull,
136: 83-99.

10

Benson, \C.We -LIGwE The birds of Aldabra and their status. Atoll
Res. Bull. 118: 63-111.

Benson, C.W. 1970a. Land (including shore) birds of Cosmoledo.
Atoll Res: Bull? L363 67-8):

Benson, C.W. 1970b. Land (including shore) birds of Astove. Atoll
Res. Bull 1363 (a'5=—126:.

Benson, C.W. and Penny, M.J. 1971. The land birds of Aldabra. Phil.
Trans.R. Soc. Lond. B 260: 417-527.

Fosberg, F.R. 1971. Preliminary survey of Aldabra vegetation. Phil.
Trans. Ri. (SOG. Lond. .B 2603 215-225.

Fosberg, F.R. and Renvoize, S.A. 1970. Plants of Astove Atoll.
Atoll Rese, Buz se 36.) pole

Braziers ws EoWvAy: Observations on sea turtles at Aldabra Atoll.
Phil, VErans.aReSOGe LEondsBeZ6OLus/s—40;

iDuguijeloly KEAieys WESII(s)- A twelve-month study of the Aldabra Fody Foudia
eminentissima aldabrana. Mors iB. M5 5—178i-
Nicoll, M.J. 1906. The birds collected and observed during the

voyage of the 'Valhalla'. Ebis,, (8) -6:, 666-712.

ROH Schi la Weslo). On the gigantic land-tortoises of the Seychelles
and Aldabra-Madagascar Group, with some notes on certain forms of
the Mascarene Group. Novitates Zool. 22: 418-442.

Smith, do. LeBo sand Smith pe Memes L963r The fishes of Seychelles.
Grahamstown: Department of Ichthyology, Rhodes University.

Sitoddari, Disa mlOm ire Settlement, development and conservation of
Aldabra. Phidy rans. Rss SOC. Hons. {Biepe26O22) 610-628

Siteddart, D.R. and Benson, C.W. 1970* An old record of a Blue Pigeon
Alectroenas species and sea-birds on Farquhar and Providence.
Atoll Ress Bulle L36e 35-367

Stoddart, D.R-.,, Benson, .C-Wa and Peake; dF. 1970: Ecological change
and effects of phosphate mining on Assumption Island. Atold Ress
Bulli 136i) V2 PA5%

wg

suoije907 ajduies x’

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}OUUO!ID assed

GEOCHEMISTRY AND MINERALOGY OF CARBONATE ROCK SAMPLES
FROM ALDABRA ATOLL, INDIAN OCEAN

Stephen T. Trudgill!

INTRODUCTION

The stratigraphy and petrology of Aldabra have been described by
Braithwaite et al. (1973) and Braithwaite (1975). Little is known about
the geochemistry of the rocks present, apart from the analyses of two
samples published by Stoddart et al. (1971). Accordingly, data are
given in this paper which may contribute to the geological study of the
island and also to the studies of soil chemistry and plant nutrition.

The samples were taken from the stratigraphic units identified by

Braithwaite et al. (1973). These units are listed in Table 1, with the
addition of beachrock. The numbers of samples within each unit are
indicated. The sampling locations are given in Table 2 and shown in
Figure l.

METHODS

The rock samples have been analysed by X-Ray Fluorescence
Spectrometry (Leake et al., (1969)) for Ca, Mg, Na, K, Fe, Sr, Cu, Pb, Al,
ny, 2G WhlLpCOp ey S aieel Sake In the presence of very high levels of
calcium the X-RF flowmeter became saturated and thus gave underestimates
of the calcium present. Therefore analyses for calcium were also
carried out by wet chemistry (acid digestion and EDTA titration; Bisque,
IUQYSIL)) Both data sets are reported. The wet digestion method also
permits analyses of the amounts of acid insoluble residue to be carried
out.

In addition, carbonate staining (Wolf et al., 1967a) was also
carried out on the hand specimens from which the samples for the X-RF
analyses were taken. Acid Alizarin Red S solutions were used to
differentiate aragonite and calcite from other minerals; a cobaltous
nitrate boiling test to differentiate between aragonite and calcite and
an alkaline Alizarin Red S solution to distinguish magnesium rich
carbonates. As staining is not always a wholly reliable method of
mineral identification, especially some shell materials, the mineralogy
of seven of the samples was checked by X-Ray Diffraction using rock
‘powders. The rocks were classified petrologically using the method of
Folk (1959). The porosity of three samples was also estimated, using a

lDepartment of Geography, University of Sheffield, S10 2TN, U.K.
(Manuscript received June 1978-- Eds.)

Atoll Res. Bull. No. 255: 11-22, 1981.

14

Table l. Stratigraphic units studied (after Braithwaite et al., 1973)
(See Figure 1 for location of samples)

a Unit Name Samples Studied
aL Esprit limestone il
2 Esprit phosphorites: oolites 2
3 Esprit phosphorites: conglomerates 3
4 Picard calcarenites Aay D
5 Picard calcarenites + "soils" -
6 Takamaka limestone 6
i] Soils filling subaerial fretting 7
8 Hard calcarenites 8a, b
9 Aldabra limestone Qa Dat C7 > Api Cre Ene
a oksre ee
10 Solution pit fillings TOaynb
11 Crab burrowed calcarenites =
12 Algal stromatolites =
153, Beachrock INS) Hols ea Cl

comparison of particle density and bulk density and a value of 2.7 for
the specific gravity of calcite.

The samples were all near-surface samples but the actual surface
was excluded from the chemical analyses (except for 13d which was a
surface sample).

RESULTS

The petrology and mineralogy of the samples are shown in Table 2.
The geochemical data are shown in Table 3a and 3b. The porosity data
are shown in Table 4.

DISCUSSION

Substantive conclusions about each stratigraphic unit are not
possible from the data presented since the samples are individuals from
units which are often petrologically heterogeneous and which show
considerable lateral and vertical facies diversity. However, some
broad interpretations can be made. In terms of mineralogy and porosity,
extreme diversity is evident. Frequently the samples appear to be
simple bioclastic deposits, cemented by high magnesium calcite or
aragonite.

Phosphates are present in small amounts in several rocks, as well
as in the Esprit phosphorites. The X-Ray diffraction results suggest
that a calcium phosphate form may be present (possibly Ca3 (PO,)>.nH 0
with peaks at d spaces of 3.446, 2.808 and 1.937 or Cay,P 09 at 2.784 and

15

2.6, HAKS)))c Phosphate in these forms is relatively insoluble at the soil
pH values encountered and this probably accounts for the formation of
residual soils with a high total phosphate content but a low available
phosphate content (Trudgill, 1978).

The main features of element distribution are summarised in Table 5
which lists those rocks in which relatively high concentrations occur.
Two conclusions can be drawn. Firstly, that the rock element
concentrations frequently bear relationship to the presence of carbonates
laid down in association with organisms known for their association with
particular elements (Wolf et al., 1976b); secondly that the soils and
solution pit fillings show markedly high concentrations of many elements,
suggesting a residual origin for these deposits.

Silicon is of interest since it appears to be absent from many
samples, apart from the pit fillings, soils and two samples of Aldabra
Limestone. This is of interest in the context of the foliar chemistry
of grasses which are known for their high silica content. Indeed
Renvoize (personal communication) reports a scarcity of silica bodies in
Aldabran grasses (Table 6). A specimen of Sporobolus virginicus from
Esprit was analysed for silica by acid digestion and molybdenum blue
spectrophotometry (Allen, 1974) and it was found to contain 0.16 mg/g
(dry weight). This is low when compared with values of 0.1 to 1.5% for
many U.K. grasses (Allen, 1974, Table A5).

These data are of use in indicating possible trends of
concentrations of elements present in the rock samples. ity etSy EO) be
hoped that further work on the inputs, outputs and storage of elements
may be made in the future because a geochemical study of a discrete
island unit may prove to be a viable and interesting topic.

ACKNOWLEDGEMENTS

I would like to thank Mr. D.I. Smith, Professor B.E. Leake, |
Professor J. Murray and Dr. D.G. Hamilton (University of Bristol) for
help with the geochemicaland mineralogical analyses. The Natural
Environment Research Council and The Royal Society are acknowledged for
financial support.

REFERENCES

Allen, S.E., 1974. Chemical analysis of ecological material.
Blackwell.

DULCE ps Istathe 5 MUS ILA Analyses of carbonate rocks for calcium,
magnesium, iron and aluminium with EDTA. Journal of Sedimentary
Petrology, 31: 113-122.

Braithwaite, C.J.R., J.D. Taylor and W.J. Kennedy, 1973. The evolution

of an atoll: the depositional and erosional history of Aldabra.
Phil. Trans. of the Royal Society, Lond., B, 266: 307-340.

16

Brawthwatte,, CedioRe, O75. Petrology of palaeosols and other
terrestrial sediments on Aldabra, Western Indian Ocean. Phil.
Trans ."“ROY. “SOC. ‘hond., “21/3: 4-32"

HOWkK;) elie; ek OSOe Practical petrographic classification of limestones.
Bull. Am. Ass. Petrol. Geol., 43: 1-38.

Leake, B., et al. 1969. The chemical analysis of rock powders by
automatic X-Ray fluorescence. Chemical Geology, 5: 7-86.

Stoddart, DER., w.D.) Taylor, HIR. Fosberg and -Gsky Farrow, Oiiee
Geomorphology of Aldabra Atoll. Phil, ‘Prans.“Roy. SOC... Lond Br
26078 S65.

mrudgqainel ; Stel =, LOTS. The soils of Aldabra. Phil . ‘Trans Roy ws SOC,
Eond.., By 2862 — 67-77%

Wolf, K.H., A.J. Easton and S. Warne, 1967a. Techniques of examination
and analysis of carbonate skeletons, minerals and rocks, in
Chilingar, G.V., H.J. Bissell and R.W. Fairbridge (Eds.).

Carbonate Rocks, Developments in Sedimentology, 9B, Elsevier,
253-341.

Wolf, K.H., G.V. Chilingar and F.W. Beales, 1967b- Elemental
composition of carbonate skeletons, minerals and sediments, in
Chilingan, “Guv., H.d- Bissell’ and R.W.* Haarbridge (disk):
Carbonate Rocks, Developments in Sedimentology, 9B, Elsevier,
23=4 9x

Table 2.
No. Location*
dL Esprit
2 Esprit
3 Esprit
4a Picard
b Picard
6 Picard
7 Dune Jean Louis
8a Cing Cases
b Cing Cases
9a Anse Var
b Anse Var
Passe Houareau
d Dune Jean Louis
e Picard
ifs Passe Houareau
g Passe Gionnet
h Dune Jean Louis
1 Dune Jean Louis
j Dune Jean Louis
k Anse Var
Anse Var
10a Picard
b Tle Malabar
13a Picard
b Picard
e Picard
d Picard

Abbreviations and key

*See Figure l.

high magnesium calcite Fe
low magnesium calcite dbo

aragonite
calcite

second mentioned,

Petrology

Calcarenite
Ooilhize

Calcirudite
Calcarenite
Calcarenite
Calcilutite
UNSoyaL ILM

Calcarenite
Calcarenite
Calcarenite
Calcarenite
Calcarenite
Calcarenite
Calcirudite,
Calcirudite,
Calcirudite
Calcirudite,
Calcirudite,

Chama shell

corals

corals

shells

Halimeda

Goniastrea coral

Platygyra coral

Calcirudite

Calcirudite
Calcirudite

Calcirudite

‘gd
|

ine)
Il

= phosphate

iron

= carbonate

Location, petrology and mineralogy of samples

aif

Mineralogy

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20

Table 4. Porosity of selected Aldabra rock samples (3%)
Sample 9a 42 Sample 9f 48 Sample 9h 10
ALS) an 16
26 aL 14
30 7 2
24 IRS) 7
al 7
8
26
mean 2822 mean ILS) GS mean LASS

standard deviation 8.66 standard deviation 16.76 standard deviation 6.24

Table 5. Rocks with high concentrations of the elements measured
(Approximate ranges of high values indicated in brackets)

Aluminium (over ca. 1000 ppm): shelly calcirudites, hard calcarenite,
solution pit filling, Esprit phosphorites, beachrock, Takamaka
Limestone

Barium: slightly higher (0.2 ppm) in solution pit fillings

Cobalt (ca. 1O-20 ppm): solution pit fillings, phosphorites, some
calcarenites, some shells and corals

Copper (ca. 1OO ppm and over): phosphorites, some calcarenites,
beachrock, solution pit fills

Iron (over ca. 2000 ppm): solution pit fills, phosphorites, beachrock,
some corals

Potassium (over ca. 1000 ppm): shelly calcirudites, beachrock, solution
pit fills, algal limestones

Magnesium (over 2 per cent): beachrock, shells, corals, solution pit
fills, algal material

Sodium (over ca. 2000 ppm): beachrock, some corals, "soils"
Nickel (over 20 ppm): "soils", solution pit fills
Lead (over ca. 100 ppm): algal material, some calcarenites, Esprit

phosphorites, solution pit fills

Sulphur (over ca. 2000 ppm): corals, shells, solution pit fills,
phosphorites

Silicon (over 2 per cent): “solution pit fills

Strontium (over ca. 1500 ppm): corals, beachrock, algal material

Zinc (over ca. 300 ppm): Esprit phosphorites, shells, solution pit fills

eu

Table 6. Silica in Aldabra grasses (S.A. Renvoize, personal
communication) (microscopal examination)

Stenotaphrum clavigerum Sparse silica bodies and silica-containing
cells, up to 8 um long

Panicum aldabrense No silica seen

Sclerodactylon macrostachyum No silica seen

Sporobolus virginicus No silica bodies seen but silica-containing
cells, 4 um long frequent

HISTORY OF GOATS IN THE ALDABRA ARCHIPELAGO

by D.R. Stoddart!

A summary of the history of goats on Aldabra was given by Stoddart
(1971, 617-619), and references to goats on other islands are included
in Stoddart (1970). In recent years further references to the early
history of goats have been found, and this note summarises present
knowledge on the history of goats in the Aldabra archipelago.

Assumption

Goats were said by Bergne (1901) to have been first introduced
to Assumption in 1867 by Capt Bidenfield, R.N., H.M.S. Wasp; this is
the earliest date at which goats are mentioned in the Aldabra group.
Abbott (1893, 763) stated that at Assumption 'Numbers of goats run
wild, having been introduced many years since from Europa Island’.
Dupont (1907, 12) stated that they had been introduced about twenty
years prior to his own visit in 1906.

Rivers in 1878 reported that there were some 500-600 goats of two
to three 'species' on Assumption: 'they all seem to be in fine
condition, they feed on shrubs such as the "Bois amanthe" [Pemphis
acidula] but greatly prefer the "affouches" [Ficus spp.?] of which they
even eat the bark'. In 1895 Baty estimated the numbers to be 300-400.
He himself saw a party of 60 goats, and some of his group captured a
further 30. He noted that 'their bellies are swollen to almost
abnormal proportions’. Tonnet (1905) captured 70 in two days. In
1906 Dupont reported 'several thousands' (1907, 12), ‘in spendid order'
even at the end of the dry season; hethought of them as a major
resource, and even suggested that a resident guardian be appointed on

the island to safeguard the population and regulate cropping. In the
same year Nicoll (1908, 112) found 20 very wild animals near the foot
of the high dunes: 'some of the males were remarkably fine animals.

They are excessively wild'.

By 1916, however, there were only 'a few' left on the island
(Dupont 1916). Vesey-FitzGerald (1942) makes no mention of goats in
the late 1930s, and in 1964 Gaymer found them to be absent. In 1968
Stoddart et al. (1970) saw none, but were told that some still
existed in the north.

1 Department of Geography, Cambridge University.
Atoll Res. Bull. No. 255: 23-26, 1981.

eu
Cosmoledo

In 1878 Rivers found 'a few goats' on Menai, together with ‘a
quantity of skeletons in bushes', which he attributed to lack of rain.
Bergne (1900) reported goats 'in numbers'; but half a dozen earlier
seen on Menai had been reported not to be doing well (Baty 1895).
Griffiths reported at second-hand a comment by James Spurs in 1892 that
there were 'large numbers of goats' at Cosmoledo (Fairfield, Griffith
and Abbott 1893, 154).

There were no goats on Menai in 1968 (Bayne et al. 1970), though
some were reported on North East Island by Piggott (1961).

Astove

Baty in 1895 drew attention to the absence of goats on Astove, and
there is apparently no other reference to them on this island.

Aldabra

The first record of goats at Aldabra is that by Rivers (1878):
"There is also a few goats found at "Ile Picard" and some have been set
loose on the other islands’. In 1895 Baty 'saw several goats'. He
stated that one female and one kid had been placed on Picard in 1890,
presumably by the lessee James Spurs, and by 1895 there were 40-50.

In 1892 Griffiths had seen 'several specimens which had been taken to
Aldabra' from Cosmoledo, again apparently by Spurs (Fairfield, Griffith
and Abbott 1893, 154). Baty (1895) released eight more goats on
Picard which had been brought from Assumption. He recommended that
"Goats should be placed on Aldabra Island, and especially on the plain
near the Takamakas. There I feel sure they would do extremely well.
It would however be necessary to shoot them when required for food'.

By 1900 it was reported that 'The goats introduced by Mr Spurs
have thriven exceedingly, and now abound in great numbers on Picard
island! < In October 1901 Bergne saw 20-30 goats on Picard, and there
were said to be several hundred. He reported Spurs's intention to
capture them all.

By the time of Dupont's first visit in 1906: ‘Goats... are
thriving. Mr Spurs took them over from Assumption in 1890 and although
they have been destroyed by the fishermen at Ile Picard I have come
across herds which are scattered on the main island from Dune Jean Louis
to the west end of Grande Terre (about 8 miles). They are extending
every day and in the course of time they will reach the South East end
where the ground and the water supply are much better' (1907, 22). In
1916 Dupont reported "hundreds of them' at Cinq Cases, though they were
very scarce in other areas, and by August 1929 'several thousands'
(Dupont 1929, 14).

More recently Travis (1959, 178-181) describes considerable herds
on the southern dunes, and Prosperi (1957, 198) also records goats at

the east end of Ile Malabar. Goats were present on Grande Terre,
Malabar and Ile Picard at the start of the Royal Society Expedition to
Aldabra in 1967.

References

Abbott, W.L. 1893. Notes on the natural history of Aldabra, Assumption

and Glorioso Islands, Indian Ocean. Proc. U.S. nat. Mus. 16:
759-764.
Baty Cors) £695. Report on the Aldabra Group of Islands. Mahe:

Crown Lands Department.

Bayne, C.J., Cogan, B.H., Diamond, A.W., Frazier, J., Grubb, P.,
Hutson, A., Poore, M.E.D., Stoddart, D.R. and Taylor, J.D. 1970.
Geography and ecology of Cosmoledo Atoll. Atoll Res. Bull. 136:
37-56.

Bergne, H.A'C. 1900. Fair record of islands in the Indian Ocean.
Manuscript.

Bergne, H.A'C. 1901. Rough notes on a voyage to the Aldabra Group.
Manuscript.

DUO we IO. Repore on a Vvisve of investigation to St Pierre,
Astove, Cosmoledo, Assumption and the Aldabra Group of the
Seychelles Islands. Mahe: Government Printing Office. 51 pp.

Dupont, R. 1916. Aldabra: its guano deposits and other resources.
Typescript, 25 pp. Seychelles National Archives: Aldabra Group,
Correspondence relating, 1889-1943: C/SS/73.

DUSOMIG, Ro wkOZQe REPOLEVON "a valiSheNOm Invest gaeron) EO) che primes pall
outlying islands of the Seychelles archipelago. Typescript,
AX0) 19) 96 Mahe: Department of Agriculture files.

ines @llel> jh4 A CGlemansgalely, UWois chayel Nolo. Violino. ditelS)sie Aldabra Islands

(correspondence) . BOULICS WULHOs IAG, IR, Joos, (Cebaso. IMaiyy AuSIS)Be
LD A=ILS S54

Nicoll, M.J. 1908. Three voyages of a naturalist, being an account
of many little-known islands in three oceans visited by the
NATO SEW) 12694 SIS London: Witherby. xxvi, 246 pp.

Piggott Cis) OGIS A report on a visit to the Outer Islands between
October and November 1960. Directorate of Overseas Surveys,
Land Resources Division, typescript (subsequently published, 1969,
UZ) js395))) 6

Prosperi, F. 1957. Vanished continent: an Italian expedition to the

Comoro Islands. Translated by D. Moore. London: Hutchinson.
2325.

|

26

Rivers, F. 1878. [Letter of Chief Civil Commissioner, Seychelles,
11 December 1878, on a visit to Aldabra on board the Flower of
Jarrow]. Seychelles National Archives: Letter book (outward),

1878-1880, B37; also copies in Mauritius Archives, TA 85, and
Gltinther Papers, British Museum (Natural History).

Stoddart, -D. Rion edaplO7Or Coral islands of the western Indian Ocean.
Atoll Res. Bull. 136: i-x, 1-224.

Sitoddaisitz,. Dake Oia Settlement, development and conservation of
Aldabra. Phil. Trans. R. Soc. Lond. B 260: 611-628.

Stoddart, D.R., Benson, C.W. and Peake, J.F. 1970. Ecological change
and effects of phosphate mining on Assumption Island. Atoll Res.

Bull. 36:74 205s

Tonnet, A. 1905. Report on visit to outlying islands and history of
Seychelles. Seychelles National Archives, C/SS/5.

PBA AES, We 23959): Beyond the reefs. New York: Dutton. “221 pp-

Vesey-FitzGerald, L.D.E.F. 1942. Further studies of the vegetation on
islands in the Indian Ocean. Je VE COLS 30-71-16.

ABBOTT'S BOOBY ON ASSUMPTION

1oy7 ID) IR. Stoddart!

Records on Assumption

Abbott's Booby Sula abbotti was first described by Ridgway (1893,
599) from a specimen collected by Abbott in 1892, apparently on
Assumption Island. Abbott's own description of Assumption (1893, 763)
does not mention boobies at all, but Ridgway (1895, 521) quotes notes
supplied by Abbott. Under S. abbotti he quotes: '"Fou boeuf". A
few breed on Assumption. Said not to be found on any other island in
these seas'. Abbott (in Ridgway 1895, 520) also records Sula sula (as
S. piscator) and S. dactylatra (as S. cyanops) from the same island.
Of the latter he comments: '"Fou general"': 'A few breed in Assumption,
laying a single egg on bare ground on sand dunes'.

In 1906 M.J. Nicoll visited Assumption. He collected S. dactylatra
(Nicoll 1906, 697) and photographed Sula sula (1908, 112), but he does
not mention S. abbotti, and in his general account of the island he does
not mention boobies at all (1908, 107-113).

J.C.F. Fryer collected two specimens of S. abbotti in 1908, which
he records ‘inhabits the large dune, never descending to low parts of
the island, and only going a few miles to sea to fish; it was never
seen on Aldabra' (1911, 433). He further notes: 'This species is
also recorded on Christmas Island. I have not compared my specimens
with those from this island but if identical the distribution becomes
even more curious'.

S. abbotti has not been seen on Assumption since.
Confusion with the Iles Glorieuses?

Gibson-Hill (1950) reviewed the literature on S. abbotti and
suggested that there had been confusion over the labelling of the type
specimen, which could have been obtained on Grande Glorieuse rather than
Assumption. His argument was based partly on the apparent unsuitability
of treeless Assumption compared with forested Grande Glorieuse for what
is on Christmas Island an arboreal species, partly on what seemed to be
ambiguities in Abbott's written accounts. Thus Abbott does not describe
his booby in his account of Assumption itself; but in his description
. of Grande Glorieuse he mentions that 'Among sea-birds there is a booby,
which seems to be peculiar to the island. They breed. in large numbers
upon the 'fouche' [Ficus?] trees, in company with frigates and common

1 Department of Geography, Cambridge University, Cambridge, England.
AGolle RES Sula mNOn Abbi if se OO.

28

boobies' (Abbott 1893, 764). However, the only boobies he collected on
the Iles Glorieuses were S. sula and S. dactylatra (Ridgway 1895, 524),
locally called 'Capucin' and 'Fou général'. Gibson-Hill (1950) suggested
that the 'peculiar' booby was in fact S. abbotti, and that Abbott's type
specimen really came from Grande Glorieuse rather than Assumption. He
adduced the further point that Nicoll did not find S. abbotti on
Assumption, which was still uninhabited and undisturbed in 1906; whereas
the fact that he did not find it on Grande Glorieuse either could be
explained by the settlement and partial clearance of woodland on that
island by the time of his visit. Nicoll (1906), however, stated that
the most common booby on Grande Glorieuse was dark-phase S. sula. The
S. sula collected by Abbott on the same island was also dark-phase
(Ridgway 1895, 524), and this morph could equally be the 'peculiar'

booby Abbott mentions, since white-phase sula are predominant on western
Indian Ocean coral islands (other than Tromelin: Staub 1970, 203-205).

One further piece of published evidence, hitherto overlooked, is
provided by R. Dupont (1907, 45), who visited Assumption in 1906. He lists
only S. dactylatra (as S. cyanops) ands. sula (as S. piscator) from
Assumption, while for the Iles Glorieuses he lists S. sula and S.
abbotti. This latter he notes "exists only in Gloriosa" and gives as
the Creole name 'Fou glorieuse'. However, there seems to be no record
of Dupont visiting the Iles Glorieuses themselves.

Nelson (1974) repeats Gibson-Hill's habitat argument, and agrees
with him that S. abbotti 'may have nested not on Assumption but on
Glorioso’.

Further evidence on Assumption

This trend of argument tends to overlook the fact that in addition
to Abbott's type, S. abbotti was also collected on Assumption in 1908
by Fryer (who did not visit the Iles Glorieuses) . Fryer” (L911; 7433)
refers to specimens, but these were not described until 1950 (Gibson-
Bobi) MILQ)Syo\s 0 7/24)).% Bourne comments (1976, 122) that 'there really seems
no need to question Abbott's specific statement... that his booby
nested on Assumption', since nesting on Assumption and the Iles
Glorieuses are obviously not exclusive possibilities. Nevertheless,
it would be useful to have further confirmatory field evidence of the
existence of S. abbotti on Assumption, and some rather fragmentary
and allusive evidence is available.

The first additional notice of sulids on Assumption is provided by
StC.E! Baty im-L895* He states that '... to the North East of the big
sand hills at about six hundred yards therefrom and close to the next
[north] and lower sand hill... There is a camp of 'boobies' on the
spot’. This appears to refer to a ground-nesting species, probably
S. dactylatra as recorded by Abbott. Baty also says: ‘Boobies or
Fous of different kinds are to be found in the trees all over the
island’. The implication of this is that there is more than one
species of tree-nesting sulid: the only two possibilities would be
De SULA and |S. abboeen: Finally, he says: ‘On the slopes of the big

sand hills the trees are much bigger, and on their branches the Frigate
birds and 'fous' make their nests'. This last reference could be to a
single species, either sula or abbotti; but we may recall that Fryer
(1911, 433) found that abbotti ‘inhabits the large dune, never
descending to low parts of the island'. Dupomts CEIOW.,. 22) ienoted that
wboobles tillock together in, a plain of SO acres, anvthe north of the
island; but he lists only S. sula and S. leucogaster (which to add to
the confusion he calls the Black Gannet and the Red-footed Gannet,
respectively), and does not mention either the high dune or S. abbotti.

Fryer's published references to Assumption boobies are slight, but
his manuscript diary adds more detail. On 6 September 1908 he records:
"There was a fresh species of Fou (S. leucogaster) and on the guano beds
[word illegible] of ordinary Fou were breeding being continually preyed
upon by a regular flock of Frigates of both species’ (Fryer, 1908, 73).
Gibson-Hill (1950, 69) has already noted that in his account of Aldabra

Fryer seems to use the name leucogaster for sula (as did Dupont); but
in the Diary passage the ‘ordinary Fou' must be sula, and the 'fresh
species of Fou' presumably dactylatra, Leucogaster has not been

recorded on Assumption; Fryer used the name 'S. capucina' for it on
Aldabra.

On 7 September 1908, Fryer writes: "A fresh kind of Fou (Fou Boeuf)
was found: it was larger than the ordinary with a grey back tinged with
pink: dark eyes and wings and long coverts black: its cry was very
like a cow (from which it takes its name)' (Fryer 1908, 75). TAGES! aS
very clearly S. abbotti, and Fryer's specimens must have come from this
population. Unfortunately his diary makes no comment on breeding.
However, his reference to voice is remarkable. Neither Abbott (1893)
nor Ridgway (1895) mention voice, though Abbott (in Ridgway 1895, 521)
uses the name 'Fou boeuf’. The only reference to voice prior to
Fryer's observation, with which Fryer was certainly unfamiliar, is
contained in an anonymous French account of Rodriguez, ca 1730, on 'The
Boeuf', said to 'bellow like a bull’. This was published by Milne-
Edwards (1875, 8); Bourne (1968) tentatively identified it as
S. dactylatra, but Nelson (1974) used the evidence of voice to determine
it as abbotti. Nelson at Christmas Island found abbotti to have 'the
deepest and loudest voice of any sulid, resonant and commanding' (1971,
436).

Comment

There thus seems little question, in spite of the ambiguities in
Abbotts account and the clear statement by Dupont, that abbotti was
definitely present on Assumption in 1892, possibly in 1895, and
- CEreninatieSilyy atia\ IUoNs}c BheNGCannot be Stated wath certainty, that the
species was breeding; but equally it is clear that its presence on
the island was not inhibited by the absence of tall forest, contra
Gibson-Hill and Nelson.

Whether abbotti formerly existed on the Iles Glorieuses remains
unclear. Benson et al. (1975) suggest that Abbott's 'peculiar' booby
(Abbott 1893, 764) is not abbotti as Gibson-Hill suggests but the

30

"capuchin' of Ridgway (1895, 532), i.e. dark-—phase S. sula. Abbott
(1893, 764) identifies 'capucin' as sula, as does Dupont (1907, 55):
Gibson-Hill (1950, 69) is mistaken in saying, when trying to identify
Fryer's 'S. capucina', that Abbott used 'capucin' for S. leucogaster.

It is certainly possible that abbotti inhabited the Iles
Glorieuses in spite of the absence of positive evidence. Bourne (1971,
188) speculated that this species would be found somewhere between
Assumption and its present breeding station on Christmas Island, possibly
in the Chagos Archipelago, and there has been some confirmation of this
by sightings at sea in, that vicinity (Bourne, 19/1); Himons et iat. Love,
Bourne and Nelson 1976). Bourne has also identified fossil material of
abbotti from Mauritius and possibly Rodriguez, and has quoted historical
evidence (1968; Nelson 1974) for the latter island. Becking (1976)
gives a record of a foraging bird from Java. Bourne (1976, 122)
particularly notes that the vernacular name (Fou boeuf) on Rodriguez
in the early eighteenth century was the same as that on Assumption in
the early twentieth, suggesting a previous wider occurrence and more
numerous population.

When did abbotti become extinct on Assumption? Vesey-FitzGerald
(1941, 521) says 1926, and Betts (1940, 501) says 1936, and these dates
are quoted by Gibson-Hill (1950) and later writers. Settlement began
at Assumption in 1908, and guano was then mined on a major scale
(Stoddant letra 97 O) is By the time that R. Dupont visited the island
for the second time in 1916, he reported that the boobies 'have been
all destroyed'; and in 1929, during a later visit, he stated that the
land and sea birds had been destroyed by 1909. It thus appears that
man and his activities, plus introduced predators, caused massive
extinctions within a very short time of the first settlement. No
boobies of any kind now breed on Assumption, though a few individuals
of dactylatra and sula have been seen in recent years (Stoddart et al.
1 /.O) es

Acknowledgements

I was able to visit Assumption on 15-16 September 1967 during the
Royal Society Expedition to Aldabra 1967-69, and the Iles Glorieuses on
9 December 1976 while a guest on board Lindblad Explorer. Literature
and archival research on the faunas of western Indian Ocean islands has
been carried out since 1966 as part of the Royal Society Aldabra
Research Programme. I thank Lady Joan Fryer for access to Sir John
Fryer's diary.

References

Abbott, W.L. 1893. Notes on the natural history of Aldabra, Assumption,

and Glorioso Islands, Indian Ocean. Proc. U.S. nati. Mus. 16:
759-764.
Batyjieos Colin UOO5.. Report on the Aldabra Group of Islands. Mahe:

Crown Lands Department.

Sil

Becking/ Wiech L976: Feeding range of Abbott's Booby Sula abbotti at
the coast of Java. busy, LS = ee 589—590 :

Benson, C.W., Beamish, H.H., Jouanin, C., Salvan, J. and Watson, G.E.
1975. The birds of the Iles Glorieuses. Atoll Res. Bull. 176:
1-34.

Betts, F.N. 1940. Birds of the Seychelles. Pt II. The sea-birds —
more particularly those of Aride Island. Ibis (14) 4: 489-504.

Bourne, W.R.P. 1968. The birds of Rodriguez, Indian Ocean. Ibjoahesy
110: 338-344.

Bourne, W.R.P. 1971. The birds of the Chagos Group, Indian Ocean.
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Bourne, W.R.P. 1976. On subfossil bones of Abbott's Booby Sula abbotti
from the Mascarene Islands, with a note on the proportions and
distribution of the Sulidae. Tpit sy, AA ak O23"

Bourne, W.R.P. and Nelson, J.B. 1976. Birds on the Chagos Bank.
Nature, Lond. 261: 452.

DupOMey wk. LIOW: Report on a visit of investigation to St Pierre,
Astove, Cosmoledo, Assumption and the Aldabra Group of the
Seychelles Islands. Mahe: Government Printing Office. 5l pp.

iDEIOOMIE, A ARSLSe Aldabra: its guano deposits and other resources.
Typescript, 25 pp. Seychelles National Archives: Aldabra Group,
Correspondence relating, 1889-1943: C/SS/73.

DupOMity eRe LIZ 9) Report on a visit of investigation to the principal
outlying islands of the Seychelles archipelago. Typescript, 20 pp.
Mahe: Department of Agriculture files.

Buyer, wJiscer. L908. Diary. Manuscript.

IEW c, ValGolva Mee The structure and formation of Aldabra and
neighbouring islands — with notes on their flora and fauna.
Trans. Linn. Soc. Lond. (2) 14: 397-442.

Grbson-Hilly CoA. L950. Ornithological notes from the Raffles Museum.
9. Notes on Abbott's Booby. Bulle RartLes Musi. 23) 165-76).

Hirons, M.J., Bellamy, D.J. and Sheppard, C. 1976. Birds on the Chagos
Bank. Nature, Lond. 260: 387.

Milne-Edwards, A. 1875. Nouveaux documents sur 1'époque de la
disparition de la faune ancienne de l'ile Rodrigue. Ann. Sci.
nat=.Zeol. Paléontol. (6)--2, art. 4: 1-20.

Nelson, J.-B. 1971. The biology of Abbott's Booby Sula abbotti. Ibis,
113: 429-467.

32

Nelson, J.B. 1974. The distribution of Abbott's Booby Sula abbotti.
pis, IuGinS6S8—369):

Nicoll, Mdisrel 906. On the birds collected and observed during the
voyage of the 'Valhalla', R.Y.S., from November 1905 to May 1906.
Epis, (8) Gs e6G— EZ

Nicoll Mawes ISOSr Three voyages of a naturalist, being an account of
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RipYeeSte London: Witherby. xxvi, 246 pp.

Ridgway, R. 1893. Descriptions of some new birds collected on the
islands of Aldabra and Assumption, northeast of Madagascar, by
DG W.L. Abbott. Proc’. U.S> nat. Must. le:t "597—6GOO.

Ridgway, R. 1895. On birds collected by Doctor W.L. Abbott in the
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Sitauboye Hs uO OF Geography and ecology of Tromelin Island. Atoll Res.
Bull.” L363) LOV7—209%

Stoddart, D.R., Benson, C.W. and Peake, J.F. 1970. Ecological change
and effects of phosphate mining on Assumption Island. Atoll Res.
Bull. 136: 121-145.

Vesey-FitzGerald, L.D.E.F. 1941. Further contributions to the
ornithology of the Seychelles Islands. Tbis; | (14)o575°5i8=53i.

PENETRATION OF THE HOST PLANT TISSUES BY THE STYLETS OF THE
COCCOID ICERYA SEYCHELLARUM (COCCOIDEA: MARGAROIDEFA) ON
ALDABRA ATOLL

oh Se Bia eore

ABSTRACT

The path of penetration of stylets of Icerya seychellarum through
tissues of Euphorbia pyrifolia Lam., Avicennia marina (Forssk.) Vierh.,
Scaevola taccada (Gaertn.) Roxb., and Casuarina equisetifolia L. was
studied microscopically. Most stylets were seen to end in the phloem,
others in the cortex, xylem or pith. Unlike other members of the
Coccoidea I. seychellarum frequently penetrates thick walled cells such
as sclerenchyma and xylem. This suggests that the anatomy of host
plants, and the distribution of thickened tissues in particular, is not
a restricting factor in host plant specificity.

INTRODUCTION

Stylet penetration has been studied in coccoids which act as
vectors for virus diseases of economic plants, notably the potato
(Smith, 1926), apple (Glass, 1944), and cocoa (Entwistle & Longworth,

IGS) ie Most coccoids feed on the phloem of their hosts but a few are
cortex feeders. Penetration is intracellular or, less commonly, within
the cell walls (Entwistle & Longworth, 1963). Smith (1926) described

the penetration of xylem vessels by Dactylopius longispinus but
otherwise penetration of thickened tissues has not been observed.

This study was intended to establish whether I. seychellarum
conforms to the known pattern, if any tissue damage is caused by the
stylets and whether host plant anatomy might influence host plant
specificity of the coccoid. The biology of I. seychellarum on Aldabra
Atoll has been studied by Renvoize (1975) and recently by Hill and
Newbery (publications in preparation). I. seychellarum infests a wide
range of plants. More than half of the one hundred andeighty flowering

'plant species on Aldabra Atoll have been reported as hosts. Renvoize
listed about thirty species which were subject to heavy infestations,

tRoyal Society Aldabra Research Station, Seychelles. Present address:
Chancellor College, P.O. Box 280, Zomba, Malawi.
MECuLiLMiness, wills Wes L559 SS—36, aWoehile

34

including all the examples discussed in this paper. In the granitic
islands of the Seychelles the coccoid is a pest of Citrus crops.

MATERIALS AND METHODS

The host plants studied were Euphorbia pyrifolia Lam., Avicennia
marina (Forssk.) Vierh., Scaevola taccada (Gaertn.) Roxb. and Casuarina
equisetifolia L. In each case a minimum of thirty stylets were studied
in situ in sections of stems or leaves and petioles. The plant
material was fixed in Formalin-acetic-alcohol, sectioned by hand or by
microtome, stained in safranine and light green and mounted. Hand cut
sections were particularly useful since with careful examination the
stylets were visible in the plant before it was cut. The sections
were examined microscopically and the length, width and path of the
stylets and the salivary sheath they secreted were recorded.

RESULTS

A number of general points were noted; these are summarized in
Table 1 and discussed below.

We Leaves and petioles were usually penetrated from below.
Penetration of the upper surface was only frequent in Ficus nautarum
and Thespesia populnea. The stylets usually entered the leaf to one

side of the midrib.

PAs The mean length of stylets was approximately 800um which is shorter
than the 988yum reported in Pseudococcus njalensis by Posnette &
Robertson (1950). The mean length of stylet inserted into the plant
was 520um.

3s Stylets were usually seen to end in the phloem but were also found
in the cortex, xylem, laticifers and pith.

4, Penetration was intracellular except in thickened cells where it
was intercellular. When the stylet first encountered such tissues it
was often withdrawn and re-inserted in a different position. If. the

stylet did not locate a path through less resistant tissues it was
usually eventually successful in penetrating the thickened cells
intercellularly.

Die Stylets usually penetrated in a transverse plane and were often
found in their entirity on one or two serial sections. Entwistle &
Longworth (1963) found that the plane of penetration in other coccoids
was related to the orientation of the insect and the stylet was rarely
to be found in so few adjacent sections.

Ge There was no evidence of tissue damage other than to the cells
penetrated. Small groups of callus cells were occassionally found in
Scaevola taccada.

Table 1. Summary of stylet penetration data for four host plants.

Euphorbia Scaevola Avicennia Casuarina
Stem Leaf. Stem Leaf Stem Leaf Stem Leaf

Mean length
Stylet 424 504 750 509 675 548 450 305
sheath, um

Mean number
branches in 1.6 0.5 2S 2 B83 Bak Lev Oss
sheath

% stylets
ending in 84 5 64 67 65 46 83 75
phloem

% stylets

penetrating 20 23 25 3 87 83 95 85
thickened

tissues

Specific results are as follows:
Euphorbia pyrifolia

The leaves have little thickened tissue except a band of collenchyma
along the midrib. Stylets enter to one side of this and frequently
penetrated to the phloem without being withdrawn and relocated. Often
several vascular bundles were penetrated in succession. The stems have
a well developed cork which was sometimes penetrated intercellularly.
Several stylets were seen to enter the stem through lenticels. Many
ended in, or passed through, the abundant laticifers of the cortex.

It is possible that the latex is an additional food source.

Scaevola taccada

Leaf penetration was similar to that in Euphorbia. The stem has
more sclerenchyma around the vascular tissues which may account for the
greater number of reinsertions of the stylet and resulting branching of
the salivary sheath.

Avicennia marina
The undersides of the leaves have a dense cover of simple trichomes.

The stylets pass between these and secrete a salivary sheath extending
slightly out beyond the trichomes. The vascular tissues are completely

{

36

surrounded by sclerenchyma and reinsertion of the stylets is more
frequent. The stems have large cortical air spaces. Some stylets
passed through the spaces but an equal number followed a sinuous course
through the surrounding cells. Additional cambia within the xylem
produce small groups of phloem within the xylem. About 20% of stylets
penetrated through to these inner phloem cells.

Casuarina eqisetifolia

The leaves or, strictly, phyllodes have angular, ridged surfaces
covered by a thick cuticle beneath which a layer of sclerenchyma
overlays the chlorophyllous tissues. Between the ridges the cuticle
is thin and the sclerenchyma absent and stylets entering here did not
encounter so much mechanical tissue. Equal numbers of stylets entered
through and between the ridges. Those passing through the
sclerenchyma produced branced salivary sheaths, the others did not.

CONCLUSIONS

I. seychellarum differs from most coccoids in the ability of its
stylets to penetrate thickened tissues. The ability of the stylets to
locate the phloem, even if this requires passage through thickened
cells, implies that the stylets are to some extent sensory.

Host plants of the coccoid on Aldabra Atoll range from fleshy
leaved plants such as Euphorbia pyrifolia to others with heavily
thickened stems such as Casuarina equisetifolia. The ability of the
insect to feed on plants with such varied anatomy suggests that host
anatomy is unlikely to be a limiting factor in the selection of hosts.

ACKNOWLEDGEMENTS

I am grateful to Professor J. Rutter and Dr. N. Waloff for their
encouragement with this project, to Dr. G. Hill and Dr. D. McC. Newbery
for the helpful discussion and to Professor E.J. Popham who kindly read
and commented on the manusscript.

REFERENCES

Entwistle, P.F. and Longworth, J.F. 1963. The Relationships
Between Cacao Viruses and their Vectors: the Feeding Behaviour
of Three Mealybug (Homoptera:Pseudococcidae) Species. Ann.

AD DRA "Blois ss) 2m SO Sole.

Glass, E.H. 1944. Feeding Habits of Two Mealybugs, Pseudococcus
comstockii Kuw. and Phenacoccus colemani Ehrh. Tech. Bull. Va
AGLIG. Exp. Stawg.o 5); a6. pp.

if

Posnette, A.F. and Robertson, N.F. 1950. Virus Diseases of Cacao in
West Africa, VI. Vector Investigations. Anna Apple BLOM.) Sil:
86S"

Renvoize, S. 1975. Icerya seychellarum on Aldabra. Unpublished
Royal Society Aldabra Research Committee Report, ALD/21(75).

Smash kKeoM., | LIZ6e A Comparative Study of the Feeding Methods of
Certain Hemiptera and the Resulting Effects upon Plant Tissues
with Special Reference to the Potato Plant. Ann. Appl. Biol.
3g LOSS). :

METEOROLOGICAL DATA FROM ULUL ISLAND, NAMONUITO ATOLL

by John Byron Thomas and Mary Durand Thomas!

Meteorological observations were conducted on Ulul Island, Namonuito
Atoll, Truk District, Trust Territory of the Pacific Islands from
November 8, 1973 to September 30, 1974. A record was maintained of
maximum and minimum temperatures, maximum and minimum relative humidity,

and rainfall. Temperatures and rainfall were recorded daily at
600" p'.mi. Relative humidity was measured twice daily at noon and
6:00 p.m.

The meteorological station was located in a cleared area permitting

unobstructed exposure to rain and sun. The maximum and minimum
thermometers were housed in a louvered wood structure and the rain gauge
was located about four meters distant. Relative humidity, obtained
with a sling psychrometer, was measured at the same location. Ala

instruments and their associated gear were supplied by the U.S. Weather
Bureau at Moen Island, Truk District.

Informants considered that meteorological conditions during the
recorded period were typical for Namonuito Atoll. The data presented
in the accompanying table indicate the climate to be uniformly warm and
humid. Extreme temperatures ranged from 20.5°C. to 35.0°C. (69°F. to
95°F.); relative humidity from 66% to 100%; and monthly rainfall,
mgm L376 7 sm co) SNOoS imim' (YEAS: sins io) sy olile ails) 6 Generally, the
daily low temperature occurred just prior to sunrise and the high during
early afternoon. Daily relative humidity varied froma low at about noon
to a high at about the time of sunrise. (Because a daily relative
humidity measurement was recorded at 6:00 p.m. rather than at sunrise,
the extreme high and average high relative humidity percentages were
probably somewhat higher than indicated in the table.) With a few
exceptions, daily rainfall was in the form of brief, random showers.

By extrapolation, the total annual rainfall would have been approximately
S2rS mm (26)..5) kms.) -

As is typical for this area of the Pacific, the greatest seasonal
climatic variation is reflected in wind velocity and direction rather
than in temperature, relative humidity and precipitation patterns.
Northeast trade winds prevailed during the winter months. Summer
months were characterized by lower velocity variable winds and
occasional thunder storms. Accurate wind velocity measurements could
not be obtained but the periodic use of a hand-held wind meter ata

IDepartment of Anthropology University of Hawaii.
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uat skeq (UW) TeIOL oHbeASAW ouIeTAX ouUerAX| uq.uUOW

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PLOT “‘O€ UNEWHLdHS — €L6T “8 AHAWHAON
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windward beach site indicated that winter winds seldom exceeded
approximately twenty knots per hour while summer winds usually ranged
between five and ten knots per hour.

The meteorological measurements were compiled during the course of
our respective anthropological researches at Namonuito Atoll. One of
us (J.B.T.) would like to acknowledge the research support provided by
Wenner-Gren Foundation for Anthropological Research Grant No. 2977 and
National Science Foundation Grant No. GS-39878; the other (M.D.T.)
would like to acknowledge the support provided by National Center for
Health Services Rearch and Development special fellowship No. 5 FO3
HS50513. We would also like to express our appreciation for the
assistance of Saul Price and the late William Tolliver of the U.S.
Weather Bureau, Honolulu and for that of Lazaro Mayipi of the Trust
Territory Weather Bureau, Truk District.

MARINE BENTHIC ALGAE OF KAYANGEL ATOLL, PALAU

by Roy T. Tsuda!

Introduction

During January 13-17, 1976, four faculty members from the
University of Guam Marine Laboratory visited Kayangel Atoll in the
Palau District to conduct a preliminary biological survey of the algae,
corals and fishes. Our original plan was to incorporate all of the
findings under one cover, but other research priorities of the faculty
directed each member to work at his own pace in getting the results in
print.

This paper reports on the 51 species of marine benthic algae
collected on the barrier reef flat, channel and lagoon of Kayangel
Atoll. All specimens cited here are deposited in the Herbarium of the
University of Guam Marine Laboratory. Previous to our visit to
Kayangel Atoll, only one paper (Lowenstam, 1955) mentions the algae from
this atoll; this study reports on aragonite needles secreted by the
calcareous green alga Halimeda. Based on the collections and
observations of Sargassum crassifolium during our visit to Kayangel
Atoll, Tsuda (1976) reported its presence on atolls and speculated why
this genus is rarely observed on atolls as opposed to high islands.
Five species of sea-grasses were also collected during our survey and
are included in another paper, presently in preparation, on Micronesian
sea-grasses.

Kayangel Atoll (8°05'N, 134°43'E), about 7 km long (N-S orientation)
and about 4 km wide (E-W orientation), is the northernmost atoll in the

Palau Archipelago. The atoll is almost completely encircled by a
barrier reef with four islets present on the eastern side. The
maximum depth of the lagoon is only 11 m. Gressitt (1952) provides a

description of the atoll, as well as the four islets.
Acknowledgements

Our sincere appreciation to Toshiro Paulis, Fisheries Officer of
the Palau District, who coordinated the logistics and provided us the
SOR Mablotk fom” the. trips Tokiwo Sumang, Chief Western District
Sanitarian, loaned us a Boston Whaler which proved indispensible. We
-are most grateful to Chief Rdechor Ruluked and Magistrate Mutsuo Delkum
of Kayangel Atoll for their permission to conduct biological studies on
the atoll, and to Usim Belesam, Principal of the Elementary School, for

IMarine Laboratory, University of Guam, P. O. EK, Agana, Guam 96910.
Atoll Res. Bull. No. 255: 43-8, 1981.

4h

giving us a replacement belt for the air compressor. Our sincere
gratitude to Frank A. Cushing, Tewid Boisek, Teruwo Remoket, Max Moras,
and Francisco Yamada for their technical assistance during the field
work. My appreciation to my colleagues Richard H. Randall, Charles E.
Birkeland and Steven S. Amesbury for calling my attention to certain
algal species. Sincere thanks to the wonderful people of Kayangel
Atoll for their hospitality during our stay on their atoll, and
especially to those cooks who provided the delicious island food each
night.

Stations

The terminology of Tracey et al. (1955) is used to describe the
different zones of Kayangel Atoll.

Station l. Lagoon reef margin, northern end of atoll, .5 m deep,
Jan-sel33;, 976% (RT 5088 - 5098).

Station 2) Lagoon reef margin, northwest end of atoll, .5 m deep,
Jans, 3, SEO76 (RT 5099).

Station 3. Inner reef flat, coral mound off northwest tip of Ngariungs

Islet, .3 m deep, Jan. 14, 1976 (RT 5100 - 5103).

Station 4. Lagoon, coral mound, 1.5 km south of main channel, 1 m deep,
Jan. Lain EOWG.: (RT 5104 = 5217)!.

Station 5. Reef flat, 1 km north of main channel, 2 m deep, Jan. 15,
LOWES (RE ISLES S2522))>,

Station 6. Lagoon shelf, southwest of Ngariungs Islet, 1 m deep, Jan.
Iss. FILS) 7ASie (RT 5124 - 5129).

Station 7 Lagoon, coral mound, 1 km northeast of channel, 9 m deep,
jan. *LE5; F976% (RE S230 = 5133).

Station 8s Lagoon, coral mound, 1 km west of southern tip of Ngajangel
Islet, 9 m deep, Jan. 15, 1976. (RT 5134 - 5140).

Station 9: Reef flat, .6 km north of Ngajangel Islet, .5 m deep, Jan.
7 or: (RT 5144 - 5149).

Station 10. Lagoon, coral mound, west of mid-section of Ngajangel Islet,
2-8 m deep, Jan. “16;,’ 1976. CRESS 5 O07) e557)

Station ll. Reef flat, 2 km south of main channel entrance, .3 m deep,
Jans 6 LOG? (RT 5158 - 5159).

Station 12. South channel, just west of Gorak Islet, 2 m deep, Jan.
AWS ALS) 7Keye (RE Si6L = 5163)"

Station 13. Slope of main channel, 1-10 m deep, Jan. 16, 1976.
(RT 5164 - 5176).

45

Station 14. Inner reef flat, between Ngajangel and Ngariungs Islets,
im deep, Jane a27), «4-9/6 (RTE 47737, 25U77 =45182)e.

Station 15. Slope of main channel, 2-9 m deep, Jan. 17, 1976.
(RT 5183 - 5188).

Station 16. Inner reef flat, between Ngariungs and Ngaraplas Islets,
im deep, wan. 17, 1976. (RETA 7A 5189).

SEaiteron 157 « Reef flat, just northwest of the northwest tip of
Ngajangel Islet, 1m deep, Jan. 15, 1976. (RE 51905 — 5198) 2

Station, 13): Lagoon shelf, sea-grass beds off Ngajangel Islet, 1m
deep, Jan. 17, 1976. (RE 52017— 5203).

Species Listing

The species are listed alphabetically under their respective
Divisions.

Cyanophyta
Calothrix confervicola (Roth) Ag. - RT 5132a (epiphytic on Ceramium
mazatlanense) .
Calothrix pilosa Harvey - RT 5162, RT 5198.

Microcoleus lyngbyaceus (Kiitz.) Crouan - RT 5145 (epiphytic on
Halimeda incrassata), RT 5203.

Schizothrix calcicola (Ag.) Gomont - RT 5lll.
Chlorophyta

Avrainvillea lacerata Harvey - RT 5095, RT 5100.
Avrainyvallealobscura ld. Age = RT S201. >:
Caulerpa antoenis Yamada - RT 5194 (on sand).

Gaullerpa racemosa (Forsskal) "ud. Ag. = RE 5108, RT 5124, RT 5153, RT 5182.

CGaullenpa serrulata (rocsskal)) S-aAg=--—- RL 5096, RT Sila Ri 5129c,
RUE SiS) ae Siley. sue bikes.

Gaulerpaytaxifolia (Vahl) €.s Ag. = RT 5138.

Caulerpa urvilliana Montagne - RT 5181.

Caulerpa vickersiae Boerg. - RT 5103 (on sand), RT 5133.
' Dictyosphaeria cavernosa (Forsskal) Boerg. - RT 5167.

Dictyosphaeria versluysii W. v. Bosse - RT 5093, RT 5113, RT 5119,
RE Silg6e

Halimeda cylindracea Decaisne - RT 5104, RT 5131, RT 5183.
Halimeda discoidea Decaisne - RT 5144, RT 5150, RT 5184b.

46

Halimeda gracilis Harvey - RT 5088, RT 5165
Halimeda incrassata (Ellis) Lamx. - RT 5149, RT 5202b.

Halimeda lacunalis Taylor - RT 509la, RT 5106, RT 5126a, RT 5134,
RT 5184a.

Halimeda micronesica Yamada —- RT 5089, RT 5105, RT 5129b, RT 5185a,
RT 5192;

Halimeda minima (Taylor) Colinvaux - RT 5151, RT 5164, RT 5186.
Halimeda opuntia (L.) Lamx. - RT 5090a, RT 5107, RT 5125, RT 5187.
Halimeda simulans Howe - RT 520O2a.

Halimeda stuposa Taylor - RT 5190b, RT 5202a.

Halimeda taenicola Taylor - RT 5126b, RT 5191.

Microdictyon okamurai Setchell - RT 5097, RT 5130, RT 5136, RT 5155.
Neomeris mucosa Howe - RT 5139.

Ratpilia Orlentalts Aw Satis. Oa) 1GCpD UI IRE Si Spee Loo.

Valonia wericularis (Roth) C= Agu wo wRiol5o7,, RE S767 RT Sro5e
Valonia’ ventricosa, J. Ag. — Riol2 7, sRreola2.

Valoniopsis pachynema (Martens) Boerg. - RT 5116.
Phaeophyta

Dictyopteris repens (Okamura) Boerg. - RT 5170b.
Dictyota bartayresii Lamx. - RT 5179.
Feldmannia irregularis (Kiitz.) Hamel - RT 5158.

Lobophora variegata (Lamx.) Womersley - RT 5094, RT 5099, RT 5110,
RT 512), Rl Sig. Rok fle.

Padina tenuis Bory - RT 5146.

Sargassum crassifolium J. Ag..— RI .4773, RD. 4774.

Turbinaria ornata,,(Turner):J.2Ag. - RI. 5092, RT, 5109, RL. 5163%
Rhodophyta

Amphiroa fragilissima (L.) Lamx. - RT 5148.

Ceramium gracillimum v. byssoideum (Harv.) Mazoyer - RT 5112b.

Ceramium mazatlanense Dawson - RT 5132a.

Claudea multifida Harvey —- RT 5161, RT 5168.
Endosiphonia spinuligera Zanard. - RT 51lOla, RT 5156.
Galaxaura oblongata WEi& Si.)r Lamx. (+ REMSLSO.
Gelidiopsis intricata (Ag<) Vickers,-— RT. 5122:

Jania capillacea Harvey - RT 5178 (epiphytic on Sargassum crassifolium) ,
REE SLOT"

Laurencia cartilaginea Yamada - RT 510lb, RT 5128.

Laurencia majuscula (Harv.) Lucas - RT 5098 (intermixed with Laurencia
cartilaginea), RT 5140 (epiphytic on Halimeda opuntia), RT 5169
(epiphytic on Halimeda lacunalis), RT 5188 (on dead coral).

Liagora pinnata Harvey - RT 5177, RT 5193.

Polysiphonia howei Hellenberg - RT 5112a, RT 5117, RT 5120, RT 5129
(mixed with Polysiphonia scopulorum), RT 5159, RT 5174.

Polysiphonia scopulorum Harvey - RT 5102, RT 5129 (mixed with
Polysiphonia howei), RT 5132b.

Discussion

Claudea multifida (see Papenfuss, 1937) is the only alga collected
from Kayangel Atoll which has not been previously reported from the
Micronesian region. The rest of the algae represents an extension of
the marine algae previously known from Palau proper or other Micronesian
islands (see Tsuda and Wray, 1977).

The prostrate green alga Microdictyon okamurai is the dominant
alga on the solid calcareous substratum of the barrier reef and on the
coral mounds in the lagoon. This species covers about 23% of the
consolidated substratum on the northern barrier reef and is equally
abundant in the lagoon. Lobophora variegata and Dictyosphaeria
cavernosa are also conspicuous in these same areas.

The calcareous green alga Halimeda is abundant in the channel and
lagoon areas. O£ the 11 species of Halimeda collected from this atoll, |
four species (H. cylindracea, H. incrassata, H. simulans, and H. stuposa)
inhabit the sandy substratum which makes up the majority of the substrata
type within the lagoon. Four species of Caulerpa (C. antoensis, C.
taxifolia, C. urvilliana, and C. vickersiae) also inhabit the sandy
areas of the lagoon. The turf community is developed on the coral
mounds in the lagoon. Polysiphonia howei, P. scopulorum, Jania
capillacea, and Laurencia majuscula are the dominant algae comprising
the turf.

The small size and shallow depth of Kayangel Lagoon makes this
atoll a perfect natural laboratory for distributional and seasonal
studies of benthic algae, especially the Halimeda species, in a lagoon
environment.

REFERENCES
Gressitt, Jol. 1952. Description of Kayangel Atoll, Palau Islands.
Atoll Res. Bull. 14:1-6.
Lowenstam, H.A. 1955. Aragonite needles secreted by algae and some
sedimentary implications. wis Sa6ls IBeeIRO 2S8 20-22

48

Papenfuss, G.F. 1937.

The structure and reproduction of Claudea

multifida, Vanvoorstia spectabilis, and Vanvoorstia coccinea.

Symbolae Botanicae

Upsalienses I1I(4) :1-66.

Tracey ,, iis," PLE Clioudy aid 1 rand eK Osi EmeryeeO 55:5 Conspicuous
features of organic reefs. Atoll Resi. Bull. A261 —37 2b icice
Tsuda (Rath OMG. Occurrence of the genus Sargassum (Phaeophyta)

two Pacific atolls.

Micronesica 12:279-282.

Tsuda, Reiley and HO Wisaya lh OWie Bibliography of marine benthic
algae in Micronesia. Mieronesica 13:85-120.

on

QUALITATIVE ASSESSMENT OF THE ASTEROIDS, ECHINOIDS
AND HOLOTHURIANS IN YAP LAGOON

by Deborah A. Grosenbaugh

INTRODUCTION

A limited amount of work has been done on the echinoderm population
of Yap. Two previous technical reports (Amesbury et al., 19/76, 1977)
list 16 echinoderms observed in two localized areas: one at the
Donitsch Island sewer outfall site and the other at a proposed dock site
in Colonia, Yap.

The only other observations of this group relate to the asteroids.
Earlier reports (Hayashi, 1938a, 1938b) mention only Culcita
novaeguineae and Fromia monilis as being identified from Yap. Clark
(1954) recorded Protoreaster nodosus as also being found there.

During the period of time when Acanthaster planci infestations were
a concern throughout the western and central Pacific, several separate
surveys were made on Yap. Beginning in July of 1969, Chesher (1969)
observed no large populations of A. planci. By late 1970 (Tsuda et al.,
1970) two small populations (one consisting of 100 - 150 individuals
and another of about 50 starfish) were identified outside of the barrier
reef. A subsequent survey in 1971 (Marsh and Tsuda, 1973) noted an
apparent increase in A. planci, but they were widely scattered with no
obvious major concentrations.

This particular paper provides a qualitative assessment of the
asteroid, echinoid, and holothurian populations in the Yap Lagoon, with
particular reference to the latter.

The author wishes to acknowledge L. G. Eldredge and C. E. Birkeland
for their aid in the identification of some of the echinoderms en-
countered in this study and for reviewing the manuscript, and R. L. Kock
for providing the data concerning the stomach contents of Mespilia
globulus and Tripneustes gratilla. Identification of most holothurians
was obtained from Rowe and Doty (1977).

Teonerineeion No. 116, University of Guam Marine Laboratory. The field
work was conducted during a general reef survey of Yap Lagoon with funds
provided by the Office of Planning and Statistics, Trust Territory of
the Pacific Islands, to the University of Guam Marine Laboratory (R. T.

Tsuda, Principal Investigator).
ACO Rese Bulkley NOMeo oro => le lOGiks

50

METHOD

The observations made here were done during the course of a general
reef survey, conducted during July 9-20, 1977, which covered a major
portion of the reef flat in the Yap Lagoon. Most observations were
made as a result of towing the observer from a small boat in a zigzag
pattern from the seagrass beds near shore to the barrier reef margin.
When unusually conspicuous populations of echinoderms (particularly
holothurians) were noted, a closer examination was made by snorkeling
in the area. Periodically, more detailed observations of randomly
chosen areas were made in order to note less conspicuous species.

Two 30-m transects were run in a Thalassia hemprichii bed just east
of Bik Island in Tomil Harbor. In this case, all echinoderms 0.5 m on
either side of a 30 m-long transect line were enumerated.

RESULTS AND DISCUSSION

Four zonal habitats (seagrass, seagrass-live coral, live coral,
and coral pavement) characterize the reef flats in the Yap Lagoon.

The seagrass zone is characterized by thick beds of Thalassia

and Enhalus adjacent to shore, becoming patchy and interspersed with
live coral heads (predominately Porites) as one progresses out toward
the reef margin. This transitional zone is referred to as the seagrass-
live coral zone in this paper. As the seagrass becomes less dominant
other corals (notably Acropora) along with the Porites form a live coral
zone. Near the reef margin, the live corals become replaced by coral
pavement. Table 1 lists the various echinoderms found in these four
habitats, as well as those found on patch reefs, holes, and channel.

The densest populations of echinoderms on the reef flat occur in
the seagrass beds adjacent to land, though not all of the seagrass beds
surveyed exhibited obvious numbers of echinoderms. Holothurians in
the seagrass beds were either absent, rare (1-2 seen per hour of
snorkeling), or unusually abundant.

Table 2 presents the results of the two 30-m long transects (lm
wide) run in the Thalassia bed. Transect A was located adjacent to Tomil
Harbor channel and southeast of Bik Island. Transect B was situated just
east of the island. The major components in both cases were the echinoid
Mespilia globulus and the holothurian Actinopyga echinites. The
unusually large numbers of M. globulus present had covered themselves
with seagrass detritus. Examination of the stomach contents of 25 M.
globulus and four Tripneustes gratilla that were also present, revealed
that Thalassia was the only food item ingested. Actinopyga echinites
seemed to be feeding primarily on the detritus. In some cases, A.
echinites could be seen feeding on the detritus that was attached to

Dil:

Table 1. Checklist of asteroids, echinoids, and holothurians observed
in various reef flat zones and other habitats in Yap Lagoon.
(1. seagrass zone, 2. seagrass-live coral zone. 3. live coral
zone, 4. coral pavement zone, 5. patch reefs in Tomil, Mil,
and Gofenu Harbors, 6. holes in reef flat, 7. Tomil Harbor
Channel).

Reef Flat Zones Other Hdbitats
SPECIES 1 2 3 4 5) 6 7

Class Asteroidea
Acanthaster planci (Linnaeus) X
Culcita novaeguineae Muller &
Troschel X X X X
Echinaster luzonicus (Gray) xX xX
Fromia milleporella (Lamarck) xX
Linckia laevigata (Linnaeus) X
Linckia multifora (Lamarck) xX
Protoreaster nodosus (Linnaeus) X

Class Echinoidea
Diadema setosum (Michelin) xX xX Xx
Echinothrix calamaris (Pallas)
Heterocentrotus mammillatus (Linnaeus)
Mespilia globulus (Linnaeus) X xX
Tripneustes gratilla (Linnaeus) X

x PN

Class Holothuroidea
Actinopyga echinites (Jaeger) X
Bohadschia argus Jaeger X Xx
Bohadschia sp. X
Holothuria atra Jaeger X X X X
Holothuria axiloga Clark xX
Holothuria edulis Lesson X
Holothuria hilla Lesson X
Holothuria flavomaculata Semper Xx
Holothuria leucospilota Brandt
Holothuria nobilis (Selenka)
Stichopus chloronotus Brandt X
Stichopus variegatus Semper X X
Thelenota ananas (Jaeger) x
Unidentified synaptid Xx

MPS PS PS
rs
PS
Pa

52

M. globulus. Actinopyga echinites exhibited the clumped distribution
referred to by Amesbury et al. (1976), being found often in "piles" of
five or more in close association. One small dark red holothurian,
which the author could not identify, was observed in Transect A.

Table 2. Density of echinoderms along two 30-m long transects
(1 m wide) in a Thalassia bed off Bik Island, Tomil
Harbor, Yap.

SPECIES A B

Actinopyga echinites 164 450
Mespilia globulus Sih 245
Holothuria edulis TIL
Protoreaster nodosus 9
Holothuria atra 4
Stichopus chloronotus 2
Holothuria hilla Al
Unidentified holothurian iL
Holothuria flavomaculata - 4

Tripneustes gratilla ~ iL

In another Thalassia bed off Pekel Island in Tomil Harbor,
Actinopyga echinites was also abundant, exhibiting the same tendency
toward clumping that was observed earlier. Also present but not as
abundant were Stichopus variegatus and S. chloronotus and a variety
of occasionally observed holothurians such as Holothuria hilla, H. atra,
and the asteroid Protoreaster nodosus, along with the scyphozoan
medusa Cassiopeia sp. Large population densities of H. atra were also
noted in an Enteromorpha bed west of Thilimad Island, near the northern
tip of Map Island. Only one synaptid was observed during the entire
survey, in a seagrass bed off the northeast corner of Map.

As one progresses out toward the reef margin into the seagrass-—
live coral zone, Holothuria edulis replaces Actinopyga echinites as the
predominant holothurian species. Stichopus chloronotus was usually
present in this zone, but, as in the seagrass zone, was not particularly
common, It was seen mostly in sandy areas between coral beds.
Holothuria leucospilota was occasionally seen with the anterior part
of its body extended out from under the Porites heads. Other
holothurians observed in the seagrass-live coral zone were an occasional
Holothuria atra, H. nobilis, and H. flavomaculata. The "cushion star-
fish" Culcuta novaeguineae was also seen.

In the live coral zone, the asteroids Linckia multifora, Fromia
milleporella, and Echinaster luzonicus, along with the echinoid Diadema
setosum, were common in among the Acropora, and under Porites heads.

DS

Holothuria leucospilota was seen, as in the seagrass-live coral zone,
extending out from under Porites heads, whereas H. flavomaculata was
common in close association with stands of Acropora.

In the coral pavement’ zone out near the reef margin, only an
occasional Holothuria atra represented the holothurians. The only other
obvious echinoderms were Linckia laevigata and Culcita novaeguineae.

The majority of the echinoderms in this zone were found in coral rubble
and under rocks. These include an unidentified ophiuroid, Echinaster
luzonicus, Echinothrix calamaris, and Heterocentrotus mammillatus.

Besides the general zonation across the reef flat, there exist
distinct echinoderm populations in the deep holes that occur on the
reef flat, and on patch reefs in the various harbor entrances. The
holes are surrounded by a Porites-Acropora type community and the
echinoderm fauna corresponds thusly. As the sides of the holes slope
downward to a sandy bottom, those holothurians which prefer sandy sub-
strate are seen, e.g., Holothuria edulis and Bohadschia argus.

Patch reefs in the three major harbors (Tomil, Mil, Gofenu)
provide a diverse environment for a variety of echinoderms. Sandy
areas with some seagrass are interspersed between areas of high coral
density. The sandy areas of these reefs include the holothurians
Holothuria edulis, H. atra, Stichopus chloronotus, S. variegatus, and
Bohadschia argus. The holothurians H. flavomaculata and H. leucospilota
were found associated with the corals Acropora and Porites, respectively.
The echinoid Diadema setosum and the asteroid Fromia milleporella were
found in the coral while Culcita novaeguineae was often observed in
sandy patches between corals. On the downward slope of these patch
reefs and along channels, the larger detritus-feeding holothurians,
i.e., Thelenota ananas, Bohadschia argus, and Holothuria axiologa,
were located.

While holothurians of major economic importance were identified
within the reef flat (Thelenota ananas, Bohadschia argus, Holothuria
nobilis), they were not observed to be present in such quantities that
would make their export profitable. The Yapese, themselves, are not
overly fond of the delicacy, but the Palauan community of Yap does
occasionally utilize the above-mentioned species as a food source as
well as Actinopyga echinites which is found in considerable abundance
in some of the seagrass areas. Since the Palauan community is situated
in the Colonia area, only edible holothurians in the Tomil Harbor area
are likely to be harvested because of the strict reef tenure on Yap
where the various villages have exclusive fishing rights on designated
areas of the reef.

54

LITERATURE CITED

Amesbury, S. S., R. D2. Esuda,, Ro. Ho Randall. i¢. EB. Birkeland, andah awa.
Cushing. 1976. Limited current and underwater biological survey
of the Donitsch Island sewer outfall site, Yap, Western Caroline
Islands, Univ. Guam Marine Lab., Tech. Rept. 24, 48 p.

Amesbury, S. S., R. T. Tsuda, R. H. Randall and C. E. Birkeland. 1977.
Marine biological survey of the proposed dock site at Colonia,
Yap: Guam Marine Lab., Tech. Rept. 35, 22 p.

Chesher, R. H. 1969. Acanthaster planci, impact on Pacific coral
reefs. Final Report to U. S. Department of Interior. Westing-

house Research Laboratories No. 96-951-Ocean-R2, 152 p.

Clark, A. H. 1954. Records of Indo-Pacific echinoderms. Pac. Sci.
8 (3): 243-263.

Hayashi, R. 1938a. Sea-stars of the Caroline Islands, Palao Trop.
Biol. Sta. Studies 1(3):418-446.

. 1938b. Asteroidea of Japan's southern islands. South

Sea Sci. 1(2):35-38.

Marsh, J. A., Jr., and R. T. Tsuda. 1973. Population levels of
Acanthaster planci in the Mariana and Caroline Islands, Atoll
RES i Buld... GO) 1—€6.

Rowe, F. W. E., and J. E. Doty. 1977. The shallow-water holothurians
of Guam. Micronesica 13(2):217-250.

Tsuda, R. i, D. P. Cheney; J.-A Marsh, Jr.,tand M. R, Struck aeelo7ior
Acanthaster planci, crown-of-thorns starfish. Resurvey of Yap,
Yap District. Univ. Guam Mar. Lab., Misc. Rept. 13 p.

ACANTHASTER IN THE CULTURES OF HIGH ISLANDS

by

Charles Bee eedaniee”

Residents of high islands in Micronesia, Melanesia and Polynesia usually
have their own special names for Acanthaster, each have similar advice on
curing the sting of the spines, and each claim that Acanthaster has been
abundant at certain times in the past. I believe that this familiarity of
Acanthaster in some high island cultures implies that outbreaks are a naturally
recurring phenomenon around high islands. An apparent lack of familiarity of
Acanthaster in the cultures of atolls implies that outbreaks of Acanthaster
are much less frequent around atolls (low islands).

In Palau (Micronesia) , the Acanthaster is called rrusech (Birkeland, 1979)
while other starfish are called btuch or tengetang. At Fiji (Melanesia) ,
Acanthaster is called bula (a homonym of "hello") while the general terms for
"starfish" are gasagasan or basage (Atelaite Smalley, pers. comm.). In the
Cook and Society Islands (Polynesia) , Acanthaster is called taramea and in
Samoa and Tonga (Polynesia), Acanthaster is called alamea (Garlovsky and
Bergquist, 1970; Birkeland and Randall, 1979; Flanigan and Lamberts, 1981).

In contrast, the languages from atolls appear to not contain terms for
Acanthaster. There is no special word for Acanthaster on Pingelap (Spensin
James, pers. comm.); the crown-of-thorns is merely referred to as 7Ssu, a term
used for all starfish. Similarly, Acanthaster is called talwalyol on Ulithi,

a general term for all starfish (Eulalia Harui, pers. comm.). Abo et al. |
(1976) list 12,000 entries with information on about 30,000 Marshallese words.

Many fishes, three groups of sea cucumbers, and other marine organisms are

mentioned, but there was no word for Acanthaster. It must not be important |
to the Marshallese.

The Gilbert Islands (Kiribati), the Ellice Islands (Tuvalu) and Fanning
Island are all atolls. Lobel (1978) presented a list of 407 names of fishes
and 95 names of marine invertebrates used by Gilbertese and Ellice Islanders
on Fanning Island, but Acanthaster was not listed. These inhabitants of atolls |
had their own specific names for many species of fishes, mollusks and
crustaceans and even distinguished between three groups of holothurians, but
all seastars came under one name. Acanthaster may have never been abundant
on these atolls.

There is an exception from the atoll of Mokil (near Ponape) at which the
people do call Acanthaster by the name Zarmz. Of course there may be other
exceptions. However, there Joes seem to be a general presence of words for
Acanthaster in high island languages and an apparent lack of words for

Acanthaster in low island languages. These tendencies suggest that Acanthaster
may be more common around high islands.

Ly
Marine Laboratory, UOG Station, Mangilao, Guam 96913.
Atoll Res. Bull. No. 255: 55-58, 1981.

56

People from atolls say they rarely, if ever, see Acanthaster and they
have never heard of them being common (Matuatua Smit, Takapoto Atoll, Tua-
motu, French Polynesia; Eulalia Harui, Ulithi, Yap District; Spensin James,
Pingelap, Ponape District, pers. comms.). Around high islands, the people
remember previous outbreaks. According to localfishermen, there was an out-
break in American Samoa in 1938 (Flanigan and Lamberts 1981). Vine (1970)
reported that fishermen in the Solomon Islands (Melanesia) remembered large
concentrations of Acanthaster about 1930, forty years previous to 1970.
Chesher (1969) reported that Micronesians remember an outbreak on Ponape just
after World War II.

Michael Parke talked to an old Palauan fisherman who described an
extensive infestation that took place just prior to World War II. According
to this fisherman, the Acanthaster soon disappeared, leaving algae in the
place of coral. Then urchins became abundant during the early years of
World War II. The fisherman felt that Acanthaster were transitory and no
real problem. The abundance of urchins that resulted were a benefit. Old
people could easily collect them for food within wading depth on the reef
flat. We have not heard of other cases of an abundance of urchins following
Acanthaster. It will be interesting to see if herbivorous urchins become
common following the present devastation in Palau. Except for areas around
artificial sea walls, breakwaters and ramps, regular urchins are remarkably
scarce in Palau at the time of this writing.

The people on high islands tell of dangers of stepping on Acanthaster
when fishing at night at times when Acanthaster is abundant (Vaolui, pers.
comm. ; Flanigan and Lamberts, 1981). A cure for injury from stepping on
Acanthaster is claimed by several high island cultures to be their own dis-
covery. When I was studying an Acanthaster outbreak in Palau, I accidentally
jabbed my knee strongly against an Acanthaster and came to the boat with a
lot of blood dripping out of six cuts in my knee. The boatman, Ngirbauliad
("Yahd'") Mineich, advised me to take one of the Acanthaster and place it
mouth down on the bloody knee. (This was tried, but was not found to be of
great help.) When asked if he heard of this cure from a Samoan or Fijian,
Yahd said it has always been common knowledge in Palau. Ramon Rechebei,
another Palauan, said that he knew of this cure since he was a boy.

Spensin James told me that this cure had worked for him when he tried
it. This cure was common knowledge among Ponapean fishermen and it works
if you are sure to use the same individual Acanthaster that harmed you as
the individual to cure you. If you are jabbed by one Acanthaster and lift
another to cure your pain, it will be of no use. (I am not sure I used the
same individual in Palau.)

Laite Smalley told me that when Fijian fishermen step on bula on the
reef flat, they turn over the same bula and put their food against the mouth
so that the bula will suck out the poison. She said this was generally known
by Fijian fishermen and there is no reason to believe it was learned from
the Palauans or Samoans. Maybe the cure was discovered in Fiji.

This same cure has been known on Tonga (Richard Braley, pers. comm.),
and as a proverb on Western Samoa (Garlovsky and Bergquist, 1970) and
American Samoa (Birkeland and Randall, 1979; Flanigan and Lamberts, 1981).

of

The Secretariat of the British Solomon Islands Protectorate (1970) noted that
this same cure by turning over and stepping on the underside of the Acanthaster
was known in the Solomon Islands,New Britain, Manus Islands, and Gambier
Islands.

The apparent history of recurring abundances of Acanthaster around high
islands but not around atolls may be explained by the causal mechanism of
Acanthaster outbreaks as suggested in Birkeland and Randall (1979) and
Birkeland (1980). Acanthaster larvae may survive in much greater abundance
following heavy rainfall. This might be because phytoplankton blooms are
triggered by terrestrial runoff and this provides an abundant food source
for Acanthaster larvae. Terrestrial runoff resulting from rains on high
islands trigger phytoplankton blooms (Marsh 1977), but it is doubtful that
terrestrial runoffs from low, sandy atolls carry an amount of nutrients into
the coastal waters adequate to trigger blooms.

Acknowledgements

I wish to thank the Samoan fishermen who told me of the folklore of the
alamea: Leuila Alaimaleata and Vaolui of Alofau, Upuese Taifane of Poloa,
Chief Faiaipa'u of Onenoa and Ierenimo Laupapa, Senior Biologists of the
Office of Marine Resources, Government of American Samoa. I am grateful to
Henry Sesepasara, Director of the Office of Marine Resources, and to the
Honorable Peter T. Coleman, Governor of American Samoa, for inviting me to
come to American Samoa to study the Acanthaster populations and for air fare
and per diem. I am indebted to Toshiro Paulis, Division of Marine Resources,
Government of Palau, for arranging for an airplane ticket to Palau. The
University of Guam provided the air fare for Tahiti (Society Islands) by way
of Rarotonga (Cook Islands).

References Cited

Abo, T., B. W. Bender, A. Capelle, and T. DeBrum. 1976. Marshallese-English
Dictionary. Pali Language Texts: Micronesia. The Univ. Press
Hawalaae Hono. S569 api.

Birkeland, C. 1979. Report. of the Acanthaster planet (rrusech) survey of
Palau, 18 to 26 May 1979. Report submitted to the Division of
Marine Resources, Palau. 30 p.

Birkeland, C. 1980. Population postulation. Glimpses of Micronesia and
the Western Pacific 20(2) :76-77.

Birkeland, ©., and R. H. Randall. 1979. Report on the Acanthaster planet
(alamea) studies on Tutuila, American Samoa. Report submitted to
the Division of Marine Resources, American Samoa. 50 p.

Chesher, R. H. 1969. Acanthaster planet: impact on Pacific coral reefs.
Final report to U. S. Dept. of the Interior, Westinghouse Res.
Lab. No. 69-951-Ocean-R2. 152 p.

aS

5B

Flanigan, J. M., and A. E. Lamberts. 1981. Acanthaster as a recurring
phenomenon in Samoan history. Atoll. Res. Bull.

Garlovsky, D. F., and A. Bergquist. 1970. Crown-of-thors starfish in
Wester Samoa. South Pacific Bull.20(3) :47-49,

Lobel, P. S. 1978. Gilbertese and Ellice Islander names for fishes and
other organisms. Micronesica 14(2):177-198.

Marsh, J. A., Jr. 1977. Terrestrial inputs of nitrogen and phosphorus on
fringing reefs of Guam. Proceedings, Third International Coral Reef
Symposium. 1. Biology:331-336.

Secretariat, Honiara, British Solomon Islands Protectorate. 1970. A note
on traditional local remedies for injuries received from the crown-
of-thorns starfish in the British Solomon Islands Protectorate. 1 p.

Vine, P. J. 1970. Densities of Acanthaster planct in the Pacific Ocean.
Nature :22 8-342.

ACANTHASTER AS A RECURRING PHENOMENON IN SAMOAN HISTORY

by John M. Flanigan! and Austin E. Lamberts”

Verbal history, linquistic evidence and proverbs indicate that |
Acanthaster planci have been long known and endured in Samoa.

The Crown-of-Thorns starfish (Acanthaster planci (L.)) has been
known to science since it was described by Rumphius in 1705. Only in
the past decade and a half has it stirred popular interest after great
numbers of these starfish were reported almost simultaneously from Guam,
the Great Barrier reef province of Australia and from other Indo-Pacific
coral reef areas. Significantly, this occurred at about the time that
face mask and snorkel were becoming standard equipment for visitors
to coral reefs.

Recently Acanthaster have appeared on the reefs of Tutuila,
American Samoa, in impressive numbers. The fact that collection
efforts which reportedly netted nearly a half a million starfish have
not significantly affected the total number attests to the host of
Acanthaster (Sesapasaro, 1978). In previous decades this animal was a
great rarity locally. Between 1967 and 1973 one of us (JF) encountered
only two of these creatures during hundreds of dives on the reefs of
Tutuila. Nevertheless, there is evidence that these animals have |
occurred in great numbers in the past.

The specificity with which objects in the Samoan environment are
named bears a relationship to the importance of those objectSin the
eulittuner Thus, Samoans have a single word to refer to all types of
branching corals ('amu) but distinctive word for the less conspicuous
massive coral (puga) and flat coral (lapa) used for burning to make
lime. The language also has a single word for nearly all starfish
(aveau) but Acanthaster alone has the highly specific name of alamea.
It is noteworthy that a similar word, taramea is used for Acanthaster
in the Polynesian Cook and Society Islands even though there had been

IDept. of Mathematics & Science American Samoa Community College Pago
Pago, American Samoa 96799.
21520 Leffingwell, N.E. Grand Rapids, Mi. 49505.

(Manuscript received 10 March 1979-- Eds.)
AtolbeRess Bully Norws25 52 "59=62, G9 sah

60

no apparent contact between these subcultures for centuries before they
were rediscovered. (The cushion starfish Culcita has a specific name,
palutu, but the Samoans do not recognize it as a starfish.)

Additional indication that Acanthaster have occurred previously in
great numbers is provided by Samoan proverbs that mention it. LEEOLO
e le alamea lana sale" or "E fofo e le alamea le alamea" meaning "the
alamea is its own doctor," or "falau a alamea" meaning ‘ture of the
alamea" refers to the belief held throughout the tropical Pacific that
the excruciating pain caused by the stingof the spines of the
Acanthaster is relieved by placing the mouth of the same animal against
the wound (Herman, 1953). One of us (AEL) missed an opportunity to
confirm the efficacy of this treatment when he was stung by an
Acanthaster before hearing of the purported cure. One would expect
that an animal would enter a culture's folklore only if its nature was
very familiar to generations of members of that culture. It seems
probable that the sting of the alamea has been known and respected by
generations of Samoan fishermen.

Several Samoan fishermen, all over fifty years old reported to one
of us (AEL) that in their youth there were huge numbers of alamea on the
reefs of Nu'uuliand Aua, Tutuila and that all of them disappeared quite
suddenly. Mary Prichard, an astute and perceptive Samoan lady of 75
recalls this vividly and dates the infestation to about 1938. She was
an avid shell collector even before this but insists that she saw not a
single Acanthaster in the intervening years. The Acanthaster appeared
in large numbers on several reefs, were present for a time, then were
not seen again until recently.

Similarly, one of us (JF) interviewed an elderly fisherman in the
village of Suano, Upalu, Western Samoa and was told that perhaps forty
years before, when the man was young, so many alamea appeared on the
reef fronting the village that it could no longer be fished in safety.
(Typically, "fishing" refers to an activity involving deep wading or
swimming near the reef edge by pushing from one coral head to another

with the feet.) Reportedly there were so many alamea that a fisherman
could not stand on the reef without stepping on one. Consequently,
fishing was done at a more distant reef. Presumably, such numbers of

Acanthaster would cause extensive damage to living corals, yet when
visited by the author, the same reefs were found to be 90% to 100%
covered with healthy corals. These reports tend to confirm the
hypothesis that Acanthaster infestations are a recurring phenomenon,
and that the resulting reef damage is temporary and of no lasting
consequence.

Generations of Samoans have modified their fishing habits to
accommodate periodic infestations of alamea. Samoan fishermen seem
less concerned about a new infestation than non-natives who view with
alarm from a background richer in formal education and concern for the
environment but poorer in traditional knowledge and experience.

Our first priority in the study of increases in the numbers of
Acanthaster should be to ask the people who have lived with the reefs

61

for the longest time, and to study the natural history of the affected
reefs with particular emphasis on the stages of coral damage, its
ecological implications, and the progressive stages of succession
during the recovery of the reef corals.

“Acknowledgement

Supported in part by a grant from the National Geographic Society,
Washington, D.C., to A.E. Lamberts.

References cited

Herman, Brother (Seringer). 1953. Proverbial Expressions of the
Samoans. Polynesian Society Memoir, Vol. 27. Wellington, N.Z.
Pratt, Rev. G. 1862. Dictionary of the Samoan Language. London

Missionary Society. 223 pp.

Sesapasaro, H., 1978. News Bulletin, Nov. 1, 1978. Office of Samoan
Information, Pago Pago, American Samoa.

Ww»

a1IS A8AINS

aBpe joay

sj} Ndeyebuoy

L

ae
ee Sep ENS
V

-— oe
= —--—-—---—-

DISTRIBUTION AND ABUNDANCE OF THE CROWN-OF-THORNS STARFISH
(ACANTHASTER PLANCI) AROUND TONGATAPU ISLAND, TONGA

by M.P. Francis!

Introduction

Acanthaster planci infestations have been reported from many
locations in the South Pacific Ocean but Tonga has attracted little
attention because of its isolation. During six man-days of searching
Weber and Woodhead (1970) found Acanthaster "rare to common" on
Tongatapu Island where "common" was defined as 6 to 25 starfish
observed per man-day. Endean and Chesher (1973) cite personal
communication with the Tongan Department of Agriculture in 1970
regarding a proposed Acanthaster control programme. The programme,
however, never proceeded. The present survey covers the situation six
years later.

Method

The coastline and several reefs to the north of Tongatapu were
surveyed between 16.4.76 and 16.6.76 by snorkel diving (see Figure l).
Survey site N was searched at depths between ten and twenty metres
using SCUBA equipment. All observed starfish were recorded and
intensive searches were made in the immediate vicinity of any feeding
scars. At each site a subjective estimate was made of the abundance
o£ live hard corals.

Tongatapu Island is surrounded by a wide shallow fringing reef.
On all but the northern (sheltered) side of the island the reef edge is
raised to form a barrier one to two metres above the level of the reef
erates Prevailing south-easterly winds hindered searches outside this
barrier, and survey sites F to M were therefore situated on the reef
flat.

liatenisi Institute, Nuku'alofa, Tonga; present address: Joint Centre
for Environmental Sciences, University of Canterbury, New Zealand.
Atoll Res Budi. WNow 255) 63-66, 198i.

66

Results

Survey Live Coral Acanthaster!
Site Abundance Relative Abundance

Abundant
Common
Rare
Absent
Abundant
Common
Rare
Rare
Rare
Rare
Absent
Rare
Common
Common (10-20 m depth)
Common
Rare
Common
Rare
Abundant
Common
Common

(oa)

(on wo

CHNUDMOVOZEZEHAGUHTOANAHMUAWD
Sic

© Oo © 0 O' © Uiiwi' © © OFOrO © CO © OC O10 OO

lRelative abundance is the number of starfish seen per
twenty minutes searching (see Pearson and Endean, 1969).

Discussion

No Acanthaster were found on the coast exposed to the prevailing
winds (sites F to M). Coral growth is inhibited inside the barrier by
exposure to air at low tide, and outside the barrier by heavy surge.

The reef outside the barrier drops slowly to a shelf at ten metres, but
live coral is stunted and scattered above this depth. Acanthaster may
occur at greater depths. The absence of Acanthaster from exposed reefs,
and their preference for sheltered reefs, is consistent with the
observations of Dana, Newman and Fager (1972) who found the starfish
usually associated with "leeward seaward reefs" and "areas of moderate

to luxuriant coral growth".

More than 70% of the sites examined had no Acanthaster and the
maximum relative abundance was 5.3. Pearson and Endean (1969) consider
a relative abundance less than ten to constitute a normal population.

By this definition Tongatapu has normal numbers of the starfish, and

the presence of luxurious growths of Acropora hyacinthus and Porites
spp. in sheltered areas suggests there has been no infestation in recent
years.

67
References
Dana, T.F., W-A. Newman and E. W. Fager (1972). Acanthaster

aggregations: interpreted as primarily responses to natural
phenomena. PBC Sis Ze SS5-37A0

Endean, R. and R.H. Chesher (1973). Temporal and spatial distribution
of Acanthaster planci population explosions in the Indo-West
Pacific Region. Broil COns CiryVs— is lI De

Pearson, R.G. and R. Endean (1969). A preliminary study of the coral
predator Acanthaster planci (Linné) (Asteroidea) on the Great

Barrier Reef. Fish. Notes Dept. of Harbours and Marine 3: 27-55.
Weber, J.N. and P.M. Woodhead (1970). Ecological studies of the coral
predator Acanthaster planci in the South Pacific. Mar. Biol.

6a" IASI 6

RATS AS AVIAN PREDATORS: DISCUSSION

by W.R.P. Bourne!

Since F.I. Norman concludes at the end of a review of the role of
"The murine rodents Rattus rattus, exulans and norvegicus as avian
predators" (Atoll Research Bulletin No. 182, 1975) that "it appears
that the rats' role as an avian predator has been overestimated... the
basic facts should be established before widespread control campaigns
are undertaken", and this conclusion may be used as an argument for
procrastinating over such campaigns, it should perhaps be made clear
that many people have no doubt at all about the harm caused to birds by
rats, and will refuse to accept such a cautious conclusion for a moment.

The situation is of course complicated, and the literature relating
TO) ALG WE\SIES Rats were introduced to many sites where the worst damage
has occurred long ago, before reliable witnesses arrived, and those
that were present commonly reacted by introducing cats, owls or other
predators, which confused the picture. Rats do most of their work by
night, and since it only takes them a few seconds to remove an egg or
bird it may be hard to obtain direct evidence of their activities.
They are inveterate scavengers, and may take abandoned eggs or birds
which died of other causes. The extent to which they prey on other
animals may vary with the relative size and numbers of the rat and its
prey, the availability of alternative foods, and the occurrence of a
seasonal climatic regime liable to lead to restrictions on the food-
supply available to the rats at some seasons which limits the numbers
able to act as predators when birds are present at other seasons.
Their activities may also affect the ecology of other species indirectly,
by competition for food, or through damage to the habitat, for example
by eating the roots of tussock grass on subantarctic islands. Ie, abs}
dangerous to make facile generalisations about the situation except to
say that a legion of witnesses have now reported for centuries that
their presence has almost invariably been considered harmful.

The situation may be illustrated by taking a few examples:-

dhe An exceptionally rich marine avifauna breeding on Amsterdam Island
in the southern Indian Ocean may have been partly exterminated by hogs
and cats at the beginning of the last century, but most of the
surviving bones were found in rat-holes (C. Jouanin and P. Paulian,
Proc. XII Intern. Orn. Congr. 368-372, 1960).

IDepartment of Zoology, University of Aberdeen, Aberdeen, Scotland.
Atoll Res. Bull. Noi/255: 69-72, 19811.

10

Oe Five native birds, Gerygone insularis, Rhipidura cervina,
Zosterops strenua, Apolonis fuscus hullianus, and Turdus xanthopus
vinitinctus disappeared from Lord Howe Island within twenty years of
the arrival of Rattus rattus in 1918 (see especially a symposium on the
island in Australian Natural History 18(2) for June 1974).

By. The havoc caused by rats among both land and sea birds in the New
Zealand area is documented by a series of contributions from the
Wildlife Service and Royal Bird and Forest Society to the forthcoming
Proceedings of the XVI World Conference of the International Council
for Bird Preservation. Norman is wrong in identifying the muttonbird
virtually exterminated on Stewart Island as Puffinus tenuirostris since
it was Puffinus griseus; the endemic Southern Saddleback Philesturnus
c. carunculatus which he mentions was threatened when Rattus rattus
reached the outer islets in 1964 was only saved by emergency
transplantations elsewhere, while the local race of bush wren Xenicus
longipes variabilis and snipe Coenocorypha aucklandica iredalei may
have been lost entirely.

4, While the petrels Pterodroma cahow and Puffinus lherminieri do
appear to have been eliminated from the main Bermuda islands and
confined to outliers by the activities of rats among other predators,
and rats may take occasional tropic-bird Phaethon lepturus eggs there,
this exceptionally aggressive species has in fact managed to hold its
own in the face of attacks by not only rats and cats but tens of
thousands of people. Norman quotes R.C. Murphy (Oceanic Birds of
South America, 1936) incorrectly in attributing the rat predation to
Rattus exulans, since this was in fact a repetition of the original
report by Gross (Auk 29:49-71, 1912) that he had seen one egg taken by

Rattus rattus. Rattus exulans does not occur in the North Atlantic.
Sy The impact of rats on birds in the British Isles is extremely
variable. They are important predators on many species but especially

those nesting on the ground inland where rat numbers are high, though

it is not always easy to distinguish their work from that of other small
mammals. They exterminate storm petrels wherever they go, but their
impact on larger burrow-nesting birds varies. They exterminated a
large Puffin Fratercula arctica colony on St Tudwal's Islands, North
Wales, and severely reduced their numbers on Lundy, Puffin Island (also
north Wales), and Ailsa Craig without exterminating them, whereas a
large colony long continued to flourish in the presence of rats on the
Shiant Islands. They exterminated the Manx Shearwater Puffinus puffinus
at its type locality on the Calf of Man, and it only recolonised when
they were controlled by poisoning. Small numbers survive in the
presence of rats at a number of other sites, a larger colony suffers .
severely from predation in some years but not others on Canna, and a
vast one in the hills of Rhum appears to escape it. Terns suffer
severely in areas heavily infested with rats but may escape their
attention elsewhere (references for some of these sites will be found
ins. Cramp, W.R.P. Bourne and D....Saunders,.. "The Seabirds of Britain

and Ireland", 1974).

fe

The situation appears to have been particularly bad on many oceanic
islands because there were no natural predators or inclement seasons to
control the numbers of introduced rats, and the native wildlife had no
innate defences against them. The petrels which bred there were
particularly vulnerable because they leave their small, defenceless
chicks alone in their burrows by day soon after they hatch. It will
be noted that while Norman found that rats took the unguarded eggs of
Short-tailed Shearwaters Puffinus tenuirostris but avoided encounters
with the large, aggressive adults (J. Zool. 162:493-503, 1970) he did
not investigate what occurred when the rats encountered small chicks.
While the situation may often have been made worse by the introduction
of alien predators in an attempt to control rats, there is an obvious
correlation between a decline of the smaller and more defenceless
seabirds in historic times and the introduction of rats throughout the
world. However, I agree that it remains difficult to assess the full
amount of harm they have caused because it is impossible to say how
many insular landbirds they have exterminated as well. A rat may only
need to eat two or three eggs or chicks a year to cause serious damage,
whatever it eats the rest of the time.

ee

LAYARD'S BIRD HUNTING VISIT TO TROMELIN OR SANDY
ISLAND IN DECEMBER 1856

by R-K. Brooke!

This paper concerns the only known visit by a biologist to Tromelin
Island in the XiX century.

That Edgar Leopold Layard, the Curator of the South African Museum,
Cape Town, travelled as naturalist on the British Royal Navy survey ship
H.M.S. Castor on a journey in the southern Indian Ocean in 1856 and 1857
was scarcely known at all until Brooke (1976) commented on some aspects
of the journey and the bird records made on it. Subsequently Brooke
(1978) reported on those birds' eggs collected on this journey which
still survive in the South African Museum. He also deduced the
approximate route and timing of the journey from the species collected,
their known breeding seasons and migrations. As will appear below,
these deductions were close to the truth. Since then I have had cause
to examine in the South African Library, Cape Town, a run of the Cape

Monthly Magazine (Series 1) published in Cape Town between 1857 and 1862.

This magazine attempted to cater to the interests of the more educated
residents of British stock by publishing scientific, philosophical and
(EK myAmMace aia.

In volume I of the Magazine the editors caused to appear on
pp. 252 - 254 a notice entitled "Mr. Layard's cruise in the "Castor'."
From this we learn that H.M.S. Castor left Simonstown on 10 October,
1856 under sail under the command of Commodore H.D. Trotter. They
intended to land first at Rodriguez but extensive fog persuaded them to
head for Mauritius where they spent five weeks and made several trips
to nearby Round Island. Sandy Island as Tromelin Island was then often
known (Staub 1970) and the focus of this paper was the next landfall
(presumably in early December 1856). Then Farquhar Island was visited
prior to reaching Africa at Fazi Island, Kenya, on 4 January 1857.
Thereafter, Lamu, Melinda, Kisilundini, Zanzibar, Cabo Delgado and Dibo
were visited before they put out to sea for a crossing to northwestern

Madagascar via the St Lazarus bank and the Comoro Islands. Anjouan
was reached on 1 February and a few days later a visit was made to
Moheli. Two days were spent in Madagascar at Boyana Bay at a Jesuit

Mission before recrossing the Mozambique Channel to Mozambique Town.
After a substantial visit they set sail for home via Durban (departed
13 March), East London, Port Elizabeth (departed 19 March) and so back
to Simonstown on 25 March 1857.

lPitzPatrick Institute, University of Cape Town, Rondebosch 7700

This paper is part of the commemoration of the 2lst anniversary of the
establishment of the Percy FitzPatrick Institute of African Ornithology

Atoll Research Bulletin No. 255: 73-82, 1981

74

Layard promised to write a full account of his journey and his
collections for the Magazine but he did not do so, probably through
pressure of work: he was only the part-time curator of the Museum
being a nearly full-time civil servant as well. However, in 1858 in
vol. III, pp. 289 - 296, appeared the following from Layard's pen:

SCRAPS FROM THE NOTE-BOOK OF A NATURALIST

BY E.L. LAYARD

"LAND, HO!" hailed the look-out man from the foretopsail yard
of Her Majesty's ship Castor, as one lovely morning the Commodore
and myself paced the deck of the good old frigate.

"Whither away?" went up the responsive query from the midshipman
of the deck, who, stepping up to the officer of the watch and
touching his gold-banded cap, reported the fact. In his turn, the
officer reported to the Commodore; and, after a few questions, the
order was given to head the vessel to the land, and we resumed our
walk.

The low shores of the island now became visible, and the clear
blue sky was dotted with birds, winging their way to and fro; some
plunging headlong into the rippling water, that vied in the intensity
of its azure with the sky above it, sought their finny prey, some
returned to their nests heavy laden with food for their young, while
others floated contentedly on the gently heaving bosom of the sea, or
rose lazily out of the path of the frigate, as she majestically
ploughed her way along, her white sails gleaming in the bright
sunlight. One by one the studding-sails came home, and the noble
ship cautiously felt her way to the rapidly rising land; and now,
while the boats are told off, we descend to have a last look at
Horsburgh's Directory, for landing instructions, hidden dangers,
currents, -cte:

For the benefit of such of my readers (and I suppose they are
numerous) who do not know where Sandy Island lies, I transcribe
the account given by the great hydrographer of its position and
appearance: -

"Sandy Island, or L'Iisle de Sable, in latitude 15° 52. s.,
longitude 54° 40' E., is a flat, sandy spot, about fifteen feet
above water, half a mile long, from N.N.W. to S.S.E., and about a
quarter of a mile broad, having a sand-bank projecting three
quarters of a mile towards the S.S.E. It was discovered by the
ship La Diane, in 1722, and in 1761, the Flute l1'Utile was wrecked
there."

1)

After the men had got their dinners, the boats were reported
ready, and the first cutter, attended by thedinghy to assist in
landing, were soon off from the ship's side, well furnished with
baskets, guns and ammunition, harpoons, fishing-lines, etc. The
vessel had stood up as near the island as the light breeze permitted
with safety, and on quitting her, she tacked and stood off, the
Commodore trying for soundings (though finding none with one hundred
fathoms of line), and practising reefing topsails and other
manoeuvres, to exercise the men. The boats pulled cheerily along,
the men eager for a run on shore and a supper of turtle steaks, the
sportsmen anticipating no end of sport, and myself luxuriating in
the idea of visiting the breeding place of the countless frigate
birds, gannets, terns, and other birds that now filled the air with
their piercing cries, and darkened the sky over the island.

As we reached the coveted shore, the water suddenly changed
from blue to green, and presently the surf line showed itself,
thundering on the sandy beach. The dinghy being signalized to
pilot the way, the two Kroomen who rowed her, amphibious fellows
equally at home in the water as on the land, urged their frail
skiff in advance, and we soon descried a spit of land, under the
lee of which the water lay comparatively smooth, and there we
determined upon effecting a landing.

The dinghy soon lay high and dry; but our attention in the
cutter was diverted by a tempting object that floated on the calm
water, a few dozen yards ahead, and which we all, at once, and
with an instinct worthy of city aldermen, knew to be turtle soup
in its raw material. An old whaleman now crept forward to the
bows and drew out the glittering harpoon, the boat's crew pulled
noiselessly, the officer in charge gave his orders in whispers,
and we stole on our prey. "Ah! old fellow, how many basins of
soup will you make?" was in many a mind, and "What a splendid shell
for the Museum," in one at least. Look at the harpooner, he stands \
upright, one foot on the gunwale, the barbed weapon poised; half-
a-dozen strokes and he is — No, not a bit of it, — wide-awake is
Master Turtle; and without a ripple, down sunk the huge bowl of
soup and we saw him no more. |

Disappointed with the failure of our first essay in turtle-
catching, we turned our boat's head to the shore, and pulled in,
avoiding with sailor's skill the huge rollers that ever and anon
broke in masses of white foam on the beach. We ran in on the top
of a wave, backed and let it break, and then pulled in. Out
jumped the men who were appointed to run the boat up, and to our
dismay, though only a few yards from the dinghy, our fellows found
no bottom. Before they could recover the confusion into which
this untoward accident threw them, and the men in the boat had
recovered their oars, a huge surf broke right over us, washing two
men Out of the boat and throwing her on her beam ends. I held on
to the guns and the benches, the rest made ready to spring out at
a favorable moment; fortunately, the next wave had a "long-shore"
course, and though it threw us about like a nut-shell, it drove us

76

into shallow water. The men still clinging to the gunwale found
footing; the rest of the crew sprang out, and the next moment saw
us in safety, though at the expense of a good ducking, and, on my
part at least, not being a swimmer, of a good fright.

After wiping our guns and spreading our superfluous clothing
on the sand to dry, we walked up to what we at once saw to be the
remains of a human habitation. We found the ruins of a hut, which
had contained three rooms; in the largest was a square enclosure
of blackened stones, which had formerly served for a fire-place;
but which now held the nests of three or four noddies (Anous
Stolidus), who, utterly heedless of our approach, pecked furiously
at our hands when we attempted to possess ourselves of their lovely
spotted eggs. We found the nests of these birds in every hole and
corner of the ruins; they had taken possession of every stone, and
while the male bird sat perched on the top, the female covered her
solitary egg by the side in a depression in the soil, lined with a
few seaweeds. The eggs were beautifully spotted with patches of
various shades of a light purple on a delicate cream-colored ground.
An average sized egg measured two inches by one and a half inches;
but hardly two were of the same size, or colored alike; still, a
practiced eye could at once separate them from that of Onychoprion
Fuliginosus, which we found breeding on another part of the same
island.

From the ruins, we walked to the east side of the island,
which appeared covered with bushes about the height of a man's
head. Up to this time, the frigate birds had kept well out of
range; but now a rustling sound was heard above us, like the
falling of some heavy body through the air, and glancing upward, I
saw a frigate bird darting down as if to attack us. As he swept
past, I leveled my gun at him, fired, and he rolled over; and now
the guns had plenty of work, for the birds poured down, and the
ground was soon strewed with dead and dying.

How many of the wished-for birds we might have slain, I know

not, when one of the men called out, "Hold hard, sir, — don't
shoot any more, — here they sit by scores on their nests." And
now the truth flashed on my mind, — we had intruded on their

"rookery," and the poor birds had lost their lives in their vain
attempts to intimidate the two-legged monsters that had suddenly
disturbed their domestic privacy. Eager to possess myself of their
eggs, I dislodged bird after bird from her nest, by the aid of a
long stick, — to approach one's hand, was the sure precursor of

a severe bite. Every one, almost, had a callow nestling under her
maternal wing; and I obtained but few eggs in a fit state for
blowing.

The Tars, who roamed about, found one place where the
incubation seemed not so far advanced, probably the nursery of the
new-married and inexperienced couples. Jack seized the prize,
and though the eggs were in most instances sat upon, they were all
devoured: so much for taste! I heard one fellow say to his chum,

1

as he gulped one down, "Dang it, Ned, there was a bone in that un."

The frigate birds (Attagen Ariel and A. ,) and the
gannets, of which there were three species, — Sula Fasciata,
S. Personata, S. Fusca, bred side by side, in patches. Their nests

were huge structures of sea-weed, dung, and fishbones, their
stratified appearance testifying that each successive year added

to their size. The stench from this spot was dreadful, the ground
being strewed with the debris of fish and young birds. I could
not account for the numbers of the latter, till I saw a huge red
and hairy hermit crab (Pagurus), inhabiting the dead shells of a
large Turbo, which lay scattered in great abundance on the ground,
deliberately descending the trunk of one of the bushes, with a
writhing squab in his claws. I then saw that the branches were
full of these cannibals, mostly laden with fish stolen from beside
the nests. I presume, the robbers only occasionally manage the
more dainty morsel of a tender chick, secured when the mother is
absent from the nest, though doubtless sufficiently often to keep
in check the rapid growth of the birds, — robbers in their turn,
for from the mouth of one I shot was disgorged no less than seventeen
fish, from three to nine inches long.

After forcing our way through the bushes, we emerged on the
side of the island, opposite to that on which we landed, and found
a reef, or ledge of coral and rock, extending along the shore, at
the outer edge of which the surf broke in fearful splendour.

A hail from one of the men in advance, and the clustering of
the lads as they reached the spot, induced us, who were leisurely
advancing with prying eyes, to hasten our steps. We found, on
arriving at the scene, that the men were busily engaged in examining
two or three heavy iron guns, that lay half buried and jammed in
between the rocks, and to seaward lay the timber of a vessel, with
her huge anchor still with one fluke in the reef. These, then,
were the remains of the ill-fated L'’Utile, the French man-of-war
alluded to by Horsburgh: the house now tenanted by the wandering
sea-fowl had been the home of the survivors during their long
imprisonment on this speck in the ocean; the fire-place in the
large room had witnessed the gay laugh of the thoughtless, the
bitter gloom of the despairing, and the high and manly thoughts of
the undaunted and brave, each, perhaps, in his own way, pondering
over the means of escape, or the question of to-morrow's food.

The commander sat in the little room, overlooking the remains of
his lost vessel: the officers crowded in the other room, and
thought of those loved ones they might never see again; and now,
how many of that band of men survived? Not one! and the place
of their long sojourn would never again hear the sound of their
voices.

We pushed onward to the northern end of the island, and
crossed an open plain that lay between the bushes and a high bank
of rolled stones, cast up by the storms of ages on that end of the

78

island. In the centre of this plain, we came upon a circle of
stones, placed round a spot, whose vivid green attested to the
presence of water beneath: this was the place from whence the
shipwrecked men drew their supplies. Near this spot we could
hardly tread for young terns (Anous Stolidus) and eggs, and on
turning over a stone, out sprang two mice; another and another
followed from every stone moved, the place literally swarmed with
them. They had probably come in the French ship, and had peopled
the island. How to account for the presence of the vast quantities
of huge black ants, that ran their galleries in all directions
under the sward, was a more difficult matter; as also was the
advent of a lizard (Gecko) that I captured, but which subsequently
made his escape.

Advancing a few yards from this spot, we entered on the domains
of another species of tern (Onychoprion Fuliginosus), of which we
were first made aware by the fearless birds striking at our faces

with their sharp pointed bills. As in the other places, we had
not seen a single nest of this species, so now we saw not a nest
or specimen of A. Stolidus. The noise of the congregated numbers

was so great that we could barely hear the shouts of our nearest
neighbor, and I was glad to escape over the high ridge of stones
and gain the quiet of the dashing surf.

This is evidently the stormy side of the island, and the
direction of the prevailing wind. The whole shore consisted of
large rolled stones; not one of which possessed an angle; all as
smooth as a cobbler's lap-stone, they irresistibly brought to mind
hundreds of jolly Crispins busily tapping soles and welts. Here
we found another wreck — a huge tree; from whence had this
floated? Madagascar probably, and was the ark in which had come the
ants, and the gecko; so are those ocean dots peopled.

Turning from the scene of speculation, we wended our way back
to our boats along a broad road, cleared of every stone, and nicely
smoothed. For what could the poor refugees have constructed this
road? Perhaps their commander, a wise man, reflecting that
idleness excited gloomy thoughts in such situations, had set his
men to work to clear this road, to convey — what? Firewood from
shipwrecked trees, stones thrown up by the waves to build their
house, — what! as a carriage road it had certainly been used, for
we found the broken wheel of a cannonade lying on a heap of stones.

While walking here, I had given my gun to the Chaplain, who
wanted to try his hand at a shot. He had fired once and
unsuccessfully, at an oyster catcher (Hoematopus); and now to my
vexation, I saw a new lovely snow white bird, much resembling a |
tern, slowly passing him. "New bird!" I shouted; "shoot, parson,
shooti" Alas, my friend was a better hand with his Hebrew books |
than with Westley Richards, and that eminent individual's death- 1
dealing tubes were leveled in vain at the lovely stranger. To |
the bright flash and loud report, the snowy creature made a

19

graceful stoop, like a damsel in white muslin at an invitation to
dance, and it floated over our heads and looked us full in our
faces with its large black eyes. I was frantic: eagerly I
crammed in a charge of powder; but alas, no shot had I, my fidus
achates, “Ned," and "curio-man's Jack," as the sailors had dubbed
him and me, was off to the boats with my pouch. But help was at
hand, — the crack-shot of the ship, Lt. ----- , was hastening to

the scene, attracted by my shouts and vain attempts to knock the
bird over with stones. Seeing another of the queer monsters
running up, my white friend turned away, and was winging his course
sea-ward. Oh agony! only known to the naturalist, who sees a

new species slip away from his grasp. Inwardly, I resolved never
more to trust my "Shooting-iron" out of my hand on these unknown
coasts. The white vision took another turn this time, land-ward —
"yun, run! — new bird! — long shot! — ah, my beauty, the parson
does not hold that steady barrel: a deadlier eye is measuring the
distance which you will fly over, ere the leaden messengers will
reach you." And now, the sharp flash! — plump down falls the
hapless bird. The reverberating report reaches my ears, and I
dashed forward and the prize is mine; and oh, how beautiful the
pure white plumage, without a speck, save the one pink spot on

the breast, from whence oozed the ebbing life blood, the brilliant
blue bill and the large dark eye now closing in death! And now,
quam mutatus ab illo! its dry and shriveled skin and blackened

bill grace the cases of the Museum, — a mockery of life; a |
burlesque on the loveliest bird I think I ever saw. Reader, you
may see it labeled Gygis Candida.

The eggs of the Onychoprion Fuliginosus are rather larger
than those of A. Stolidus, being about 2 inches and 2 lines long,
by 1 inch and 7 lines broad, and more thickly dotted with smaller |
dots at the obtuse end. They are also generally darker, and
may easily be distinguished by any one who has ever taken the two,
though a written description would suit for either species. |

The eggs of the frigate birds, Attagen Ariel and A. ;
are very Similar in size and shape to those of the gannets; but
are smoother and thinner in texture, and free from the thick
incrustation of lime which at once distinguishes those of the
latter. They are both pure white when fresh laid; but get sadly
soiled amidst the filth in which they lie. They measure about
2 inches 7 lines long, by 1 inch 9 lines broad.

Just as we commenced our arrangements for the night, the old
frigate which had been working up for the island, suddenly backed
her topsails, ran up her recall flag, and fired a gun. Jumping
into the dinghy, I went on board to ascertain the reason of this
change of plan, while the cutter was launched and preparations
made to return, if the recall should prove general.

"What sport?" shouted the Commodore, as we got within hailing
distance. "Glorious:" was my reply. "What likelihood of
turtle?" was the next question. "Very little! no marks on the

80

land of the breeding ones." The little dinghy shot under the
tall sides of the ship, when again thundered her signal gun, and
the cutter's recall, in lanterns, ran up to her mast-head; and
just as total darkness fell on us, the cutter dashed along side,
and in another moment was swinging at her davits; the old ship
filled her sails, bowed to the night breeze, and we stood off from
the island.

"T don't like the current hearabout," said the Commodore, as
I finished my narration of our afternoon's adventure. "We must
not lay the Castor's bones alongside those of L'Utile."."

The rest of what Layard did, collected or saw on that journey is
lost save for the fragmentary allusions in his 1863 paper, his 1867
book (brought together in Brooke 1976) and what little can be learnt
from the surviving eggs discussed in Brooke (1978).

It is possible now to present a picture of the bird life on
Tromelin Island as it was in December 1856, 98 years before Brygoo's
(1955) visit, believed to be the first ornithological visitor by

Staub (1970). As Mr. A.S. Cheke has pointed out (in litt.) Brooke
(1976, 1978) was mistaken in assuming that Layard's Sandy Island was
the Sandy Island off Rodriguez. The Rodriguez records in those

publications were all made on Tromelin Island (Layard 1858).

Sula dactylatra: a youngster and adults brought back to Cape Town
(Brooke 1976), younginthe nest (Layard 1858 sub nom. S. personata).
It still breeds here (Staub 1970). The nest site fidelity described
by Layard should be noted.

Sula sula: a pair and two youngsters brought back to Cape Town
(Brooke 1976), young in the nest (Layard 1858 sub nom. S. fasciata

for normal birds and S.fusca for dark phase birds). I know. of no
other usage of S. fasciata, let alone who proposed the name if, indeed,
it is not just a misconception in Layard's mind. It still breeds here
(Staub 1970).

Fregata ariel: adults collected, most nests held young but a few held
eggs, usually well incubated (Layard 1858). Staub (1970) is uncertain
whether it still breeds on Tromelin.

Fregata minor: as for F. ariel (Layard 1858). It still breeds here
(Staub 1970).

Haematopus ostralegus: shot at but missed (Layard 1858). Not recorded
on any Indian Ocean island by Watson et al. (1963) and there are no
subsequent records (A.S. Cheke in litt.).

Larus cirrocephalus: an adult brought back to Cape Town (Brooke 1976).
He gives reasons for not rejecting the record even though the specimen

no longer survives and there are no later records.
a vagrant from Madagascar nearly 400 km to the west.

Presumably it was

Sterna fuscata: eggs collected (Layard 1858 sub nom. Onychoprion
fuliginosus) . Not recorded by Staub (1970) whose visit was in late
August when the birds would not have been breeding.

Sterna dougallii: Staub (1970) suggests that the terns described by
Morris (1964) as "with light grey mantles and black crowns" were this
species. They might equally have been S. bergii thalassina since no
indication of size is given. Layard (1858) makes no suggestion that
either species was present during his visit.

Gygis alba: an adult brought back to Cape Town (Brooke 1976). IEIE
does not appear from Layard (1858) sub nom. Gygis candida that it was
breeding at the time of his visit. Not recorded by Staub (1970).

Anous stolidus: a youngster and adults brought back to Cape Town
(Brooke 1976) as well as eggs (Brooke 1978). They had both eggs and
young in their nests (Layard 1858). Not recorded by Staub (1970).

Anous tenuirostris: adults brought back to Cape Town (Brooke 1976) but
it does not appear from Layard (1858) that he realised that two species
of Anous were present. Not recorded by Staub (1970).

As for the fauna other than birds it appears from Layard (1858)
that Mus musculus was by then well established and it has remained so
(Staub 1970) . Layard's belief that they had come from the wrecked
L'utile is likely enough. He does not mention Rattus norvegicus, the
other well established rodent noted by Staub. Hermit crabs Pagurus
sp. were common both in the 1850s and 1950s: Layard's remarks on their

predation on the young of Sula dactylatra should be noted. Layard's
gecko (Reptilia, Gekkonidae) was probably, as he said, a drifted
vagrant since there is no mention of lizards in Staub. I am unable to

equate Layard's (1858) "huge black ants" with Skaife's (1953)
description of Pheidole megacephala, the only ant recorded by Staub
E97.)

References
Brooke, R.K. 1976. Layard's extralimital records in his Birds of
South Africa and in the South African Museum. JatolILihig Jehaulian “(Okaalc

Club 96: 75-80.

Brooke, R.K. 1978. XIX century Indian Ocean seabirds eggs in the
South African Museum. Buel Brae Orn CLUDE IO aio COE

BY GOO yEie LoD Ds Observations sur les oiseaux de Tromelin. Nat.
Malgache 7: 209-214.

82

Layard, .E). le U858r Sport on Sandy Island. Cape Monthly Magazine
ser, Vly vols 32549289-=2968

Layard, E.L. i863. Ornithological notes from the antipodes. Ibis
863: -241—2 50

Layard, Ee be L867! The birds of South Africa. Juta, Cape Town.
Morris, R.O. 1964. Tromelin Island. Sea Swallow 16: 76-77.
Skai fei='SoHe “19538 African insect life. Longmans Green, Cape Town.

Staub, H-L970- Geography and ecology of Tromelin Island. Atoll
Res. Bull. 136: 197-209.

Watson, GtE2, -Zusit? ReLe & Storer, eRowW.) 1963: Preliminary field
guide to the birds of the Indian Ocean. Smithsonian Inst.,
Washington.

A SUBMERSIBLE, RECHARGEABLE, BLECTRIC DRILL

by W. H. Easton!

Abstract

Melight, werghit,. condliess; = rechargeable, cllectric hand drill
mounted in a submersible plastic case is linked by ashaft through
themhousing) to “an external bit extensions Tungsten carbide masonry
bits or small diamond core drills can be attached to the end of the
bit extension without opening the underwater housing. The assemblage
has been used to set survey points used in measuring shoreline erosion
and to take core samples for radiometric dating and for determining
growth rates of corals by X-radiography. Other applications have
been suggested.

Design

The prototype model (Figure 1) was designed to accommodate the
smallest rechargeable drill in order to reduce weight and volume taken
on flights to distant islands. A submersible housing was constructed
of 3/8" and 1/2" acrylic plastic to fit two inter-changeable models
ofdrills — the Craftsman Rechargeable 1/4" Model No. 135.111100, and
Skilshop Model 1702 Type 1 Cordless 1/4" Reversing Drill. The
assemblage weighs 3 kg (6.7 lbs) and will float. General information
and materials used in the construction of plastic housings can be
obtained from hobby shops, plastics salesrooms, and Toggweiler (1970).
Prefabricated control glands with O-rings, and levers, shafts, and
stud retainers with studs and nylon wing nuts for closing housings
are available from some dive shops and Toggweiler.

A few special requirements must be met in designing the apparatus.

The acrylic plate in front of the drill chuck must be at a right angle
to the axis of the chuck, because the control gland containing an
O-ring through which the shaft of the bit extension passes also must
be aligned perfectly with the axis of the chuck after the gland has
been cemented to the front plate of the housing. The rear of the
drill case must just touch the inner surface of the rear plate of the
housing so that a thrust applied to the housing by the operator will

IDepartment of Geological Sciences University of Southern California
Los Angeles, California, USA 90007.
Contribution No. 371. Department of Geological Sciences, University

of Southern California, Los Angeles, California, 90007
AGOLL RES = Bull, Nog 255% S390, Weld.

8h

O./39"=3.5.3mM SOLID ROUND NEOPRENE
EXTRUSION IN ROUTED CHANNEL

S/16"= 8mm

COVER, INSIDE

61/4" *+/5.8CM
5 /a"=/4 850M

21/4 "= 57M

BASE, INSIDE

FRONT PANEL, PROFILE

: 7 ttached.
; : ith bit extension a
ible housing wit
Bugure: 1. Submers

85

Tapped for 10-32 set screw— 2

Burnished surfdce

kW

%

Silver soldered joint

Diamond Core Bit : |

Figure 2. A. Specifications for the bit extension.
B. Specifications for the diamond core bit assemblage.

86

be transmitted to the electric drill. Lateral wobble of the drill
handle is controlled by the oval receptacle in the base and by cushions of
vinyl foam cemented to the two side walls of the housing and to the
inside of the cover. The chuck key is stored inside the housing.

A short bevelled piece of shaft brazed to the long trigger release
shaft at a right angle provides the contact with the trigger switch.
If the case is designed with the greatest economy of inner space, the
trigger release shaft will be too long to be inserted through the
control gland from inside the housing. In this event a small hole
can be drilled laterally through the plastic trigger on the drill and
a thin diameter screw can be threaded through the hole and extended
about 3/8" (=9.5 mm) to the right side where it will be engaged by the
bevelled flange brazed on the control shaft. No provision was made
for external activation of the on/off safety switch or the reversing
Switch on the motor case.

The bit extension (Figure 2) is fabricated from stainless steel
stock, including the two set screws. The shaft must be 1/4" in
diameter in order to pass through the prefabricated control gland.
However, the hole for the reception of the shank of the masonry bit or
the diamond core bit may be of different diameter, depending upon the

size and brand of masonry bit. Some bits are of uniform diameter from
the tip, along the twist, and on the shank. Others are of smaller
diameter behind the tip, so the bit must be selected before the
receptacle in the bit extension is bored. Furthermore, the shank

of stainless steel brazed on the diamond core bit must be the same
diameter as the shank of the masonry bit if these tools are to be
interchangeable. Machine shop experience indicates that it is
advisable first to turn the 1/4"-deep recess on the shank assemblage
and braze it into the rear opening of the diamond core bit before
machining the 1/4" shank. This ensures that errors of alignment
causing wobble when the apparatus is operated will be eliminated.

The diamond core bit is Type SICD, Serial N. 55955, without
collet, 7/8" 0O.D., available through Felker Operations, Dresser
Industries, Inc., 1900 South Crenshaw Boulevard, Torrance, California,
90509. Other core bits are available in sizes as small as 1/8" and
larger than the one selected, so there is a wide selection of sizes
enabling a choice of bits calculated to provide the volume of material
required. In ordering any brand of core bit one must specify that
the bit must not have anything fastened to the open end opposite the
cutting edge.

Uses

The apparatus was designed to facilitate collecting coral samples
for three lines of research being conducted on various islands in the

Pacific Ocean. Growth rates of some corals (for example, species of
Porites) can be determined by X-radiography of slices taken normal to
the direction of growth (Knutson, Buddemeier, and Smith, 1972). The

length of the cores (about 7.5 cm or 3") and the facility with which
variously oriented and positioned samples can be obtained are ideal for

thats line or esearch. Growth rates also could be determined by the
228Ra, 90sr, and 7!°pph methods if two or more cores of 7/8" diameter
were taken, or if a larger core bit were utilized (Moore, Krishnaswami,
and Bhat, 1973).

Cores 7/8" in diameter weigh about 25 grams and contain enough
material for either !*c or uranium series (23 0pp /238y and 234 /238y)
analyses (Barnes, Lang, and Potratz, 1956; Thurber, et al., 1965).
Radiometric dating of samples by these techniques is used to determine
ages of raised shorelines containing corals, ages of certain reefs, and
changes of sea level.

Masonry bits were used to drill holes in coral, limestone, and
several kinds of igneous rocks. Bronze nails driven into plastic
molly fasteners (hollow wall hangers) inserted in the drill holes serve
as reference points against which amounts of erosion can be measured
from time to time alone shorelines.

The fully charged prototype assemblage drilled as many as 20 1/4"
hotesmor, £our Sly cores in coral. Four drill holes or one core one

inch long can be cut in hard, siliceous limestone or in moderately hard
igneous rock. A spare motor was carried in order to accomplish more
work and to ensure against motor failure in remote places. Although
the electric drill lacks the power of the commercial pneumatic drills
and impact wrenches, it has the advantages of light weight, portability,
low cost, and replacement of parts from hardware stores. Furthermore,
it can be recharged almost anywhere that electricity is generated.

Biologists and geologists who observed the drill in operation
suggested additional uses. In the tropics a drill must be insulated
from normal high humidity, rain, and water splashed around boats, even
if it is not used underwater. This light drill could be equipped with
circular core saws or scroll saws to cut plugs of wood encrusted with
lichens, mosses, algae, and other plants. Plugs are transplanted from
one tree to another when studying recovery of vegetation, adaptations
of plant communities, and certain environmental problems.

Attachments such as screw drivers, routers, grinding wheels, mills,
and facing plates could be employed, particularly if an adjustable drill
chuck were added to the bit extension.

Operation and Maintenance

Both the masonry bit and the core bit cut most effectively if the
hnollesmare) “Kept free trom Cuceings). Otherwise grooves in the twist of
the masonry bit become solidly packed with fine debris, preventing the
cutting edges of the bit from biting into any hard ground. The core
bit does not become jammed as readily as the masonry bit. Debris can
be flushed from holes quite effectively using a rubber ear syringe.

It is difficult to maintain a force against a solid object when
thrusting underwater, but bits cut most effectively when a steady
pressure is maintained. Instructions with cutting tools warn against

88

letting them rotate ineffectually against the surface to be cut, lest
the cutting edge of the tool be damaged. Fortunately, the drill is so
light or buoyant that borings can be completed in awkward situations by
holding on to the work surface with one hand and operating the drill
with the other.

Heat created by the motor warms the air inside the housing, causing
a pressure increase. This is a beneficial development to a certain
extent because the pressure differential tends to prevent leakage
around the O-rings in the control glands and the cover. me. als:
possible, although not personally observed, that long-continued
operation of the motor might raise the interior air pressure to
unacceptable levels. Intermittent running of the motor seems to
permit adequate radiation of heat through the acrylic housing when the
apparatus is submerged. It is probable that careful attention should
be paid to the O-rings in cold water, for if air should leak out under
high pressure, then possibly water might bypass the O-rings when the
apparatus cools off. So far I have not observed any leakage in either
direction in shallow water.

The O-ring around the drive shaft is subject to increased wear if
it becomes dirty. Drilling generates fine abrasive material, so it is
beneficial to maintain an excess of thick lubricant such as silicone
grease between the shaft and the control gland. Evidence of wear
appears aS a grey or black stain in the lubricant near the O-ring.
Periodic replacement of the O-ring and grease in the drive shaft
control gland is advisable.

The steels from which both bits are made rust quickly so they
should be given protective coatings of grease when not in use.
Moreover, rust-prone shafts should not be left in the receptacle of
the bit extension.

Modifications

The two models of drills used in the prototype require from 16 to

20 hours for total recharging. However, a later model, Skil Model
2006, is 3/8" cordless drill requiring only one hour for complete
recharging. Furthermore, it has much more torque capability, greater

battery capacity, and other special features, yet it has the same
dimensions as the 1/4" models used in this project and therefore will
fit the same housing.

Other slightly larger models of various brand names may be
provided with removeable battery packs so that spare packs can be
attached to the motor unit. If. two. drills, are tobe provided, 2c
is important to verify their dimensions, because dimensions of the
plastic motor cases of one model are significantly different, yet they
both bear the same model number.

Some persons who have examined the drill indicate that they might
tap the motor with leads that could be fastened to battery packs
either in an enlarged submersible housing or carried externally.

89

Commercial belt packs of batteries are available for under-water
photography and might be adaptable for the drill. In any case it

would be necessary to ascertain the electrical engineering specifications
of any motor being modified.

Acknowledgements

John Wilson fabricated the bit extension and core bit shank.
Anna Dillon drafted Figure l. The manuscript benefitted from
constructive criticism by A. C. Hine and D. Hubbard. This |proyiece
is an outgrowth of National Science Foundation grant EAR 77-13680.

References

Barnes, WieW., Lang, E.J.-, and Potratz, HA. 1956. Ratio of ionium
to uranium in coral limestone. Science 124: 175-176.

Knutson, D.W., Buddemeier, R.W., and Smith, S.V. 1972. Coral
chronometers: seasonal growth bands in reef corals. Science
UWS BUOLPAWAS

Moore, W.S., Krishnaswami, S., and Bhat, S.G. 1973. Radiometric
determinations of coral growth rates. Bulletin of Marine
sezence 23: | 157-176.

Thurber, D.L., Broecker, W.S., Blanchard, R.L., and Potratz, R.L. 1965.
Uranium-series ages of Pacific atoll coral. Science 149: 55-58.

Toggweiler, M. 1970. How to build your own underwater camera housing. |
Hydrotech Co., P.O. Box 14444, Long Beach, California, USA, |
90814. (Prtcesa US $1595).

Addendum

Skil Corporation recently introduced two models of 3/8 inch recharge-
able drills that are interchangeable with the 1/4 inch drill originally
used. The newer drills have much more torque and much faster recharge
times than the 1/4 inch drill. Skil model 2003 can be recharged in three
hours, whereas model 2006 can be recharged in only one hour and also has
a ready light that indicates when charging is complete. The dimensions

of all three drills are identical.

ARTIFICIAL REEFS IN DISCOVERY BAY, JAMAICA

by Michael J. Risk!

Introduction

Many studies have been done of artificial reefs placed in the
marine environment. In some cases, the emphasis has been on
augmentation of nearshore fisheries (for example, see Turner et al.,
1969). In other studies, the motivation has been largely to gain
insight into the ecology and ethology of reef fishes (Randall, 1963;
Luckhurst, 1972).

Present-day coral reefs are not only highly productive in
themselves, but frequently act as a rampart or barrier, protecting
coastal lagoons and bays which in turn may be highly productive (Odum,

U7) e The submarine vegetation in these embayments (usually
Thalassia, in the Caribbean) is typically under-utilized by herbivorous
fishes and invertebrates due to lack of refugia from predators. Where

small patch reefs occur in lagoons, they are commonly surrounded by
"halos" of bare sand from which the vegetation has been completely
grazed. These halos have been attributed to grazing by fishes
(Randall, 1963, 1965; Luckhurst, 1972) and by sea urchins (Ogden
Cea 973) The scarcity of hiding-places in lagoons is probably
a result of the relative inability of corals to attach and grow on
soft substrates in murky water (for a review of factors affecting
corals, see Endean, 1976).

As one of the many unpleasant things which the future has in store
for us is undoubtedly increasing shortages of high-protein food in
Third World countries, many of which are located in or on tropical
oceans, it would be valuable to investigate the possibility of
utilizing more fully the vegetation in shallow coastal areas.
Artificial reefs which attract fish populations may be a means of
essentially converting the high-carbohydrate plant material into high-
protein fish. The total productivity of the system would, of course,
not be changed, but less carbon would be incorporated into the
sediments.

The topography of reefs (both natural and artificial) dictates
that modern high-yield fishing methods cannot be used; the common
Antillean fish pot "is still the most effective way to catch bottom
fish around the Caribbean" (Brownell, 1972, p. 29). Several designs

I Department of Geology, McMaster University, Hamilton, Ontario, Canada

L8S 4M1
Atoll Res. = Budde Noss eo 5c Ol=100s NOS.

92

of trap exist, of which the Jamaican "Z-trap" seems to be the most
effective (Crossland, 1976). Trapping by individual local fishermen
would seem, therefore, to be a reasonably efficient way of "harvesting"
artificial reefs.

As a necessary preliminary, the morphology of various types of
artificial reefs should be investigated, as should the relative cost
(both in money and in labour) of building different types. This paper
is a preliminary report on three such reefs, constructed in Discovery
Bay, Jamaica, W.I.

Methods
(a) Reef construction

The artificial reefs had to fulfil two criteria: the construction
materials had to be locally abundant and relatively cheap, and the
finished reefs had to present a wide variety of available interstices
and crevices. Therefore, the reefs were constructed out of various
combinations of beach boulders and cinder blocks.

Three reefs were constructed on December 10, 11 and 12, 1973, on
the east side of Discovery Bay, Jamaica (Fig. 1), equidistant from each
other, in 4.3 m of water. The precise location is 100 m directly
offshore from a concrete retaining wall and dock painted pink, and is
marked with a small white buoy. (The author's last visit to the area
was in early 1975, so both landmarks may have changed or disappeared.)
The reefs are readily visible from the surface, and are located in a
large Thalassia bed. A description of each of the three reefs
follows (from south to north):

Reef One (Organized Reef) was constructed solely of concrete
blocks, placed in such a way as to maximize the amount of holes and
spaces available (Plates 1 and 2). Approximately 35 blocks were used
in construction. The finished reef was rectangular, long dimension
parallel to the shoreline, and 2.8 m long, 1.0 m wide, and O. 7 m high.

Reef Two (Rock Reef) was constructed of subrounded boulders of
Pleistocene beachrock, average diameter approximately 25 cm, placed so
as to maximize the porosity. The finished reef was elongate parallel
to the shoreline, 2.2 m long, 0.9 m wide, and 0.7 m high (Plate 3).

Reef Three (Veneered Reef) combined both construction materials:
there was a core of beach rock, over which was placed a layer or veneer
of about 15 concrete blocks. The finished reef was elongate parallel
to the shoreline, 2.5 m long, 1.8 m wide, and 1.0 m high (Plate 4).

Cost of concrete blocks was $20 Canadian (1973), and total time
for construction of all three reefs was two man-days' shore time
(loading and transporting materials), and 6-8 man-hours’ underwater
labour. All rocks and blocks were placed by hand, underwater; the
original research plan called for construction of a fourth reef by
simply throwing beachrock overboard, but a family emergency forced the

author to return to Canada before this could be accomplished.
(b) Reef monitoring

Return visits to the reefs were in late June, 1974, and late
February, 1975 (or at 6- and 14-month intervals). Fishes were
identified and counted on both occasions. On each visit, several
censuses were taken of each reef on successive days, sometimes with
two operators. Estimates of the length of the more abundant fishes
were made by comparison with the length of the concrete blocks (20 cm).
Fish were identified with the aid of Randall (1968), and the boring
sponges were identified using Pang (1973) and Rtitzler (1974). Major
invertebrates in and around the reefs were also identified, in some
cases to major group only.

Data were supplemented with observations made by biologists
working at the Discovery Bay Marine Laboratory.

Results and Observations
A summary of the fish census data is given in Table l.

The day after construction, Reef One had attracted two fish: a
small Dusky Damselfish (EHupomacentrus fuscus) and a Spotted Moray
(Gymnothorax moringa) . Within a few weeks, all three reefs had
attracted fish, and had grown a covering of algae (Paul Sammarco,
personal communication).

Two spiny lobsters were seen on Reef Three in June, 1974.

After 14 months, the reefs had attracted a wide variety of
organisms other than fish, including calcareous and fleshy algae,
sponges, hydroids, anemones, sabellid polychaetes, hammer oysters,
arrow crabs, Stenopus, tunicates and urchins (Diadema antillarum,
Eucidaris tribuloides, Tripneustes ventricosa and Lytechinus variegatus).
In addition, the concrete blocks of Reefs One and Three supported small
colonies of the corals Porites astreoides and Dichocoenia stokesii, |
and colonies of Millepora alcicornis up to 20 cm high. Coral colonies |
occurred on the beachrock, but much less frequently.

Both the concrete blocks and the beachrock were infested with
boring algae and boring sponges (Siphonodictyon coralliphagum, Cliona
lampa, Cliona laticavola). Infestation was highest in the beach rock;
amount of material removed from the exposed outer part of some rocks
was estimated to be about 8%. Some samples of beach rock contained
fossil (Pleistocene) Montastrea annularis. Sponges boring into these
fossil corals exhibit the same boring pattern as does Cliona vermifera
in living Montastrea annularis: the sponge colony advances up the
vacated corallite, cutting back the septa to the septathecal wall
(Ward and Risk, in press). Boring pattern is evidently a response
to substrate hardness.

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25

Although no grazing halo was developed around any of the reefs,
comparison of 6-month and 14-month bottom photographs suggests a
decrease in length and density of Thalassia blades in the immediate
vicinity of the reefs. Grazing marks were common on the concrete
blocks, but not on the beachrock.

Trapping by local fishermen around the reefs began after about
six months.

Discussion

Most of the fishes on the reefs were carnivores or microcarnivores,
an observation made previously by Randall (1963) and Luckhurst (1972).
It is likely that the reefs were exploited more for shelter from
larger predators than for the fact that there is some in situ food
production.

Reef Three (beachrock core, block veneer) supported the highest
density and the largest number of species of the three reefs at both

6-month and 14-month visits. The average size of the dominant
species of fish was greater on Reef One (concrete blocks), however;
Reef Two (beachrock) was intermediate in all respects. Reef One had

fewer nocturnal fishes, perhaps due to its open construction, and
also supported more large carnivores.

The relationships among reef size, crevice size and shape, fish
density and fish diversity remain obscure, due largely to the limited
scope of the experiment and the short observation span. It is likely,
however, that most of the fish attracted to the reefs are those which
are nocturnal transients over the sand and grass flats of tropical
lagoons. These fish would readily take refuge in the large cavities
in the concrete blocks. The open construction of Reef One may account
for the lower density of smaller fish (lack of small holes) and the
larger average size. The relationship between fish populations and
reef morphology has been emphasized by several authors (Bardach, 1959;
Hiatt and Strasburg, 1960; Randall, 1963; Risk, 1972).

Longevity of artificial reefs also remains unknown. Rates of
infestation by boring organisms seem comparable to rates observed in
live corals. Undermining by Callianassa was also observed to cause
some foundering. Bioerosion and bioturbation, therefore, would seem
to set an upper limit on the age of the reefs. On the other hand,
settlement and growth of corals on the reefs could prolong their life
as shelters. Coral growth on artificial reefs is rarely observed
(Luckhurst, 1972). Growth of corals on the reefs under discussion may
be a result of relatively clear lagoon waters and a high resident
diversity and density of corals (Goreau and Wells, 1967).

Artificial reefs may be constructed relatively quickly and
inexpensively. Results of this study suggest only that the
construction of an artifical reef affects the fishes attracted to it.
Further monitoring of the reefs in Discovery Bay is necessary. Ie

96

would also be important to undertake a major experiment, based on these
preliminary results, using larger reefs in sets of replicates, in order
to determine the degree in which fish size and species may be influenced
by the type of reef construction.

The author regrets that financial restrictions terminated his site
VilESadttsr Persons visiting Discovery Bay are invited to observe the
reefs, as this is the only way in which further information may be
obtained.

Acknowledgements
Peter Woodhead, Eileen Graham, Paul Sammarco and Brian Keller,
all of the Discovery Bay Marine Laboratory during the time of this
study, aided in the initial phases of the project and provided periodic

reports on the state of the reefs.

Diving assistance was provided by David Kobluk and Fran and

Morty Ross. Bob Cavan identified some of the boring sponges, and
produced the infestation estimates. Cindy Matthews typed the
manuscript.

Supported by the International Development and Research
Corporation.

References

Bardachy vite L959 The summer standing crop of fish on a shallow
Bermuda reef. Limnol. Oceanogr. 4:77-85.

Brownell, W.N. 1971. Fisheries of the Virgin Islands. Comm. Fish.
Rev. 33:23-30.

Crossland, Jd. W976: Fish trapping experiments in northern New
Zealand waters. N.Z. J. Mar. Freshwater Res. 10:511-516.

Endean, R. 1976. Destruction and recovery of coral reef communities:
in O.A. Jones and R. Endean (eds.), Biology and geology of coral

reefs III, Biology 2:215-254.

Goreau, T.F. and J.W. Wells. 1967. The shallow-water Scleractinia of
Jamaica: revised list of species and their vertical distribution
range. Bulle Mabe, SCL. AGU le Cann Dp ly sta 2—A5 3%

Hiatt, RJW. and DiW. Strasburg, 1960. Ecological relationships of
the fish fauna on coral reefs of the Marshall Islands. Ecol.
Monogr. 30:65-127.

Luckhurst, Brian. 1972. Reef fish populations on small coral heads,
with special reference to the territoriality of Eupomacentrus
fuscus. M.Sc. Thesis, Marine Sciences Centre, McGill University,
HOSS ps2

Oclonn, ilsavs IS. Primary production measurements in eleven Florida

springs and a marine turtle-grass community. Limnol. Oceanogr.
AO Sa) S)7}/ 6
Ogden, J.C., R.A. Brown and.N. Salesky. 1973. Grazing by the echinoid

Diadema antillarum Philippi: formation of halos around West
Indian patch reefs. Science VS2 757 li.

PEM, Role IVS. The systematics of some Jamaican excavating sponges
(Porifera). ro genitile, Nik, 7S" jX¢

Randall, J.E. 1963. An analysis of the fish populations of artificial
and natural reefs in the Virgin Islands. Carib. TI o Clie 2 Oo Ares

IRGUNCENLIE A, Wolke ILS) 4 Grazing effect on sea grasses by herbivorous
reef fishes in the West Indies. Ecology 46,:255-260.

Randall, J.E. 1968. Caribbean reef fishes. T.F.H. Publications
Inc., Jersey City, 318 p.

Risk, M.J. 1972. Fish diversity on a coral reef in the Virgin
Islands. Atoll. Res. Bull. 153:1-4.

RUutzler, K. 1974. The burrowing sponges of Bermuda. Smithson.
Gontr. Zool 65): 1-32).

Turner, C.H., E.E. Ebert and R. Given, 1969. Man-made reef ecology,
The Resources Agency of California, Dept. of Fish and Game, Fish
Bulletin 146: 1-221. \

Ward, P. and M.J. Risk (in press). Pattern of Cliona vermifera
boring in Montastrea annularis. J. Paleontol.

ES A TS LT te Re an RESINS ny wr
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ald 0D Y3SIVY ee cae
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Plate 1:

Plate 2:

Reef One in February, 1975;
in encrusting organisms.

note increase

-—

Plate 3: Reef Two in June, 1974.

Plate 4: Reef Three in June, 1974.

DISTRIBUTION OF THE DECAPODS BRACHYURA AND ANOMURA
(EXCLUDING PAGURIDEA) OF THE CRYPTOFAUNA IN THE
REEFS NEAR TULEAR

By Mireille BeysaoeCLaeecis |

This study of the distribution of the Decapods Brachyura and
Anomura (excluding Paguridea) is a part of a broader work about the
whole cryptofauna of the reef flats of Tuléar. The Great Reef of
Tuléar, two zones of which were studied in detail, is the principal
field of my investigations. Some samples were also extracted from
the reefs of Sarodrano, Nosy Tafara, and Songoritelo (map 1).

Cryptofauna consists in all the small fauna of mobile invertebrates

which shelter in the little holes on the organic tracts of the reef
flats; these holes are O.5mm to 5 centimeters in diameter. The
cryptofauna was extracted from the blocks of hard hollowed-out
substrate, the volume of which is lam?. This volume of ldm? has been
defined as the minimum in which it is possible to obtain almost all the
species present in the biota studied (Clausade, 1970 ). For every
station studied, 5 samples of lam? have been extracted.

Altogether 34 stations, which represent 170dm? of hollowed-out
substrate containing cavities, subsequently broken into pieces, yielded
84 species of Brachyura and Anomura. The determination of these
species has been confirmed by Mr. Seréne. Those which appeared
particularly interesting from a systematic point of view formed the
subject of a separate paper (Peyrot-Clausade and Seréne 1976).

The areas studied on the Great Reef are shown in map l. In the
northern area (here called Sector 1), 9 stations have been studied.
Fig. 1 shows their localities (for clarification of terminology, refer
Eounecnuysi suppl, 27), 1972) In the area situated in the center of the
Great Reef (Sector 2) at the place named Antseteky, 12 stations have
been studied (fig. 2): 4 in the outer reef flat, 3 in the boulder
Eractrandyoiitn theyanner Geer tlat: Samples have been studied on the
other reef flats where some formations have appeared new in relation
to those of the Great Reef.

Istation marine d'Endoume, 13007 Marseille, France
(Manuscript received April 1975, revised figures June 1979 --Eds.)
INGO LIL) Tsxetsrs:. OUI » QS Spa. 9y TOM AaLNGe. IeYS ak

ee

102

The reef flat of Sarodrano is subjected to the alluvial deposits
of the river Onilahy, rich in clay particles in suspension. Moreover,
it is situated in an unstable zone and is tilting more and more towards
the open sea (J. Picard, personal information). The different
formations present on this reef, in order to stay at the degree of
dampness which is suitable to them, counterbalance this tilting by an
upper growth or elevation. Thus, the little Vermetus Dendropoma sp.,
which form only simple veneerings at the base of the blocks of the
boulder tract of the Great Reef, forms at Sarodrano thick pads rich in
Cavities, on the upper part of the blocks. For this reason, I have
studied two kinds of samples in this zone: the first among the blocks
of the boulder tract, the second among the Dendropoma sp. formations.
At the back of the reef flat, micro-atolls are present. Five samples
have been taken at the base of these formations which are widely
covered with Algae and rich in sandy particles.

The reef of Songoritelo results from the junction of two initial
shields (J. Picard, personal communication). The boulder tract,
present on the old shields, is absent from this junction Zone. This
zone is used as an out-fall through which the water of the channel
Situated between the reef and the Mangrove flows out at ebb-tide.
This sea water, rich in sandy earthy particles coming from the river
Fiherenana, promotes colonization and growth of the Sabellariids
Idanthyrsus pennatus. Between this area of Idanthyrsus and the
boulder tract are found layered Melobesiae formations. They are
calcareous Algae, more or less intermingled, and creating interstices
partially filled up by sand. It is in this zone that the boulder
tract later forms, when the reef has reached a certain degree of
evolution. Seven sations have been studied at Songoritelo: two on
the outer reef flat (one on the spur upper platform, one on the outer
moat rich in Algae) and two others around the outer creeks (on the
edge and towards the back among madreporarian coral colonies covered
by Zoantharian colonies). The last third are distributed in the
following manner: one in the boulder tract,one in the Idanthyrsus
formations and the last one in the layered Melobesiae.

Nosy Tafara can be distinguished from other reefs by its boulder
tract which is a boulder-rampart.

At each station, the abundance and average dominance of the
species present have been determined and is shown in table 1* From
this table, I have constructed tables 2, 3, 4 in which is recorded the
dominance of the different Brachyura and Anomura families, for each
station.

Figs. 3 and 4 show in the two sectors of the Great Reef, the
evolution of the settlements from the spur upper platform to the
blocks of sea grass bed basins. The population of each station is
represented by a circle in which each family is represented by an
arc ina direct ratio to its average dominance.

*Table 1 not reproduced here, available from author.

POPULATION OF THE OUTER REEF FLATS

For this study, I deal with 60 samples from 12 stations: 8 from
the outer reef flats: 2 from Sector 1, 4. from Sector 2, and 2 from
SOngorueevon (Graig) >). The last 4 come from the flat around the outer
creeks of Sector 1 and Songoritelo. For all the outer stations, the
total populations vary from 209 for the settlement of the station of
the glacis in Sector 1 to 47 in the station on the edge of the outer
creek in this same Sector. The average population is 17.8 individuals
for lam? of hollowed-out substrate; the number of species varies
considerably: 4 in the station of the spur upper platform in Sector 2,
and 24 for the one in the glacis of Sector l.

The Sector 2 shows an increase in the diversity of species of
Brachyura aS we move away from the front of the reef. The number of
species varies from 4 to 19. Indeed, in station 12, 97% of the
settlement belong to the single sub-family of Chlorodinae and are
Liocarpilodes integerrimus (Dana 1852) 71%, Pilodius paumotensis
(Stimpson 1858) 18%, and Chlorodiella laevissima(Dana 1852) 8%. Daira
perlata constitutes the remaining 3%.

In the next station (no. 13), ten species have been collected.
The Xanthidae family still represents 85%, the dominant species are
Chlorodiella laevissima and Pilodius paumotensis. Liocarpilodes
integerrimus is absent but I find Liomera rugata (H. Milne Edwards
1834) and Zozymus aeneus (Linné 1758) that will be found again in the (
next two stations. This is true also, for the two species of Anomura

Pisidia delagoae(Barnard 1955)and Pachycheles penicillatus(Heller 1862).

Station 14, established on the outer moat, contains 13 species
but they are not abundant. Indeed, in the Sam3, I number only 58
individuals. Among Xanthidae, Liocarpilodes integerrimus and
Pilodius paumotensis are not very abundant. The dominant species is
still Chlorodiella laevissima, and near the species already mentioned, {
I collected Pilumnus purpureus(A. Milne Edwards 1873),the dominance of

which will increase twofold in the next station as will the Pilumninae

ind. The Porcellanids are also enriched by a new species: Petrolisthes

lamarckii (Leach 1820). I note for the first time the presence of

Portunidae with Thalamita at a juvenile stage.

Station 15 is on the outer reef flat just in front of the boulder
tract, in an area of layered Melobesiae; 19 species or 101 individuals
were collected. Porcellanidae (with essentially Petrolisthes
lamarckii) have an increasing dominance at the expense of Xanthidae.

In this last family the two more abundant species are Chlorodiella
laevissima and Actaeodes tomentosus (H. Milne Edwards 1839). I note
an increase of Liomera rugata and a complete disappearance of
Liocarpilodes integerrimus and Pilodius paumotensis. Portunidae are
represented by Thalamita gloriensis (Crosnier 1962)and Grapsidae by some
individuals of Pachygrapsus minutus(A. Milne Edwards 1873).

If I compare the population of the stations of the outer reef flat
of Antseteky (Sector 2) with that of the station of the outer reef flat

104

of Sector 1, I see a real Similitude of settlement between the station
14-15 and the stations 1-2. Thus, in station 11 on the spur upper
platform, I found 133 individuals belonging to 17 species. In station
2, on the reef glacis rich in Algae Turbinaria, there are 24 species
and 209 individuals. Twelve species are common to these two stations
but their dominances vary. Liocarpilodes integerrimus (16%) and
Pilodius paumotensis (18%) are dominant in station 1, whereas Pilumnus
purpureus (23%) is more abundant in the 2nd station. In this last
station, I find also Liomera rugata and Actaeodes tomentosus. Three
species are collected solely on the outer reef flat of Sector 1l.

They are: Actumnus elegans (de Mann 1888) ,Domecia glabra(Alcock 1899) and
Pachycheles garciaensis (Ward 1942).

The settlement of the outer reef flat of Songoritelo appears quite
different from those previously studied. Indeed, the diversity is
more important in the station of the spur upper platform than in that
realized on the outer moat. Contrary to previous recordings, the
abundance and average dominance of Liocarpilodes integerrimus also
increase in the inner station. Chlorodiella cytherea (Dana 1852) and
C. laevissima are abundant, but Pilodius paumotensis and Pilumnus
Purpureus are completely absent.

The station around the outer creeks have settlements which
resemble more closely those of the outer reef flat stations. Indeed,
at Songoritelo, Xanthidae represent in the two stations more than 96%
of the individuals collected. Chlorodinae are the most abundant
with essentially the three following species: Liocarpilodes
integerrimus,Chlorodiella cytherea and C. laevissima. In sector l,
Xanthidae constitute only 70-78% of the settlement of these stations
around the creeks. The outer station contains: Dromiacea, Oxyrhyncha,
Grapsidae and Porcellanidae, the inner one has no Grapsidae nor
Dromiacea but 8% of Portunidae.

All the settlements of the outer reef flat show a certain
homogeneity, despite some variations. Among Xanthidae, Chlorodinae
dominates with three species more particularly abundant: Liocarpilodes
integerrimus, Pilodius paumotensis and Chlorodiella laevissima.

Fig. © shows the dominance of. these species in the different stations
OF the OUECr Teer rracs. Although the extension of Chlorodiella
laevissima is not limited to the outer reef flats, the dominance
increase from the outer station to the inner one and is non-existant

in the boulder tracts. At Antseteky, after a peak of dominance in the
dead madreporian colonies, it is clear that this species diminishes in
importance up to the boulder tract. The dominance of Pilodius
paumotensis diminishes regularly as we move from the front to the back
of these outer reef flats.

Liocarpilodes integerrimus, is, of these three, the most abundant
species. It is seen on fig. 6 that in all the biota studied, its
abundance and its dominance decrease towards the back of the outer
flat, except on the reef of Songoritelo. On this reef, the dominance
of Liocarpilodes integerrimus is lower on the outer station that in the
inner one (although its value is similar to that recorded on the other

outer stations); but on the inner station (situated on the outer moat)
the value of dominance is greater than on homologous stations (about
twice as great as that on homologous stations). An explanation of
this abundance can be found in the fact that, the front of this reef
being in a phase of very fast growth, the outer moat is not very old.
This outer moat is very rich in burrowing organisms (small sipunculids).
The size of the network and of cavities is very well suited to the
Liocarpilodes integerrimus which are very small individuals. This
area is between two phases of colonisation: the first colonisation
characterizes zones exposed to the beating of the waves (at present in
the outer station, on the spur upper platform) and the second
characterizes a well degraded substrate (found in Sector 1 of the

Great Reef of Tuléar, in the zone rich in Algae Turbinaria). Among
the "no carpilodea", Liomera rugata seems also to characterize quite
well the settlement of the outer reef flats. This species is absent
from the biota the most exposed to the beating of the waves. Zozymus
aeneus (Zozymoidea) is never abundant, but is present in almost all the
stations studied. Pilumnus purpureus and Actumnus elegans are also
linked to this zone of the reef. They are still more abundant in the
calmer biota. Among the Anomura, I find essentially Porcellanidae
with Pachycheles natalensis (Krauss 1843), Petrolisthes lamarkii,

P. penicillatus, Pachycheles garciaensis and P. pisoides (Heller 1865).
The last third are solely recorded on the outer reef flats.

A problem is posed by the Oxyrhyncha. Some of them are known to
live in the thallus of Algae but Acanthonyx quadridentatus (Krauss
1843) alone is limited to the biota very rich in Algae, thus species is
not included in the cryptofauna.

POPULATION OF THE BOULDER TRACTS AND OF THEIR BIOTA SUBSTITUTES

As has been seen in the first part of this paper, the structure of
the boulder tracts varies on the reef flats; the various stations were
established according to these variations in structure. SOjpeLn

Sector 1, samples were collected on the crags. In Sector 2, over

and above the station identical to that of Sector 1, two others have
been established: one in the gravel tail and the other in the
filtering dike. On the reef of Songoritelo, I have three stations:
one on the crags, one on the Idanthyrsus formations and the last in the
layered Melobesiae. On the Sarodrano reef flat, I studied the
settlement of the crags and of the little Vermetus Dendropoma sp.
formations. At Nosy Tafara, one Station is in the gravel sheet at

the back of the boulder rampart.

All these stations in the boulder tracts are qualitatively poor.
'From 4 to 13 species are found. The two stations on the crags of

the Great Reef have very similar settlement. Grapsidae are dominant

(56 and 44%) with only two species: Pachygrapsus minutus(A. Milne
Edwards 1873) and Nanosesarma minutum (De Mann 1887). Among

Xanthidae, I find Zozymodes xanthoides (Krauss 1843) Actaeodes tomentosus,
Eriphia scabricula (Dana 1852) and Pilumnopeus trispinosus (Sakai 1965).
This last species represents from 16 to 18% of the settlement.

106

Porcellanidae, with only one species — Petrolisthes lamarckii —
constitute 5 to 9% of the individuals present. The station on the
filtering dike differs from those of the crags by a considerable
decrease in Grapsidae and increase in Xanthidae (Pilumnopeus
trispinosus 32% and Actaeodes tomentosus 24, 5%) and in Porcellanidae
(Petrolisthes lamarckii 17%). On the gravel tail, 71% of the
settlement is constituted by Porcellanidae (still Petrolisthes
lamarckil). Pachygrapsus minutus constitutes only 1% of the
individuals collected on this station. Chlorodiella cytherea is, of
Xanthidae, the most abundant species (15%). If I compare this
settlement to that of the gravel sheet of Nosy Tafara, I see that they
are quite identical. On this station, Petrolisthes lamarckii
represents 67% of the population, Chlorodiella cytherea 22%, Pachygrapsus
minutus 6% and Pilumnopeus trispinosus 4%.

On the reef of Songoritelo, the crags have settlements similar to
those of the crags of the Great Reef, yet with a greater diversity of
species. Some species come from settlement of the outer flat
(Pilodius paumotensis, Liomera rugata, Liocarpilodes integerrimus) and
some species come from the inner reef flat such as Pilodius spinipes

(Heller 1861). None of them have a high dominance. In these’ crags;
Grapsidae are represented only by Pachygrapsus minutus 27%. Nanosesarma
minutum (de Mann 1887) is absent. It has been senn supra that at
Songoritelo, in absence of the boulder tract, we find Idanthyrsus
formations and layered Melobesiae. These formations have very

different populations. In the cavities between the tubes of the
Idanthyrsus pennatus, Grapsidae are very abundant and constitutes 49%

of the individuals collected. Porcellanidae are less abundant:
Petrolisthes lamarckii does not reach 5,5% of settlement.

The population of layered Melobesiae differs by the very feeble
proportion of Grapsidae (1,4%), the presence of Portunidae (14%) and of
Ocypodidae-Macrophtalmus boscii Audouin(et Savigny) 1825 7%. Xanthidae
family is the most important (68% of the individuals) with essentially
three species: Chlorodiella cytherea, Actaeodes tomentosus and
Pilodius areolatus (H. Milne Edwards 1834).

The last two series of samples are those of the boulder tract of
Sarodrano, among the crags and the formations of little Dendropoma sp.
In these two stations, the most abundant species is Pilumnopeus
trispinosus which constitutes 54% of the settlement of the first and
87% of the second. The crags of the boulder tract are characterized
by the absence of Pachygrapsus minutus and by the quite important
number of Macrophtalmus boscii.

Among the boulder tracts, four species have an important part in
the settlement. They are: Pachygrapsus minutus, Nanosesarma minutum,
Pilummopeus trispinosus and Petrolisthes lamarckii. In examining the
first three species, I note that Pachygrapsus minutus is a very mobile
species whereas Nanosesarma minutum and Pilumnopeus trispinosus are
more or less non-mobile. The study of table 1 shows that Nanosesarma
minutum is present only on the Great Reef and more abundant in the
biota in the highest part of the mid coast-level. Its abundance

ILO}

decreases in the filtering dike which is lower. In the same three
stations, Pilumnopeus trispinosus has an opposite behaviour and is more
abundant an "they filtering dike. When this species reaches its maximum
of dominance in the stations of the reef of Sarodrano, I see the
diminution and absence of Pachygrapsus minutus. I think that this
absence can be explained by the salinity of the sea-water around
Sarodrano reef being often very feeble, owing to the river Onilahy.

On the hard substrate on the coast Plante (1964) did not find this
Pachygrapsus where it would normally be collected, and it is also
absent from the hard substrate of the mangrove. ifia

the absence of Pachygrapsus minutus, Pilumnopeus trispinosus, which is
no doubt more resistant to the variations of salinity, inhabit all the
little cavities. Porcellanidae — Petrolisthes lamarckii — have an
abundance and dominance which increase as we move from the crags to

the gravel sheet or tail of the boulder tracts, this species goes deep
into the chips of dead Madreporian as the sea-water level gets lower,
and so protects itself from drying up at low tide. Apart from
Pilumnopeus already cited, I find also Actaeodes tomentosus in all the
stations. Zozymodes xanthoides is only collected on the crags.

POPULATION OF THE INNER REEF FLATS

It has been possible to study these settlements by means of the
eleven stations (five in every Sector of the Great Reef) and one in
the zone of micro-atolls of the reef of Sarodrano. The average number
of individuals collected in 1dm? of hollowed-out substrate is about

IS However, if I suppress the two series of samples from the branched
Melobesiae which constitute a special biota by the abundance of very
little cavities between the thallus of these calcareous Algae, there
are only 10 individuals per dm? in the other biota. The number of
species is situated between © and 19 and reaches 23 in the branched
Melobesiae. All the families found on the outer reef flats and on
the boulder tracts are represented in this part of the reef flats, but
it seems that Xanthidae are more abundant. Anomura, particularly
Galatheidae, play in some biota a very important in the inner reef
flats and particularly with the species Macrophtalmus boscii, in the
compact reef flat in the two sectors studied. This species is also
found abundantly in the blocks of the sea-grass bed basins of Sector 2.
Among Xanthidae, the two most important sub-families are Actaeinae and
Chlorodinae. In the first sub-family, Actaeodes tomentosus in the
dominant species. It has its maximum abundance in the branched
Melobesiae, but they are chiefly individuals at a juvenile stage.
Actaea cavipes (Dana 1852) is also frequent but not very abundant in
Sector 2. In Chlorodinae sub-family, Chlorodiella cytherea,

C. laevissima and C. barbata and Pilodius spinipes are the most
‘abundant. The last two species are only recorded on the inner reef
flats and have their maximum average dominance among the branched
Melobesiae. The "no carpilodea" are also represented by some species
gathered only in the inner reef flats. They are: Liomera bella

(Dana 1852), L. cinctimana (White 1857), L. semigranosa (de Mann 1888)
and L. monticulosa (H. Milne Edwards 1873). Among Anomura, one can
notice a gradual replacement of Porcellanidae by Galatheidae, as we

en

108

move off the boulder tracts towards the sea-grass bed basins. The most
abundant Porcellanids are: Pisidia delagoae(Barnard 1955) which reach
their maximum in the branched Melobesiae. Galatheidae are represented
by several species. The most frequent is Galathea humilis(Nobili 1905)
which is present in 7 of the eleven stations studied, the maximum
dominance is recorded in the micro-atolls of Sarodrano. Galathea
affinis (Ortmann 1892)is the species dominating in the biota of the
blocks in the sea-grass bed basins of Sector l. Galathea platycheles
(Miyake 1933),like many other species, reaches its maximum dominance in
the branched Melobesiae.

This study of the population on the different reef flats allow us
to make certain inferences about the distribution of the families even
of some species of brachyura and Anomura. The Dromiacea are
represented by only one species in these samples; it is Dynomene
hispida(Desmarest 1825) exclusively found on the outer reef flats.

It is also on the outer reef flats that the majority of Oxyrhyncha are
recorded: Menaethiops natalensis (Barnard 1955) ,Hyastenus aff.
elongatus (Ortmann 1893),Daira perlata, Elamena matthei (Desmarest 1825)
and more particularly among the places rich in Algae. Among Portunidae,
Thalamita gloriensis is the only species determined. The three biota
in which it is the most abundant are, in an increasing order: the
layered Melobesiae of the outer reef flats of Sector 2, the madreporian
colonies on the edge.of the outer creeks.ofSector 1 and layered
Melobesiae of the reef of Songoritelo where it represents 13.7% of

the settlement. In almost all the stations, juvenile forms of
Thalamita have been collected, and above all on the spur upper platform
of the outer reef flat of Songoritelo. Xanthidae family is the most
important qualitatively and quantitatively of all the families of
Brachyura in these cavities.

(a) no Carpiloidea

Liomera rugata seems to be particularly well adapted to the outer reef
flats just behind the spur upper platform, the other species of
Liomera — L. bella, L. monticulosa, L. semigranosa, L. cinctimana, all
came from the inner reef flats and more especially from the branched
Melobesiae.

(b) Zozymoidea

Five species belong to this group: Zozymus aeneus, Zozymoides
xanthoides, Atergatis aff. subdentatus(de Mann 1835), Platypodia
cristata (A. Milne Edwards 1865) and Platypodia anaglypta (Heller 1861).
The first species of this list, which is found above all in the outer
reef flat biota, like Zozymoides xanthoides settled only on the boulder
tracts, are the most abundant.

(c) Xanthoidea

The five species of this group are never abundant. The best
represented is Leptodius nudipes (Dana 1851) which come from the gravel

109

tail of the boulder tract of Sector 2.
(ad) Galenoidea

Actaeinae and Chlorodinae are the most frequent and abundant.

(1) Actaeinae includes 9 species:

Actaea quadriareolata which come from the boulder tract is not abundant.
Excluding Actaea consobrina and aff. Pseudoliomera varialosa

(Borradaile 1902), which are only samples on the outer reef flats, the
other species are collected in the different biota. The most abundant
is Actaeodes tomentosus which is found at all the stages of growth from

a few mm to several centimeters, in all the stations, the branched
Melobesiae having in their cavities an important concentration of
juvenile forms.

(2) Chlorodinae are, with 12 species, the richest group.

It has been seen that Liocarpilodes integerrimus and Pilodius paumotensis
are quite localized exclusively on the outer reef flats. On the other
hand Liocarpilodes armiger has been collected only in the inner reef
flats. The most abundantly represented genus in all the reef flat
formations, except on the boulder tracts, is the genus Chlorodiella

with three species. Chlorodiella laevissima although present in all

the stations, dominates especially on the outer reef flats where it

can represent up to 56% of the settlement (such as in the Zoantharian
zone of Songoritelo creek) Chlorodiella cytherea is the one of these
three species which resist best to drying-up, for it is the one

collected in the gravel tail and sheet. Chlorodiella barbata, never
sampled on the outer reef flats, is abundant on the inner reef flats

and particularly among the branched Melobesiae. In the Chlorodinae, |
the genus Pilodius is also well represented. With Pilodius paumotensis

collected in the outer reef flats, Pilodius areolatus present in the
inner reef flats and reaching its maximum abundance in the Idanthyrsus
formations and the layered Melobesiae, there is also Pilodius spinipes
which plays an important role in the settlements of the biota of the
inner reef flats and more especially in the blocks in the sea-grass-bed
basins and reef flat with scattered coral growth. Pilodius pugil

Dana 1852 is nowhere abundant. Among the Phymodius, only one species
— Phymodius ungulatus(H. Milne Edwards 1834) is quite well
represented in the inner reef flats. Paretisus globulus (Ward 1933),
not abundant, is collected in 4 different biota.

(e) Cymoidea

Eriphia scabricula seems to characterize the settlement of the boulder
ERAGES

(f) In the Hyperomista, only the Pilumnidae are abundant.

Domecia glabra(Alcock 1899),on the spur upper platform is the only
Mennipinae collected. Among the Piluminae, only two species have been
determined: Pilumnus purpureus and Actumnus elegans. Both occupy only
the cavities in the outer reef flats. The extension of Actumnus
elegans is however limited to Sector l. Pilumnopeus trispinosus is

110

among the Heteropanopeidea, the most abundant species. If some rare
individuals are on the outer reef flats, it reaches very high dominance
on the boulder tracts particularly in the reef of Sarodrano on the
Vermetus Dendropoma sp. where there are 18 individuals per dm? Lybia
leptochelis (Zehntner 1894) is essentially present in the inner reef
flats, but always in very feeble number.

Ocypodidae family is represented by one species — Macrophtalmus boscii —
which is particularly abundant in the inner reef flats, in calm biota
where colloidal suspension can form a deposit in which this species
makes its hole. This Macrophtalmus is also recorded in the boulder
tract of Sarodrano, where as we have seen the water of the river Onilahy
covers the reef.

Grapsidae. Pachygrapsus minutus is the most abundant of this family.
It occupies all levels of the mid-coast level and takes shelter in the
highest part of the reef constituted by the boulder tracts. As it is
a very mobile species, at low tide, it moves away and so can be

collected in the biota surrounding the boulder tracts. In the
absence of boulder tracts at Songoritelo the highest biota is the
Idanthyrsus formations where the Pachygrapsus are abundant. The

samples realized at Sarodrano show that this species does not like
fresh water, but reappears when the degree of salinity increases (as
at Nosy Tafara). Nanosesarma minutum lives at the high part of the
mid coast-level and is collected only on the Great Reef.

Anomura

Two families are abundant in the cryptofauna: Galatheidae and
Porcellanidae.

(a) Porcellanidae family, is, with 11 species, the most important.
A succession of species is clearly found moving from the front to the

back of reef flats. Petrolisthes penicillatus is strictly localized
on the outer reef flats, as are Pachycheles garciaensis, P. pisoides
and P. natalensis. Petrolisthes lamarckii, which is found also on

the outer reef flats, reaches its maximum dominance in the boulder
tracts and particularly in the gravel tail and sheet where it
constitutes 67% of the settlement. Pisidia delagoae is recorded on
all the reef formations and more abundantly in the inner reef flats.
It is also in that zone that Polyonix triungulatus(Zehntner 1894) and
P. aff. maccullochi(Haig 1965) are found.

(b) Galatheidae. Six species are collected essentially in the inner
reef flats, and their dominance increases as we move from the boulder
tracts to the sea grass bed basins. The most two abundant species

are Galathea humilis and G. affinis. The lst is frequent on the whole
of the Great Reef, and very abundant in the micro-atolls of Sarodrano;
the 2nd dominates the settlement of the blocks in the sea grass bed
basins in Sector 1 representing more than half of the local cryptofauna.

Hacil

From this study, some settlement appear quite typical of certain
reef zones. Thus, for a stabilized reef, it is possible to define
an outer reef settlement which increases in diversity as we move away
from the breaking waves. In the outer stations, Dynomena hispida,
Liocarpilodes integerrimus, Pilodius paumotensis, Actumnus elegans,
Pilumnus purpureus and Petrolisthes penicillatus are sampled.

Liomera rugata, Zozymus aeneus are added to the species already cited
in calmer zones. In the boulder tract, it can be said that the
classic population of the crags is composed of: Pachygrapsus minutus,
Zozymodes xanthoides, Eriphia scabricula, Petrolisthes lamarckii and
Pilumnopeus trispinosus. This last species increases very much when,
because of the weak salinity of the sea water, Pachygrapsus minutus
disappears almost completely. In the gravel zone, Petrolisthes
lamarckii and Chlorodiella cytherea are the dominating species. In
the inner reef flats, Pilodius spinipes, Chlorodiella barbata,
Actaeodes tomentosus, Galathea humilis play an important part in the
populations. It is in this zone that the highest density of
individuals per dm? is found among the branched Melobesiae which are
also quantitatively very rich.

REFERENCES
Clausade M. 1970. Importance et variations du peuplement mobile des
cavités au sein des formations épirécifales, et modalités
d'échantillonnage en vue de son évaluation. REC era Vjem Oleale! Ina Te.

Endoume, fasc. hors série suppl. 10: 107-109.

Clausade M., Gravier N., Picard J., Pichon M., Roman M.L., Thomassin B.,
Vasseur P., Vivien M., and Weydert P. 1971. Morphologie des
récifs coralliens de la région de Tuléar (Madagascar): Eléments

de terminologie récifale. Tethys, suppl. 2: 1-73.

Plante, R. 1964. Contribution a l'étude des peuplements de hauts
niveaux sur substrats solides non récifaux dans la région de
Tuléar. Madagascar. Rec. Trav. Sta. mar. Endoume, fasc. hors
série suppl.2 Trav. Sta. mar. de Tuléar: 205-316.

Peyrot-Clausade, M. and Seréne R., 1976. Observations sur quelques
espéces de Brachyoures (Crustacés Décapodes) de Madagascar.
Bull. Mus. Hist. Nat. Paris 3 serie (416) Zool. (293): 1339-1372.

2

Table

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Average dominance of the different families on the boulder
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Average dominance of the different families in the inner
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Map 1: The coral reef in the vicinity of Tuléar.

@ Spur upper platform

@) Glacis

Boulder tract
Compact reef flat

Branched melobesiae

Reef flat with coral alignements
and sandy couloirs

Reef flat with scattered coral growth

©28O © ©

Sea-grass-bed basin

Outer creek

Zoantharian zone

© @ ©

Inala, dug Localities of the stations in Sector l.

OUTER REEF FLAT

12 Spur upper platform {
13. Outer moat
14 Glacis

15 Layered melobesiae \
BOULDER TRACT

16 Crag
22 Filtering dike
38 Gravel tail |

INNER REEF FLAT

17 Compact reef flat

18 Reef flat with coral alignments
and sandy couloirs

19 Branched melobesiae

20 Reef flat with scattered coral
growth

21 Sea-grass-bed basin

OUTER REEF BOULDER TRACT INNER REEF FLAT
FLAT Wily Dromiacea [7 Ocypodidae

Oxyrhyncha EEEERE Grapsidae
- SECTOR 1 - [-JPortunidae [See] Porcellanidae
Xanthidae [7,2,] Galatheidae

lane pe Vip Distribution of Brachyura and Anomura in the different
stations in Sector l.

13

FLAT
”y) OUTER REEF 38

ey Dromiacea
Oxyrhyncha
[_] Portunidae
Xanthidae
[OD Ocypodidae
23335 | Grapsidae

Porcellanidae

(oar) Galatheidae

19 11

INNER REEF FLAT

- SECTOR 2-

Fig. 4: Distribution of Brachyura and Anomura in the different
stations in Sector 2.

OUTER REEF FLAT

29 30

OUTER CREEK

SONGORITELO
i

OUTER CREEK

SECTOR 1
ee Dromiacea
Oxyrhyncha Xanthidae Porcellanidae
[]Portunidae [EBB] Grapsidae [<=] Galatheidae

Imaliejg “Se Distribution of Brachyura and Anomura in the outer reef flat
and outer creek of Songoritelo and of outer creek of
Sector l.

ae Liocarpilodes integerrimus

(o]
Pilodius paumotensis
Chlorodiella laevissima
50
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Pilg. 16's) Dominance of the three more abundant species of Chlorodinae

in all the stations of the outer reef flats.

SECTOR 1 SECTOR 2

24

Oxyrhyncha
[XJ Oxystomata
[| Portunidae
SARODRANO [ZZ=] xanthidae

4] RRB TT) Ocypodidae

5 : FRR Grapsidae
EEE==J Porcellanidae

38

SECTOR 2 NOSY-TAFARA
33

SONGORITELO

lmal@jg 78 Distribution of Brachyura and Anomura in the different
stations on the boulder tracts.

April 1982

“ ATOLL RESEARCH
BULLETIN

256. Cays of the Belize Barrier Reef and Lagoon
by D.R. Stoddart, F.R. Fosberg and D.L. Spellman

257. Ten years of change on the Glover’s Reef Cays
by D.R. Stoddart, F.R. Fosberg and M.-H. Sachet

258. Plants of the Belize Cays
by F.R. Fosberg, D.R. Stoddart, M.-H. Sachet
and D.L. Spellman

259. Floristic observations on South Water and Carrie Bow
Cays, Stann Creek District, Belize, in 1979-1980
by James S. Pringle.

Issued by

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

ATOLL RESEARCH BULLETIN

os. 256-259

256.

257.

258.

259.

Cays of the Belize Barrier Reef and Lagoon
by D.R. Stoddart, F.R. Fosberg and D.L. Spellman

Ten years of change on the Glover’s Reef Cays
by D.R. Stoddart, F.R. Fosberg and M.-H. Sachet

Plants of the Belize Cays
by F.R. Fosberg, D.R. Stoddart, M.-H. Sachet
and D.L. Spellman

Floristic observations on South Water and Carrie Bow
Cays, Stann Creek District, Belize, in 1979-1980
by James S. Pringle.

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

April 1982

ACKNOWLEDGMENT

The Atoll Research Bulletin is issued by the Smithsonian
Institution, as a part of its activity in tropical biology, to place
on record information on the biota of tropical islands and reefs,
and on the environment that supports the biota. The Bulletin is
supported by the National Museum of Natural History and is produced
and distributed by the Smithsonian Press. Publication of this issue
on the Belize Cays is made possible by a special grant. The editing
is done by members of the Museum staff and by Dr. D. R. Stoddart.

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

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

Editors

F. R. Fosberg
Tan G. MacIntyre
M.-H. Sachet

Smithsonian Institution
Washington, D.C. 20560

Di Reo Stoddart

Department of Geography
University of Cambridge
Downing Place
Cambridge, England

ATOLL RESEARCH BULLETIN
No. 256

CAYS OF THE BELIZE BARRIER REEF AND LAGOON
by

D.R. Stoddart, F. R. Fosberg and D. L. Spellman

Issued by
THE SMITHSONIAN INSTITUTION
Washington, DG, U.S.A.
April 1982

CONTENTS

page
List of Figures i
List of Plates Li
Abstract HE
ae Introduction 2
De Structure and environment 5
Bis Sand cays of the northern barrier reef 9
St George's East Cay 9
Paunch Cay 9
Sergeant's Cay 10
Curlew Cay iE
Goff's Cay ale)
Seal Cay IZ
English Cay 12
Sandbore south of English Cay IES}
Samphire Spot 14
Rendezvous Cay 14
Jack's Cays 15
Skitpensand 15
Cay Glory 16
Tobacco Cay 16
South Water Cay 18
Carrie Bow Cay 20
Curlew Cay Zyl
Dis Sand cays of the southern barrier reef 23
Silk or Queen Cays DS
North Silk Cay 23
Middle Silk Cay 24
Souiehy SasikiCay, ZI5)
Samphire Cay 25
Round Cay 25
Pompion Cay 26
Ranguana Cay 20)
North Spot 28
Tom Owen's Cay DDS)
Tom Owen's East Cay DE)
Tom Owen's West Cay 30

Cays between Tom Owen's Cays and
Northeast Sapodilla Cay Sal
The Sapodilla Cays Sil

Northeast Sapodilla Cay Sul

thal

Frank's Cays
Nicolas Cay
Hunting Cay
Lime Cay
Ragged Cay
Seal Cays

Cays of the barrier reef lagoon

A. The northern lagoon

Ambergris Cay

Cay Caulker

Cay Chapel

St George's Cay

Cays between Cay Chapel and
Belize

Moho Cay

Stake Bank

Spanish Lookout Cay

Water Cay

B. The Southern Triangles

Robinson Point Cay
Robinson Island
Spanish Cay

C. Cays of the central lagoon

Tobacco Range
Coco PlumeCay,
Man-o'-War Cay
Water Range
Weewee Cay
Cat Cay

Lagoon cays between Stewart Cay

and Baker's Rendezvous
Jack's Cay
Buttonwood Cay
Trapp's Cay
Cary Cay
Bugle Cay
Owen Cay
Scipio Cay
Colson Cay
Hatchet Cay
Little Water Cay
Laughing Bird Cay
Placentia Cay
Harvest Cay

page

32
34
35
37
38
38

40

40

40
42
43
44

45
45
45
46
46

46

46
47
47

47

48
49
50
50
51
yl

52
yi
52
52
53
53
54
54
54
55
56
56
57
Di

ababal

page

D. The Punta Gorda cays 5 7/

East Snake Cay 58
Middle Snake Cay 58

West Snake Cay 59

South Snake Cay 60

Wild Cane Cay 60
Frenchman's Cay 61

Moho Cay 62

6. Summary and conclusions 63
A. Types of islands 63

B. Topographic change 65

C. Flora and vegetation 66

ie References 69

Index of islands 75

iv

LIST OF FIGURES
Location of the Belize reefs and cays following page 4
Reefs and cays of the central barrier reef lagoon
Sergeant's Cay 1972 following page 22
Goff ts (Cay 1972
English Cay 1972
Rendezvous Cay 1972
Reef topography between Columbus Cay and Channel Cay
Tobacco Cay 1972
North-south profile of Tobacco Cay 1960
South Water Cay 1972
Carrie Bow Cay 1972
North Silk Cay 1961 and 1972 following page 39
Middle Silk Cay 1961 and 1972
South Silk Cay*l96il and 1972
Round Cay 1960 and 1972
Pompion Cay 1960 and 1972
Profiles of Round and Pompion Cays 1960
Ranguana Cay 1960 and 1972
North Spot 160 “and 97/2
Reefs and cays of the southern barrier reef

Topographic change on the southernmost barrier reef,
between the surveys of Owen (1830) and those of 1960-1972)

Tom Owen's East Cay 1960 and 1972
Tom Owen's West Cay 1960 and 1972
Northeast Sapodilla Cay 1960 and 1972

Frank's Cays 1972

26.

Zils

2335

22K.

310)6

ils

36

333

34.

359

36.

S76

38}

38)5

40.

Nicolas Cay 1960 and 1972
Hunting Cay 1960 and 1972
Lime Cay 1960 and 1972
Ragged Cay 1960 and 1972
Seal Cay 1972

Buttonwood Cay 1972
Hatchet Cay 1972

Little Water Cay 1972
East Snake Cay 1961
Middle Snake Cay 1961

West Snake Cay 1961

West Island West Snake Cay 1961

South Snake Cay 1961
Wild Cane Cay 1961

Moho Cay 1961

following page 39

following page 62

2s

22's

23

24,

LIST OF PLATES

Paunch Cay: aerial view from the southeast 1961 following page 22

St George's East Cay: aerial view from the northwest 1961
Sergeant's Cay: aerial view from the east 1961
Sergeant's Cay: aerial view from the east 1962

Goff's Cay: aerial view from the east 1961

Goff's Cay: aerial view from the east 1962

English Cay: aerial view from the southeast 1961
English Cay: aerial view from the southeast 1962
English Cay from the south 1961

English Cay from the south 1962

Rendezvous Cay from the south 1960

Rendezvous Cay: aerial view from the southeast 1962
Rendezvous Cay from the east 1962

Rendezvous Cay from the west 1972

Rendezvous Cay from the southwest 1972

Cay Glory: aerial view from the east 1961

Rendezvous Cay: erosion on the east shore 1972
Rendezvous Cay: aggradation on the southwest shore 1972
Tobacco Cay: aerial view from the northwest 1960
Tobacco Cay: aerial view from the east 1962

Tobacco Cay from the south, seen from tne reef-crest rubble
ridge 1960

South Water Cay: aerial view from the south 1961
South Water Cay: aggradation at the southwest point 1972

South Water Cay: erosion on the northwest shore 1972

ZS) 6

26.

Zils

28.

2K

310)6

Sik

32.6

335

34.

315)6

36.

Sie

3K8)6

IDs

40

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

Carrie Bow Cay: aerial

Carrie Bow Cay: aerial
Carrie Bow Cay from the
Carrie Bow Cay from the
Norm chyisitlic Cay eaerial:

South Silk Cay: aerial

view from

view from

southwest

southwest

view from

view from

Walab

the northwest 1960

the east 1962

1965

WY

the east 1962

the south 1962

Round Cay: aerial view from the southeast 1962

Pompion Cay: aerial view from the southeast 1962

Pool on North Silk Cay 1961

Middle Silk Cay 1961

South Silk Cay 1961

following page 22

following page 39

Ranguana Cay: aerial view from the northeast 1962

North Spot: aerial view from the southeast 1962

North Spot from the south 1972

Tom Owen's Cays: aerial view from the northeast 1961

Hymenocallis littoralis under coconuts at Pompion Cay 1972

Ranguana Cay: erosion on the north shore 1972

Ranguana Cay seen from the shingle islet at the west end 1972

Tom Owen's East Cay from the southwest 1972

Nowth Spot 1972

Tom Owen's West Cay: north shore 1972

Tom Owen's Cays from the west 1972

Tom Owen's East Cay: aerial view from the south 1972

Tom Owen's West Cay: aerial view from the southwest 1962

The Sapodilla Cays 1972

Northeast Sapodilla Cay:

aerial view from the east 1962

following page 39

Sie

0

on

54.

5D.

56.

Dilis

Soe

SS)

60.

Gis

627

637

64.

65'.

66.

672

68.

69:.

IO

gale

72%

ey

74,

Nicolas Cay: aerial view from the northeast 1962

Northeast Sapodilla Cay: northeast shingle ridge 1972
Northeast Sapodilla Cay: eroding northwest shore 1972
Frank's Cays: aerial view from the south 1961

Hunting Cay: aerial view from the north 1961

Hunting Cay: aerial view from the southeast 1962

Nicolas Cay: '‘rootrock' exposed on the eroding west shore 1972
Hunting Cay east bay from the north 1972

Lime Cay: aerial view from the south 1962

Lime Cay: aerial view of the west end from the north 1962
Lime Cay: aerial view of the east end from the north 1962
Ragged Cay: aerial view from the south 1962

Ragged Cay from the west 1972

Seal Cay: aerial view of the cay and faro from the east 1962
Seal Cay: east rampart and moat 1972

St George's Cay: aerial view from the north 1962 (channel A)

following page 62

St George's Cay: aerial view from the north 1962 (channel C)

St George's Cay: aerial view from the east 1962 (channels D and E£)

Tobacco Range: view inland from the northeast sand ridge 1972

Tobacco Range: Rhizophora killed by Hurricane Hattie in 1961,
invaded by Batis, at the northeast point 1972

Cary Cay: hurricane-deposited shingle on the east side 1962
Scipio Cay: hurricane-deposited shingle ridge 1962

Colson Cay: hurricane-deposited shingle on the southeast shore
1962

Water Range (Twin Cays): Thrinax thicket, south shore 1972

UD

76.

Ut

iss

VQ

80.

ix
Little Water Cay: shingle rampart on the southeast shore 1972
following page 62
East Snake Cay: aerial view from the southeast 1961
West Snake Cay: aerial view from the southeast 1961
South Snake Cay: aerial view from the southeast 1961
East Snake Cay rampart and moat from the north 1961

West Snake Cay rampart and moat from the south 1961

CAYS OF THE BELIZE BARRIER REEF AND LAGOON

IOS7/ ID) S18%c Stoddart, ! Beets Fosberg, andy Davis Spellman?

ABSTRACT

25 aslands of the Bell:ze barrier reek are mapped in this paper,
together with 10 islands of the barrier reef lagoon. A further 4
barrier reef and 28 lagoon islands are described. For 22 islands
comparative maps are presented showing topographic change between
LIGO-6il and 9.7/2’: Lists of vascular plants recorded from each
island are given. Eleven types of island are recognised. Most of
the islands have undergone erosion during the period of the surveys,
notably during individual catastrophic hurricane events. The flora
of the cays comprises 164 species. Vegetation units are briefly
described, with special reference to the vegetation of dry mangrove
cays and mangrove ranges, and to plant succession following major
hurricane damage.

Department of Geography, Cambridge University, England.
2 smithsonian LISeLeUclon Washington, sD Go, WS.
3 Missouri Botanical Garden, Saint Louis, Missouri, U.S.A.

IMSWE (Investigations of Marine Shallow Water Ecosystems)
Contribution No. 12.

Manuscript received January 1981 -- Eds.

1. INTRODUCTION

"The whole extent of the main reef is ... studded with
numerous islets of every variety of size and solidity:
first the spot of sand, just raised above the water; then
the more extended beach, which nurtures into trees the
cocoa-nut cast upon its shores; next those places
composed of sand and mud, where the mangrove springs

out of the water, and which are in this neighbourhood
called by the appropriate name of ‘drowned cays'; and
lastly, the island, a mile or more across, whose soil
produces the grapewood tree, button-wood tree (very hard
and valuable for cogs of wheels), and other species of
trees, with a girth of more than 3 feet" (Allen 1841,
PwS)E.

"... so deceitful are the different keys found from the
general resemblance they bear to each other, that the most
experienced seaman, when placed amongst them, often becomes
fatally perplexed. ... A more particular description of
these spots, so numerously scattered over the whole of this
coast, would extend far beyond any moderate limits; nor
could any material value be affixed to it, from the little
characteristic difference that would be found connected
with them" (Henderson 1811, pp. 20, 24).

This paper presents an account of the cays of the barrier reef and
coastal shelf of Belize (British Honduras), Central America. It is
based on data collected during five expeditions between 1959 and 1972.
The first of these studied most of the barrier reef islands between
St George's East Cay and Ragged Cay during the Cambridge Expedition to
British Honduras 1959-60, when the field party consisted of Stoddart
and J.D. Poxon. In 1961 Stoddart and S.P. Murray re-surveyed several
of these islands, and also visited the Punta Gorda cays in the southern-
most Gulf of Honduras, during a season supported by the Office of Naval
Research and the Coastal Studies Institute, Louisiana State University.
Following the catastrophic Hurricane Hattie of 30-31 October 1961,
Stoddart remapped the barrier reef islands as far south as Carrie Bow
Cay and also studied the numerous islands of the central coastal. shelf
between Placentia and Gladden Spit, during March-April 1962; this visit
was supported by the Royal Society. The barrier and central shelf
islands were again mapped, with Royal Society support, in March-April
1965, primarily to establish the nature of recovery following catastro-
phic hurricane damage. Finally, in June-July 1972 a comprehensive
study was undertaken of the barrier and shelf cays (including the
southernmostbarrier reef cays only briefly visited previously in 1960)
as part of the Smithsonian Institution's IMSWE (Investigations of Marine
Shallow Water Ecosystems) Programme centred on Carrie Bow Cay. The

field party consisted of Stoddart (responsible for mapping and geomor-
phology), Fosberg and Spellman (vegetation and flora), and Henry Pelzl
(ornithology) .

Some of the information presented here has appeared in preliminary
form in earlier papers. The regional setting and environment were
described, together with general features of the reefs, in an account
of parallel studies of the Belize offshore atolls (Stoddart 1962a).
Hurricane effects on the northern and central barrier reef cays were
considered in a paper on Hurricane Hattie (Stoddart 1963). Island
recovery in the northern and central areas was considered in an account
of the 1965 re-survey (Stoddart 1969). A preliminary account has
appeared (Stoddart 1974) of changes on five of the cays monitored in
UTZ « The southern barrier reef cays (south of the Silk Cays) and
those of the Punta Gorda area in the Gulf of Honduras have received
only incidental mention in the published literature (Stoddart 1962b,
1965), and studies on them appear here for the first time. With the
pUublaication Of a tull account of “the vascular plants of the cays
(Hosbexg eb al. 1981), ehe nomenclature used in previous papers is
here revised, and lists are given of species recorded from each of the
islands during succesSive surveys. An analysis of plant species
diversity on the cays has been published by Stoddart and Fosberg (1981).

With a few exceptions, all of the islands here mapped were
surveyed by pace-and-compass methods as described by Stoddart (1962a,
Do LAS) Maps from succesSive surveys were related to each other by
reference to prominent objects. Islands such as Rendezvous Cay and
Carrie Bow Cay were also mapped by tape-and-compass methods, and the
results used to calibrate the scales of the other maps. Oblique low-
level air photographs were taken of many of the islands in 1960, 1961,
1962 and 1965. All the maps of cays in this report are orientated
conventionally (i.e. with magnetic north at the top).

When this study began in 1959 virtually nothing was known of any
of the islands of the Belize coast, or indeed of the reefs themselves.
The first detailed hydrographic charts were made during the 1830s by
Richard Owen, Edward Barnett and Bird Allen, but their ships carried
no scientific men and only brief accounts were published (Owen 1838,

Allen 1841). Darwin utilized their information and called the Belize
barrier "the most remarkable reef in the West Indies" (1842, p.201l),
but this led to no further field investigation. Both Agassiz (1894,

p-.-162) and Vaughan (1919, p.300) called on evidence from Belize in
SUppOre Of their own theories of reef development, but neither visited
the area. Not until 1957 were the cays investigated, in a pioneer
reconnaissance study by Vermeer (1959). Three major series of
investigations have, however, subsequently made the Belize shelf and
barrier reef one of the best known reef provinces in the world. The
Rice University studies of shelf sediments directed by Edward G. Purdy
during 1960-67, recently summarised by Wantland and Pusey (1975), led
to Purdy's classic papers (1974a, 1974b) on reef structure and
morphology. Deep-diving submersible and geophysical studies by James
and Ginsburg (1960) have led to new insights into the form and evolution

of the central barrier reef. Finally the IMSWE investigations
directed by Klaus Ruetzler at Carrie Bow Cay have greatly extended
knowledge of reef biota, ecology and growth. The present paper is
thus concerned only with the most recent and ephemeral accumulations
of clastic carbonate sediments on the top of these massive reef
structures; with their vegetation cover; and with their short-term
variability in form and location in response to environmental events.

We thank the British Department of Scientific and Industrial
Research, the Royal Society of London, the Office of Naval Research,
and the Coastal Studies Institute of Louisiana State University for
support of the investigations in 1959-1965, and the Smithsonian IMSWE
Programme for that in 1972. The support of the Government of Belize
has greatly assisted this work from the time of its initiation, and
nothing would have been achieved without the practical assistance of
many Belizeans. We are especially grateful to Mr M. Young and Mr R.
Coe, of the Department of Geography, Cambridge University, for their
cartographic and photographic skills over many years.

Several of the diagrams within this paper have been prepared from
copies of charts and surveys provided by the Hydrographer of the Navy
and with the sanction of Her Majesty's Stationery Office. EES Saa
pleasure to thank successive Hydrographers, notably Rear Admiral Sir
Edmund Irving and Rear Admiral D.W. Haslam, both surveyors of the
Belize coast and reefs, for their interest in this work.

!

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2. STRUCTURE AND ENVIRONMENT

Belize is located in the southeastern part of the Yucatan
Peninsula, central America. Themsoughern part Or the countny, (consists
of an uplifted block of Palaeozoic metamorphosed sediments and
instrusives, rising in the Maya Mountains to 1110 m above sea-level.
The block is partly surrounded by hills of Cretaceous limestones, which
are themselves overlain by low, flat-lying Cenozoic limestones which
make up the greater part of the Yucatan Peninsula. The northern part
of the east coast of the Peninsula, between Cabo Catoche and Ambergris
Cay, is remarkably straight, formed by low cliffs with a fringing reef.
South of Ambergris Cay, however, the Belize coastal shelf forms a major
indentation up to 37 km wide. It is edged by a broadly arcuate
barrier reef more than 200 km long. North of the city of Belize the
shelf is generally less than 5 m deep; immediately south of Belize it
inereases to 10-12 m, and deepens southwards to depths of 45-55 m in
the Gulf of Honduras. Seaward of the barrier reef the sea floor falls
very steeply to more than 300 m, and depths of 1500 m are found within
6 km of the reef edge, except in the lee of the offshore atolls (Figure
1) (Dillon and Vedder 1973; James and Ginsburg 1980). Pronounced
topographic lineations (notably connecting the southern barrier reef,
the eastern side of Glover's Reef and the east side of Lighthouse Reef;
the central barrier reef and the eastern side of Turneffe; and the
coastline of Ambergris Cay) are clearly fault blocks aligned en echelon
along the northern margin of the Cayman Trough, a major tectonic
structure up to 7 km deep.

Purdy et al. (1975) have drawn attention to the very different
characteristics of the shelf-edge reefs north and south of Belize City.
To the north, where the shelf is shallow, there is a prominent chain of
linear islands (Cay Caulker, Cay Chapel) located 2.5-4 km landward of
the shelf edge; the edge itself is fringed by a discontinuous reef which
lacks any well-developed reef flat. Purdy etwas LOS .0s)e Comment on
the resemblance between this situation and that of the Bahama Banks.
There are no true sand cays on these reefs north of latitude 17°33'N
(the site of St George's East Cay which disappeared in 1961).

south of Belize City the deeper coastal shelf is edged by a

baraier plattormmn, in the terminology of Purdy et al. (1975). This has
quite different characteristics north and south of Gladden Spit in
latitude 16°30'N. In the northern sector, 80 km long, the shelf floor

reaches depths of 20-24 m at distances of 8-15 km from the mainland
coast; the shelf edge platform is 3-5 km wide and its upper surface

4-5 m deep. At its outer edge there is a well-developed reef flat
450-550 m wide, and there are numerous small patch reefs in its lee.
South of Gladden Spit the same units are recognisable but with differing
degrees of development. The shelf here has a constant width of 30 km,

and its maximum depth increases southwards from 35 to 77 m. The
barrier platform maintains its depth on its upper surface of 4-5 nm,

but it is less than 2 km wide, is dissected by deep channels, and
carries little surface reef. In spite of these differences in
dimension, however, the volume of the barrier platform remains constant
along the entire shelf edge (Stoddart 1977, p.47).

Within the coastal shelf there are three main groups of islands,
in addition to those of the shelf edge reefs, which are considered
here. The reefs of the Southern Triangles are developed at the inner
end of the Belize Deepwater Channel. They are scattered isolated
patch reefs, many of which have small mangrove cays on their upper
SUBEACEeS. In the latitude of Gladden Spit there is a complex array
of lenticular reefs, mostly slightly submerged (Stoddart 1963, figs.
35-36), fully described by Purdy (1974b), termed 'shelf atolls' by
Purdy et al. (1975, p.15) and ‘rhomboid platform atolls or faroes" by
James and Ginsburg (1979). Surface reefs are irregularly developed
on these structures, but where they do occur islands are common.
Finally, small patch reefs rise from shallow water near the mainland
coast in the Gulf of Honduras, north of Punta Gorda, and carry sand
and mangrove cays.

Purdy (1974b) has shown that the present surface reefs at the
Belize barrier are relatively thin Holocene structures capping an
older antecedent and possibly karst-eroded reef topography. The
Holocene reefs are 8-20 m thick, and average 12 m. The positive
features on which they grow are presumed to be reefs of last inter-
glacial age. In contrast to many other parts of the Caribbean there
are no last interglacial reefs standing above present sea-level anywhere
on the coast of Belize, though they are described at the usual heights
of 2-4 m, with ages of 120-125,000 years, in northeastern Yucatan and
Cozumel by Szabo et al. (1978). Their greater depth beneath the
Belize reefs must indicate long-continued subsidence connected with
the horst structures north of the Cayman Trough.

Modern reef growth on the Belize shelf must have post-dated the
flooding of the antecedent topography on which the present reefs stand
during the Holocene transgression, about 6000-8000 yr B.P. The deeper
surface of the southern shelf would have flooded earlier, and Purdy
(1974b, 854) has obtained dates of 6000-10,000 yr B.P. for shelf floor
peats. Halley et al. (1977) have documented the sequence of Holocene
reef growth on a pre-existing Pleistocene limestone prominence for a
small patch reef of the central barrier reef. This and other Belize
reefs have grown in the Holocene at rates of 1-2 m per 1000 yr (Purdy
VOTAa SiG ie. The date at which reef flat development close to low-
tide level could provide a platform on which clastic sediments could
accumulate to form cays will thus be a function of the depth and
dimensions of the reef foundations, governing both the time at which
they were flooded by the Holocene transgression and the volume of
constructional reef growth required for the reef to reach the present
sea-level. The actual age of reef flat formation has yet to be
determined for Belize reefs.

The Belize reefs lie in the zone of Northeast Trades, which blow
with great constancy, especially during the summer months of April-
September. At Belize, for the period 1917-1949, 56% of winds at O600
hours came from the east and southeast and 23% from the northwest (means
for the years), while at 1800 hours 75% came from the east and northeast.
The Trades are interrupted during the months of November to February
by 'northers', 4-5 day periods of northerly winds and low temperatures
associated with the southerly extension of the North American high
pressure system.

Rainfall over the east coast of the Yucatan Peninsula is about
1500 mm. June-September are the wettest months, and there is a
pronounced dry season in February-April. Annual totals increase
markedly southwards, to nearly 1900 mm at Belize, 2400 mm at Mango
Creek, and 4200 mm at Punta Gorda (Table 1); the dry season is also
very much less pronounced in these wetter areas. There are no rainfall
records for any of the islands of the coastal shelf, but it is
reasonable to presume that annual totals on the cays mirror those of
coastal localities. Certainly there is evidence in the southernmost
cays of much more humid conditions than in the north.

Minimum temperatures at Belize vary from ROO shin January and
December (monthly means), with a mean maximum of LZ sin August.
Mean temperatures range from 24° in winter to 27° in summer. Extreme
temperatures recorded at Belize during 1951-60 were 9.4° and 36°
(Ponevg. 19/6), p..4 70).

Hurricanes occur during the months of July to October. Especially
destructive storms have been recorded in 1787, 1813, 1827, 1831, 1931,
1942, 1945, 1955 and 1961; for an annotated list of recorded storms
Scessitoddant) (L963) ippienl 27 — 130). The most recent significant storm
to affect the reefs was Hurricane Greta in September 1978; this post-
dated all the surveys reported here. The effects of the lesser
Hurricane Fifi of October 1974 have been documented for Carrie Bow Cay
by Miller and Macintyre (1977, p.27).

Mean tidal range at springs is only O.2 m, and water level may
be more affected by wave set-up and wind than by the astronomical tides.
The main Caribbean current which flows from east to west from the Lesser
Antilles is diverted northwards through the Yucatan Channel by the
Yucatan Peninsula. Water movement on the coastal shelf is mainly to
the south, into the Gulf of Honduras (Purdy et al. 1975, p.13).

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3. SAND CAYS OF THE NORTHERN BARRIER REEF

This chapter describes the sand cays of the shelf-edge reefs
between St George's East Cay in the north and Curlew Cay, south of
Carrie Bow Cay, in the south, a distance of some 90 km. In each case,
citations are given of previously published accounts, together with a
list of plants recorded from the island. Since some of these cays
have been treated in previous papers, the descriptions in this chapter
are abbreviated.

SEG CorgelismHasit iCayseCl7ie33UNiSSOSW) Plate 2

When mapped in 1960, this was an arcuate shingle ridge 110 m long

with a maximum altitude of about 1 m. he tvadvan areavot Oj3 hae The
vegetation cover consisted of herbs and vines, with taller shrubs of
Tournefortia, Conocarpus and Borrichia at the northeastern end. The
island disappeared completely during Hurricane Hattie in 1961, and had
not re-formed when the site was visited in April 1962. For a fuller
account see Stoddart (1963, pp. 40-41). Plants recorded are:

Sesuvium portulacastrum 60st Conocarpus erectus 60S

Canavalia sp. 60S Ipomoea sp. 60S

Euphorbia mesembrianthemifolia 6©OS Tournefortia gnaphalodes 60S

Rhizophora mangle 6OS (seedling) Borrichia arborescens 60S
Paunch Cay (17°24'N, 88°02'W) Plate l

Charted as Punchgut Kay by Speer (1771) and as Paunchgut by

Jefferys (1775). It was vegetated in 1830 and in 1896-7 had coconut
palms lOmm tal: The island was destroyed in the 1931 hurricane.

When mapped in 1960 it was an unvegetated sandbore 45 m long, with three
parallel lines of beachrock up to 45 m long on its south side. The
sandbore disappeared but the relict beachrock survived the 1961
hurricane. No land was present in 1962 and the site has not been
visited since. See Stoddart (1963, p.4l and fig. 21). The only

+ In the plant lists the numeral refers to the year of record, and S
or C refer to sight records or collections respectively. Species
preceded by an asterisk are not native to the Belize cay flora,

though this does not necessarily imply that they have been deliber-

ately introduced on the cays where they occur.

10

plant recorded is:

*Cocos nucifera 1896S

Sergeant's Cay (17°23%'N, 88°02'W) Figure 3, Plates 3-4

This island was charted by Speer (1771) and Jefferys (1775); at
the time of the survey by H.M.S. Rambler in 1896-7 it had palm trees
P5emy talks It was first mapped in 1960, when it was a triangular sand
cay with greatest length of 100 m and width of 45 m. Its area was
3680 sq m. The island was occupied by a large clubhouse, and the
ground surface beneath the coconut palms was cleared. The cay was
almost completely destroyed during Hurricane Hattie and was reported
awash a few days later. In March 1962 a triangular sand cay had re-
formed, with longest dimension of 70 m and area of 2000 sq m. The
surface was colonised by patches of Portulaca oleracea, together with
Sesuvium portulacastrum and Euphorbia mesembrianthemifolia. By 1965
the island was more elongate, but only 68 m long and with an area of
1640 sq m. The pioneer Portulaca had disappeared. A central area,
occupying about 25 per cent of the surface and persisting from before
the hurricane, had a continuous cover of Ipomoea, Wedelia and Euphorbia;
Sesuvium was also absent. Shrubs were represented by three Conocarpus
up to 3 m tall, six Suriana up to,l.5 m tall, and three Journetortzva
less than 1 m tall; there was a single Coccoloba seedling. Euphorbia
and Ipomoea were colonising fresh sand areas. The number of plant
species had risen from 4 in 1962 to 13 in 1965. For further details,
see Stoddart (1963, pp. 41-42, fig. 22; 1969, p.8, fig. 4) and Vermeer
(LOSS eepp.. 7LaANvs

The island was re-mapped on 11 July 1972, when it was 90 m long
and 25 m wide, with an area of 1750 sq m. The vegetated area was still
largely limited to that part of the surface inherited from before the
hurricane: it was covered with a mat of Euphorbia, Ipomoea, Vigna, and
Sesuvium. All the 1965 shrub species were present, but the Coccoloba
was in danger of being undermined as the south shore continued to be
eroded. Fresh sand to the west of the main vegetated area was being
colonised by Portulaca. A new coloniser was Spartina spartinae, the
only grass, forming a conspicuous clump near the centre of the island.
Three species had become extinct since 1965: Batis maritima, Euphorbia
blodgettii and Eustoma exaltatum. 14 species were present in 1972,
bringing the total recorded for the island to 20:

Spartina spartinae 72C Canavalia rosea 72C

Cocos nucifera 1896S 60S Suriana maritima 65S 72C
Coccoloba uvifera 65S 72C Euphorbia blodgettii 65S

Batis maritima 65C Euphorbia mesembrianthemifolia
Philoxerus vermicularis 65S 72C O2C 65S" 72E

Portulaca oleracea 62C 72C
Sesuvium portulacastrum 62C 72C Rhizophora mangle 62S
Cakile lanceolata 65C 72C (seedling)

itil

Conocarpus erectus 65S 72C Tournefortia gnaphalodes 65S 72C
Eustoma exaltatum 65C Avicennia germinans 60S
Ipomoea macrantha 65S 72C Wedelia trilobata 65S 72C

Ipomoea pes-caprae 65S 72C
Curlew Cay (17°23'N, 88°02'W)

Curlew Cay is described by Speer (1765, 19) as "very low", with

"only a few bushes", and comparable to Paunch Cay. The Honduras
Almanack for 1830 stated that "there are no trees on Curlew, but bush,
such as bay cedar [Suriana maritima] and lilly-root grass". It was
charted by Owen in 1830 and by H.M.S. Rambler in 1896-7, but disappeared
some time after that date, possibly in the 1931 hurricane. When
visited in 1960 it was a small sandbore 20 m long and 0.6 m high; it
was not seen in 1962. Plants recorded:

Gramineae indet. 1830S Suriana maritima 1830S
Cones Cay (7 -2iUN /1887O2"W) Figure 4, Plates 5-6

In February 1960 Goff's Cay was triangular, with sides 65-75 m
long, built of grey sand, with a fresh sandspit at the north end. ast
was charted by Speer (1765, p.19 - a "Small round Kay ... not so big
as English Kay") Jefferys (1775), and H.M.S. Rambler in 1896-7, when
it had coconuts 14 m tall. Coconuts dominated the vegetation in 1960.
There were a few Coccoloba and Avicennia trees. The ground beneath
the coconuts was sparesely covered with Euphorbia, Ipomoea and
Canavalia; the fresh sand was being colonised by Sesuvium, Ipomoea and
Canavalia. The area of the island was 2920 sq m, and 7 plant species
were recorded. The island suffered severe erosion during Hurricane
Hattie. The old island surface was reduced to about 800 sq m. By
March 1962, however, marginal fresh sand accumulation had increased the
surface area to 2280 sq m. Surface sand was stripped from the old
island area; two or three coconuts and some broken Coccoloba survived.
The only new coloniser was Portulaca oleracea. Relict beachrock, not
previously visible, was exposed on the south and east sides of the cay.
For further details of the island in 1960 and 1962, see Stoddart (1963,
SOG CSU aealopy W46))ie By 1965 slight recession of the south coast had
been balanced by considerable aggradation on the north side, increasing
the total area to 2870 sq m, approximately that of the pre-1961 island.
Dense vegetation - a mat of Euphorbia, Sesuvium, Ipomoea and Ernodea -
was limited to the surface area surviving from before the storm, though

vines if Ipomoea were colonising fresh sand. Eighteen coconuts planted
since 1961 were up to 1m tall. In both 1962 and 1965 there was a small
unvegetated sandbore about 400 m north of the main cay. For details of

the island ain A965) seer stoddart (9697, ps8, sbrg. 24):

12

Goff's Cay was re-mapped on 11 July 1972. The core island
remained, with some marginal erosion on the east side, but the marginal
sand accumulations had considerably increased to give a total island
area of 3190 sq m. The herb mat consisted of Portulaca, Sesuvium,
Ipomoea, Ernodea and Cakile, and 17 coconuts survived. The eastern
sand spit was being colonised by Cakile. A substantial concrete jetty
had been built on the leeward side (the island is much used for
recreational purposes), and a deep depression near the centre was
presumably the sand source for the concrete. 10 plant species were
recorded in 1972, compared with 2 in 1962 and 4 in 1965. No species
became extinct either between 1962 and 1965 or between 1965 and 1972
except for the broken Coccoloba which did not survive. New colonisers
in the first three years were Cocos (planted), Sesuvium, Ipomoea and
Euphorbia; after 1965 they included two grasses (Andropogon, Eragrostis),
a sedge (Cyperus planifolius), and Cakile. A total of 13 species is
now recorded from the island:

Andropogon glomeratus 72C Cakile lanceolata 72C
Eragrostis prolifera 72C Canavalia sp. 60S

Cyperus planifolius 72C Euphorbia mesembriantihemi-
Cocos nucifera 1896S, 60S 65S 72S folia 72C

*Coccoloba uvifera 60S 62S Euphorbia sp. 60S 65S
Portulaca Qleracea 72C Ipomoea pes-caprae 72C
Portulaca’ Esp. 625 Ipomoea sp. 60S 65S

Sesuvium portulacastrum 60S 65S 72C Avicennia germinans 60S
Erithalis fruticosa 65S 72C

Seal Cay (17°21'N, 88°02'W)

Seal Cay was noted by Speer (1766, p.19) as "a very low, small,
sandy Kay". It was not charted by Owen in 1830, but the Ramoler survey
noted a sandbore awash at high water immediately on the north side of
the entrance to the Belize Deepwater Channel. It was not seen during
our Own surveys. The seal in question was presumably the West Indian
Monk Seal Monachus tropicalis, though the animal itself does not appear
ever to have been recorded from the Belize coast, which is not included
in King's (1956, p.217) map of the range of this species. That it was
reasonably common in the eighteenth century is shown by the presence of
other 'Seal Cays' on the barrier reef.

English Cay (17°20'N, 88°03'W) Figure 5, Plates 7-10

English Cay was charted by Speer (1765, p.19: "a short, round,
DIME Kay 71771) and Jetfiexys. (1775)< It has been a lighthouse and
pilot station for many years; the present lighthouse was built in 1935.
In 1960 the island was triangular, with sides 75-90 m long, and with an
area of 6510 sq m. The eastern shore was largely artificial, being
partly a masonry wall and partly a ridge of conch shells. A leeward

eS

sand spit seen in January 1960 was evidently a seasonal feature

(Vermeer 1959, p.75). The vegetation was limited to coconuts 12-15 m
tall, witn the ground surface kept clear; there were two mature
Rhizophora and one Coccoloba tree. During Hurricane Hattie in 1961

the island suffered severe marginal erosion and stripping of surface
sand; its area was reduced to 3390 sqm. 90 of the 98 coconuts
disappeared together with all the houses, but the lighthouse remained
standing. Rhizophora survived but the Coccoloba was killed. No new
plant species had colonised the island by March 1962 (see Stoddart 1963,
pp. 44-45, fig. 24). By 1965 there had been virtually no change in the
morphology of the island. 18 coconuts had been planted, and there was
a patchy ground cover of Sesuvium, Portulaca and Euphorbia. The area
Gimtnencay was stabie atl 3520 sqm) (Stoddart 1969), 0.9, fig. 6). By
1972, however, walls had been built round almost the entire periphery
of the cay, close to the position of the 1960 shoreline, and low-lying
sand had accreted within this area. There were two main areas of
Euphorbia, Sesuvium, Portulaca and Ipomoea, and several smaller patches;
a strip of littoral Sporobolus; a single small shrub each of
Tournefortia and Suriana; and 25 coconuts. The total dry-land area
had increased to 4940 sq m, and the number of plant species from 2 in
IQG2 io) 6 sid IQS ehevel IS} sin WY) 7/ 2, Two species recorded by Dwyer, Elias
and Maxwell in 1967 (Ernodea littoralis, Eclipta prostrata) were not
EOuUNG setsehvers, nO 6S oman 97/2 = Since the island is subject to
continuous human interference it is clear that the vegetation is also
much disturbed, notably by periodic clearing of the ground cover.
Species recorded from English Cay are:

Paspalum disticinum 72C Euphorbia sp. 65S
Sporobolus virginicus 72C Phyllanthus amarus 72C

* Cocos nucifera 60S 62S 65S 72S *Carica papaya 65S
Coccoloba uvifera 60S wObherabulinnis, ileigenebis 71/ Xe
Philoxerus vermicularis 65S 72C Rhizophora mangle 60S 62S
Portulaca oleracea 67C 72C Tpomoea pes-caprae 72C
Portulaca sp. 65S Ipomoea stolonifera 72C
Sesuvium portulacastrum 65S 6©7C 72C (seedling)

*Tamarindus indica? 72C (seedling) Tournefortia gnaphalodes 72C
Suriana maritima 72C Ernodea littoralis 67C
Euphorbia blodgettii 72C Hedyotis corymbosa 72C
Euphorbia mesembrianthemifolia Eclipta alba 72C

67C 72C Eclipta prostrata 67C

*Euphorbia thymifolia 72C

Sandbore south of English Cay (17°19%'N, 88°03'wW)

H.M.S. Vidal charted this sandbore in 1957-8; it was not charted
by H.M.S. Rambler in 1896-7 or earlier surveys. in ilQ6Ohittawase ss) Mm
long and up to 1 m high, and unvegetated. It has not been seen since.

14

Samphire Spot (17°17'N, 88°024'W)

This was charted as Saphire Kay by Jefferys in 1775 but omitted
by him in 1792 and 1800; it was not noted by Speer or Owen in 1765 and
1830, but appears as Samphire Spot following the 1896-7 Rambler survey.
In 1959-61 it was a small unvegetated sandbore; it presumably
disappeared during Hurricane Hattie, but in 1962 was again a small
sandbore 23 m in diameter.

Rendezvous Cay (17°15'N, 88°03'w) Figure 6, Plates 11-15, 17-18

During the 1920s and 1930s this island was inhabited by a local
fisherman, and in consequence is still locally known by some as ‘Brown's
Cay' It was the headquarters of the Cambridge Expedition to British
Honduras in 1959-60. At that time it was a Slightly arcuate island,
aligned north-south, 90 m long and 9-23 m wide. It was formed entirely
of sand and its greatest height was less than 1 m. The leeward shore
had been extended by accumulations of conch shells, giving a crenulate
outline and forming small-boat harbours. The total area of the cay was
2610 -sgunie Speer (1765, p.19) described it as "a low sand Kay, with
only one bush on it". It was also charted by Jefferys in 1775 and Owen
aisgy als} {0} In 1921-22 H.M.S. Mutine recorded coconuts 14 m high. In
1960 the taller coconuts reached almost 20 m; there were mature
Coccoloba and Avicennia trees; and frequently cleared patches of
Sesuvium, Euphorbia and Sporobolus. Rhizophora was represented by
numerous seedlings round the shore. The island was greatly damaged
during Hurricane Hattie. The house disappeared and all the coconuts
were destroyed. The overall dimensions remained much the same, but
the island shifted slightly to the west and the area was reduced to
2180" Sq im The Avicennia was killed and the Coccoloba much broken.

By March-April 1962, however, there was a varied though patchy cover of
herbs, grasses and sedges. Portulaca was the most common species,
together with Cyperus planifolius and Sporobolus. Sesuvium, Euphorbia
mesembrianthemifolia and Ipomoea pes-caprae were also widespread.

Shrub species were represented by seedlings of Suriana, Conocarpus and
Tournefortia. Stoddart planted over 40 young coconuts on the cay in
April 1962. For further details, see Stoddart (1963, fig. 4; 1663,
ppes 47-49, «fig. 28)

In 1965 the cay was of similar dimensions but had continued to
migrate lagoonwards: the seaward shore in 1965 stood 10-15 m westwards
of its 1962 position, and the area had declined slightly to 2000 sq m.
The planted coconuts were 3 m tall, and in the north there were low

shrubs of Tournefortia and Suriana. The ground surface was covered
with a dense mat of Ipomoea and Euphorbia, with areas of Wedelia and
Sesuvium; Portulaca had become rare. There was a Single Casuarina 2 m

tall at the north: end (Stoddart 1969, pp. 9-10). fig. 7s

BL)

In 1972 these trends had continued. The cay had maximum
dimensions of 92 x 32 m, and an area of 1840 sq m. Both its seaward
and lagoonward shores were located up to 10 m westward of their 1965
positions (and hence up to 16 m westward of their 1960 locations).

Some of the coconuts were already bearing. The ground cover was
dominated by Euphorbia, Ipomoea, Philoxerus, Sesuvium, Portulaca,
Cyperus, Sporobolus and other grasses. One clump of Coccoloba still
survived from before the hurricane, but the Casuarina had disappeared
and the Suriana appeared dead. There were several clumps of Conocarpus,
Rhizophora and Avicennia. A small hut had been built and appeared to
be intermittently occupied. There was evidence that the conch fishery
had been resumed.

The total number of species recorded from Rendezvous Cay is now
DAN 7 were recorded in 1960, 16 in 1962, 16 in 1965 and 16 in 1972.
The uniformity of these figures disguises considerable turnover,

however, as the detailed records show. The following species are
mecorded::
Paspalum distichum 62C 72C Cakile lanceolata 62C 65S
Sporobolus sp. 60S 62S 65S Suriana maritima 65S 72C
Cyperus ligularis 62C Euphorbia mesembrianthemifolia
Cyperus planifolius 72C 626 7/2
Cyperus Sp. 65S Euphorbia sp. 60S 65S
Fimbristylis cymosa 62C 72C Rhizophora mangle 60S 62S 65S 72C
*Cocos nucifera 20S 60S 65S 72S cConocarpus erectus 65S 72C
*Casuarina equisetifolia 65S Laguncularia racemosa 72C
Coccoloba uvifera 6OS 62S 65S Ipomoea pes-caprae 62S 72C
72C Ipomoea sp. 65S

Philoxerus vermicularis 62C 72C fournefortia gnaphalodes 65S 72S
Portulaca oleracea 62C 65S 72C Avicennia germinans 60S 72S

Portulaca sp. 62S Solanum campechiense 62C
Sesuvium portulacastrum 60S 62C *solanum lycopersicum 62C
65S 72C Ageratum sp. 65S

Wedelia trilobata 62S 65S
Jack's Cays (17°14'N, 88 °O2%'W)

These are intermittent sandbores 20-40 m long. One was often
visible in 1959-60, the other was seen in 1962. They are unvegetated.

Skiff Sand (17°13'N, 88°03'W)

An ephemeral sandbore on the outer barrier, charted by H.M.S.
Mutine in 1921-22, and occasionally visible; it is unvegetated.

16

Cay Glory (17°06'N, 88°0O1'W) Plate 16

Cay Glory was mapped in 1960: it was 105 m long, up to 21 m wide,
and had an area of 1430 sq m. There was a single young coconut 1.2 m
tall, and a small vegetated area covered with Sesuvium, Euphorbia,
Philoxerus, Cakile, Paspalum and Ipomoea. The island was built of
well-sorted beach sand (Stoddart 1964, figs. 4, 6) and its outline was
clearly very variable. The island was previously larger. H.M.S.
Mutine noted coconuts 21m tall in 1922, but these must have been
destroyed by either the 1931 or 1945 hurricanes. Beachrock relics of
this older island are widespread north and west of the 1960 cay. The
island disappeared entirely during Hurricane Hattie in 1961, and no land
was seen on its site in 1962, 1965 or 1972. For further details, see
Stoddart, (196377 pps 52=53 jeri sO) re The following 7 plant species
have been recorded:

Paspalum distichum 60C Cakile lanceolata 60C
*Cocos nucifera 22S 60S Euphorbia mesembrianthemifolia 60C
Philoxerus vermicularis 60C Ipomoea asarifolia 60C

Sesuvium portulacastrum 6O0C

Tobacco Cay (16°544'N, 88°03'W) Figures 8-9, Plates 19-21

Tobacco Cay lies on the north side of one of the most important
entrances in the central barrier reef. It was charted by Speer in
1771 and Jefferys in 1775, and there is an unverified suggestion that
the island is named after tobacco cultivation during the early years of
the English settlement in 1630-40 (Winzerling 1946; Caiger 1951, 28-29).
The most recent hydrographic chart of the area was made by Owen in 1834.
The cay was mapped in 1960, 1961 (in July after Hurricane Abby), 1962
(April, after Hurricane Hattie), 1965, and 1972.

In 1960 the island was roughly triangular, with a greatest north-
south dimension of 275 m and a greatest width of 140 m; its area was
2.74 ha. The whole island was built of sand, with a flat featureless
surface at about 1.2 m above sea-level. There were ridges of fresh
sand along the south shore, and relict strips of beachrock 35 m offshore.
The cay was inhabited and densely vegetated. In 1834 Owen had noted
coconut trees 20 m tall, and Smith (1842, p.732) described a fig tree
20 m tall near the northeast point. In 1960 many of the coconuts were
old, reaching up to 25 m, and there were also mature trees of Terminalia,
Coccoloba and Cordia. There was a dense ground cover of Stachytarpheta,
Wedelia, Ipomoea pes-caprae, Ipomoea stolonifera, Sesuvium, Hymenocallis,
Euphorbia, Canavalia and Vigna. Hurricane Abby on 15 July 1960 removed
the fresh sand ridges and blew down a number of coconuts; the inhabitants
also cleared a good deal of the ground vegetation.

During Hurricane Hattie there was severe marginal erosion,
stripping of surface sand, and deposition of fresh sand along the south
and west shores. At the southeast point these new deposits reached a

LY
height of 2 m, and were up to 14 m wide. 7O per cent of the coconuts
were felled, the trunks being aligned N40°E. Coccoloba and Cordia were
much broken, though Terminalia was less damaged. Since the cay was not

inundated by the storm surge the ground vegetation survived with little
change, though shade-seeking Wedelia had given way to Ipomoea in many
places. The fresh sand ridges were also colonised by Ipomoea. By
1965 the fallen trees had been cleared and Wedelia had greatly expanded.
Ipomoea and Sesuvium had expanded on the sand ridges, where Tournefortia
had also appeared. The general ground cover, however, was similar in
Composition, = choughsnOem nl pateerny, eto) that seen any 1960. The cay had
been reduced in size to 2.25 ha immediately after Hurricane Hattie; by
1965 it had increased to 2.30 ha. For further details, see Stoddart
(L963, jo BID5_ sven Sikp ISD 7 joe IMO)s sseatofa 18) 6

The island was re-mapped on 24 June 1972. Its area had increased
to 2.50 ha, largely because of the presence of a fresh sand spit at the
north point; the southern sand ridges had disappeared. The area of the
vegetated cay has, however, remained very largely constant since 1960.
Many new coconuts were growing, together with tree species (Terminalia,
Coccoloba, Cordia) surviving from before Hurricane Hattie. The ground
cover beneath the coconuts again consisted of Wedelia, Stachytarpheta,
Sesuvium, Euphorbia, Vigna, Hymenocallis, Portulaca, sedges and grasses,
though the central part of the island had been recently: cleared and was
bare. The southern sand ridge was colonised by Tournefortia, Suriana,
Euphorbia and Ipomoea. The prominent rubble ridge on the reef flat to
the southeast of the cay supported small patches of Philoxerus and
Sesuvium.

The number of plants species recorded in 1960 was 8, in 1961 12,
1962 13, 1965 21 and 1972 41; the total recorded for the island is 44.
Undoubtedly this increase results from more thorough collecting rather

than from new colonisation. There is only one certain extinction over
this period of record: a single Sophora tomentosa, O.7 m tall near the
south point, had disappeared by 1972. The plants recorded are as
follows:

*Fleusine indica 62C 72C Coccoloba uvifera 61S ©2S 65S 72C
Eragrostis ciliaris ©2C Philoxerus vermicularis 72C
Paspalum distichum 72C Portulaca oleracea 60C 61S 62S
Phragmites cf. australis 72C 658: WAC
Spartina patens 72C Sesuvium portulacastrum 61S 65S
Sporobolus virginicus 65S 72C 72C
Sporobolus sp. 65C Caesalpinia bonduc 72C (seedling)
Cyperus ligularis 72C Canavalia rosea 6O0C 72C
Cyperus planifolius 72C Sophora tomentosa 65S
Cyperus sp. 615 65S Vigna luteola 60C 62C 65S 72C

*Cocos nucifera 60S 61S 62S 65S Suriana maritima 72C

728 Euphorbia blodgettii 62C 72C
Crinum amabile 72C Euphorbia glomerifera 72C
Hymenocallis littoralis 61S 62S Euphorbia mesembrianthemifolia

65S a7 2e 62E57/2E

Ficus sp. 1842S Euphorbia prostrata 72C

18

‘Euphorbia sp. 60S 65S Cordia sebestena 60C 61S 65S 72C
Phyllanthus amarus 72C Tournefortia gnaphalodes 65S 72C
*Carica papaya 65S 72S Avicennia germinans 61S 65S 72C

Rhizophora mangle 62S 65S 72C Stachytarpheta jamaicensis 60C
Conocarpus erectus 61S 65S 72C 61'S 265Siei2e
*Terminalia catappa 61S 62S 65S Spermacoce assurgens 72C

12E Ageratum littorale 72C
Bumelia retusa 65C 72S Bidens cynapiifolia 72C
*Catharanthus roseus 72C Melanthera nivea 72C
Ipomoea pes-caprae 62C 65S 72C Pluchea symphytifolia 72C
Ipomoea stolonifera 6O0C 72C Wedelia trilobata 6OC 615 62S
Ipomoea sp. 65S 658), 7.2€

Purdy (1974b, pp. 841, 853) drilled a core hole 16.4 m deep on
Tobacco Cay, the upper 14 m of the section being Holocene in age.

South Water Cay (16°49'N, 88°05'W) Figure 10, Plates 22-24

South Water Cay is the largest inhabited sand cay on the barrier
reef. It is prominently marked on Jefferys'schart in 1775, and in
1830 Owen noted coconut trees 15 m tall. The island was mapped in
1960, visited again in 1961, and mapped again in 1962, 1965 and 1972.
In addition to the collections made during these visits a large
collection made by J. Pringle in 1979-80 considerably increased the
recorded flora.

In 1960 the island had a north-south extent of about 640 m; its
width varied from 70 to 180 m; and it had an area of 8.18 ha. The
northern part, closest to the reef, was covered with a dense palm
thicket and had an eastern shore of coral rubble. The rest of the
island was settled, covered with open coconut woodland, and formed of
sand. In the northern palm thicket the ground cover consisted of
Wedelia, Euphorbia, Ipomoea, grasses and sedges, with Sesuvium,
Tournefortia and Suriana along the coast. Both Borrichia and Coccoloba
were common along the seaward margins of the woodland, and extended
southwards along the east side of the island. The rest of the cay had
a varied vegetation of planted exotics and weedy species which were
frequently cleared.

During Hurricane Hattie the cay was subject to considerable marginal
erosion, especially in the southwest bay and at the northeast point.
There was considerable wind damage to coconuts, the mean direction of
fall being N30-40°E, but the island was not overtopped by the storm

surge. In the centre of the island about 80 per cent of the trees were
felled. Along the shores Coccoloba, Avicennia and much Borrichia
survived the storm. All the jetties and several houses were destroyed.

For detailssee Stoddart (1963), pp. '55=5 7), eblge 232)i-

19

By 1965 physiographic change had been slight. The seaward
beaches had retreated slightly, but the leeward beaches had accreted;
there had been continuing erosion at the south point. The vegetation
of the northern part of the island was little changed, with Wedelia,
Ipomoea, Batis, Euphorbia .and Sesuvium, and thriving Thrinax and
Coccoloba under the coconuts. In the centre of the island there was
a dense growth of Euphorbia, Ipomoea, Stachytarpheta, Ambrosia and
Cassytha, with some Hymenocallis and Coccoloba. Along the eastern
shore there was bushy Borrichia with much Cassytha, a few Tournefortia
seedlings, and a ground cover of Sesuvium, Ipomoea, Euphorbia and
Sporobolus. Many young coconuts planted in the southern part of the
cay after the hurricane were 3-4.5 m tall. Two juvenile Casuarina
were growing by the lagoon shore, but there were many fewer Rhizophora
seedlings round the shores and especially along the east coast than in
SNE) A jetty and several new buildings had been erected and the
effects of the hurricane were no longer obvious. For further details,
see Stoddart (1969, p.1l).

In 1972 the trends of erosion and accretion had continued. There
was retreat at the south point, along the southwest bay and at the
north point. The island had a maximum length of 660 m, a width
varying from 55 to 175 m, and an area of 6.49 ha. Masonry and rubble
walls had been inadequate to prevent continuing erosion both in the

southwest bay and at the northeast point. Apart from a sector in the
south, recently cleared, there was a dense ground cover of herbs, vines
and grasses under the open coconut woodland. Hymenocallis, Ipomoea

stolonifera, Ambrosia and Wedelia were all common and conspicuous.

Borrichia was the most common shrub colonising the east shore, with
Sesuvium aS ground cover; a single seedling of Morinda was noted on the
beach crest.

No attempt was made during the earlier visits to make a complete
collection of plants from South Water Cay, and by 1965 only 26 species
had been recorded. 68 were recorded in 1972, bringing the total to
69, and Pringle's collections in 1979-80 raise the total to 80. This
is the highest number for any of the Belize cays, but clearly reflects
the continuous and prolonged human disturbance of the vegetation. The
following species have been recorded:

Andropogon glomeratus 61C 72C Sporobolus virginicus 72C 79C

79C Sporobolus sp. 60S 65S
Anthephora hermaphrodita 79C Cyperus ligularis 72C
Cenchrus incertus 72C 79C Cyperus peruvianus 72C 79C
Distichlis spicata 79C Cyperus planifolius 72C 79C
Eragroseis Cll Janis 26 s/9C Cyperus sp. 60S
Eustachys petraea 72C 79C Fimbristylis cymosa 72C 79C
Panicum virgatum 79C *Cocos nucifera 60S 61S 62S 65S
Paspalum blodgettii 72C 72S
Paspalum distichum 72C Thrinax radiata 65S 72C
Phragmites cf. australis 72C Agave sp. 72C
Spartina patens 72C 79C Crinum amabile 72C

Spartina spartinae 72C

20

Dracaena sp. 72C 79C Conocarpus erectus 72C 79C
Hymenocallis littoralis 65S Laguncularia racemosa 72C 79C
TAC TILE *Terminalia catappa 72C
*Casuarina equisetifolia 65S *Psidium guajava 72C
12EM1IE Polypremum procumbens 72C 79C
Coccoloba uvifera 60S 65S 72C 79C*Nerium oleander 79C
Batis maritima 65S 72C 79C *Plumeria rubra 72C
Philoxerus vermicularis 72C 79C Ipomoea macrantha 65S 79C
Boerhavia coccinea 79C Ipomoea pes-caprae 72C 79C
Portulaca oleracea 72C 79C Ipomoea stolonifera 65C 72C 79C
Sesuvium portulacastrum 60S 65S Ipomoea sp. 60S 65S
(2E7719E Jacquemontia havanensis 72C 79C
Cassytha filiformis ©OS 65S Cordia sebestena 72C 79C
VAGASIE Tournefortia gnaphalodes 60C
Cakile lanceolata ©O0S 72C 79C 6S *65S7 7/2 79
Chrysobalanus icaco 72C Avicennia germinans 60S 72C 79C
Canavalia rosea 72C 79C Lippia nodiflora 72C 79C
Crotalaria retusa 72C 79C Stachytarpheta jamaicensis 65S
Desmodium incanum 72C 79C 12E IC
Desmodium tortuosum 79C *Solanum lycopersicum 79C
Erythrina sp. 72C Eri thalis*frutrcosa 726579
Sophora tomentosa 65S 72C 79C Ernodea littoralis 72C 79C
Vigna luteola 72C 79C Hedyotis corymbosa 72C
Suriana maritima 60S 79C *Hedyotis lancifolia 79C
Euphorbia blodgettii ©OS 72C Morinda citrifolia 72C
19E Spermacoce assurgens 72C 79C
Euphorbia mesembrianthemifolia Ageratum littorale 72C 79C
VAIS STIS Ambrosia hispida 65S 72C 79C
Euphorbia trichotoma 72C 79C Borrichia arborescens 60S 65S
Euphorbia sp. 60S 65S 12E AIC
Phyllanthus amarus 72C 79C Conyza canadensis 72C
*Hibiscus rosa-sinensis 72C Eclipta alba 79C
Waltheria indica 72C 79C Emilia sonchifolia 79C
Passiflora suberosa 72C 79C Melanthera nivea 72C
Rhizophora mangle 60S 65S Wedelia trilobata 60S 65S 72C
127 79E 19€
Carrie Bow Cay (16°48'N, 88°05'W) Figure 11, Plates 25-28

Carrie Bow Cay, named after the Bowman family of Stann Creek, was
charted by Owen as "Jack Ellin's Cay" and appears on charts as "Ellen
Cay". Owen noted "tops of bushes 20 feet", but by 1960 the cay was
covered with an open coconut woodland surrounding large houses; the
ground surface was kept cleared. The island was then 140 m long and
32-40 m wide, with an area of 5450 sq m. rt was -buaitiofwsand; Les
surface about 1m above sea level, with several strips of relict beach-
rock 25-30 m off the east shore. During Hurricane Hattie there was
some beach retreat, especially at the northern and southern ends,
exposing fresh beachrock. Many coconuts were felled in a generally
northerly direction. Immediately after the storm the main coloniser

Oye

was Euphorbia. HOG. detamlish Seer StoddanesGlIG6s) i ipp.. Sii—oo aug SS)! .
By 1965 a number of young coconuts had been planted, and low
Tournefortia bushes were growing along the northern part of the seaward
shore. The ground cover comprised sparse Euphorbia, Sesuvium and
Ipomoea, but was repeatedly cleared (Stoddart 1969, p.1l).

By 1972 erosion had continued at the south point and along the
southern end of the leeward shore, where the shore had retreated up to
15 m since 1962. Seasonal sandspits at both the northern and southern
points partly mask the intensity of this continuing erosion. As a
result the area of the island declined from 5450 sq m in 1960 to 4260
Sqamean) 1965) and 3940%sq m in 1972. A number of new coconuts had been
planted, but the ground surface was Kept bare, apart from two clumps of
Tournefortia on the seaward shore. Further erosion occurred at the
northern and southern points during Hurricane Fifi in December 1974,
and the island was remapped by K. Ruetzler and H. Pulpan (in Miller and
Macintyre 1977, p.27); its area was reduced to 2810 sq m. In spite of
the fact that vegetation was regularly cleared from the cay surface,
many seedlings germinate. Some were recorded in 1972, and others in
a collection by J.D. Ferraris in May 1978. 22 species have now been
recorded from Carrie Bow Cay, at least 9 of them only as seedlings.

The list is as follows:

Paspalum distichum 72C Cakile lanceolata 72C 78C
Cyperus planifolius 72C (seedling)
Cocos nucifera 60S 62C 65S Bursera simaruba 72C (seedling)
72S Euphorbia blodgettii 72C 78C
Eichhornia crassipes 78C Euphorbia mesembrianthemifolia
(seedling) V2E" WIC
Hymenocallis littoralis 78S Euphorbia sp. 60S 62S
(seedling) 79C * Terminalia catappa 72C (seedling)
* Casuarina equisetifolia 78C Ipomoea. pes-caprae 72C (seedling)
Coccoloba uvifera 72C 78C Ipomoea sp. 65S
(seedling) Tournefortia gnaphalodes 65S
Suaeda linearis 78C 72C 78C (seedling)
Philoxerus vermicularis 72C 78C Eclipta alba 78C
Portulaca oleracea 72C 78C Wedelia trilobata 72C
Sesuvium portulacastrum 65S
12E WiSE

Salvin visited South Water Cay in May 1863 and collected birds
(Coues 1864, p.391).

Curlew Cay (16°474'N, 88°05 'Ww)

Owen (1830) noted bushes 6 m tall here, at a time when Curlew Cay
must have closely resembled Carrie Bow Cay. In 1960 and 1961, however,
the island was a low crescentic sandbore 36 m long and up to 9 m wide.
It was unvegetated apart from two Rhizophora seedlings. There is
extensive beachrock east of the present site of the cay. The date of
destruction of the vegetated island is unknown; it may have been during
the 1945 hurricane. The sandbore disappeared after Hurricane Hattie,

22

but had reappeared by April 1962. It was present in 1972, but clearly
still ephemeral and of fluctuating dimensions. Its area in 1960 was
370 sq m. Only one species of plant is recorded:

Rhizophora mangle 60S

For further details see Stoddart (1963, p.58, fig. 34). Salvin visited
this island in 1863 and collected birds (Coues 1864, p.391).

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Plate 2. St George's East Cay: aerial view from the northwest 1961

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Plate 12. Rendezvous Cay: aerial view from the southeast 1962

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Plate 14. Rendezvous Cay from the west 1972
Plate 15. Rendezvous Cay from the southwest 1972
Plate 16. Cay Glory: aerial view from the east 1961

Plate 17.

Plate 18.

Rendezvous Cay: erosion on the east shore 1972

Rendezvous Cay: aggradation

on the southwest shore 1972

Plate 19.

Plate 20.

Tobacco Cay: aerial view from the northwest 1960

Tobacco Cay: aerial view from the east 1962

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Plate 21.
ridge 1960

Plate 22.

Tobacco Cay from the south, seen from the reef-crest rubble

South Water Cay: aerial view from the south 1961

23. South Water Cay: aggradation at the southwest point 1972

Plate 24. South Water Cay: erosion on the northwest shore 1972

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23

4. SAND CAYS OF THE SOUTHERN BARRIER REEF

South of Gladden Spit (16°31'N, 87°59'W) the barrier reef extends
in a northeast-southwest direction for 55 km to Ragged Cay, though
surface reef patches are discontinuous and small. There are several
scattered cays in the northern part of this reef, and a group of
islands known as the Sapodilla Cays in the south. At its southernmost
extent the barrier reef curves sharply west and north, and terminates
aemsealiCay.. The islands are described from north to south.

Silk or Queen Cays

The Silk Cays are situated 9 km southwest of Gladden Spit, ata
channel through the barrier reef carrying 6 m of water. They are
charted as "Queen Cays", a name applied to them by the officers of the

1830 survey. Winzerling (1946) states that the name derives from a
seventeenth century local vessel Queen of Bohemia, but we have seen no
evidence for this. Smith (1842, p.732), however, states that "the
Queen cays ... were formerly named the Seal Cays, now altered to prevent

confusion [with other islands of this name in 17°21'N and 16°08'N]".
The name Silk Cays is used locally and should be retained, though the
Admiralty usage is well-established on charts and in pilot books, and
has been used by James and Ginsburg (1979) for one of their reef survey
Sites. The group consists of three small vegetated islands standing
on patch reefs; for a survey of their location see the survey by
Barnett in 1840, inset to Admiralty Chart 1797.

North Silk Cay (16°274'N, 88°0O2)'W) Figure 12, Plates 29 and 33

North Silk Cay is an oval-shaped island built of concentric shingle
ridges enclosing a small shallow pool, and with a leeward sand beach
masking the shingle. It was mapped in 1961 and 1972. During this
interval the distribution of the shingle ridges changed slightly; the
greatest dimension increased from 62 to 70 m; the width remained the
same at 48 m; and the area increased from 0.24 to 0.25 ha. Pumice lay
thickly on the pool margins in 1960 but was no longer conspicuous in
LQ) 7/2 In 1960 the centre of the island was covered with a low thicket
of Suriana, Tournefortia, Conocarpus and Avicennia, with several
Rhizophora round the pool margins, and patches of Ipomoea, Sesuvium,
Erithalis and Cyperus on the shingle surface. There were no coconuts.
In June 1972 the vegetation was more extensive, but still dominated by
Rhizophora and Avicennia. around the pool. The main shrubs were again

24

Tournefortia and Suriana, and the ground cover included Sesuvium,

Ipomoea, Portulaca and Hymenocallis. There were three young coconuts.
No beachrock was seen during either survey. Altogether 13 species of
plants have been recorded, 9 in 1961 and 12 in 1972. New records in
the second survey included Portulaca, Hymenocallis, Cocos and Paspalum;
Erithalis disappeared. The species recorded are as follows:

Paspalum distichum 72C Suriana maritima 61S 72C

Cyperus planifolius 72C Rhizophora mangle 61S 72C

Cyperus sp. 618 Conocarpus erectus 61S 72C

*Cocos nucifera 72S Ipomoea macrantha 72C

Hymenocallis littoralis 72C Ipomoea sp. 618

Portulaca oleracea 72C Tournefortia gnaphalodes 61S 72C

Sesuvium portulacastrum 61S Avicennia germinans 61S 72C

XS Erithalis fruticosa 61S

Middle Silk Cay (16°28'N, 88°02 'W) Figure 13, Plate 34

Middle Silk lies about 300 m southwest of North Silk Cay. In 1961
it was oblong in shape, with maximum dimensions of 80 x 45 m and an area
of 0.43 ha. It was built of concentric sand ridges rising about 1m
above sea level, with sand spits extending to southeast, northeast and
northwest. It stands on a small patch reef, the windward crest of
which is outlined by emergent rubble. Relict beachrock off the
southern side of the island indicated northerly migration of at least
50 m. The vegetation cover was restricted to a cluster of
Tournefortia and Suriana bushes at the eastern end of the cay, some
scattered shrubs of Suriana, and a few coconuts, with a ground cover of
Sesuvium, Ipomoea and Euphorbia. By June 1972 there had been
considerable changes. The island was now triangular, with a cliffed
southern shore and aggradation on the northern side; one line of old
beachrock was visible, passing beneath the cay sands. The area was
reduced to 0.40 ha, but the vegetation cover had substantially increased.
It was dominated by Tournefortia, Suriana and Conocarpus bushes, with
three tall coconuts and about 20 recently planted juveniles. The
ground surface was covered with Euphorbia, Sesuvium, Ipomoea, Cyperus,
Wedelia and Erithalis. A temporary hut had been constructed in the
middle of the island. Six species of plants were seen in 1961 and 7
more in 1972, making a total flora of 13 species:

Eragrostis prolifera 72C Conocarpus erectus 72C

Paspalum distichum 72C Ipomoea macrantha 72C

Cyperus planifolius 72C Ipomoea pes-caprae 72C

*Cocos nucifera 61S 72S Ipomoea sp. 6158 ,

Sesuvium portulacastrum 61S Tournefortia gnaphalodes 61S
W2E TAS

Suriana maritima 61S 72C Erithalis fruticosa 72C

Euphorbia mesembrianthemifolia Wedelia trilobata 72C
Xe

25

South Silk Cay (16°274'N, 88°03'W) Figure 14, Plates 30 and 35

South Silk Cay is situated 460 m southwest of Middle Silk Cay, and
like it is built wholly -of sand. In 1961 it was a lens-shaped island
90 m long and up to 32 m wide, with an area of O.17 ha. Its surface
stood about 1 m above sea level. It was covered by a scrub of
Suriana, with Hymenocallis and Sesuvium, and less than a dozen
scattered coconuts. There was a Single Coccoloba tree at the eastern
end. By June 1972 the island was of similar shape but had changed its
orientation and location. The vegetated area had been eroded and
cliffed at its eastern end, and then extended by a broad sandspit.

The cay was then 110 m long and 30 m wide, with an area of O.23 ha.
The single Coccoloba tree survived, and as before the dominant

vegetation was of Suriana, Hymenocallis and Sesuvium. There were six
coconuts. No beachrock was seen during either survey. The extent of
Hymenocallis is of particular interest on this cay. Five™Specites) Of

plants were noted in 1961 and a further four in 1972, giving a total of
9 for the island:

Eragrostis prolifera 72C Portulaca oleracea 72C

Cyperus planifolius 72C Sesuvium portulacastrum 61S 72C
*Cocos nucifera 61S 72S Suriana maritima 61S 72C
Hymenocallis littoralis 61S 72C Euphorbia mesembrianthemifolia
Coccoloba uvifera 61S 72C 72C

Samphire Cay (16°264'N, 88°03'W)

This was charted by Barnett in 1840, 1.6 km south of South Silk
Cay, but had disappeared by 1913. It has not been seen since. It was
apparently vegetated and must have resembled the Silk Cays. "Samphire"
usually denotes Sesuvium portulacastrum in this area.

Round Cay (16°25'N, 88°05'W) Figures 15 and 17, Plate’ 31

Round Cay stands on a patch reef at the inner edge of the barrier
platform, about 1.6 km from the shelf edge. In 1960 it was a lens-
shaped island 120 m long, up to 45 m wide, and with an area of 0.25 ha.
The northern and southern shores consisted of ridges of shingle and
rubble, with coral boulders up to O.3 m in diameter. The ridges rose
to 1.8 m above sea level; the centre of the island, of sand and fine
shingle, stood at about 1.5 m. The eastern end of the island was
formed by a broad sand beach, overlooking a zone of broken intertidal
rubble. The island was covered with a dense coconut woodland with
marginal Tournefortia and Suriana. Under the coconuts there was a
thick cover of Hymenocallis and grasses. There were two lines of
beachrock along the southern shore. By 1972 the island had extended
to 110 x 40 m, and its area increased to O.31 ha. The southern shingle

26

shore had extended to cover the beachrock, and the eastern sand beach
to largely cover the rubble zone. The island continued to be covered
with a coconut woodland with Hymenocallis beneath. The marginal
Tournefortia was much less extensive, however, presumably as a result
of substrate erosion; and there was a large clump of Conocarpus at the

south point. Nine species were noted in 1960 and 15 in 1972, giving a
total flora of 16 species; only one (Cakile lanceolata) had disappeared
between the surveys. Species recorded are:

Paspalum distichum 72C Cakile lanceolata 60S

Paspalum sp. 60S Suriana maritima 60S 72C

Cyperus planifolius 72C Euphorbia mesembrianthemifolia

Cyperus sp. 60S ZC

*Cocos nucifera 60S 72C Euphorbia sp. 60S

Hymenocallis littoralis 60S 72C Conocarpus erectus 72C

Ficus sp. 7J2€ Ipomoea macrantha 72C

Rivina humilis 72C Ipomoea pes-caprae 72C

Portulaca oleracea 72C Tournefortia gnaphalodes 60S 72C

Sesuvium portulacastrum 60S 72C Morinda citrifolia 72C

Pompion Cay (16°24'N, 88°06%'W) Figures 16 and 17, Plates 32 and 40

Pompion Cay, like Round Cay, is situated on a reef patch rising
from the inner edge of the barrier platform, 3.2 km southwest of Round
Cay. It stands about 1.6 km from the seaward edge of the barrier
platform, and with the exception of four small patch reefs nearby it is
the only reef to reach the surface between Round and Ranguana Cays, a
distance of 1.5 km. When the island was mapped in 1960 it was of
simple outline, 115 m long and up to 50 m wide, with an area of 0.35 ha.
The eastern end of the cay was low and sandy; the northern and western
shores are built of shingle and rubble rising to 1.3 m above sea level.
There was no beachrock. On the northwest side the beach crest was
colonised by a mat of Sesuvium with scattered Tournefortia. To the
northeast there is a clump of Coccoloba, and along the south shore
mature trees of Bursera and Cordia. The main part of the cay was
under coconut woodland with a ground cover of Hymenocallis, Erithalis
and Cakile. By 1972 the island had expanded, both by sand accretion
at the east end and shingle accretion at the west end, to a total length
of 123 m; it was slightly narrower (45 m maximum), and had an area of
0.42 ha. The Coccoloba clump and trees of Bursera and Cordia could
still be located. The area of Hymenocallis beneath the coconuts had
expanded and coalesced; the rest of the vegetated surface was covered
with Sesuvium, Cyperus and Euphorbia. The area of coconut woodland
was very Similar to that of 1960. Hydrographic charts mark a prominent
"White Rock" to the west of the cay. This is a small shingle island,
40 m in diameter and 1 m above sea level, built of Acropora sticks in
concentric ridges, Situated about 500 m to the west of the cay. Tt..is
not vegetated and owes its colour to guano staining. Eleven plant
species were noted on Pompion in 1960 and 19 in 1972, giving a total of
21 species. Of those noted in 1960, Cakile and Erithalis were not
FOUN ans The species recorded are as follows:

x]

Sporobolus virginicus 72C Bursera simaruba 60S 72C
Cyperus planifolius 72C Euphorbia mesembrianthemifolia
*Cocos nucifera 60S 72C 72C

Hymenocallis littoralis 60S 72C Euphorbia sp. 60S

Ricus Sp. /2€ Rhizophora mangle 72C (seedling)
Coccoloba uvifera 60S 72C Conocarpus erectus 72C
Philoxerus vermicularis 72C Ipomoea pes-caprae 72C

Rivina humilis 72C Cordia sebestena 60S 72C
Portulaca oleracea 72C Tournefortia gnaphalodes 70S
Sesuvium portulacastrum 60S 72C W2€

Cakile lanceolata 60S Erithalis fruticosa 60S

Suriana maritima 60S 72C Morinda citrifolia 72C (seedling)

Ranguana Cay (16°20'N, 88°093'w) Figure 18, Plates 36, 41 and 42

Ranguana Cay stands on the north side of a channel (Ranguana
Entrance) through the barrier platform, carrying 6.7 m of water. IG Ey aS)
located on a patch reef at the lagoonward side of the barrier platform
about 1.8 km from the seaward edge. Speer (1766, p.24) noted the
island, “called by the Baymen Renegado Kay", and Jefferys marks both
names on his 1775 chart. Barnett's survey in 1840 noted trees 15 m
ieeLIL

In 1960 the island was 255 m long, with a constant width of 55 m,
and had an area of 1.33 ha. About 80 per cent of the total area formed
a core of older darker sand with cliffed margins, extended southeast-
wards by a fresh sand spit with pioneer vegetation. The whole of the
northeast shore was cliffed, with fallen coconuts; the island surface
here is at its highest, reaching 1.5 m. The vegetated area is
dominated by coconut woodland. Suriana, Coccoloba and more rarely
Tournefortia were found along the northeast shore, with Hymenocallis,
sedges and grasses beneath the coconuts. The fresh sand spit was
colonised by young Tournefortia, Euphorbia and grasses. By LOT 2ethe
cay had been much eroded at its eastern end, though it still consisted
of an older core area and a southeastern sand spit. Its length was
reduced to 240 m and its area to 1.12 ha. Much of the margin of the
core area was cliffed. The coconut woodland had a ground cover of
Euphorbia, grasses and sedges, with patches of Hymenocallis. There
were several clumps of Coccoloba near the north point, one collapsed
into the sea as a result of shore erosion. The sand spit was colonised
by Sesuvium. In both 1960 and 1972 there was a rubble ridge about 25 m
long located 45 m northwest of the north point of the cay; in 1972 this
was colonised by Philoxerus.

In 1972 there were 35 pelicans at the cay (on 3 July), 30 boobies,
6 frigates, 16 sooty terns, and several shorebirds. 11 species of
plants were noted in 1960 and 22 in 1972, giving a total of 24. In
addition, one moss was collected (Bryum coronatum Schwaegr., Spellman
and Stoddart 124). The plants recorded are:

28

Eragrostis prolifera 60S 72C
Eustachys petraea 72C
Paspalum distichum 72C
Paspalum sp. 60S

Sporobolus virginicus 72C
Sporobolus sp. 60S

Cyperus planifolius 72C
Cyperus sp. 60S

*Cocos nucifera 57S 60S 72S
Hymenocallis littoralis 60S 72C
Coccoloba uvifera 60S 72C
Philoxerus vermicularis 72C
Portulaca oleracea 72C
Sesuvium portulacastrum 72C

Sophora tomentosa 6OC

Vigna luteola 72C

Suriana maritima 60S

Euphorbia blodgettii 72C

Euphorbia mesembrianthemifolia
VE

Euphorbia sp. 60S

Rhizophora mangle 72C (seedling)

Terminalia catappa 72C
(seedling)

Ipomoea macrantha 72C

Tournefortia gnaphalodes 60S
12

Stachytarpheta jamaicensis 72C

Cassytha filiformis 72C Wedelia trilobata 72C

Cakile lanceolata 72C

North Spot (16°15'N, 88°11'W) Figure 19, Plates 37, 38 and 44

North Spot lies 10 km southwest of Ranguana Cay,on the south side of
a channel through the barrier platform which carries 6.4 m of water. It
stands on a patch reef about 360 m wide. When mapped in 1960 the
island was a ridge of Acropora shingle 85 m long, with a maximum width
of 27 m and.an area of 0.05 ha- It was orientated northeast-southwest.
It was covered with a thick mat of Sesuvium, Portulaca and Ipomoea, with
a Single juvenile coconut and some grasses. Barnett's survey in 1840
noted that the island was "very low sand with grass and clump of cocoas".
Presumably the coconuts had been destroyed during a hurricane and the
cay re-formed. In 1972 the geomorphology was transformed. North Spot
was then a near-circular island, with greatest dimensions of 53 x 35 nm,
and an area (0.13 ha) double that of 1960. The surface was covered
with a mat of Sesuvium over coarse rubble, with patches of Ipomoea.
In addition there were single trees of Rhizophora and Avicennia, and a
clump of Tournefortia. The surviving coconut was about 5m tall.
There was a flock of about 100 sooty terns on the island on 4 July 1972.
Three species of plants were recorded in 1960 and 7 in 1972, giving a
EGtail of 78:

Rhizophora mangle 72C
Ipomoea macrantha 72C
Ipomoea pes-caprae 72C
Tournefortia gnaphalodes 72C

*Cocos nucifera 60s 72S
Philoxerus vermicularis 72C
Portulaca sp. 60S
Sesuvium portulacastrum 60S 72C

Red Rock and Black Rock (16°14'N, 88°114'W; 16°13'N, 88°11}'W) of
charts are unvegetated rubble islets between North Spot and Tom Owen's
Cays.

29

"Tom Owen's Cay" (16°12'N, 88°13'W) Figures 20 and 21

South of 16°15'N the history and nomenclature of the barrier reef
cays becomes complex. Owen in his 1835 survey charted three islands
near the major channel which intersects the barrier platform in latitude
IS STUN One on the north side he named Tom Owen's Cay; it was
described as having "one Cocoa N. Tree & Shrubs". Two small un-named
islands on the south side were described as having "one large tree and
high shrubs" (on the west) and "grass and bushes" (on the east)

(Admiralty Ms L45). The northern cay has now disappeared, though
still marked on Admiralty Chart 1573, and the name Tom Owen's Cays is
now given locally to the two southern islands. Both are conspicuous

islands, the eastern cay with a cluster of tall coconuts, and the
western with a larger coconut woodland.

The earlier nomenclature of these islands is uncertain. Speer in
his sailing directions (1766, p.24) mentions "a Kay called Cohune's Kay;
and a very remarkable small Kay, that has many tall Palmetoe Trees on
it; which look like so many round balls hanging in the air". He states
that "Cohune's Kay, and the easternmost of the Sappatilla Kays |[i.e.
Northeast Sapodilla Cay], are N. by E. and S. by W. of each other;
distance about 5 miles". This would bring Cohune's Kay to the position
of the former Tom Owen's Cay; Speer carefully described the deep channel
lying between the two. "Cohune's Kay", he continues, "is full of
bushes and trees. A little to the eastward of Cohune's Kay there are
two Sand Boors; but they join to one another, and to Cohune's Kay by a
Reef. You leave them on your starboard side; and on your larboard
side, you leave a Sand Boor that has breakers all round it, and no
bushes on it. And about two cable's length to the N.W. of this Sand
Boor there is a small spot of Rocks, which breaks when it blows fresh.
And to the eastwards of that Sand Boor there is another Boor. There is
a channel between them of 2 l-half, and 3 fathom water" (1766, p.24).
This would suggest the identity of Cohune's Kay with the present Tom
Owen's West Cay, and that in the eighteenth century the present Tom
Owen's East Cay was one of the sandbores joined to Cohune by a reef.

In this case the "very remarkable small Kay" with palmettoes was
probably old Tom Owen's Cay, now disappeared. Jefferys marks Cohune
Kay on his 1775 chart, and immediately north of it, across a main
channel, "Halls Key". Thus Hall's may also be tentatively identified
with old Tom Owen's. We have no information on when, after 1835, this
island disappeared; there is certainly no trace of it now.

Tom Owen's East Cay (16°114'N, 88°134'W) Figure 22, Plates 39, 43, 46
and 47

Tom Owen's East Cay is the smaller of the two. When mapped in
1960 it was triangular in shape, with sides 35-45 m long. It abuts on
its north side on the reef-crest rubble zone, and the northwest and
northeast shores were built of shingle. Most of the cay was built
of sand, however, rising up to 1 m above sea level. There were four

30

tall coconuts on the island, and a number of smaller ones, together
with a scrub of Tournefortia and Coccoloba and extensive Hymenocallis
and Sesuvium. The island had changed considerably by 1972. The old
island had been eroded and then extended southwards by a new sand spit.
The area had increased from 0.10 to O.12 ha. The area of Coccoloba
had greatly increased, and there was also a patch of Conocarpus.
Otherwise the vegetation resembled that in 1960. No beachrock was
seen in either survey. In 1972 the area of rubble along the north
shore was much more extensive than in 1960, and probably represented a
hurricane event.

Ten plant species were recorded in 1960 and 11 in 1972, giving a

totals ofois: The following are recorded:
Paspalum distichum 72C Sesuvium portulacastrum 60S 72C
Sporobolus virginicus 72C Bursera Simaruba 60S
Cyperus planifolius 72C Euphorbia mesembrianthemifolia
Cyperus sp. 60S 72C
*Cocos nucifera 60S 72S Euphorbia sp. 60S
Hymenocallis littoralis 60S 72S Conocarpus erectus 72C
Coccoloba uvifera 60S 72C Tournefortia gnaphalodes 60S
Philoxerus vermicularis 72C Erithalis fruticosa 60S
Portulaca oleracea 72C Wedelia trilobata 60S

Portulaca sp. 60S

Tom Owen's West Cay (16°11%'N, 88°14'W) Figure 23, Plates 45, 46 and 48

In 1960 Tom Owen's West Cay was about 90 m long and 35 m wide; it
had an area of 0.26 ha. It was fringed along its northern side by a
zone of coral rubble, and both north and south shores were formed of
shingle. The interior was sandy with occasional blocks of coral
rubble, and there were sand beaches at both east and west ends. The
general surface stood about 1m above sea level. The island was
covered with coconut woodland, especially dense at the western end, with
an undercover of Wedelia, Cakile, Ambrosia, Sesuvium, Euphorbia, sedges
and grasses. There was some marginal Coccoloba, Conocarpus and
Tournefortia. No beachrock was seen. The island remained very much
the same in 1972, with rather minor changes in outline; the area had
decreased to 0.24 ha. It was still covered with coconut woodland, but
Conocarpus and Caesalpinia appeared much more extensive. Cordia
survived from 1960 on the south shore. Portulaca was extensive on the
windward beach crests. Fifteen species of plants were recorded in 1960
and: 17 inbil972,, soinvang a total jo£ 22:

Paspalum distichum 72C *Cocos nucifera 60S 72S
Paspalum sp. 60S Coccoloba uvifera 60S 72C
Sporobolus virginicus 72C Iresine diffusa 72C
Sporobolus sp. 60S Rivina humilis 72C
Cyperus planifolius 72C Portulaca oleracea 72C

Cyperus sp. 60S Portulaca sp. 60S

31

Sesuvium portulacastrum 60S 72C Ipomoea pes-caprae 72C

Cakile lanceolata 60S Tournefortia gnaphalodes 60S
Caesalpinia bonduc 72C 72C

Canavalia rosea 60S Lippia. nodiflora 72C
Thespesia populnea 72C Erithalis fruticosa 60S
Rhizophora mangle 60S 72C Ambrosia hispida 60S
Conocarpus erectus 60S 72C Wedelia trilobata 60S

Ipomoea macrantha 72C

One of the present Tom Owen's Cays was visited by the Marshall
Field Expedition in 1923, when K.P. Schmidt and L.L. Walters collected
two lizards (Aristelliger georgeensis and Anolis sagrei) (Schmidt 1941).

Cays between Tom Owens Cays and Northeast Sapodilla Cay Figures 20 and 21

Owen in 1835 (Admiralty MS L45) charted two small islands lying
respectively 3.2 and 4 km southsouthwest of the present Tom Owen's Cays.
The northern island had "dry sand, one Cocoa N. Tree and Shrubs"; the
other "Dry Sand, 2 Cocoa N. Trees, Shrubs & bushes". They were
situated on the south side of the main channel between Tom Owen's and
Northeast Sapodilla Cays, and are probably the "Sand Boors" referred to
by Speer (1766, p.24; quoted above). Both islands have now disappeared,
though still marked on charts; when they were destroyed is not known.

The Sapodilla Cays Figures 20 and 21, Plate 49

The southern six kilometres of the barrier reef consists of a
series of large shoal patches intersected by deep channels. The six
islands now existing on these patches are termed the Sapodilla Cays
(Admiralty MS L45, 1835) or Zapotilla Cays (Chart 1573), and there is
some informal suggestion of a territorial claim to them by the Republic
of Honduras (Aguilar Paz 1954). The main islands are here referred to
as Northeast Sapodilla, Frank's, Nicolas, Hunting, Lime and Ragged Cays.
Speer (1766, 24) refers to them as "six large Champaign Kays" [champaign:
"an expanse of level, open country. Unenclosed or common land"

(Shorter Oxford English Dictionary, I (1933), p.289)].

Northeast Sapodilla Cay (16°O8'N, 88°15'W) Figure 24,Plates 50,52 and 53

Northeast Sapodilla Cay, the most northerly, is a rectangular
island with greatest north-south and east-west dimensions of 300 m. In

1960 its area was 4.79 ha. The seaward shore was formed by a shingle
ridge rising 2 m above sea level, highest near the northeast point and
declining southwards. The northwest and southwest beaches are formed

by low sand ridges. The centre of the cay is low-lying and poorly
drained. The main dry land areas were to leeward of the seaward

32

shingle ridge and at the west end of the cay. In the southeast bay
there are 8, possibly 9, lines of relict beachrock, standing up to

30 m from the present beach; the outer lines carry up to 1 m of water.
In 1960 there was a further patch of beachrock, 15 m long, poorly

cemented, on the southwest shore.

The low-lying centre of the island

was occupied by hundreds of young coconut palms, surrounded by
Acrostichum aureum. The sandy areas were occupied by taller coconut
woodland. Coccoloba and Thrinax grew along the seaward beach crest.

A large patch of rubble at the eastern point was colonised by Sesuvium.
Little morphological change had occurred by July 1972 except for
erosion along the northwest bay. This averaged 10 m and reached 20 m
at one point, leaving a cliffed shore with fallen coconut trees. The
southwest sandy shore had been covered with shingle for its entire
length and the small beachrock exposure hidden. On the northeast
shore the beach itself was formed of sand overlying the face of the
high shingle ridge, and low beachrock was exposed for part of its
length. The vegetation was observed in greater detail than in 1960.
In the low-lying coconut woodland with Acrostichum, the dominant woody
species beneath the coconuts was Erithalis. Chrysobalanus was common
round the margins of the area on slightly higher ground, and individuals
were scattered through the woodland itself. Coccoloba, Conocarpus and
Suriana formed a littoral hedge on the beach crest round much of the
windward side of the cay, but Tournefortia was surprisingly absent.
Trees in the coconut woodland included Ficus, Terminalia, Hibiscus
tiliaceus, and Citharexylum caudatum, this last forming trees 3-6 m
tall near the south point. The island was inhabited in 1960 but had
been abandoned by 1972, when the main house was in ruins and the jetty

broken.

26 plant species were recorded in 1972, which with the six noted
in 1960 brings the total recorded to 28:

Acrostichum aureum 60S 72C
Eragrostis prolifera 72C
Eustachys petraea 72C
Paspalum blodgettii 72C
Paspalum distichum 72C
Sporobolus jacquemontii 72C
Stenotaphrum secundatum 72C
Cyperus planifolius 72C
*Cocos nucifera 60S 72S
Thrinax radiata 60S 72C
*Musa sapientum 72C

Bicus (Spe AZ

Coccoloba uvifera 60S 72C
Sesuvium portulacastrum 60S

Frank's Cays (16°O8'N, 88°15'wW)

Cakile lanceolata 60OC
Chrysobalanus icaco 72C
Sophora tomentosa 72C
Suriana maritima 72C
Euphorbia blodgettii 72C
Hippocratea volubilis 72C
Hibiscus tiliaceus 72C
Passiflora suberosa 72C
Conocarpus erectus 72C
*Terminalia catappa 72C
Citharexylum caudatum 72C
Stachytarpheta jamaicensis 72C
Prithalus prueba cosagH2©
Wedelia trilobata 72C

Figure 25, Plate 54

Owen in 1835 charted a single island 0.5 km south of Northeast
Sapodilla Cay, named "Grass Cay", and this is marked on Admiralty Chart

38

D730 Three islands now exist at this point and are known as Frank's
Cays: a main island (Frank's Cay) and two smaller ones (Frank's Cay
East, Frank's Cay West). Whether these have formed by fragmentation

of the earlier island or whether the small cays have formed since 1835
is not known.

Frank's Cay in 1960 was 230 m long and up to 100 m wide. It was
low and formed entirely of a dark sand. The beaches were low and
uneroded. At the eastern end of the cay there was a dense belt of
mangroves (Rhizophora, Conocarpus, Laguncularia, Avicennia) . The
rest of the island was covered with a dense coconut; woodland with herbs
and grasses. Conocarpus, Coccoloba and Thrinax were found around the
shores. There were two patches of poorly cemented beachrock at the
eastern end of the south shore. By 1972 much of the north shore was
cliffed and a substantial area of mangroves at the northeast point had
been destroyed. The area of the cay had declined from 1.94 ha to
Lo 76 I0e\- The eastern mangrove fringe was much narrower and consisted
mainly of Conocarpus and Laguncularia. Chrysobalanus, Pithecellobium
and Thrinax were common round the shores. The south shore beachrock
was still exposed.

Frank's Cay East is separated from the main cay by a shallow
channel 25 m wide. In 1960 it was an arcuate island 120 m long, but
by 1970 the western embayment had been infilled and the shape of the
island was more regular; the area had increased from O.5 to O.57 ha.
None of the shores were undercut in 1960, but the northeast and

southeast shores were cliffed in 1972. Mangroves fringe the west
Side of the cay. The rest is covered with coconut woodland, with much
Conocarpus, Laguncularia and Thrinax. A continuous belt of Coccoloba

along the northeast shore in 1960 had been reduced to two clumps in
1972, presumably by shore erosion.

Frank's Cay West lies 75 m southwest of the main cay, and in 1960
was a lens-shaped sandy island 70 m long with an area of O.11 ha. ste
was covered with a thicket of Conocarpus and coconut; a single
Avicennia was prominent on the northwest shore. thas tree steal
existed in 1972 but the northwest shore was cliffed; the southeast
shore had been colonised by Rhizophora immediately offshore. Thrinax,
Ficus and Laguncularia were noted in addition to Cocos and Conocarpus.

31 plant species have been recorded from the main island (13 in

IQEO, SO) aliay IDV) In addition the moss Calymperes richardii C. Mull.
was collected (Spellman and Stoddart 129a). Plants recorded are:
Andropogon glomeratus 72C Stenotaphrum secundatum 72C
Eragrostis prolifera 72C Cyperus peruvianus 60C 72C
Eustachys petraea 60C 72C Cyperus planifolius 72C
Paspalum blodgettii 72C Fimbristylis cymosa -72C
Paspalum sp. 60S *Cocos nucifera 60S 72S
Spartina patens 72C Thrinax radiata 60S 72C
Sporobolus virginicus 72C ICUS So VAC

Sporobolus sp. 60S Coccoloba uvifera 60S 72C

34
Cakile lanceolata 60S Rhizophora mangle ©OS 72C
Chrysobalanus icaco 72C Conocarpus erectus 60S 72C
Gliricidia sepium 72C Laguncularia racemosa 60S 72C
Pithecellobium keyense 72S *Terminalia catappa 72C
Sophora tomentosa 72C Avicennia germinans 72C
Euphorbia blodgettii 72C Lippia nodiflora 72C
Euphorbia mesembrianthemifolia Stachytarpheta jamaicensis 72C
VAS Erithalis fruticosa 72C
Euphorbia sp. 60S Wedelia trilobata 60S 72C
Passiflora suberosa 72C
Seventeen species are recorded from Frank's Cay East (12 in 1960,
Ibi walsh AUS) 7/2) They include the uncommon orchid Brassavola nodosa from
the Rhizophora at the easternmost point. Species recorded are:
Acrostichum aureum 72C Cakile lanceolata 60S
Andropogon sp. 60S Pithecellobium keyense 72S
Eustachys petraea 60S Euphorbia sp. 60S
Paspalum sp. 60S Rhizophora mangle 60S 72S
* Cocos nucifera 60S 72S Conocarpus erectus 60S 72S
Thrinax radiata 60S 72S Laguncularia racemosa 60S 72S
Brassavola nodosa 72C Avicennia germinans 60S
Coccoloba uvifera 60S 72S Erithalis fruticosan?2ZsS

Twelve species are recorded from Frank's Cay West (9 in 1960 and
6-1an L972). They are:

Andropogon sp. 60S Canavalia rosea 60S
Paspalum sp. 60S Euphorbia sp. 60S
Sporobolus sp. 60S Rhizophora mangle 60S 72S
* Cocos nucifera 60S 72S Conocarpus erectus 60S 72S
Thrinax radiata 72S Laguncularia racemosa 72S
FUCUS “Spee 125 Avicennia germinans 60S
Nicolas Cay (16°O7'N, 88°16'W) Figure 26, Plates 51 and 57

Nicolas Cay lies 2 km southsoutheast of Frank's Cays, on the
northern side of a channel carrying 11m of water. It is oval-shaped,
and in 1960 had maximum dimensions of 310 x 190 m. There was some
beach erosion at the north and southwest ends of the cay, but most of
the shore is formed by a broad sand beach. No beachrock was exposed.
By 1972 there had been considerable erosion, averaging 10-15 m, on all
Sides of the cay except the south and southeast. Shores were cliffed
on the west and northeast sides, and a soft root filled lithified
material ("root rock') was exposed on the west shore. The area had
declined, from ~4.53.to 3°86 cha. The cay is densely vegetated with
coconut woodland, with an understorey of Psychotria nervosa. On the
seaward side of the cay the beach crestis lined by Coccoloba,
Conocarpus, Cordia and Sophora, with Erithalis and Chrysobalanus,

35

partly fronted by a strip of herbs and grasses with germinating drift
coconuts. A patch of Tournefortia growing at the east point in 1960
had disappeared as a result of beach retreat by 1972, and this removed
the species from the island. The littoral hedge on the west shore is
largely formed by Erithalis and Conocarpus. Other species noted
include Terminalia and a large Ficus. There are temporary fishing
huts on the cay, but it is uninhabited. Sherds of red Maya pottery
were eroding from superficial midden deposits in the area of root rock
exposure on the west shore in 1972 (Stoddart 1980, p.162). When
mapped by Owen in 1835 the island was inhabited and had fresh water.

32, species have been recorded from the cay (4 in 1960, 31 in 1972):
In addition four mosses were collected in 1972: Taxithelium planum
(Brid.) Mitt. (Spellman and Stoddart 125b), Callicostella depressa
(Hedw.) Jaeg. (Spellman and Stoddart 125c), Calymperes erosum C. Mull.
(Spellman and Stoddart 126BO), and Calymperes richardii C. Mull.

(Spellman and Stoddart 127a). The plant species recorded are:
Nephrolepis multiflora 72C Pithecellobium keyense 72C
Polypodium lycopodioides 72C Sophora tomentosa 72C
Polypodium polypodioides 72C *Citrus aurantiifolia 72C
Andropogon glomeratus 72C Euphorbia mesembrianthemifolia
Eragrostis prolifera 72C 72C
Eustachys petraea 72C Passiflora suberosa 72C
Paspalum blodgettii 72C Conocarpus erectus 60S 72C
Paspalum distichum 72C *Terminalia catappa 72C
Sporobolus virginicus 72C Pouteria campechiana 72C
Cyperus peruvianus 72C *Nerium oleander 72C
Cyperus planifolius 72C Tournefortia gnaphalodes 60S
Fimbristylis cymosa 72C Lippia nodiflora 72C

*Cocos nucifera 60S 72S Diodia serrulata 72C
Thrinax radiata 72C Erithalis fruticosa 72C
Ficus sp. 72C Ernodea littoralis 72C
Coccoloba uvifera 60S 72C Psychotria nervosa 72C

Chrysobalanus icaco 72C

Hunting Cay (16°07'N, 88°16'W) Figure 27, Plates 55, 56 and 58

Hunting Cay lies 0.6 km south of Nicolas Cay, on the south side of
the channel which divides the barrier platform at this point. When
mapped in 1960 it had a maximum north-south length of 440 m and was up
to 220 m wide; its area was 5.79 ha. The west and south coasts were
formed by broad sand beaches. The east coast was more complex, with
two projections separated by a wide sandy bay facing a gap in the reef.
The northeast shore was built of shingle but the other seaward shores
of sand. Both in the bay and on the southern headland beachrock
indicated beach retreat. There waS some undercutting near the north-
west point. By 1972 there had been erosion on the west and south
shores as well as in the east bay. The west shore was cliffed, with
fallen coconuts, for much of its length. More beachrock had been
exposed on the eastern side.

36

The island is covered with a dense coconut woodland. Coccoloba
formed an extensive littoral hedge along the east shores in 1960 and
again an 972): Grasses and juvenile Casuarina were colonising the
fresh sand accumulations in the south in 1960 and 1972. Beach retreat
in the east bay had destroyed the only colony of beach-crest
Tournefortia between the two surveys, and this species was not found
an, L972 Hymenocallis is particularly common in the southeast part
of the island. Vermeer (1959, p.90) mentions Manilkara zapota in the
coconut woodland, but this record has not been confirmed. The cay is
inhabited and has a lighthouse. Craig (1966) mentions it as one of
the islands with Maya remains, but these were not seen during our
surveys.

58 species of plants have been recorded, 7 in 1960 and 57 in 1972.
They are:

Acrostichum aureum 72C Euphorbia blodgettii 72C
*Eleusine indica 72C Euphorbia mesembrianthemifolia
Eragrostis ciliaris 72C 12Z€

Eragrostis prolifera 72C Phyllanthus amarus 72C
Eustachys petraea 72C Hippocratea volubilis 72C
Paspalum blodgettii 72C Hibiscus tiliaceus 72C
Paspalum distichum 72C Passiflora suberosa 72C
Sporobolus virginicus 72C *Cucumis melo 72C

Sporobolus sp. 60S Conocarpus erectus 60S 72C
Cyperus ligularis 72C *Terminalia catappa 72C
Cyperus peruvianus 72C Eugenia sp. 72C

Cyperus planifolius 72C [Manilkara zapota Vermeer (1959,
Fimbristylis cymosa 72C p.90) |
*Cocos nucifera 60S 72S *Nerium oleander 72C

Thrinax radiata 72C *Plumeria rubra 72C
Hymenocallis littoralis 60S 72C Rhabdadenia biflora 72C
Brassavola nodosa 72C Tournefortia gnaphalodes 60S
*Casuarina equisetifolia 60S 72C Citharexylum caudatum 72C
BiCuSs;, Spi) ZC Lippia nodiflora 72C
Coccoloba uvifera 60S 72C Stachytarpheta jamaicensis 72C
Portulaca oleracea 72C *Russelia equisetiformis 72C
(seedling) Enallagma latifolia 72C

Cakile lanceolata 72C Diodia serrulata 72C
*Kalanchoe pinnata 72C Erithalis fruticosa 72C
Chrysobalanus icaco 72C Ernodea littoralis 72C
Canavalia rosea 72C *Txora coccinea 72C
Gliricidia sepium 72C Spermacoce assurgens 72C
Pithecellobium keyense 72C Spermacoce prostrata 72C
Sophora tomentosa 72C Ageratum littorale 72C

Vigna luteola 72C Bidens pilosa 72C
*Citrus aurantiifolia’ 72C Conyza canadensis 72C

*Codiaeum variegatum 72C

S7/

Lime Cay (16°O06'N, 88°164'W) Figure 28, Plates 59-61

Lime Cay, the "Low Cay" of charts, lies nearly 1 km south of
Hunting Cay. In 1960 the island was 300 m long and up to 110 m wide;
it had an area of 1.83 ha. The northeastern point was built of
shingle and abutted on a semicircular reef-crest rubble zone; the other
shores were sandy. Much of the northwest-facing shore was cliffed,
and beachrock offshore indicated past beach retreat. The shingle hook
at the northeast point was apparently a recently formed and perhaps
still mobile feature. In 1972 the hook had disappeared and the north-
eastern and northwestern shores were cliffed and eroding. The width
of the island in this sector had decreased from 75 to as little as 35 m.
Conversely there was a great deal of fresh sand forming two large spits
at the southwest end (in July 1972), and as a result the island had
increased in length to nearly 330 m and in area to 2.07 ha.

The island is covered with a coconut woodland, which is most dense
in the southwest. In the centre, on low-lying poorly-drained ground,
there is an area of Hymenocallis, Thrinax and Acoelorrhaphe. Beach-
crest vegetation consists of discontinuous Erithalis, Coccoloba,
Conocarpus and Sophora; in particular the small patch of Conocarpus
noted on the shingle hook in 1960 had greatly expanded by 1970. En
1970 there was also a small patch of Sesuvium on the seaward rubble.

Thirty species of plants have been recorded from Lime Cay, 5 in
1960 and 28 in 1972. The Rhizophora seedlings colonising inside the
shingle hook in 1960 had disappeared by the later survey. Species
recorded are:

Nephrolepis multiflora 72C Hymenocallis littoralis 72C

Andropogon glomeratus 72C Coccoloba uvifera 60S 72C
Anthephora hermaphrodita 72C Sesuvium portulacastrum 60S 72S
Cenchrus incertus 72C Pithecellobium keyense 72C
Eragrostis prolifera 72C Sophora tomentosa 72C

Eustachys petraea 72C Euphorbia mesembrianthemifolia
Panicum pilosum 72C 72C

Paspalum blodgettii 72C Passiflora suberosa 72C
Sporobolus virginicus 72C Rhizophora mangle 60S (seedlings)
Stenotaphrum secundatum 72C Conocarpus erectus 60S 72C
Cyperus peruvianus 72C *Terminalia catappa 72C

Cyperus planifolius 72C Lippia nodiflora 72C
Fimbristylis cymosa 72C Stachytarpheta jamaicensis 72C
Acoelorraphe wrightii 72C Diodia serrulata 72C

*Cocos nucifera 60S 72S Erithalis fruticosa 72C

Thrinax radiata 72C

In addition the moss Calymperes richardii C. Mull. was also collected

(Spellman and Stoddart 128a).

38

Ragged Cay (16°05'N, 88°174'W) Figure 29, Plates 62-63

Ragged Cay, the southernmost on the barrier reef, appears on
charts as "South Cay" and "Zapotilla Cay". It stands on a small reef
patch with deep channels to east and west. In 1960 it was triangular,
with sides 40-70 m long, and with an area of 0.24 ha. It was built
entirely of concentric ridges of coarse shingle, reaching about 1 m
above sea level. By 1972 it had changed considerably, and was an oval
island, orientated north-south, 75 m long and 35 m wide, with an area
OLORZ2 "har: It was still built of shingle, with a small southern sand
spit. Owen in 1835 noted a "Small cluster of cocoanut trees, grasses
and bushes". In 1960 there were five Casuarina trees, one a mature
specimen 15 m tall, together with 10 coconuts. The ground surface was
covered with a thick mat of Sesuvium with bushes of Conocarpus and
Rivina. In 1972 the vegetation remained dominated by the Casuarina
and the coconuts, with Sesuvium and Rivina. A feature of this island
is the great number of coenobitid hermit crabs inhabiting it. Only
seven plant species have been recorded from Ragged Cay (5 in 1960 and
5 in 1972); they are:

Cyperus planifolius 72C Portulaca oleracea 72C
*Cocos nucifera 1835S 60S 72S Sesuvium portulacastrum 60C
*Casuarina equisetifolia 60S 72C 12C
Rivina humilis 60C Conocarpus erectus 60S
Seal Cays (16°104'N, 88°20'W) Figure 30, Plates 64-65

At Ragged Cay the barrier reef trends westwards for 3.7 km and
northwards for 9 km, forming a narrow hook. Where not occupied by
surface reefs this carries 4-5 m of water over it; inside the hook
the water depths are 29-33 m, and outside it 55 m. The reef is
uninterrupted by channels in its north-south limb. Admiralty Chart
1573 shows Seal Cays on this reef at 16°O8'N: these are the
"Observation Seal" Cays mapped by Owen in 1835. They then comprised
two islands on an arcuate reef, one with a "cluster of cocoa-nut trees",
the other with a "grove of cocoa-nut trees, huts". Only an
unvegetated sandbore occupies this site now; the cays are said to have
been destroyed in the 1945 hurricane. At the northern end of the reef
there are two large reef patches. The southernmost possessed a cay in
1835, with "one bushy tree"; this, too, is now an unvegetated sandbore.
The other forms a perfect faro or circular reef enclosing a lagoon,

about 150 m in diameter, at the end of the barrier reef. It has a cay
at its northeast point, and Owen noted a "small cluster of cocoa-nut
trees" in 1835. This is the present Seal Cay. A group of small

sandbores lies) north of “the cay, bearing 335~, 003°, ,O13° and:021°
frOM “ES snoneny polnt.

Seal Cay is a small sandy island sheltering in the lee of a rubble
ridge which occupies the reef top on the windward side of the faro.
It is arcuate in shape, about 65 m long and only 15-20 m wide, with an

39

area when mapped in 1972 of 0.14 ha. Its main vegetation is
Rhizophora, with single trees of Cordia, Coccoloba, Avicennia and
Cocos. There are shrubs of Concarpus at the west point. The ground
cover consists of Hymenocallis, Ipomoea, Sesuvium and Rivina. The
island is uninhabited but is used as a fishing station: the leeward
beach has been modified by conch shells to form small boat harbours.
Seal Cay was approached but not visited in 1960; there is a note on

it in Vermeer (1959, p.90). Sixteen species of plants are recorded:
Paspalum distichum 72C Thespesia populnea 72C
Cyperus planifolius 72C Rhizophora mangle 72C
Cocos nucifera 1835S 57S 60S 72S Conocarpus erectus 72C
Hymenocallis littoralis 72C Laguncularia racemosa 72C
Coccoloba uvifera 72C Ipomoea macrantha 72C
Philoxerus vermicularis 72C Cordia sebestena 72C
Rivina humilis 72C Avicennia germinans 72C

Portulaca oleracea 72C Lippia nodiflora 72C

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Figure 20. Reefs and cays of the southernmost barrier reef. Based on
original plots by R. Owen, 1835

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Figure 27.

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dense,open/
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Figure 28.

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Plate 40. Hymenocallis littoralis under coconuts at Pompion Cay 1972

Plate 41. Ranguana Cay: erosion on the north shore 1972

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Plate 42.

Ranguana Cay seen from the shingle islet at the west end 1972

Plate 43.

Cay from the southwest 1972

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Plate 44. North Spot 1972

Plate 45. Tom Owen's West Cay: north shore 1972

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Plate 50. Northeast Sapodilla Cay: aerial view from the east 1962

Plate 51. Nicolas Cay: aerial view from the northeast 1962

Plate 52. Northeast Sapodilla Cay: northeast shingle ridge 1972

Plate 53. Northeast Sapodilla Cay: eroding northwest shore 1972

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Plate 55. Hunting Cay: aerial view from the north 1961

Plate 56. Hunting Cay: aerial view from the southeast 1962

Plate 57.

Plate 58.

Nicolas Cay: 'rootrock' exposed on the eroding west shore 1972

Hunting

Cay east bay from the north 1972

—

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Plate 63. Ragged Cay from the west 1972
Plate 64. Seal Cay: aerial view of the cay and faro from the east 1962
Plate 65. Seal Cay: east rampart and moat 1972

40

5. CAYS OF THE BARRIER REEF LAGOON

The barrier reef lagoon contains islands of many different types
and widely differing sizes. The great majority are mangrove islands,
and these have been very little studied. Others are of more complex
form and vegetation, but only a few of these have been studied in the
same detail as the barrier reef cays. The lagoon cays fall into
several distinct groups, and these are described from north to south.

A. The northern lagoon

North of latitude 17°30'N the coastal shelf has an average width
of 24 km, and between Belize and the Bulkhead Reef (which extends as a
shallow bar from the mainland to Ambergris Cay) is 2.5-4.5 m deep.
The Bulkhead itself carries only O.5-1.4 m, and encloses the 90 km
long Bahia de Chetumal, a shallow embayment with depths of 2-6 m. The
cays of the northern lagoon consist mainly of a series of linear
mMangrove-sand cays (mangrove islands with a sand ridge on their
windward sides) and mangrove cays located 3.5-4 km landward of the
barrier reef and separated from it by a depression termed the "reef
lagoon" by Ebanks (1975, p.238) and less than 4.5 m deep. In the
southern part of the area, large mangrove cays (Hicks Cays, Hen and
Chickens, Peter's Bluff, North Drowned Cay, the Drowned Cays) extend
across the shelf towards the mainland.

Ambergris Cay

This is the largest island of the Belize coastal shelf, extending
for 40 km southwards from the largely artificial cut Boca Bacalar Chico
which forms the Belize-Mexico boundary. It varies in width from 6.5
km to less than 90 m. Jefferys noted in his 1775 chart that there
were "plenty of deer in it and Ambergrease often found on its Beach".
Henderson (1811, p.24) described it as "of considerable size,
abounding with extensive fresh water lakes, and at most seasons ...
plentifully stocked with many kinds of game. This Key is likewise
said to produce Logwood, and the more valuable kind of Dye-wood, named
Brasiletto." Allen (1841, p.80) also stated that the cay was named
"from the produce of its shores". It was visited by Stoddart in 1960,
1962"and 1965 and briefly described “C1963, .pp.: 31-33)7tiga 6.5 1969",
DD 9-6) po and, by (Hew Pelzi inves 727 Sediments and geology have been
studied in detail by Ebanks (1975) and Tebbutt (1975). Stoddart's
vegetation map (1963, fig. 16) may be compared with Ebanks's geological

4l

map (1975, fig. 3): there is a broad correspondence between vegetated
supratidal flat of Ebanks and mangrove swamp of Stoddart; between
Pleistocene limestone and high and low woodland; and between coconut
woodland and windward beach ridge. Ebanks distinguishes four main
components to the island: .

(a) Windward beach ridge, a linear sand body along the east
side of the island, about 3 m thick above bedrock, reaching 1.2-2.4 m
above sea level (with a maximum of 6.6 m at San Pedro), and 90-460 m
wide. Sediment analyses of 19 samples showed that 90.8% of the
sediments on average are of skeletal origin (Halimeda 35.5%, molluscs,
18.0%, coralline algae 14.8%, corals 8.6%) (Ebanks 1975, p.247). Most
of our floristic information about Ambergris Cay comes from this unit.

(b) Supratidal flats, formed of fine sediments with dolomitic
crusts and extreme environmental conditions (mean salinity 76.3°/oo0,
range 41.5-123.8°/oo; maximumwater temperature 42°C). ism unas aS
vegetated by Rhizophora and Avicennia on the flats, with some shelly
windward sand ridges carrying a palmetto-salt grass-mangrove assemblage
(Ebanks 1975, p.253); these ridges reach O.3-1.3 m above sea level.

(c) Intra-island lagoons: oval-shaped ponds, often with arcuate
and smooth lee shores and irregular windward shores, the bigger ones
1.3-2 m deep, the shoaler areas less than 0.6 m deep, occupying about
one-third the total area of the cay (Ebanks 1975, p.265).

(d) Limestone surface. This is of uncertain age: north of Boca
Bacalar Chico Butterlin and Bonet (1960a, 1960b) map the limestones in
the Xcalak area as belonging to the Carrillo Puerto Formation of Upper
Miocene-Pliocene age; Sapper (1896) mapped those on Ambergris Cay as
Quaternary; and Flores 1952, p.409) by implication states that they

are Miocene to Pliocene. Tebbutt (1975, p.300) believes them to be
Pleistocene, and broadly correlative with the Key Largo Limestone of
Florida, of Last Interglacial Age. Reef facies in this limestone

outcrop at Reef Point on Ambergris Cay in a manner comparable to that
of the widespread "ironshore" of other Caribbean localities, but no
absolute dates are available. The existence of Last Interglacial reef
limestones further north along the Yucatan coast has already been noted
(Szabo et al. 1978). The deep well drilled by Gulf-Shell at Basil
Jones revealed 425 m of Tertiary rocks (post-Eocene) and 1790 m of
Lower Cretaceous. The vegetation of the Ambergris limestone area

is clearly an extension of that of neighbouring Quintana Roo. Wagner
(1964, p.223) simply and misleadingly maps this as Tropical Rain Forest,
but from casual observation it is more realistically classed in Beard's
(1955; 1949, p.82) categories of Dry Evergreen Woodland, Dry Evergreen
thvekee,, and bvergneen Bushland (cis Stoddart LI8O)," pp. 55-58) . This
is the view taken by Romney et al. (1959, p.28). No collections have
been made on the surface limestone on Ambergris Cay and the flora is
thus unknown.

The windward sand ridge is vegetated by a coconut woodland with a
mainly weedy ground vegetation, and a beach-crest vegetation of shrubs

—

42

and pioneer grasses and herbs. On the leeward side the ridge is
abutted by mangroves. At the small village of San Pedro there are
introduced trees, shrubs and herbs. It is worth noting that the
vegetation of Ambergris Cay has been influenced by man since pre-
European times (Stoddart 1980, pp. 160-161). It has several Maya
sites, two of which were located by Romney et al. (1959, fig. 10) and
Hammond (1977, p.48). Several were excavated by Gann (1926, p.59).
He found them to be "for the most part kitchen middens, composed of
enormous numbers of conch, cockle, whelk and oyster shells, turtle
carapaces, crab and crayfish shells, and vast quantities of such fish
as inhabit the surrounding waters. Amongst these were numerous
potsherds, all of the coarse, red and gray, domestic varieties, broken
flint spear-heads in great numbers, with a few broken obsidian knives,
and greenstone celts, many stone and pottery net sinkers, and a few
broken hand corn mills of Esquipulas stone. There were also found
the bones of a few small mammals, including the gibnut and armadillo,
but none of deer or wild hog, such as are usually found at most Maya
sites on the mainland." Later Gann mentioned "innumerable burial and
kitchen-midden mounds along the coast" (1927, p.95).

Only 22 species of plants have been recorded from Ambergris Cay
so far; undoubtedly a complete list would be very much larger. Those
recorded are:

Distichlis spicata West(1977) Terminalia catappa 62S 65S

Sporobolus sp. 62S 65S Ipomoea macrantha 65S
Cyperus sp. 62S Ipomoea sp. 65S

*Cocos nucifera 57S 60S 62S 65S Cordia sebestena 65S

Thrinax radiata 62S Tournefortia gnaphalodes 65S

Hymenocallis littoralis 65S Avicennia germinans 60S
Sesuvium portulacastrum 62S Solanum blodgettii 65C

Sophora tomentosa 65S Hamelia patens 72C
Suriana maritima 65S Ambrosia hispida 65S
Euphorbia sp. 65S Borrichia arborescens 65C
Rhizophora mangle 60S Flaveria linearis 65C

Bucida spinosa 72C

Mangrove cays in the lee of Ambergris Cay (Deer Cay, Swab Cay,
Blackadore Cay, Mosquito Cay) have only been observed from the air.
There are small mangrove islands at the southern tip of Ambergris Cay,
and also Cay Cangrejo (17°505'N, 88°03'W), a mangrove island.

Cay Caulker

Cay Caulker (the Cay Corker of Jefferys (1775) and Owen (1830)) is
a large mangrove-sand cay 8 km south of Ambergris Cay and 1.5-2.4 km

from the edge of the coastal shelf. It has been described briefly by
Vermeer’ (1959, ‘pp. 55=58), Stoddart (1963) "pp! 35-35, 81g.) 1s) and
Stoddart, (19699 p. 6). It is 7.2 km long and 60 m to 1.2 km wide.

Its windward side is formed by a sand ridge generally less than 1m

43

high, but rising at the village site to over 2 m. In places there

are clumps of mangroves on the seaward side of the ridge. On the
western side there are extensive mangroves, with in places a low
leeward sand ridge. Palm thicket occupies the interior areas not
cleared for coconut woodland. During Hurricane Hattie there was much
marginal erosion and surface sand-stripping and scouring on the seaward
side of the windward sand ridge. Some coconuts and other trees were
felled near the seaward shore, and exposed mangroves were defoliated.
But in general, away from the cleared area of the village, physiographic
effects were negligible and vegetation damage moderate. 34 species of
plants have been recorded (30 of them in 1965), and this is undoubtedly
@\ joeuetealeuil isis. They are:

Eragrostis prolifera 65C Hibiscus tiliaceus 65C
Eustachys petraea 65C Sida acuta ©5C
Cyperus ligularis 65C Turnera ulmifolia 65C
Fimbristylis cymosa 65C Rhizophora mangle 60S
*Cocos nucifera 57S 60S 61S Bucida spinosa 65C

62S 65S *Terminalia catappa 61S 65S
*Musa paradisiaca 57S *Catharanthus roseus 65C
*Casuarina equisetifolia 62S 65C Ipomoea sp. 62S
Coccoloba uvifera 62S 65C Cordia sebestena 65C
Philoxerus vermicularis 65C Tournefortia gnaphalodes 65C
Crotalaria verrucosa 65C Lantana involucrata 65C
Pithecellobium keyense 6©5C Stachytarpheta mutabilis 65C
Vigna luteola 65C Solanum blodgettii ©65C
Suriana maritima 65C Ernodea littoralis 65C
Euphorbia blodgettii 65C Spermacoce suaveolens 65C
Euphorbia mesembrianthemifolia Ageratum littorale 65C

65C Borrichia arborescens 6©5C
*Mangifera indica 57S 60S Wedelia trilobata 65C

Cay Chapel (17°41'N, 88°02'W)

Cay Chapel lies 1.6 km south of Cay Caulker and 2.5 km from the
shelf edge. It is 3.6 km long and up to 460 m wide, and built of
parallel sand ridges rising to 2 m above sea level. The leeward
mangrove is not well developed, and most of the sand ridges are
covered by coconut woodland: mechanical clearing has destroyed not
only the original vegetation but also the surface topography. This
clearing has taken place since 1775 when Jefferys noted a single "large
cocoa tree"; the greater part, however, took place in 1960-61. During
Hurricane Hattie it is said that some 60% of the coconuts were felled,
and there was much marginal erosion and surface sand stripping. For
idetarlsssee stoddare d963) uppewo>-so, fig. 17): By 1965 the windward
shore had been extensively colonised by Conocarpus, Borrichia, Suriana
and Tournefortia, with herbs and grasses beneath the coconut woodland.
For an account of recovery from Hurricane Hattie see Stoddart (1969,
D6 G7) Eighteen species of plants have been recorded, and many
more must be present. ThepAsesitea rs:

44

Cenchrus incertus 62S 65S Euphorbia sp. 62S 65S

Cyperus sp. 628 Rhizophora mangle 62S 65S

*Cocos nucifera 1775S 62S 65S Conocarpus erectus 65S

Thrinax radiata 62S Merremia dissecta 65C

Coccoloba uvifera 62S 65S Tournefortia gnaphalodes 62C 65S
Rivina humilis 65C Ernodea littoralis 65S

Cassytha filiformis 65S Ageratum sp. 65S

Cakile lanceolata 65S Conyza canadensis 65S

Suriana maritima 62S 65S Wedelia trilobata 65C

Craig (1906, p.21) describes a mound faced with shaped stone
blocks, with broken pottery and conch shells, on this island.

St. George's Cay (17°334'N, 88°04'W) Plates 66-68

St. George's Cay is an arcuate mangrove-sand cay standing 2.5 km
from the shelf edge. The western part is about 1.6 km long and 90-
180 m wide, and consists entirely of mangrove. At the northeast end
is a low sandy ridge 1.2 km long and 40-90 m wide, with some mangrove
on its lee side. The sandy area was cleared and settled at the end
of the eighteenth century and was entirely devoted to residential use
at the time of Hurricane Hattie. It was overtopped by the storm
surge during the hurricane and five channels were cut through the
ridge. Most coconuts were felled and mangroves defoliated, though
ground vegetation survived in patches. By 1965 two of the scoured
channels had been blocked, some of the mangrove was still living, and
there was a luxuriant growth of herbs and grasses. Detailed accounts
of the island before and immediately after Hurricane Hattie, and in
1965, are given by Stoddart (1963, pp. 37-40, figs. 18 and 19; 1969,

PPro pein aS) Romney et al. (1959, fig. 10) describe the cay as
a shell midden site. 30 plant species are recorded (16 during the
earlier surveys and 21 by Spellman in 1972). They are:
Eustachys petraea 72C Conocarpus erectus 65S 72C
Paspalum sp. 62S Eustoma exaltatum 72C
Sporobolus sp. 65S Echites umbellata 72C
Cyperus sp. 62S 65S Ipomoea pes-caprae 62S

*Cocos nucifera 60S 62S 65S 72S Ipomoea stolonifera 72C
Hymenocallis littoralis 62S 65S Ipomoea Sp. 65S

Coccoloba uvifera 62S 65S Cordia sebestena 72C
Batis maritima 65S 72C Tournefortia gnaphalodes 72C
Philoxerus vermicularis 72C Stachytarpheta jamaicensis 65S
Portulaca oleracea 72C 72C
Sesuvium portulacastrum 60S ErLeChalis trutetcosa 12
65S'772€ Ernodea littoralis 72C
Cakile lanceolata 62S Spermacoce verticillata 72C
Canavalia rosea 72C Ageratum sp. 62S 65S
Euphorbia mesembrianthemifolia Melanthera nivea 72C
H2Gi Synedrella nodiflora 72C
Euphorbia.sp. 62S 65S Wedelia trilobata 62S 65S 72C

Rhizophora mangle 62S 65S

45

Cays between Cay Chapel and Belize

There are numerous large mangrove islands on the coastal shelf
between 17°30'N and 17°45'N. They include Long Cay (the northernmost
of that name), Hicks Cays, Montego and Frenchman's Cays, Hen and
Chicken Cays, Rider's Cays, and the northernmost Drowned Cays. Apart
from a narrow seaward sand ridge with coconut woodland on Long Cay
and on the Drowned Cays north of Gallows Point, these islands consist

wholly of mangrove. Thus the Hicks Cays consist of a dozen islands,
four of them large, separated by deep meandering channels through
which tidal currents set with great rapidity. The Drowned Cays too

are intersected by a number of east-west tidal channels or 'bogues',
namely Shag Cay Bogue, Bannister Bogue, Farls Bogue and Goring Bogue,
most of which carry 5-7.5 m of water. For further details on these
islands, none of which have been studied in detail, see Vermeer (1959,
PPD o>) yand Stoddart (1963, pp. 36-37):

Moho Cay (17°315'N, 88°12'W)

Moho Cay is a small island immediately north of the Belize delta.
Gann (1925, p.19) mentions "innumerable flint spear-heads ... with
thousands of circular pottery rings, probably used as net sinkers."
He suggested that the island had been used as a Maya fishing station:
"Many tons of manatee bones have been washed out from the northern end
of the cay." Craig (1966, p.20) found habitation sites exposed by
wave erosion for 125 m along the shore, with "numerous pottery sherds,
abundant chert cores, flakes, and stone artifacts, including jadeite

celts, and large (25 cm) fine-wrought projectile points." In addition
to manatee bones, Craig also found oyster and conch shells, and two
human burial sites. The island was not visited during our own surveys.

Stake Bank (17°28'N, 88°074'w)

This is a mangrove island approximately 400 m long and 120 m wide,
standing on a shoal area 2.3 km long and 1.4 km wide. There is a
widespread tradition that the shoal was built with ballast deposited
by ships in the harbour (Anderson 1958, p.95), though this could only

account for a small part of the shoal volume. In the survey of
Belize harbour by Anthony de Mayne in 1828 the Bank had only one or
two mangrove seedlings. The bank was described as "dry in some parts"

by Speer in 1765, and was marked by Jefferys in 1775.

46

Spanish Lookout Cay (17°24'N, 88°04'wW)

This very small island at the southern end of the Drowned Cays

was visited briefly in 1965. It is largely artificial, built of
conch shells collected by fishermen, and surrounded by a coral block
wall. It has no natural vegetation.

Water Cay (17°23'N, 88°03%'W)

This large mangrove island is immediately south of the Drowned

Cays; it has a sand ridge along its windward side. J. Dwyer collected
Euphorbia mesembrianthemifolia, Lantana involucrata and Conocarpus
erectus there in 1967. It was not visited by us.

B. The Southern Triangles

Though this name no longer appears on charts it is convenient to
use it for the groups of islands at the inner end of the Belize
Deepwater Channel: it was used on the charts of Speer (1765, 1771) and
Jefferys (1775) and also in the Anglo-Spanish Treaties of 1753 and
KS Altogether there are some two dozen islands standing on the
flat tops of steep-sided shoals. On the north side of the channel
Robinson Point and Robinson Island both have dry-land vegetation in
addition to mangroves, but the other islands (Grennels Cay, Ramseys
Cay, One Man Cay) are entirely of mangrove. On the south side
Spanish Cay is a sand cay and the others (Long, Crayfish, Simmonds and
other unnamed cays) are mangrove.

Robinson Point Cay (17°22'N, 88°114'W)

This island, totalling about 29 ha, has the largest dry land area
in the Southern Triangles. It is described by Vermeer (1959, pp. 75-
719, Lig. 2l6)reand Stoddart m(1963;, spp. G46=4 7) Eg. 26) i There is a low
narrow sand ridge along the east and southwest sides of the cay; it is
nowhere more than 50 m wide and about 900 m long. Before Hurricane
Hattie it was covered with a dense thicket of coconuts, Thrinax and
other strand species, except at the small settlement near the light-
house, built at the western point in 1939. During the storm most of
the dry-land vegetation was destroyed and the mangroves defoliated, and
in 1962 the island had been abandoned by its inhabitants. The
following plant species were noted in 1961 before the storm:

Cyperus sp. 61S Hymenocallis littoralis ©1S_
*Cocos nucifera 57S 61S Coccoloba uvifera 61S
Thrinax radiata 61S Suriana maritima 61S

47

Euphorbia sp. 61S Cordia sebestena 61S
Rhizophora mangle 61S Avicennia germinans 618
Conocarpus erectus 615 Stachytarpheta jamaicensis 618

Robinson Island (17°224'N, 88°10%'W)

There is a less well-developed sand ridge along the south side of
Robinson Island. The palm thicket here was badly damaged during
Hurricane Hattie, and the mangroves defoliated (Stoddart 1963, p.47).
Only two species were noted in passing in 1961:

*Cocos nucifera 6185S Rhizophora mangle 61S
Spanish Cay (17°21'N, 88°09%;'W)

Spanish Cay when mapped in 1961 was a low-lying sandy island
100 m long and 10-18 m wide. The southern end of the cay was fringed
with Rhizophora, and there were several tall Avicennia trees. Ie
appeared that mangroves had previously been more extensive but had
been cleared. There were several coconuts and a ground cover of
sedges, grasses and ornamental plants. Much of the vegetation
disappeared during Hurricane Hattie; the island has not been revisited
since (see Stoddart 1963, p.47, fig. 27). Its total area in 1961 was
1260 sq m. The following plant species were recorded, but the list
is Certainly partial:

*Cocos nucifera 61S 62S Conocarpus erectus 61S 62S
Euphorbia sp. 61S Avicennia germinans 61S 62S
Rhizophora mangle 61S 62S

C. Cays of the Central Lagoon

For 60 km southwards from the Southern Triangles to the latitude
of South Water Cay the coastal shelf has an extremely simple topography.
The floor shelves gradually eastwards from the mainland coast to depths
of 20-24 m, and the shelf is edged by a barrier platform 5-8 km wide
and 4-6 m deep. There are no patch reefs or cays within the lagoon.
Near the inner edge of the barrier platform there is a series of linear
Mangrove islands, aligned north-south, and generally about 1.8 km from
the lagoon edge and 3.4 km from the seaward edge of the platform.

These islands include Middle Long Cay, Bluefield Range, Alligator Cay
and Colson Cays. Others, such as Southern Long Cay, Columbus Cay and
Cross Cay, are less regular islands closer to the seaward margin of the
platform. All are mangrove islands; none has been investigated in
deteanmike The linear islands occupy the same relative position as do
the mangrove-sand cays of the northern barrier reef (Cay Caulker, Cay

48

Chapel), but they lack the windward sand ridge and strand flora.

South of 17°N the mangrove islands near the inner edge of the barrier
platform become more complex. Instead of simple linear cays they are
complexes of smaller mangrove islands, often arranged in an ellipse,
and known as "ranges". These include Fly Range, Tobacco Range, South
Water Range and Blue Ground Range. Some of these have been visited
and will be described below. For further brief notes on this sector
of the reef, see Vermeer (1959, pp. 59-71 and 79) and Stoddart (1963,
PP eel 52))s

South of South Water Cay the barrier reef swings southeastwards
in a broad continuous arc to Gladden Spit and then turns sharply
southwestwards. There are no sand cays on the southeast-trending
sector, and only small patch reefs with small cays (already described)
to the south of it. At Gladden Spit the coastal shelf reaches its
maximum width of nearly 40 km, and the greatest lagoon depths reach
S5yme The barrier platform on the edge of the shelf becomes narrower
and less regular in this sector, and lacks mangrove islands. Within
the lagoon the topography is extremely intricate, with either long
arcuate ridges or elongate elliptical ring-like structures with deep
central depressions, the largest being up to 13 km long. These
ridges rise to within 5 m of the surface; they are interrupted by deep
gaps; and their upper surfaces are often pitted with deep holes; active
coral reefs reaching the surface are comparatively rare. The inference
from morphology that these structures resulted from karst erosion of
older reef limestones (Stoddart 1963, p.60) has been amply confirmed by
Purdy (1974a, 1974b), who terms them "Shelf atolls". This intricate
topography provides a wide range of substrates and exposure conditions
for island development: all types of island from true sand cay to
mangrove island are found.

The islands of this central lagoon area are described from north
to south, beginning with the Ranges of the northern area.

Tobacco Range (16°53'N, 88°05'W) Plates 69-70

Tobacco Range is an elliptical array of mangrove islands with a
shallow central lagoon, standing near the middle of the barrier
platform at Tobacco Entrance; it is roughly 4.8 km long north-south
and 1.8 km wide. The largest northeastern island was visited in 1972
at its northern and southern ends. At its northern end there is a
seaward sand ridge 0.6 m high and 18 m wide; a zone 60 m wide of low
boggy ground with coconuts and no other ground vegetation; and a zone
90 m wide of Batis maritima, open pools, and standing dead mangrove
trunks. These are 4-4.5 m tall and date from Hurricane Hattie's
effects in 1961. The mangrove zone in the lee is low and presumably
regenerated since that time. Beach shrubs include Tournefortia,
Sophora, Suriana, Borrichia and Laguncularia, with some Rhizophora on
the seaward shore. The ground cover includes Spartina, Ageratum,
Cyperus and Euphorbia, with carpets of Sesuvium. Pithecellobium takes

49

the place of Conocarpus, and Pluchea symphytifolia and Echites
umbellata are also unusual additions to the flora. Both north and
south from the point of survey the coconut zone narrows until it is
only 2 m wide, separating the Batis zone from coastal Sesuvium. At
the southern end of the island the seaward sand ridge is 20 m wide,
with Rhizophora and Avicennia clumps on its seaward side. Inland
there is a zone of mangroves 20-30 m wide and then open standing water.
The sand ridge itself supports a dense scrub of Thrinax, Conocarpus
and coconuts, with Cyperus, Batis, Spartina, Sesuvium and Borrichia.
Coconut boles 3-4 m offshore and outcropping lightly-cemented root
rock on the shore indicate beach retreat. At both. of the sites
visited the effects of Hurricane Hattie are still very apparent.

Before the storm tall mangrove presumably extended to the beach ridge. |
The death of a zone of mangrove up to 100 m wide, indicated by the
standing dead trees, has allowed colonisation by Batis maritima, and

because of the high level of the substrate, apparently rarely flooded,

there is no opportunity for recolonisation by mangroves. One might

expect colonisation in due course by sedges, grasses, and shrubs such

as Conocarpus and Pithecellobium.

Thirty species of plants were recorded in 1972:

Eragrostis prolifera 72C
Eustachys petraea 72C
Paspalum distichum 72C
Phragmites cf. australis 72C
Spartina spartinae 72C
Cyperus planifolius 72C
*Cocos nucifera 72S

Thrinax radiata 72C

Crinum amabile 72C

Batis maritima 72C
Salicornia perennis 72C
Sesuvium portulacastrum 72C
Pithecellobium keyense 72C
Sophora tomentosa 72C
Suriana maritima 72C
Euphorbia blodgettii 72C

Euphorbia mesembrianthemifolia
I2E
Passiflora suberosa 72C
Rhizophora mangle 72C
Conocarpus erectus 72C
Laguncularia racemosa 72C
Bumelia retusa 72C
Echites umbellata 72C
Tournefortia gnaphalodes 72C
Avicennia germinans 72C
Erithalis fruticosa 72C
Ernodea littoralis 72C
Ageratum littorale 72C
Borrichia arborescens 72C
Pluchea symphytifolia 72C

Coco Plum Cay (16°53'N, 88°07'W)

Coco Plum Cay is a linear mangrove island near the inner edge of
the barrier platform, immediately west of Tobacco Range. Like Tobacco
Range it has a low beach-ridge area with similar vegetation. Seventeen
plant species are recorded:

Paspalum distichum 72C
Spartina spartinae 72C
Sporobolus virginicus 72C
Cyperus planifolius 72C

Fimbristylis spadicea 72C
Thrinax radiata 72C

Batis maritima 72C
Suriana maritima 72C

50

Euphorbia mesembrianthemifolia Tournefortia gnaphalodes 72C

72C Avicennia germinans 72C
Rhizophora mangle 72C EE Chalss 2oucicosa 126
Conocarpus erectus 72C Pluchea symphytifolia 72C
Laguncularia racemosa 72C Wedelia trilobata 72C

Phillips Petroleum drilled a hole 91.5 m deep through Coco Plum Cay;
the sub-Holocene unconformity was reached at 9.2 m depth (Purdy 1974b,
Pps O41, . ODS)ic

Man-o'-War Cay (16°52%'N, 88°07'W)

This island, one of several of the same name on this coast (e.g.
at 16°39'N, 88°12'W) lies at the southern end of Coco Plum Cay. It
is a small mangrove island, with spartina and succulents. Six plant
species are recorded:

Spartina spartinae 72C Rhizophora mangle 72C
Batis maritima 72C Laguncularia racemosa 72C
Sesuvium portulacastrum 72C Avicennia germinans 72C

Water Range (16°50'N, 88°06'W)

This is the group of mangrove islands (the Twin Cays of charts
and Figure 1) 2.2 km northwest of South Water Cay. There is a small
sand patch measuring 100 x 18 m at the south end on the seaward side,
and another of shell sand on the lee side. Both support a scrub of
Thrinax, Conocarpus, Laguncularia, Suriana, Erithalis and Borrichia,
with grasses, sedges and succulents. On their inner sides they pass
into Rhizophora swamp through Laguncularia and Avicennia. Twenty
plant species are recorded:

Eustachys petraea 72C Euphorbia mesembrianthemifolia
Paspalum distichum 72C 72C

Sporobolus virginicus 72C Rhizophora mangle 72C

Cladium jamaicense 72C Conocarpus erectus 72C

Cyperus planifolius 72C Laguncularia racemosa 72C
*Cocos nucifera 72S (juvenile) Avicennia germinans 72C
Thrinax radiata 72C Erithalis fruticosa 72C
*Casuarina equisetifolia 72C Ageratum littorale 72C

Batis maritima 72C Borrichia arborescens 72C
Salicornia perennis 72C Wedelia trilobata 72C

Suriana maritima 72C

5

Weewee Cay (16°46'N, 88°083'W)

Weewee Cay stands on an isolated patch reef 6.5 km southwest of
South Water Cay. It is a small triangular island with sides 90-140 m
long, surrounded by dense Rhizophora, with a low-lying central sandy
area about 70 m in diameter. The dry-land vegetation is limited to
coconuts with an undercover of grasses and sedges. It is used asa
fishing station. Hurricane Hattie in 1961 killed some Rhizophora at
the south end and felled many coconuts; the island has not been visited

since 1962. For a brief account of it at that time see Stoddart
GUS6S pao) Six species of plants are recorded:
Eustachys petraea 62C *Cocos nucifera 62S
Cyperus peruvianus 62C Batis maritima 62C
Fimbristylis cymosa 62C Rhizophora mangle 62S

Cat Cay (16°40'N, 88°12'w)

This is one of the small constituent islands of the Pelican Cays,
the northernmost shelf atoll of the central lagoon. It was visited in
IIo2 me (Stoddart 1963) si p102))i It has a narrow fringe of Rhizophora on
its north and east shores and much of the west shore. The centre of
the island is low-lying and sandy, with a woodland of coconuts,

Thrinax and Thespesia. Ospreys were nesting here in 1962. Eight
plant species are recorded:

Cyperus sp. 6258S Thespesia populnea 62C
*Cocos nucifera 62S Conocarpus erectus 62S
Thrinax radiata 62S Avicennia germinans 62S
Euphorbia sp. 62S Wedelia trilobata 62S

Lagoon Cays between Stewart Cay (16°46'N, 88°10'W) and Baker's
Rendezvous (16°34'N, 88°12'W)

These are briefly noted by Stoddart (1963, pp. 61-62). They are
mainly mangrove islands and have not been studied in detail. Peter
Douglas Cay (16°43'N, 88°10'W) and the adjacent Little Peter Cay both
have an area of palm thicket with Sophora tomentosa. There are
coconuts on Peter Douglas Cay and Baker's Rendezvous.

Jack's Cay (16°33'N, 88°0O4'w)

Jack's Cay is a mangrove island on an isolated reef patch in the
lee of the Gladden Spit reef, 5.5 km from the reef edge. Age, aL}
dominated by mature Rhizophora trees. Seven plant species are
recorded:

SZ

Sporobolus virginicus 72C Rhizophoramangle 72C
Brassavola nodosa 72C Laguncularia racemosa 72C
Batis maritima 72C Avicennia germinans 72C

Sesuvium portulacastrum 72C
Buttonwood Cay (16°325'N, 88°03'W) Figure 31

Buttonwood Cay lies 1 km southeast of Jack's Cay. It was mapped
in 1L96lAa(Stoddart 19638)! p..64i,; firg 71389)" anditagain sone 9772): Tees
surrounded on its eastern side by Rhizophora, with a low-lying central
sand area measuring 95 x 75 m. The sand area is covered with a coconut
woodland with a ground cover of grasses and herbs; there is a discon-
tinuous fringe of Conocarpus, Laguncularia and Coccoloba between the
coconut woodland and the Rhizophora. The island was mapped in 1961
before both Hurricane Anna and Hurricane Hattie: the northeastern
mangrove area has since decreased in area, and the Sand area has
accreted to leeward. The island is a fishing station with accumula-
tions of conch shells along the shores. Its area in 1972 was 0.72 ha.
26 species of plants have been, recorded:

Andropogon glomeratus 72C Euphorbia mesembrianthemifolia 72C
Eragrostis prolifera 61C 72C Euphorbia sp. 61S

Eustachys petraea 61C 72C Hibiscus tiliaceus 72C
Spartina spartinae 72C Rhizophora mangle 61S 72C
Cyperus ligularis 61C 72C Conocarpus erectus 61S 72C
Cyperus planifolius 72C Laguncularia racemosa 72C
Fimbristylis cymosa 72C Avicennia germinans 61S 72C
*Cocos nucifera 61S 72S Lantana involucrata 72C
Coccoloba uvifera 615 72C Hedyotis corymbosa 72C
Batis maritima 72C Eclipta alba 72C

Protulaca oleracea 72C Melanthera nivea 72C
Sesuvium portulacastrum 72C Pluchea symphytifolia 72C
Vigna luteola 61C 72C Wedelia trilobata 72C

Euphorbia blodgettii ©1C 72C

Trapp's Cay (16°305'N, 88°10'W)

Trapp's Cay (the Moho Cay of charts) stands on an isolated patch
reef in the deepest part of the lagoon. It was mapped after Hurricane
Hattie in 1962 (Stoddart °1963/) pp.s63=64, fig). 38): It is a sandy
island with maximum dimensions of 240 x 190 m, and an area of 4.54 ha.
The north and east sides of the cay were blanketed with fresh shingle
up to 0.6 m thick and 9-23 m wide during Hurricane Hattie. The island
is covered with a woodland of coconuts and Thrinax, with Coccoloba and
Thespesia. The vegetation was not studied in detail and only 10
species are recorded:

53

Cyperus sp. 62S Sesuvium portulacastrum 62S
*Cocos nucifera 62S Phyllanthus amarus 62C
Thrinax radiata 62S Thespesia populnea 62S
Hymenocallis littoralis 62S Rhizophora mangle 62S
Coccoloba uvifera: 62S Conocarpus erectus 62S

Cary Cay (16°314'N, 88°12'W) Plate 71

Cary Cay stands on one of the longest reef ridges in the central
lagoon: it is 460 m long and has a total area of 4.29 ha. In the
north there is an extensive mangrove swamp up to 180 m wide; in the
south there is a narrow tapering sand and shingle ridge. In the centre
of the island there is an Acrostichum marsh with standing water. The
dry sand area has a palm thicket with Cocos, Thrinax, Coccoloba and
Cordia. Much fresh shingle was deposited along the east and southeast
sides of the island during Hurricane Hattie, but vegetation damage was
minor. For details see Stoddart (1963, pp. 62-63, fig. 37); the

island has not been visited since. Only 11 plant species are recorded:
Acrostichum aureum 62S Cordia sebestena 62S
*Cocos nucifera 62S Avicennia germinans 62S
Thrinax radiata 62S Stachytarpheta jamaicensis 62S
Hymenocallis littoralis 62S Ageratum sp. 62S
Coccoloba uvifera 62S Wedelia trilobata 62S

Rhizophora mangle 62S

Bugle Cay (16°294'N, 88°19'W)

Bugle Cay is a mangrove island only 5.5 km from the mainland coast
at Placentia Point. The western end of the island is an arcuate strip
of low sand 150 m long and 15-45 m wide. The island is a lighthouse
station, and the sand area is planted to coconuts. The island was much
damaged by marginal erosion and surface sand-stripping during Hurricane
Hattie, and many coconuts were destroyed. In 1965 new coconuts had
been planted, and the ground surface was being colonised by herbs and
grasses. The sand sector had an area of O.5 ha in 1962. Only seven
plant species have been recorded:

Cyperus sp. 65S Euphorbia sp. 65S
*Cocos nucifera 62S 65S Rhizophora mangle 62S 65S
Batis maritima ©5S Avicennia germinans 62S

Sesuvium portulacastrum 65S

For further details, see Stoddart (1963, pp. 67-68, fig. 43; 1969, p.1l).

54

Owen Cay (16°295'N, 88°15}'W)

Owen Cay, ©.5 km east of Bugle Cay, is a sand and shingle island
165 m long and up to 35 m wide, with an area of 0.49 ha. In the
centre of the island there are clumps of Rhizophora and Avicennia, with
a ground cover of Batis. The sand areas are covered with coconuts,
Thrinax, Cordia and Hymenocallis. Fresh shingle was deposited round
the whole margin of the cay in 1961, forming ridges up to 1.5 m high.
For details see Stoddart, 1963, pp. 65-66, fig. 41). Only seven plant
species are recorded:

*Cocos nucifera 62S Rhizophora mangle 62S
Thrinax radiata 62S Cordia sebestena 62S
Hymenocallis littoralis 62S Avicennia germinans 62S

Batis maritima 62S

Scipio Cay (16°28%'N, 88°18'W) Plate 72
Scipio Cay resembles Owen Cay 4 km to the eastnortheast. ibe, als
250 m long and up to 90 m wide, with an area of 1.67 ha. Abe als
covered by coconut woodland, with a central Avicennia swamp, and with
stands of Thrinax on the beach ridges. During Hurricane: Hattie fresh

shingle was deposited round almost the entire periphery of the cay.
By 1965 these ridges had moved inland and been colonised by Sesuvium,
Sporobolus and Euphorbia. For details see Stoddart (1963, pp. 66-67,

panto wy SCE USCIS) fojoja du IDA aesitos 5" 62) Only 11 plant species have been
recorded:
Sporobolus virginicus 65S Rhizophora mangle 62S 65S
*Cocus nucifera 62S 65S Ipomoea sp. 62S
Thrinax radiata 62S 65S Cordia sebestena 62S
Sesuvium portulacastrum 65S Tournefortia gnaphalodes 65S
Sophora tomentosa 65S Avicennia germinans 62S 65S

Euphorbia sp. 65S

Collison. Cay. (l6228N 7.882 19UW) Plate 73

Colson Cay closely resembles Scipio Cay, only 1.6 km away. Abe, abs
triangular, with sides of 140-175 m and an area of 2.03 ha. It is low
and sandy, with a dense vegetation of coconuts and Thrinax, and with a
central Avicennia swamp. Fresh shingle was deposited all round the cay
during Hurricane Hattie in 1961. These ridges had eroded and moved
landward by 1965, and had been colonised by Sesuvium, Euphorbia,
Sporobolus, Cassytha and two small Tournefortia bushes. For details
see Stoddart (1963) p.G7, -£1¢., 427.1969 sp alZ,, £1Ge el) 11 plant

species have been recorded:

55

Sporobolus sp. 65S

*Cocos nucifera 62C 65C
Thrinax radiata 62S 65S
Sesuvium portulacastrum 65S
Cassytha filiformis 65S
Sophora tomentosa 65S

Euphorbia sp. 65S

Rhizophora mangle 62S 65S
Cordia sebestena 62S 65S
Tournefortia gnaphalodes 65S
Avicennia germinans 62S 65S

Purdy (1974b, pp. 841, 853) drilled a 19.2 m hole through Colson
Cay. The entire section was in Holocene material. A date of
55504140 yr from a sample at 6.4-7.3 m indicated a Holocene reef growth
rate of 1.2 m/1000 yr.

Hatchet Cay (16°28'N, 88°04};'W) Figure 32

Hatchet Cay lies 3.3 km lagoonward of the Silk Cays, close to the
edge of the coastal shelf south of Gladden Spit. It is a low-lying
sandy island 240 m long and 45-85 m wide, with an area when mapped in
IDN72 O32 ILG SO) levers It is almost completely surrounded by a zone of
Rhizophora woodland from 5 to 40 m wide. The interior of the cay is
occupied by coconut woodland. It thus closely resembles Weewee,
Buttonwood and Trapp's Cays. Hatchet is inhabited, and has two wells.
38 species of plants have been recorded, and this more accurately
reflects the floras of this type of island than the collections made on
the other three islands mentioned. The species are:

*Cymbopogon citratus 72C *Citrus aurantiifolia 72C

Eragrostis ciliaris 72C
Eragrostis prolifera 72C
Eustachys petraea 72C
Paspalum laxum 72C

Spartina spartinae 72C
Sporobolus jacquemontii 72C
Cyperus peruvianus 72C

Euphorbia blodgettii 72C

Euphorbia mesembrianthemifolia
72C

Hibiscus tiliaceus 72C

Rhizophora mangle 72C

Conocarpus erectus 72C

Laguncularia racemosa 72C

Cyperus planifolius 72C
Fimbristylis cymosa 72C
Fimbristylis spadicea 72C

*Terminalia catappa 72C
*Psidium guajava 72C
*Nerium oleander 72C

*Cocos nucifera 72S

Thrinax radiata 72C
Hymenocallis littoralis 72C
Batis maritima 72C
Philoxerus vermicularis 72C
Portulaca oleracea 72C
Sesuvium portulacastrum 72C
Cakile lanceolata 72C

Vigna luteola 72C

Ipomoea pes-caprae 72C
Avicennia germinans 72C
Lantana involucrata 72C
Lippia strigulosa 72C
Ageratum littorale 72C
Melanthera nivea 72C

*Vernonia cinerea 72C

Wedelia trilobata 72C

A discontinuous ridge of rubble extending along the reef crest to

the southwest of the cay, about 35 m offshore, may date from Hurricane
Hattie in 1961.

56

Little Water Cay (16°274'N, 88°06'W) Figure 33, Plate 75

Little Water Cay lies 3.7 km southwest of Hatchet Cay. Lt: isa
triangular island 255 m long and up to 170 m wide, with an area in
INT CE DEO lovers Most of the island is covered with a dense coconut
woodland, with an interior area of sedge marsh and narrow fringes of
Rhizophora at the west end and along the east shore. There are some
patches of Conocarpus along the shore, especially on the north side,
but generally the coconut woodland extends to the beach. The whole
of the south shore is cliffed and eroding, with fallen coconuts.

There is an extensive zone of rubble on the reef crest at the east end

uninhabited.

of the cay, which may date from Hurricane Hattie.
29 plant species are recorded:

Eragrostis prolifera 72C
Eustachys petraea 72C
Paspalum blodgettii 72C
Paspalum distichum 72C
Spartina spartinae 72C
Sporobolus virginicus 72C
Stenotaphrum secundatum 72C
Cyperus ligularis 72C
Cyperus planifolius 72C
Fimbristylis cymosa 72C
*Cocos nucifera 72S

Batis maritima 72C
Philoxerus vermicularis 72C
Portulaca oleracea 72C
Sesuvium portulacastrum 72C

The island is

Vigna luteola 72C
Suriana maritima 72C
Euphorbia blodgettii 72C
Euphorbia mesembrianthemifolia
c2C
Hibiscus tiliaceus 72C
Rhizophora mangle 72C
Conocarpus erectus 72C
Laguncularia racemosa 72C
*Terminalia catappa 72C
Avicennia germinans 72C
Lippia strigulosa 72C
Eclipta alba 72C
Melanthera nivea 72C
Wedelia trilobata 72C

Laughing Bird Cay (16°27'N, 88°12'W)

Laughing Bird Cay is the southernmost of the islands in the central
lagoon. It stands on an elongate reef ridge 23 km from the coast and
14 km from the shelf edge: water depths to the southeast reach 45 mn,
but Purdy has shown that the Holocene sequence beneath this island is
18.3 m thick, giving a pre-Holocene relief above the present lagoon
floor of about.20 m (Purdy. 1974b, p.841). The island itself is an
elongate sand ridge about 460 m long and 9-40 m wide. During Hurricane
Hattie rubble and shingle were deposited at the northeast and southwest
ends of the cay, and along the southeast shore. When mapped in 1962
the area of the cay was 1.36 ha; it has not been visited since
(Stoddart 1963, .365,,2fig.. 40); The island is covered with coconut
woodland, with scattered coastal mangroves. Only seven plant species
have been recorded:

*Cocos nucifera 62S
Hymenocallis littoralis 62S
Sesuvium portulacastrum 62S
Euphorbia sp. 62S

Rhizophora mangle. 62S
Ipomoea sp. 628
Avicennia germinans 62S

3) 1/

Placentia Cay (16°31'N, 88°21'W)

This is a mangrove island with a small area of strand vegetation
and coconuts located 1 km east of Placentia Point on the mainland
coast. It was mapped as True Point Patience Kay of Speer (1966, p.23)
and as Patience Brother Island of Jefferys (1775). Jefferys also
mapped the small mangrove islets at the mouth of Placentia Lagoon as
The Virgins, and Speer commented on these "Several small Kays ... at
the mouth of a large lagoon, where there is plenty of Turtle passes
morning and evening" (1766, p.23). No plant collections have been
made on these islands, which are part of the coastal quartz and
carbonate sand accretional sequences.

Harvest Cay (16°29'N, 88°24'W)

Harvest Cay, 5.5 km to the southwest, is the "Hobbe's Kay" of
Speer (1766, p.23). This and other nearshore islands to the south,
such as Palmetto Cay, are properly barrier beaches built of terrigenous
quartz sand, backed by mangrove swamp, rather than true coral cays.
Harvest Cay has a dense vegetation of coconuts, Thrinax and mangroves,
with a tall undergrowth of Cladium, together with succulents and grasses.

The island was briefly visited in 1962. Four species were noted:
Cladium jamaicense 62C Thrinax radiata 62S
*Cocos nucifera 62S Batis maritima 62S

D. The Punta Gorda Cays

The cays of the Port Honduras Bight between Punta Gorda and Punta
Ycacos, in the southern part of the barrier reef lagoon, stand on
numerous steep-sided knolls, rising from a shelving lagoon floor in
from 8-24 m of water. The innermost cays, from Stuart Cay at the
mouth of the Rio Grande to Bedford Cays at the mouth of the Ycacos
Lagoon, all surrounded by water less than 10 m deep, are wholly mangrove.
Further lagoonward there are some islands with dry land in addition to
Mangrove, notably Frenchman's Cay and Wild Cane Cay, but most of the
islands are un-named and wholly of mangrove. On many of them the
mangrove in 1961 was less than 2 m tall, suggesting recent growth
following hurricane damage. The Moho Cays are all mangrove, except
for the outer one, which is a sand cay. The outermost islands, the
Snake Keys, stand on patches rising from a lagoon floor 20-30 m deep;
they are sand cays with shingle ridges; only these patches have signifi-
cant amounts of coral on them. All of these islands come under the
influence of freshwater discharge from the Deep River and the Rio Grande;
the rainfall in this area (4239 mm at Punta Gorda) is much higher than
elsewhere on the coastal cays. The published chart of the Gulf of
Honduras dates from surveys in 1835-1841 and is rather generalised, and

58

the islands are known only through a brief reconnaissance visit in

TMS vibes They would repay further investigation.
East Snake Cay (16°12%'N, 88°30%'W) Figure 34, Plates 76 and 79

East Snake Cay consists of a densely vegetated sand cay wholly
surrounded by shingle ridges which enclose a shallow moat containing
mangroves. The total area is 1.94 ha, of which the main core,
excluding the moat and rampart, comprises 1.04 ha. The whole complex
is 260 m long and up to 110 m wide, and the reef patch on which it
stands is only slightly larger. Vegetation on the shingle rampart is
limited to patches of Ipomoea and Rhizophora. On the sand cay there
is a dense woodland of coconuts, Thrinax, Cordia, Coccoloba, Terminalia
and Ficus. There is an automatic lighthouse on the island. Nepean e)
brief account, see Stoddart (1965, p.138, fig. 4). 20 species of
plants have been recorded:

Cyperus sp. 61S Rhizophora mangle 61S

*Cocos nucifera 57S 61S Conocarpus erectus 61S

Thrinax radiata 61S *Terminalia catappa 61C
Hymenocallis littoralis 61S Ipomoea sp. 61S

Brassavola nodosa 61C Cordia sebestena 61S

EIcus#Sp.. 61E Tournefortia gnaphalodes 61S
Coccoloba uvifera 61S Avicennia germinans 61S
Erythrina sp. 61C Stachytarpheta jamaicensis 61S
Sophora tomentosa 61C Erithalis fruticosa 61S
Euphorbia sp. 61S Wedelia trilobata 61S

Purdy (1974b. p.841) drilled a hole 39.7 m deep through East Snake Cay.
Recent reef material gave way to pre-Recent terrestrial sediments at a
depth of 25.9 m.

Middle Snake Cay GiGZI2U NEN SS238uw) Figure 35

Middle or Ragged Snake Cay comprises two islands built of shingle,
each having an area of O.66 ha. The northeastern island is about
275 m long but is everywhere less than 40 m wide. It is built of two
linear shingle ridges with an intervening partly water-filled depression.
The outer ridges, presumably hurricane-formed, include coral blocks 1 m
in diameter. The vegetation is limited to succulents and grasses, with
low shrubs of Coccoloba, Suriana, Tournefortia and mangroves. 7 plant
species were noted in 1961:

Coccoloba uvifera 61S Conocarpus erectus 61S
Sesuvium portulacastrum 61S Ipomoea sp. 61S
Suriana maritima 61S Avicennia germinans 61S

Rhizophora mangle 61S

5)

The southern island is also built of shingle ridges, but is
triangular, with sides of 75-110 m- The interior is more sandy, and
the concentric ridges enclose linear pools. There is a dense growth
of Avicennia and Rhizophora along the north side, with a woodland of
coconuts, Thrinax and Coccoloba on the rest of the cay. 15 species of
plants were noted in 1961:

Cyperus sp. 618 Sophora tomentosa 618
*Cocos nucifera 61S Suriana maritima 61S
Thrinax radiata 618 Euphorbia sp. 618
Hymenocallis littoralis 61S Conocarpus erectus 61S
Brassavola nodosa 618 Ipomoea sp. 61S

Coccoloba uvifera 615 Tournefortia gnaphalodes 61S
Sesuvium portulacastrum 61S Wedelia trilobata 61S

Cakile lanceolata 61S
West Snake Cay (16°11)'N, 88°34'w) Figures 36 and 37, Plates 77 and 80

West Snake Cay is a complex island with a leeward sand area, a
central mangrove swamp, a moat, and a windward shingle ridge, with a
totale area Of 5.35 ha. It has been described briefly by Stoddart
(1965, p.137, fig. 4), and recalls the features of low wooded islands
of the northern Great Barrier Reef and of Djakarta Bay. The shingle
rampart follows the eastern edge of the reef patch for about 370 m,
rising to 1 m above sea level. The moat enclosed by the shingle ridge
is about 200 m long and up to 100 m wide: it is less than O.6 m deep;
and with its enclosing shingle ridges has an area of 2.5 ha. There
are many Rhizophora seedlings in the moat and along the inner margin of
the ridge, and the shingle ridge itself is patchily covered with
Sesuvium, Tournefortia, Coccoloba, mangroves and grasses. The most
extensive vegetation unit is the mangrove swamp, consisting of
Rhizophora, Avicennia and Acrostichum, but much of it is open standing
water. The leeward dry sand area is 15-30 m wide and rises to 1 m
above sea level. It is under coconut woodland with Coccoloba and
Tournefortia and a diverse ground cover. 13 species of plants were
recorded in 1961:

Acrostichum aureum 61C Euphorbia sp. 61S

Sporobolus virginicus 61C Rhizophora mangle 61S
Cyperus sp. 61S Ipomoea macrantha 61C

*Cocos nucifera 61S Ipomoea pes-caprae 61C
Coccoloba uvifera 618 Tournefortia gnaphalodes 61S
Sesuvium portulacastrum 615 Avicennia germinans 61S

Cakile lanceolata 61C

150 m northwest of the main island is a small shingle island,
West Island. It is 140 m long, up to 23 m wide, has an area of 800
sq m, and rises 1 m above sea level. Most of the islet is covered
with Sesuvium and Ipomoea, with some shrubby Coccoloba, Suriana and
Conocarpus. Six species were recorded in 1961:

60
Sporobolus sp. 61S Suriana maritima 61S
Coccoloba uvifera 61S Conocarpus erectus 61S
Sesuvium portulacastrum 61S Ipomoea sp. 618

South Snake Cay (16°11'N, 88°34'W) Figure 38, Plate 78

South Snake Cay is very similar to West Snake Cay, but the
eastern shingle rampart is uninterrupted and adjoins the main cay at
each end. The mangrove area is smaller and the leeward sand area
forms the largest part of the complex (Stoddart 1965, pp. 137-138,
sBaltopa, <4)))"e The shingle rampart is 20 m wide and 1.5 m high at its
southeast end; it becomes narrower and lower northwards. The moat
is shallow and completely enclosed, and contains much stranded pumice;
the area of the moat and enclosing rampart is 1.05 ha. The rampart
itself is not vegetated and there are no Rhizophora seedlings in the
moat. The dry sand area has maximum dimensions of 180 x 90 m, has an
area of 1.7 ha, and is strongly undercut along its southwest shore,
where the surface stands at 1 m above sea level. Poorly cemented
beachrock outcrops along this shoreline between the small bluffs formed
by the boles of coconut trees. The sand area is covered with a dense
thicket of coconuts and Thrinax. ll species of plants were noted in
LOGUE:

Andropogon sp. 61S Sophora tomentosa 61S
Cyperus sp. 61S Suriana maritima 61S
*Cocos nucifera 61S Rhizophora mangle 61S
Thrinax radiata 61S Conocarpus erectus 61S
Hymenocallis littoralis 61S Wedelia trilobata 61S

Dalbergia ecastophyllum 61C

Wild Cane Cay (16°124'N, 88°38'W) Figure 39

This is a mangrove island 7.5 km southwest of Punta Ycacos. The
whole island is about 1 km long. At its southern end there is a
rectangular dry area measuring 300 x 100 m, with an area of 2.0 ha.
Mangroves are growing at several points along its shores. The dry
area is only about 0.6 m above sea level; the soil is dark and humic
and parts are boggy. The vegetation consists of coconut woodland with
marginal Conocarpus. The island is inhabited and there are several
introduced food and ornamental plants. The most striking feature of
the cay, however, is the presence of large stone-faced Maya burial
mounds. One of these constituted Mound 39 in Gann's excavations
CLOUGZ pe US5=—S56)s. This was a mound 3 m high consisting of sand
and blocks of reef limestone covering a surface burial. Gann found
pottery, obsidian knives, the femur of a deer with an incised design,
and copper ornaments (rings, gorgets, studs) which he suggested showed
Spanish influence. In another account (1927, p.238) Gann speaks of
"great numbers of interments in four large mounds."

61

When mapped in 1961 the four mounds still existed, the largest
being 65 m long. They have, however, been much disturbed. iba AUS}
Gann noted that they were used as a source of stone, and in 1927 (p.238)
he stated that "when the Cabral family needs a little ready money, they
excavate a small area and sell the beads, pottery, spear-heads, etc.,
which invariably turn up." In 1961 the ground surface was littered
with obsidian flakes up to 4 cm long, and with fragments of coarse red
pottery. A drilled jade bead and a projectile point 16 cm long were

also found. Other remains included grey pottery net sinkers and small
animal figures. The most remarkable object from Wild Cane Cay,
however, is an animal-shaped jar 13.7 cm high of Plumbate Ware. aTKte

is described by Bray (1970, p.32, plate 27): "The head is frog-like,
but the feet and stumpy tail belong to a different kind of animal.

The creature wears a necklace, each end of which terminates in a disc
ornamented with six pits. The body of the pot is further decorated
with broad line designs incised through the greenish-brown slip."
Plumbate Ware was manufactured in Guatemala and widely traded: the use
of zoomorphic shapes began about 1000 A.D. and this serves as a time-
marker for the early Postclassic period. Bray dates this particular
object between 1000 and 1200 A.D.

Fifteen plant species were noted in 1961; undoubtedly more are
present.

Pityrogramma calomelanos 61C *Mangifera indica 61S

Sporobolus jacquemontii 61C Sida acuta 61C

Fimbristylis spadicea 61C Opuntia sp. 61S

*Cocos nucifera 61S Rhizophora mangle 61S

Laelia tibicinis 61C Conocarpus erectus 61S
Coccoloba uvifera 61S *Psidium guajava 61C

TCLERUST SPEEOlltsS Stachytarpheta jamaicensis 61C

*Pedilanthus tithymaloides 61C

Frenchman's Cay (16°115'N, 88°38'W)

Frenchman's Cay is very similar to Wild Cane Cay, though almost
the whole shoreline is lined by mangroves. The sandy interior is
covered with coconut woodland, and the centre is low-lying and swampy.
There is a freshwater well, several huts, and an area of cultivated
plants. There are several burial mounds faced with limestone blocks,
seen in 1961. Gann (1927) reported the excavation of gold ornaments
and other artefacts from these sites. Only five species of plants
were noted in 1961; many must remain to be recorded. They are:

*Saccharum officinarum 61S *Mangifera indica 61S
*Cocos nucifera 61S Rhizophora mangle 61S
*Musa paradisiaca 61S

62

Moho Cay (16°305'N, 88°10'W) Figure 40
Apart from coastal mangrove islets, Moho Cay is the southernmost

cay in the Punta Gorda group, and the only sand cay. It is a narrow

island 160 m long, with an average width of 18-35 m, and an area of

0.54 ha. Owen in 1835 noted a solitary coconut at its northern end,

but since then the vegetation has been cleared and the island is now
covered with coconut woodland, except for an area of dense woodland at
the south end. There is a varied ground cover of herbs, grasses and
sedges beneath the coconuts. The dense woodland includes coconuts,
Thrinax and some mangroves. The whole island is sandy and low-lying,
and the south and southeast shores are undercut and eroding. There is
a freshwater well and a small hut. Twenty species of plants were
recorded in 1961:

Eragrostis prolifera 61C Thrinax radiata 618
Eustachys petraea 61C Hymenocallis littoralis 61S
Paspalum paniculatum 61C Coccoloba uvifera 61S
Sporobolus virginicus 61C Sophora tomentosa 61C
Stenotaphrum secundatum 61C Vigna luteola 61C

Cyperus ligularis 61C Euphorbia blodgettii 61C
Cyperus peruvianus 61C Bidens cynapiifolia 61C
Fimbristylis cymosa 61C Conyza canadensis 61C
Fimbristylis spadicea 61C Melanthera nivea 61C

*Cocos nucifera 61S Wedelia trilobata 61C

[TIL [] Coconuts (pen)

Se
ee
SOOO

OOP 2

Qk
Q

CSS
SBS

222k
Ses

Soe

sn a

ax SS

SST

Dense GOCOmuUts
with herb mat
and grasses

Coccoloba
Rhizophora
Avicennia
jLaguncularia
Hibiscus
s1 CONOCarpuSs

050505 0s ee hat
o50ee
C5505
OR55255

revere
>

Portulaca,Vigna,

Euphorbia

Vv

Melanthera

Wedelia
TTT (under coconuts)

mun LJndercut shoreline

e***eConch shells

[-. |Fresh sand

Buttonwood Cay 1972

Figure 31.

Dense coconuts

TTA with grasses

and herbs

Rhizophora

Q

Avicennia

Terminalia

T

Hibiscus

Musa

¢

Thrinax

conch-shell

Y)
>
Q.
C
o
V
O
Cc
O
O

iA Ay menocallis

Batis

v

**e*e Conch shells

80

metres

Hatchet Cay 1972

Figure 32.

CLOT ACD TOIEM STIITI “EE SANHTy

OOL souyow

DUI[BUOYS INDUOPUN) wow

si}Dq

ysuowW s6bpes

uuniAnses [.--

Sndup50u05 |

snol4
DIUUSSIAY

puoudoziyy
Squey puDd

SINUODOD BSUZsq

sesspu6 yim ( i

4

uN

f a5

fo}

Growing coral in lagoon entrance
Porites porites P astreoides and
Siderastrea radians

Coccoloba
Rhizophora

Avicennia

Cordia
Terminalia
ThArinax

Erithalis

SS)

©
\

Sophora

Conocarpus

O
4 .
ae eee LL Braet ole
senae NA Ipomoea

Ww Wedelia

w Euphorbia

FFF Woodland

uit, Beachridge-crest

Figure 34. East Snake Cay 1961

E24) Sand
[°°] Shingle

O m 50
Lato I

‘Aft Diploria
block

o-
6°%>5 Bio) Orie 02:
° “Ho 36

© O ices te
3 eGo © fe:

aso Vora

Coccoloba
g Rhizophora
o Avicennia

Pp ihirinax

3 1 Tournefortia
® Suriana

; 9 Conocarpus
» Sophora
[i] Sesuvium

VA, Aymenocallis
MQ! pomoea

& “Goconuit

w Euphorbia

Ej Grasses

Figure 35. Middle Snake Cay 1961

T96T ASD SYyeUS ASeM “°9E SANHTA

a|6ulus [2° 0] 7 winiansaes fee
puos [27] sndup2.0u0D oo
ADOIYIDIG NYY paowod| KA

DiquoJsusuNo, 1

ud O
DUOYUdOZIYUN B&B

A

J910M Bulpudiys

eaaouBudWw
A A UYONW UM

dwioms eaoubudw

T96T ASD SyeUS ASEM PUeTST ASEM “LE SANHTA

3$94u9-a6piu yoDag mn DaOwWOd| WN

ai6uius [5 6] WINIANSasS es

DUDIUNS 9°

sndudd0u07
DyuOUdOoZIYY

DqGo|OdD05

Sy0|q
[@)

floating

pumice-
covered

Coceoloba

ThArinax

Conocarpus

Sophora

a Hymenocallis

W Wedelia

TL Dense coconut thicket

Ug Beac hrock

mT Beach ridge-crest

www Undercut shoreline
22:1 Sand

F2] Shingle

Figure 38. South Snake Cay 1961

moa
FEHECEESHEH Mavadti
Pr Mound

Coccoloba

Rhizophora

Citrus \ |
Mango

Conocarpus

Opuntia

issaaai Coconut woodland
with grasses

E CoOcont
| v_v |Mudhole
WU’, Beachrock

[°. °.| Shingle
O m
cee seedmore comantebineten 2

Figuxe 39. Wild Cane Cay 1961

FET Thick mangrove
Coccoloba

Rhizophora
Thrinax
Sophora

Wii Hymenocallis

Cc Coconut

F——] Cyperus and grasses
awww Undercut shoreline

(asitaseeis

SRS SGCCSG aoe (ee eee es, r
{1G ieo esses ——__——
2 ean oi ue ie
A500 ee aoe weores
ESeas

4 GB
paseeseaeeTaeeeseeeeeaeeeee
/AUISOS00SGeResoEcoo
itis Geen

2 alelotelalal sete a EEE
Sek tot tcteaieletela ie etetet stele ete
Se OoUdecodoeee

HEEPeeCReeeloua | Cl KS
ia ef ex n| nfiaaf 7: Cp
Aa A) of We

Figure 40. Moho Cay 1961

—

Plate 66. St. George's Cay: aerial view from the north 1962 (channel A)

Plate 67. St. George's Cay: aerial view from the north 1962 (channel C)

Plate 68. St. George's Cay: aerial view from the east 1962 (channels D and E)
(For the location of these channels see Stoddart 1963, fig. 19)

Plate 69. Tobacco Range 1972: view inland from the northeast sand ridge
Plate 70. Tobacco Range 1972: Rhizophora killed by Hurricane Hattie in
1961, invaded by Batis, at the northeast point

alee
Was
U3} 6
1962

Cary Cay:
Scipio Cay:
Colson Cay:

eee

hurricane-deposited shingle on the east side 1962
hurricane-deposited shingle ridge 1962
hurricane-deposited shingle on the southeast

Plate 74. Water Range (Twin Cays): Thrinax thicket, south shore 1972

Plate 75. Little Water Cay: shingle rampart on the southeast shore 1972

78

Plate 76. East Snake Cay: aerial view from the southeast 1961
Plate 77. West Snake Cay: aerial view from the northeast 1961
Plate 78. South Snake Cay: aerial view from the southeast 1961

Plate 79.

Plate 80.

nis
we

att

East Snake Cay rampart and moat from the north 1961

West Snake Cay rampart and moat from the south 1961

63

6. SUMMARY AND CONCLUSIONS

A. Types of islands

This consideration of the cays of the Belize barrier and shelf has
TGucdted they vartabi lity Of eet stand types! in such an) extensive
province. In a previous paper (Stoddart 1965, pp. 134-142), nine
types of island were distinguished on the Belize reefs as a whole and
their distribution mapped. This classification can now be extended
and the following types recognised:

1. Unvegetated sand cay. These are ephemeral islands, usually
less than 50 m long and often overtopped by swash, and lacking contemp-
orary beachrock. Many on the Belize reefs are second-generation

features which have re-formed following the destruction of an earlier
larger island by hurricane damage; the location and extent of the former
island can be recognised by relict beachrock. Examples are Paunch

Cay and Curlew Cay.

2. Vegetated sand cay. These are larger islands with a
vegetation of strand scrub and strand woodland, though in most cases
the woodland has been removed for coconut plantations. Examples are
Nicolas and Tobacco Cays. Smaller vegetated sand cays with maximum
dimensions of about 100 m are still subject to major modification or
even destruction during hurricanes. Most vegetated islands, however,
persist or rapidly re-form in the same locations (e.g. Sergeant's,
Goff's, English and Rendezvous Cays). The largest vegetated islands
to disappear completely in the 1961 hurricane were St. George's East
Cay GHOnmeliong Oss aha)mandsCaya Gillonyat(lOS=m long,) Onl Sieha)):

3. Unvegetated shingle cay. These are small, ephemeral, and
located in exposed situations. Often they may be reef-top ridges
adjacent to a larger cay. They may from time to time be colonised by

patches of Philoxerus or Sesuvium and mangrove seedlings.

4. Vegetated shingle cay. These are also small, located in
exposed situations, often on small patch reefs, and are topographically
mobile. Good examples showing substantial changes during 1960 and

1972 are North Spot and Ragged Cays.

5. Sand and shingle cays. These are larger and more stable
islands, in more exposed situations than simple sand cays, and consist
of islands with a windward shingle ridge and leeward sand area, with
or without an intervening depression. Such islands are lacking on the
barrier reef in the lee of the offshore atolls, where only simple sand

64

cays are found. Because of their size and stability these islands are

always vegetated. Strand woodland survives on Nicolas Cay, but else- |
where has been replaced by coconuts. Examples are Lime, Hunting and

Northeast Sapodilla Cays, the latter with a large interior depression.
This type is also typical of the windward sides of the atolls. Most
such islands have both modern and relict beachrock, the latter
indicating beach retreat on their windward shores.

6. Mangrove cays. On the Belize shelf these are dominated by
Rhizophora mangle; Jack's Cay is an example. Ebanks (1975, p.278)
describes them as "little more than mud mounds colonised by vegetation".
They are found only within the lagoon and not on the barrier reef
itself.

7. Shelf stands: This term is introduced by Ebanks (1975, pp.
277-278) for islands which "form by sediment accretion on a partly
submerged topographic prominence" on a wide shallow shelf. These
islands are located on the shallow shelf north of Belize, 3.5-5 km
back from the reef edge. They are elongate islands, parallel to the
shelf edge, with a windward sand ridge of variable height and width,
generally less than 2 m above sea level but exceptionally reaching 6 m.
Their leeward sides are formed by mangroves, bays and intertidal flats.
Examples include Ambergris Cay, Cay Chapel and Cay Caulker, and Ebanks
draws attention to similar shelf islands in the Bahamas and Florida.
Previously they had been included in the general category of mangrove-
sand cay by Vermeer (1959, pp. 35-39) and Stoddart (1965, p.139).

8. Mangrove cay with dry sand areas. Many mangrove islands have
low-lying sand areas, often but not exclusively on the windward side,
and usually flat and featureless. These cays are found in fairly
protected situations, and may be a late stage in the evolution of
ordinary mangrove cays. Frequently the dry-land areas have been
cleared and planted to coconuts. Examples are Weewee, Buttonwood,
Hatchet, Wild Cane and Frenchman's Cays.

9. Mangrove range. These are extensive and complex arrays of
mangrove islands, often with intermittent low sand ridges on their
windward sides. They probably owe their location and arrangement to

pre-existing topographic variations in the substrate. The islands are
separated by partly enclosed bays and lagoons, rather than by inter-
tidal flats, and also by sinuous, deep, narrow tidal channels known as
"bogues'. Tobacco Range and Water Range are typical mangrove ranges;
bogues are particularly well seen in the Drowned Cays, as well as in
the extensive mangroves of Turneffe.

10. Moat islands. This term has been applied to characteristic
associations of leeward sand area, interior mangrove swamp, reef-top
shallow lagoon or moat, and windward-reef-edge shingle ridge formed on
small patch reefs in the southern shelf lagoon (Stoddart 1965). East
and West Snake Cays are good examples. Topographically these islands
have features in common with reef islands on the Queensland coast
(Steers 1937) and in Djakarta Bay (Umbgrove 1928). The Snake Cays owe

65

their distinctiveness to their relatively exposed situation in deep
water facing the open southern shelf, which allows the formation of
shingle ridges on the windward reef edges; the other features have
formed in the lee of these ridges.

11. Coastal barrier islands. On the Belize coast the term 'cay'
is applied to any island, not only those of reef origin. Many islands
near the mainland coast are barrier beaches or detached cuspate head-
lands, formed of terrigenous sediments (High 1975). Harvest Cay is
an example.

B. Topographic change

These surveys have shown considerable topographic change on almost
all islands of the coastal shelf and barrier reef for which we have
comparative data. Of 24 islands mapped in 1960-61 and again in 1972,
1l decreased in area and 13 increased. The greatest percentage loss
was 31 per cent at Rendezvous Cay, and the greatest percentage increase
was 160 per cent at North Spot. Aggregating all these islands
together, the total area decreased from 35.7 to 33.3 ha, a decrease of
6.6 per cent (cf. the changes on the islands of Glover's Reef, where
the total area of six islands declined from 33.6 to 31.8 ha, a decrease
ef 5.4 per cent: Stoddart et al. , 1981). As might be expected, the
percentage changes are most marked on the smallest cays. Four modes
of topographic change may be distinguished:

1. Catastrophic change caused by a Single event, in this case
Hurricane Hattie of October 1961, which affected barrier reef islands
in northern Belize. Small vegetated sand cays were spectacularly
eroded and two (Cay Glory and Paunch Cay) disappeared. Recovery took
place over the ensuing decade, so that comparative figures between
1961 and 1972 give a very conservative indication of change: Goff's
Cay gained 10 per cent, and Sergeant's, English and Rendezvous Cays
lost 54, 25 and 31 per cent respectively.

2. Severe marginal erosion by an unknown event, but probably by a
specific hurricane. This is widespread on the cays of the southern
barrier reef, e.g. Frank's, Lime and Nicolas Cays. This must be a
fairly common consequence, of hurricanes of lesser severity than Hattie.

3. The re-working of small islets, notably of shingle islands in
exposed situations. This may be accomplished by ordinary storms over
periods of years, or it may result from single hurricanes. A moderate
example is given by North Silk Cay, and more extreme examples by North
Spot and Ragged Cay.

4. The accumulation, probably mainly on a seasonal basis, of sand
in the form of spits, lobes and bars during ordinary non-catastrophic
conditions. This mostly results in ephemeral topographic features,
but some become vegetated and stabilized.

66

There is some suggestion in the number of cays which have
disappeared since Owen's surveys in 1830, as well as in the widespread
erosion during 1960-1972, that reef islands in the Belize province may
be undergoing a period of diminution. It is not possible to say
whether this relates to the results of fluctuating sea levels,
increased frequency of catastrophic storms, or both, but it has been
argued (Stoddart 1964) that an important factor in the present dis-
equilibrium is the destabilising effect of the replacement of natural
strand woodland by coconut woodland during the past century.

C. Flora and Vegetation

A total of 178 species of vascular plants has been recorded from
the Belize cays, including 32 non-native species (18 per cent) (Fosberg
et al. 1981). Of these, 90 species are found only on the barrier reef
and shelf islands, and not on the atolls; these include 28 non-native
species or 31 per cent; 15 are found only on the atolls and not on the
barrier reef and shelf (no non-native species); and 73 are common to
the barrier and shelf cays and to the atolls (3 non-native species, or
4 per cent).

The total flora of the barrier reef and lagoon islands comprises
164 species, including 31 non-native species (19 per cent). Of these,
61 are found only on the barrier reef islands (17 of them non-native,
or 1l per cent), and 22 only on the shelf islands (4 non-native, or

7 per cent). 37 species are extremely widespread, and include strand
grasses and sedges, strand shrubs, common weedy species of coconut
woodland, and mangroves. They are:

Acrostichum aureum Euphorbia mesembrianthemifolia

Eragrostis prolifera
Paspalum distichum
Spartina patens
Sporobolus virginicus
Cyperus ligularis
Cyperus planifolius
Fimbristylis cymosa
Cocos nucifera

Thrinax radiata
Hymenocallis littoralis
Coccoloba uvifera

Batis maritima
Philoxerus vermicularis
Portulaca oleracea
Sesuvium portulacastrum
Canavalia rosea

Sophora tomentosa

Vigna luteola

Rhizophora mangle
Conocarpus erectus
Laguncularia racemosa
Terminalia catappa
Ipomoea macrantha
Ipomoea pes-caprae
Ipomoea stolonifera
Cordia sebestena
Tournefortia gnaphalodes
Avicennia germinans
Stachytarpheta jamaicensis
Erithalis fruticosa
Ernodea littoralis
Ageratum littorale
Borrichia arborescens
Eclipta alba

Wedelia trilobata

67

It is clear that the native woodland of the cays has been greatly
disturbed by planting of coconuts, and on most islands has entirely
disappeared. Its main components probably included Thrinax radiata, {
Coccoloba uvifera, Ficus sp., Neea choriophylla, and Bursera simaruba.
Strand shrubs are represented mainly by Tournefortia gnaphalodes,
Suriana maritima, Conocarpus erectus and Borrichia arborescens. The
complete absence of Scaevola plumieri from these islands is remarkable
(it occurs on the north coast of Yucatan, on Alacran and Cayo Arcas,
and in the Cayman Islands to windward), as is the rarity of
Chrysobalanus icaco.

The plants of the dry mangrove cays and the mangrove ranges are
quite distinctive, with many more grasses, sedges and succulents than
the sand cays of the reef edge. Species which are characteristic of
the dry mangrove cays (but most of which are very widespread) are
Eragrostis prolifera, Eustachys petraea, Paspalum distichum, Spartina
spartinae, Fimbristylis cymosa, Fimbristylis spadicea, Thrinax radiata,
Batis maritima and Pluchea symphytifolia. Two species (Salicornia
perennis, Lippia strigulosa) are found only on these islands.
Conversely, some species which are common on the reef-edge sand cays
are absent from the dry mangrove cays. These include Cakile
lanceolata, Canavalia rosea, Bursera simaruba, Pouteria rivicoa,
Morinda citrifolia and Ambrosia hispida.

It is also instructive to compare the plants recorded on the two
Mangrove ranges visited, Tobacco Range and Water Range. 33 species
are recorded in total, with 15 in common. Apart from the mangroves,
these are Eustachys petraea, Paspalum distichum, Cyperus planifolius,
Thrinax radiata, Batis maritima, Salicornia perennis, Suriana martima,
Euphorbia mesembrianthemifolia, Erithalis fruticosa and Borrichia 4
arborescens. A larger sample would undoubtedly include as character- i
istic plants of these islands Eragrostis prolifera, Phragmites cf.
australis, Spartina spartinae, Cladium jamaicense, Sesuvium portulacas-
trum, Pithecellobium keyense, Ernodea littoralis, and others. ADS ALS}
particularly interesting to observe the long term effects of hurricane
damage on Tobacco Range, where mortality of mangroves in 1961 has led
to their replacement by Batis maritima, even though the dead tree
trunks still stand.

We have already noted the existence of a fairly steep rainfall
gradient from north to south along the Belize coast. The effects of
this appear in the distribution of ferns and orchids. Ferns other y
than Acrostichum aureum occur only on Nicolas Cay (3 species), Lime
and Wild Cane Cays. Orchids have been recorded only on Frank's,
Hunting, Jack's, East Snake and Middle Snake Cays (as well as on North-
east Cay, Glover's Reef). The climbing vine Rhabdadenia biflora is
only recorded from the southern shelf (Hunting Cay) and Glover's Reef
(Southwest Cay).

Finally, the catastrophic damage caused to some of the northern
sand cays by Hurricane Hattie has allowed the observation of success-
ional processes over the ensuing decade. The primary coloniser on

68

bare sand after the storm was Portulaca oleracea, together with
Sesuvium portulacastrum and Euphorbia mesembrianthemifolia; these were
well established as scattered colonies within five months of the storm.
By 1965 Portulaca had greatly declined in importance, and the main
species present were Sesuvium, Euphorbia and Ipomoea pes-caprae. By
1972 many more species were present, including shrub species such as
Tournefortia, Conocarpus and Suriana. The following table gives the
number of species recorded on the most damaged islands following
Hurricane Hattie:

Island 1962 L965 LOW
Sergeant's Cay 4 12 14
Goff's Cay 2 4 10
English Cay 2 5 18
Rendezvous Cay 16 16 Li

The opportunity clearly exists to monitor vegetational changes on these
islands in the future, as well as topographic change in general, to
establish the long-term response of islands to normal as well as to
extreme events. .

We do not wish in this paper to consider the floristic biogeography
of the cays in greater detail. A preliminary paper has been prepared
(Stoddart and Fosberg, in press), and a more detailed treatment is in
preparation. We can, however, note that there is a very close
association between island area and number of species of vascular plants
present, and that area appears to be the major control of floristic
diversity on islands of the Belize barrier reef and shelf.

69

7. REFERENCES

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(al

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72

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73

Wagner, P.L. 1964. Natural vegetation of Middle America. In R.
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Appendix: Manuscript and published charts of the Belize shelf

De Mayne, A. 1829. Coast of British Honduras, from Belize to Bluff
Point, south of Punta Ycacos. Recd. from Mr de Mayne, Sept. 25th
(18)29. Hydrographic Department, Ministry of Defence, Taunton,

No E992.

Douglas, H.P. 1922. Southern Inner Ship Channel to Belize, Rugged Cay
to Middle Long Cay. Ibid. No C8925.

Frembly, J. 1830a. A diagrammatic survey of the harbour of Belize on
the coast of British Yucatan. Ibid. No G207.

Frembly, J. 1830b. A plan of the anchorage at the Southern Triangles on
the coast of British Yucatan. Ibid. No G208.

Glennie, R.W. 1921. British Honduras, Turneffe Islands, Cay Bokel
Anchorage. Ibid.

Hydrographic Department, Ministry of Defence. 1839. West Indies from
Belize to Cabo Catoche. Surveyed by Commanders R. Owen and E.
Barnett 1830 and 1837, with additions and corrections to 1931.
london): edition WW, S39); edastiion) 2), 1l9lSie) vAdmiralty Chart No 1204.

Hydrographic Department, Ministry of Defence. 1844. West Indies,
Honduras Gulf with the Zapotillos Cays (surveyed 1835-41).
London: edition 1, 1844; new editions 1868, 1871, 1896, 1897,
LOM Lo 22) Aamaimaley Charme No) d57/3).

Hydrographic Department, Ministry of Defence. 1898. British Honduras,
Belize Harbour (surveyed 1896-7). London: edition 1, 1898; new
edition lO2ZAr Chart eNo i522)

Hydrographic Department, Ministry of Defence. 1929. British Honduras,
Approaches to Belize (from surveys 1896-7 and 1921-2). London;
edition in 929 ey Admiralty, Chart (No 959.

Hydrographic Department, Ministry of Defence. 1929. British Honduras,
Southern Inner Channel to Belize, Ranguana Cay to Columbus Cay
including Glover Reef (surveyed 1830-41 and 1922). London:
edition 1, 1929; revised 1948. Chart No 1797.

74

Hydrographic Department, Ministry of Defence. 1960. Central America,
British Honduras, Belize Harbour (surveyed 1957-8). London:
edition 1, 1960. Admiralty Chart No 522.

Irving, E.G. 1957-58. Central America, British Honduras, Belize
Harbour. 1:25,000. Hydrographic Department, Ministry of Defence,
Taunton, Chart No K2254-55E.

Owen, R. 1830a. Part of the coast of British Yucatan. From Jonathan
Point to Ambergris Kay. October 1830. Ibid. No H57.

Owen, R. 1830b. Part of the coast of British Yucatan. The Gulf of
Honduras and kays adjacent. November 1830. Ibid. No H6l.

Owen, R. 1834a. (Cay Columbus to South Water Cay). June 1834. Ibid.
No L83.

Owen, R. 1834b. (South Water Cay to Quamino Cay). July 1834. Ibid.
No L84.

Owen, R. 1835a. Sapodilla and Seal Cays in the Bay of Honduras.
September 1835. Ibid. No L458.

Owen, R. 1835b. The narrow part of the Main Channel to the southward
of Belize, between Point Placentia and Monkey River. September
PES5~ Lbids) NowEA59.

Smith, T. 1840a. Victoria Channel, Honduras. August 1840. Ibid. No
L2481.

Smith, T. 1840b. Part of the coast of British Yucatan between
Zapodilla Cays and Monkey River. August 1840. Ibid. No L2474.

INDEX OF ISLANDS

Page citaclons ace jtOu the! main, remrerence! Eo) cach Of the islands in the

Petes

Alligator Cay 47
Ambergris Cay 40
Baker's Rendezvous 51
Bedford Cays 57
Blackadore Cay 42
Blue Ground Range 48
Bluefield Range 47
Brown's Cay 14

Bugle Cay 53
Buttonwood Cay 52
Cangrejo Cay 42
Carrie Bow Cay 20
CanyeCay, 5S

Cate CEy7y Sil

Caulker Cay 42

Cay Cangrejo 42

Cay Caulker 42

Cay Chapel 43

Cay Glory 16

Chapel Cay 43

Coco Plum Cay 49
Cohune's Kay 29
Colson Cay 54

Colson Cays 47
Columbus Cay 47
Crayfish Cay 46

Cross Cay 47

Curlew Cay (north) 11
Curlew Cay (south) 21
Deer Cay 42

Drowned Cays 45

East Snake Cay 58
Ellen Cay 20

English Cay 12

Fly Range 48

Frank's Cay 33
Frank's Cays 32
Frank's Cay East 33
Frank's Cay West 33
Frenchman's Cay (south) 61
Frenchman's Cays (north) 45
Glory Cay 16

Goines Geyy dal

Grass Cay 32

Grennels Cay 46
Halilivisni€ay, 29

Harvest Cay 57

Hatchet Cay 55

Hen and Chickens 45
Hick's Cays 45

Hobbe's Kay 57

Hunting Cay 35

Jack Ellin's Cay 20
Jack's Cay (south) 51
Jack's Cays (north) 15
Laughing Bird Cay 56
Lime Cay 37

Little Peter Cay 51
Little Water Cay 56
Long Cay (north) 45
Long Cay (Southern Triangles) 46
Low Cay 37

Man-o'-War Cay 50
Middle Long Cay 47
Middle Silk Cay 24
Middle Snake Cay 58
Moho Cay (north) 45
Moho Cay (south) 62
Moho Cay (= Trapp's Cay) 52
Montego Cay 45

Mosquito Cay 42

Nicolas Cay 34

North Drowned Cay 40
Nowehisiilk ay, 23

North Spot 28

Northeast Sapodilla Cay 31
Observation Seal Cay 38
One Man Cay 46

Owen Cay 54

Patience Brother Island 57
Paunch Cay 9

Paunchgut Kay 9
PeikitcanmCay smo

Peter Douglas Cay 51
Peter's Bluff 40
Placentia Cay 57
Pompion Cay 26

Punchgut Kay 9

Queen Cays 23

Ragged Cay 38

Ragged Snake Cay 58
Ramseys Cay 46

UD

76

Ranguana Cay 27 Southern Long Cay 47
Rendezvous Cay 14 Southern Triangles 46
Renegado Kay 27 Spanish Cay 47

Rider's Cays 45 Spanish Lookout Cay 46
Robinson Island 47 Stake Bank 45

Robinson Point Cay 46 Swab Cay 42

Round Cay 25 Tobacco Cay 16

St George's Cay 44 Tobacco Range 48

St George's East Cay 9 Tom Owen's Cay 29
Samphire Cay 25 Tom Owen's East Cay 29
Samphire Spot 14 Tom Owen's West Cay 30
Sapodilla Cays 31 Trapp's Cay 52

Scipio Cay 54 True Point Patience Kay 57
Seal Cay (north) 12 Twin Cays 50

Seal Cay 38 Virgins, The 57

Seal Cays 38 Water Cay 46
Sergeant's Cay 10 Water Range 50

Silk Cays 23 Weewee Cay 51

Simmonds Cay 46 West Island West Snake Cay 59
Ski fissandeks West Snake Cay 59
South Cay 38 White Rock 26

South Silk ‘Cay ‘25 Wild Cane Cay 60
South Snake Cay 60 Zapotilla Cay 38

South Water Cay 18 Zapotilla Cays 31

ATOLL RESEARCH BULLETIN
No. 257

TEN YEARS OF CHANGE ON THE GLOVER'S REEF CAYS
by q

D.R. Stoddart, F. R. Fosberg and M.- H. Sachet

Issued by
THE SMITHSONIAN INSTITUTION
Washington, D. C., U.S.A.
April 1982

List of Figures

Location and submarine topography of Glover's Reef.
Dillon and Vedder (1973).

Topography of Glover's Reef. From Stoddart (1962,

Legend to the detailed physical and vegetation maps of the cays.

Northeast Cay 1961 and 1971: topography.
Northeast Cay 1961 and 1971: vegetation.
Long Cay North 1961 and 1971: topography.
Long Cay North 1961 and 1971: vegetation.
Long Cay 1961 and 1971: topography.

Long Cay 1961 and 1971: vegetation.

Middle Cay 1961 and 1971: topography.
Middle Cay 1961 and 1971: vegetation.
Southwest Cay I 1961 and 1971: topography.
Southwest Cay I 1961 and 1971: vegetation.
Southwest Cay II 1961 and 1971: topography.

Southwest Cay II 1961 and 1971: vegetation.

aval

LO%

ele

ae

Aig He

14.

Sis

Gi.

dhe

LS.

List of Plates

Northeast Cay: aerial view from the east 1962

Northeast Cay: storm block and shore erosion on the east side
1961

Long Cay North: Suriana scrub on the north shore 1961

Long Cay North: the south shore in 1971

Long Cay from the southeast 1962

Southwest Cay II: aerial view from the southeast 1962

Long Cay: Tournefortia on the northeast spit 1961

Long Cay: upper surface of the windward shingle ridge 1961

Long Cay: sandy island surface with backslope of the windward
shingle ridge 1971

Long Cay: mud hole and standing water at the east end, seen from
the windward shingle ridge 1961

Middle Cay: windward shingle ridge with hedge of Tournefortia
AUS) 9/At

Middle Cay: low lagoon shore with Conocarpus 1971
Middle Cay: Rhizophora at the east end of the lagoon shore 1971
Southwest Cay I: seaward rubble shore 1971

Southwest Cay I: mangrove pool at the southern end of the island
Ion.

Southwest Cay II: exposed 'rootrock' on the retreating lagoon
shore 1971

Southwest Cay I: open mangrove 1961

Southwest Cay II: beachrock with landward dip on the seaward
shore 1961

TEN YEARS OF CHANGE ON THE GLOVER'S REEF CAYS

lOSY | Dion dsta Stoddart, ! Po IRs Fosberg” and M.-H. Sachet?
ABSTRACT

Scientific studies on the atoll of Glover's Reef, Belize, since
an initial survey in 1961 are reviewed. Comparative maps are presented
of each of the six islands on the atoll, showing morphology and
vegetation, between the 1961 survey and a re-survey in 1971. Most of
the islands show shoreline erosion and remodelling of windward shingle
ridges as a result of hurricane activity. The recorded terrestrial
plants are catalogued for each of the islands, and some extinctions are
noted. Species-numbers on the cays are related to area, and are much
higher than those for other Caribbean and Gulf of Mexico coral islands,
presumably because of higher rainfall.

INTRODUCTION

Glover's Reef (16°45'N, 87°50'W) is one of three atolls off the
mainland coast of Belize (British Honduras), central America. When
visited in 1961 it was one of the least-known reefs in the Caribbean,
in spite of its typical atoll features. It was delineated on charts
(Admiralty Chart 1797 of 1929) in the most rudimentary manner, and
published scientific literature about it was limited to brief reports
on visits by the ornithologist Salvin in 1862, by the Pawnee Expedition
in 1925, and by the Mandel Caribbean Expedition in 1940. The account
of the atoll published by Stoddart (1962, pp. 17-30, 83-98) was thus
necessarily of a preliminary nature.

1Department of Geography, Cambridge University, England
2Smithsonian Institution, Washington, D.C., U.S.A.
Manuscript received January 1981 -- Eds.

IMSWE (Investigations of Marine Shallow Water Ecosystems) Contribution
No. 75

Since then, however, there has been a great deal of scientific
investigation on Glover's Reef. The CITRE project (Comparative
Investigations of Tropical Reef Ecosystems) included the atoll as one of
its potential study sites, and reconnaissance visits were made to it in
January and June 1971, followed in November 1971 by a planning workshop
for 40 scientists held at Long Cay (Smith, in Sachet and Dahl (1974),
lists participants). Descriptions of the reefs resulting from CITRE
studies were given by Dahl et al. (1974, pp. 55-58), and marine algae
and sea-grasses collected during the project were reported by Tsuda and
Dawes (1974). In 1972 N. James, R. N. Ginsburg and associates carried
out studies of the deep reefs using the deep-diving submersible Nekton
(James and Ginsburg 1980). Finally, reef structures within the atoll
lagoon have been studied by Hughes (1973), Schafersman (1972), Wallace
(1974), and Wallace and Schafersman (1977). Apart from studies carried
out during the CITRE workshop, terrestrial studies on the cays have
been limited to plant collections made by Pippen in 1972 and by Linhart
and students in 1973 (Linhart 1980; Fosberg et al. 1981).

This paper reports on the re-survey of the cays carried out during
the CITRE workshop in 1971. Comparative maps of all the islands in
1961 and 1971 are presented, together with complete lists of terrestrial
plants recorded from each cay (including those reported by Linhart
(1980)). All of the islands were mapped by pace-and-compass methods.
They were also photographed from the air in colour and black-and-white
in L961 and 1962, and mores briefly in 1971. The maps of each of the
islands are orientated conventionally (i.e. magnetic north at the top
of the diagram).

HISTORY

The early history of the atoll is obscure, as indeed is that of

most of the reefs in the western Caribbean. It can possibly be
identified as the "Ilbob" or "Ylbob" of maps by Herrera dated 1601-1615
and of De Laet in 1625, and derivative charts (Vindel 1955). elite

appears as "Longorif" in the Plano de la Costa de Honduras of 1756; as
"Arrecife de Guaneros" in a map by J. J. de Castillo, dated 1753
(reproduced by Craig 1966, p. 38); and as Glover's Reef by Speer

(1771) and Arrecife Largo, Glover's Reef or Long Reef by Jefferys

(LITE) 6 Speer's map marks three un-named cays; Jefferys's map has
five un-named cays in the south and "The Two Spots" at the northern end.
The first Admiralty survey by Richard Owen in 1830 adopts the name of
Glover's Reef, which then became standard, and also for the first time
names Northeast, Long, Middle, and Southwest Cays.

The only event of any significance in the recorded history of the
atoll is the wreck of the schooner Susan, Captain Harry Maury, which
sailed from Mobile on 4 December 1858 with emigrants and filibusters
bound for Omoa, Republic of Honduras, wrecked 10-12 miles from the cays
on 16 December (Doubleday 1886; Scroggs, 1916) . The vessel was found
by a fishing boat which had taken fish, green turtle and coconuts to
the Belize market, and the passengers and crew were taken to Middle Cay.

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This was said to be an island of 30 acres (12 ha), with a "continuous
grove of cocoa palms ... [and] entire freedom from underbrush"
(Doubleday 1886, p. 212). Here they lived on "conch and green-turtle
soup, fish in great variety, cocoa-nuts, yams, plantains, and bread-
fruit, all in great abundance", until rescued by H.M.S. Basilisk,
sent from Belize, on Christmas Day 1858. There are brief accounts of
this incident in Harper's Weekly, 8 January 1859, p. 22, and 15 January
1859, p. 39, and in the Mobile Mercury at the same time [not seen], but
the most circumstantial account is contained in letters by Captain
Maury published in The Mobile Daily Register, vol. 32 (no. 1329), p. 1
for 4 January 1859. We are grateful to Mr M. M. Weinstein, Library of
Congress, for copies of these documents.

The present study began with a visit to Glover's Reef by Stoddart
and ‘Si. Ph Murray Jin 96Uk: This was a brief reconnaissance supported
by the Office of Naval Research and the Coastal Studies Institute of
Louisiana State University as part of a wider study of the British
Honduras cays, and we are grateful to the late Professor Richard J.
Russell and Miss Evelyn Pruitt for making it possible. The second
visit took place as part of the CITRE workshop on 1-13 November 1971.
The islands were re-mapped and plants collected by Stoddart, Fosberg
and Sachet, and we are grateful to Dr Stephen V. Smith, Principal
Investigator, for the opportunity to participate in the CITRE planning
sessions. We also thank Mr M. Young and Mr R. Coe, of the Department
of Geography at Cambridge University, for much cartographic and
photographic work, and Mr M. M. Weinstein, Library of Congress, who
obtained newspaper records relating to the wreck of the Susan.

STRUCTURE AND FORM

The structure of the east coast of the Yucatan Peninsula is
controlled by a series of northeast-southwest trending fault blocks,
arranged en echelon on the northwestern flank of the Cayman Trench.
One of these lineaments underlies the Belize barrier reef south of
Gladden Spit, Glover's Reef, and Lighthouse Reef; another the central
barrier reef, Turneffe, and Banco Chinchorro; anda third the mainland
coast north of Ambergris Cay. Of these, the first is best developed,
and in places (as on the east side of Lighthouse Reef) has a seaward-
facing scarp nearly 3000 m high. At Glover's Reef itself, the sea
floor falls sharply to over 1000 m on the east side of the atoll, but
only to shallow saddles of 300-400 m between the atoll and the barrier
reef and the atoll and Lighthouse Reef. Wells drilled through the
offshore reefs have reached metamorphic rocks probably equivalent in
age to the mainland Santa Rosa Group (Carboniferous-HZarly Permian) at
between 777 and 959 m on Glover's Reef, and extrusive volcanics of
Similar age at 1692 m on Turneffe.

The surface reefs, therefore, stand on fault-block structures, as
originally inferred from topography and fully confirmed by seismic
investigations (Dillon and Vedder 1972, 1973; Uchupi 1973). The deep
well on Glover's Reef showed 260 m of calcareous siltstone of Late

Cretaceous age, overlain by 560 m of Tertiary reef facies; similarly,
on Turneffe, 660 m of shale with limestone is overlain by 1030 m of
reef facies (Dillon and Vedder 1973, p. 2721). These sequences
indicate reef upgrowth on subsiding fault blocks at rates (1-2 cm/1000
yr) and over time periods (ca 60 million yr) comparable to those of
upgrowth on open Pacific subsiding volcanoes such as Bikini and
Enewetak.

Glover's Reef itself is an elongate rectangular atoll up to 32 km
long and 12 km wide, with an area of 260 sq km. It is surrounded by a
shallow reef flat, generally about 450-700 m wide but which reaches
1200 m on the eastern (windward) reefs. Living reef reaches the
surface at the outer edge of the reef flat for about 150 m on the
windward reefs. There is no algal ridge directly comparable to those
of some Indo-Pacific atolls, but rather an algal pavement with
rhodoliths of Goniolithon boergesenii, clearly the functional equivalent
of the algal ridge. The leeward reefs are less well-defined and lack
the algal-pavement zone (Stoddart 1962, pp. 23, 25-26). There are
three gaps through the peripheral reefs; one in the northeast, 1300 m
wide; one in the south, 1600 m in width and up to 12 m deep; anda
smaller exposed channel at the southeast point.

The lagoon enclosed by the reefs has two components: a shallow
shelf or platform 6-8 m deep and generally 1-2 km wide, and a central
basin. The platform is similar to the "barrier platform" of the
coastal shelf, and is probably a reef surface of last interglacial age
on which the present surface reefs have grown. The central basin is
probably about 20 m deep (no bathymetric survey has been undertaken),
and contains some 700 patch reefs. Variation in patch reef ecology
and sedimentology has been studied by Schafersman (1972), Hughes (1973),
Wallace (1974), and Wallace and Schafersman (1977).

The coral fauna of Glover's Reef is known only in outline. York's
(1971) list of the corals of the southern Belize barrier lists 18
genera. With the addition of Mussa and Meandrina this gives 20 genera
recorded from the Belize reefs (Stoddart 1962, p. 19). 13 genera have
been recorded from Glover's Reef (ibid), but unpublished CITRE records
will undoubtedly increase this total. There are few other marine
faunistic records from Glover's Reef. Devaney (1974) records 3
asteroids, 15 ophiuroids, one holothurian and 4 echinoids from
collections made in 1969. Collections of crustaceans, molluscs, es
echinoderms, coelenterates and fishes made during the Pawnee Expedition a
are listed by Boone (1927, 1928a, 1928b, 1928c) and Breder (1927).
Tsuda and Dawes (1974) list four sea-grasses and 100 species of marine
algae from the CITRE investigations. Terrestrial records are limited
to two reptiles (Anolis sagrei, Ctenosaura similis) collected by the
Mandel Caribbean Expedition in 1940 (Schmidt 1941) and a hummingbird if
(Ancathothorax prevosti) listed by Todd (1942). e

There are no environmental records from Glover's Reef. The atoll
is under the influence of the Northeast Trades. There is a marked
rainy season from June to September, and mean annual rainfall is probably
about 2500 mm. Mean tidal range at springs is probably about O.2 m.

The atoll is frequently affected by hurricanes, and reference to their
effects will be made in the accounts of the cays.

There are six small islands on the southeastern reef of the atoll.
Their combined area is about 32 ha, and it is with the changing
morphology and vegetation of these cays that this paper is concerned.

THE CAYS OF GLOVER'S REEF

There are at present six cays on Glover's Reef, all on the
southeast side. Five of these (Northeast, Long, Middle, Southwest I
and Southwest II) are sizeable islands 360-600 m long; the sixth (Long
Cay North or Small Cay) is an islet 160 m long northeast of Long Cay.
In addition to the main islands, Jefferys in his 1775 chart marked
"The Two Spots" at the north end of the atoll: these were presumably
ephemeral sandbores and no longer exist. Admiralty Chart 1797 of 1929
also marks a further cay lying within the lagoon on the west side of
the atoll, 7 km from the northwest corner; it is described as having

trees 6 m tall (West Indies Pilot, 1956, pp. 459-460). This island
did not exist in 1961, though an ephemeral sandbore near its location
was known as "Bushy Spot" or "Bushy Cay". Owen in his 1830 chart

(Admiralty MS chart H57) also shows no cay at this location. Owen in
1830 also noted two small sandbores between Middle Cay and Southwest
Cays. Each of the existing islands was mapped in 1961 and 1971; they
are described from north to south.

Northeast Cay (16°45'N, 87°45%4'W) Figures 4-5, Plates 1-2

Northeast Cay is situated at the south end of the unbroken eastern
reef of Glover's Reef, on the north side of the entrance between it and

Long Cay. It is semicircular in shape, with maximum dimensions of

360 x 200 m. Its southern and southwestern shores, facing the reef and
the channel, are formed by a shingle ridge with a maximum height of
about 2 m. In 1961 there was evidence of recent shore erosion on the

east Side of the cay, indicated by shore cliffing, surface sand
stripping, and coral blocks on the cay surface; by 1971 the shingle
ridge had extended northwards to cover much of this area. A small
area of beachrock on the south shore in 1961 had also increased in
length to about 90 min 1971, and a further small exposure had appeared,

indicating slight retreat of this shore. The northern shore is low
and sandy, and was stable during 1961-1971. The interior of the island
is also low and sandy, and there is a small mudhole. The area of the

island was 4.73 ha in 1961 and 4.48 ha in 1971. The island is covered
with coconut woodland, which is most dense and contains tall trees of
Bursera simaruba, Neea choriophylla and Thrinax radiata near the south
shore. There is a luxuriant undercarpet of Wedelia, Euphorbia,
Ambrosia and grasses. The most extensive shrub on the seaward side of
the island is Tournefortia gnaphalodes with scattered Conocarpus,
Coccoloba and Cordia. There were extensive mats of Sesuvium along the
lagoon shore. For further details of the island in 1961 see Stoddart
(1962,-pp..' 87=88;, :figse38-ands39)..

Forty species of plants have been recorded from the island (21 in
IMSL, I2 sim UV, eiacl SZ loyy inalolmewere alin US) 7/3})) - Of the species collected
in 1961 the orchid Brassavola nodosa was not found in 1971 or by Linhart
in 1973, and the single shrub of Suriana maritima on the southwest
shore haddisappeared by 1971. In the following species list, C denotes
a collection and S a sight record; the 1973 records are those listed
bya linha (9380) pp. o2—68)) Species marked with an asterisk are not
native to the Belize cay flora, but this does not necessarily imply
that they have been deliberately introduced on all the islands on
which they occur.

Acrostichum aureum 71C 73 *Terminalia catappa 71C 73

Cenchrus incertus 73
Eragrostis prolifera 61C 71C 73
Eustachys petraea 71C 73
Paspalum distichum 71C 73
Paspalum laxum 61C 73
Sporobolus virginicus 71C
Sporobolus sp. 61S
Cyperus ligularis 73
Cyperus planifolius 61C 71C

is

Fimbristylis cymosa 71C 73
*Cocos nucifera 61S 71S 73
Thrinax radiata ©1C 71S 73
Crinum amabile 71C
Hymenocallis littoralis 71C
Brassavola nodosa 61C
ERCUSMS pen Oley /AlC) 7/3
Coccoloba uvifera 61S 71C 73
Neea choriophylla 61C 71C 73
Sesuvium portulacastrum O61C
TCS 73
Chrysobalanus icaco 71C 73
Mucuna sp. 71S

Bumelia retusa 71C 73
Pouteria campechiana 71C 73
Cordia sebestena 61C 71C 73
Tournefortia gnaphalodes 61S
CaS
Stachytarpheta jamaicensis 73
Pithecellobium keyense 73
Suriana maritima 618
Bursera simaruba 61S 71C 73
Euphorbia blodgettii ©61C 71C
Euphorbia mesembrianthemifolia
Ge WAS 7/3
Euphorbia sp. 61S 73
Passiflora suberosa ©1C 71C 73
Conocarpus erectus O61C 71C 73
Laguncularia racemosa 73
Erithalis fruticosa 61C 71C 73
Ernodea littoralis 71C
Ambrosia hispida 6©1C 71C 73
Borrichia arborescens 71C
Wedelia trilobata 61C 71C 73

Long Cay North (16°45'N, 87°46'W) Figures 6-7, Plates 3-4

This is a small shingle cay on a detached reef patch in the main
channel between Northeast Cay and Long Cay; it was so named by
StoddaresG@oG2e ppruce Co) mieaG. 40)\achough)) Dahih ectval 2) (197/45 Fon) 56)
call it Little Cay and Fosberg et al.(1981) Small Cay. In 1961 it was
mainly composed of sand with some shingle in the central part and along
the east shore; it formed an island 145 m long and up to 48 m wide,
with an area of 0.44 ha. It was probably not more than 1 m above sea-
level. It was covered with a scrub of Suriana maritima 3 m tall with
Borrichia, Tournefortia and Conocarpus and a ground cover of Sesuvium,
Euphorbia and grasses. There were a few juvenile coconuts. By IES) 7b
there had been some erosion on the south shore and considerable
accretion on the north, and the greater part of the island was formed
of shingle. The position of the group of coconuts on the south shore
enables the relative locations to be plotted accurately. The

vegetation is still dominated by Suriana scrub, but Conocarpus and

Tournefortia have greatly increased.
coconuts.

There were several more juvenile
The length of the island remained at 145 m, but the maximum

width had increased to 63 m and the area of 0.53 ha.

22 species of plants have been recorded from the island (9 in 1961,

VeEsain Wy vandwk9s by) Linhartaampel 7s)

Sophora tomentosa, which

existed as a single plant on the south shore in 1961, had disappeared

Dygetowys The species recorded are:
Paspalum distichum 71C 73
Sporobolus virginicus 71C 73
Cyperus ligularis 73
Cyperus planifolius 71C 73
Cyperus sp. 61S

*Cocos nucifera 61S 71S 73
Crinum amabile 71C
Hippeastrum puniceum 73
Hymenocallis littoralis 73

*Casuarina equisetifolia 71S 73
Coccoloba uvifera 73
Batis maritima 73

Long Cay (16°45'N, 87°46'W)

Philoxerus vermicularis 71C

Sesuvium portulacastrum 61S 71C 73

Sophora tomentosa 61S

Suriana maritima 61S 71C 73

Euphorbia blodgettii 71C

Euphorbia mesembrianthemifolia
TGS

Euphorbia sp. 61S 73

Conocarpus erectus 61S 71C 73

Ipomoea macrantha 71C 73

Ipomoea pes-caprae 71C 73

Tournefortia gnaphalodes 61S 71C 73

Borrichia arborescens 61S 71C 73

Figures 8-9, Plates 5, 7, 8, 9 and1lO

Long Cay is situated at the northeastern extremity of the

continuous southeast reef of the atoll.

When first mapped in 1961 it

was 620 m long and varied in width from 70 m in the centre to 150 m at

each end;
peninsula.

its eastern end was prolonged lagoonward by a low sandy

The area of the cay was 6.69 ha.
island was formed by a massive double shingle ridge.
reached up to 2 m above sea level and the inner ridge 3 m;
complex was 30-40 m along the whole seaward shore.

The seaward side of the
The outer ridge
the shingle
The inner ridge was

formed of older and coarser material, with blocks up to 0.6 m in

diameter.

There was an abrupt junction between the inner margin of

the shingle and the low-lying sandy surface of the rest of the cay,
especially at the eastern end where there was a depression with

standing water.
bay carpeted with sea-grasses.
quite considerably by 1971.

The north shore is low and sandy and faces a shallow
The morphology of the cay had changed
The seaward shingle ridge had been

reworked, especially at its western end where it had been pushed back

over the sand surface and had become a triple ridge.

Retreat of a few

metres had occurred along much of the seaward shore, increasing the

length of a beachrock ridge exposed from 180 to 360 m.

The greatest

change, however, had taken place at the easterm end of the cay, where
the shoreline had retreated from 15 to 25 m along its entire length.
The northeasterm peninsula had also narrowed and moved westwards. As
a result the total length of the island had decreased to 600 m, and the

width now varied from 65-130 m;

the area in 1971 was 5.85 ha.

In 1961 Long Cay was covered with a coconut plantation. The
island was inhabited, and the ground beneath the coconuts was kept
cleared. Most of the seaward shingle ridges were bare, except for
scattered Tournefortia and Coccoloba, and an extensive area at the
eastern end of Hymenocallis with Conocarpus. The northeastern
peninsula was covered with a low scrub of Suriana, Conocarpus,
Tournefortia and Laguncularia, and indeed resembled Long Cay North.
The general aspect of the cay was much the same in 1971. The extensive &
ridge-crest area of Hymenocallis remained,: but all the Coccoloba and
Tournefortia of the seaward coast had disappeared. The northeastern
peninsula was covered with Tournefortia, Laguncularia, Conocarpus and
Sesuvium.

In 1971 the mudhole area was found to be partly surrounded by a
surface outcrop of phosphate rock, previously overlooked. This has a
Po0Os5 content of 21.7% (Scoffin and Stoddart, in prep.). Long Cay is
no longer a bird island. Salvin, who visited it on 12 May 1862, found
a few terms and some pelicans (Salvin 1864, p. 384; Coues 1864). The
date of formation of the phosphate rock is not known. The physiographic
changes between 1961 and 1971 must be attributed to hurricanes.
Possibly much of the coastal change resulted from Hurricane Hattie in
1961. Antonius (1972) was on Long Cay at the time of Hurricane Laura
on 20 November 1971. This had winds of 35 m/sec, for a time reaching
more than 50 m/sec, but although some coconut palms were felled the
effects were negligible. Long Cay was occupied by a local family in
1961 to look after the plantation. By 1971 an American-owned tourist
resort had opened on the cay, with a series of holiday chalets at the
east end. These were damaged in Hurricane Laura.

28 plant species have been recorded from the cay (10 in 1961, 21
in 1971, and 22 by Linhart in 1973); in addition there are some
collections made by Pippen in 1972. One species found in 1961 (Sophora
tomentosa) has not been seen since. In addition to the species listed
below, the sea-grasses Thalassia testudinum and Syringodium filiforme
occur in shallow water on the lagoonward side of the cay. In the
following list species marked 72 were collected by Pippen in that year. €
For further notes on this island, see Stoddart (1962, pp. 89-91, figs.
41, 42 and 44).

Psilotum nudum 72 Ficus sp. 73

Acrostichum aureum 73

Spartina patens 71C

Sporobolus virginicus 71C 72
13,

Cyperus ligularis 73

Cyperus planifolius 71C 72

Cyperus sp. 61S

Fimbristylis cymosa 71C 72 73

Cocos nucifera 61S 71S 73

Thrinax radiata 71C 73

Hymenocallis littoralis 618
TAC

Coccoloba uvifera 61S 71C 73

Sesuvium portulacastrum 71C 73

Mucuna sp. 71C

Sophora tomentosa 61S

Suriana maritima 61S 71C 73

Euphorbia blodgettii 73

Euphorbia mesembrianthemifolia
Uae 78

Euphorbia sp. 61S 73

Rhizophora mangle 73

Conocarpus erectus 61S 71C 72 73

Bumelia retusa 71C 72 73

10

Cordia sebestena 71C 73

Tournefortia gnaphalodes 61S
Wke. 73}

brichalissErueicosan iG, W738

Middle Cay (16°44'N, 87°48'W)

reef of Glover's Reef.
the trend of the reef.
and up to 175 m wide, with an area of 6.03 ha.

Ageratum littorale 61C 71C 73

Ageratum (abnormal) littorale
TAKE

Borrichia arborescens 71C 72 73

Melanthera nivea 71C

Figures 10-11, Plates ‘11-13

Middle Cay is located near the centre of the unbroken southeast

It is aligned northeast-southwest, parallel to
When mapped in 1961 it was about 415 m long

In 1971 erosion at the

southwest point had been counterbalanced by aggradation at the northeast
spit, and the overall dimensions and area (5.97 ha) remained the same.
The seaward shore is formed by a shingle ridge up to 35 m wide and 2 m
beachrock outcrops at the foot of this ridge, more extensively

high;
in 1961 than in 1971.

Most of the rest of the cay is low-lying and

sandy, with a low leeward sand beach, but at the southwestern end there
is an extensive swamp area with standing water about 170 m long and

45 m wide.

In 1961 this was separated from the sea by a belt of tall

Rhizophora, but by 1971 this had been greatly reduced and the shore

showed signs of erosion.

The sand spit at the northeast end of the cay

had expanded lagoonward during this interval and increased in size.

Most of the island is covered with coconut woodland.

In 1961 the

seaward shingle was largely bare, except for extensive Hymenocallis and

Tournefortia at the eastern end.

This had almost completely disappeared

by 1971, but Tournefortia had colonised afresh along most of the seaward

shore.

There had also been much growth of Laguncularia, Conocarpus,

Rhizophora and Borrichia along the lagoon shore and on the northeast

spit.

There is a luxuriant ground cover beneath the coconuts.

These

were the dominant vegetation as early as 1858 when Maury found a
"continuous grove of cocoa palms" and "entire freedom from underbush"

(Doubleday 1886, p. 212).

That the swampy area was also then in

existence is shown by Salvin's record of Acrostichum aureum in May 1862
(Salvin 1864).

1961, has not been seen since.

42 plant species have been recorded from the cay (11 in 1961, 40
invelL97A), Vande32 by -Linhart 1nwug7s)*.

Acrostichum aureum 1862S

(le ag s}
Andropogon glomeratus 71C 73
Cenchrus incertus 71C
Eragrostis prolifera 71C 73
Eustachys petraea 71C 73
Paspalum distichum 71C 73
Paspalum laxum 71C
Sporobolus virginicus 71C
Cyperus ligularis 71C 73
Cyperus planifolius 71C 73
Fimbristylis cymosa 61C 71C 73

Hibiscus tiliaceus,
Plants recorded are:

collected in

*Cocos nucifera 61S 71S 73
Thrinax radiata 71C
Hymenocallis littoralis 61S

C73
Ficus’ sp." \Jles+73
Coccoloba uvifera 71C 73
Neea choriophylla 71C

Sesuvium portulacastrum 71C
Pithecellobium keyense 71C 73
Sophora tomentosa 71C 73
Suriana maritima 71C 73
Euphorbia blodgettii 71C

Ea

Euphorbia mesembrianthemifolia Tournefortia gnaphalodes 61S

TMC 13 We. V3
Euphorbia sp. 61S 73 Stachytarpheta jamaicensis
Hibiscus tiliaceus 61C SILS GAUC. 7/3}
Passiflora suberosa: 71C 73 Diodia serrulata 71C
Rhizophora mangle 61S 71C 73 Hel Ehalws = arueTcosa (iG. 73
Conocarpus erectus 61S 71C 73 Ernodea littoralis 71C 73
Laguncularia racemosa 71C 73 Ageratum littorale 71C
*Terminalia catappa 71C Ageratum maritimum 73
Bumelia retusa 71C 73 Ambrosia hispida 61S 71C 73
Pouteria campechiana 71C 73 Borrichia arborescens 61S 71C 73
Cordia sebestena 71C 73 Wedelia trilobata 71C 73

There is in addition the casual record in 1858, already mentioned, of
Dioscorea sp., Musa sp., and Artocarpus altilis (Doubleday 1886,
Do BIZ) o

For further details of Middle Cay, see Stoddart (1962, pp. 92-93;
figs. 43-45).

Southwest Cay I (16°42'N, 87°49%4'W) Figures 12-13, Plates 14, 15 and 17

Southwest Cays are located at the southwest extremity of the
southeast reef of the atoll, immediately east of the broad southern
entrance to the lagoon. Southwest Cay I is the easterly of the two.

It is a trapezoidal island with maximum dimensions in 1961 of

SCORx, 27Omm: The seaward shore is formed by a low carpet of rubble and
shingle rather than by a distinct shingle ridge, as on the other cays,
though there was a sector of shingle ridge up to 2 m high at the southern
point of the cay. This seaward rubble zone is backed by a zone of
Swamp and standing water measuring some 230 x 130 m at the eastern end
of the island, with a second smaller zone at the western end. Only in
the lee of the shingle ridge is the land surface continuous to the lee
shore. The lagoon shore is low, sandy, and in 1961 was undercut, with
a number of fallen coconuts. By 1971 there were only minor
physiographic changes. There was some rearrangement of rubble on the
seaward side, and the seaward shingle ridge was much smaller and eroded.
The gross dimensions of the island remained the same, and the area was
only slightly smaller (8.70 ha compared with 9.02 ha).

The vegetation of the island consists of coconut woodland and
mangroves. The western shore has a stand of tall Rhizophora closing
the entrance to the western marsh. Rhizophora was also found at the
eastern marsh entrance, together with Avicennia, and there was a large
clump of Avicennia on the seaward shore. The larger marsh was fringed
with Conocarpus and other shrubs, and many coconuts had recently been
planted in it. Changes by 1971 were relatively minor. Much of the
mangrove at the entrance to the larger marsh had disappeared, as had
most of the Avicennia on the seaward shore. There had, however, been
extensive colonisation of the shingle ridge by Cordia at the south
point.

2,

40 species of plants are recorded from the cay, and this total
doubtless reflects the diversity of habitat on the island. 16 were
recorded in 1961, 24 in 1971 and 34 by Linhart in 1973: Of the species
recorded in 1961 only Euphorbia trichotoma has not been seen since, but
this suggests oversight rather than extinction. In particular the
island was less thoroughly collected, except for the west end, than
the other Glover's Reef cays. Species in the following list marked
72 were collected by Pippen in 1972. In the following list Ageratum
maritimum must be considered a dubious record, probably a
misidentification of A. littorale.

Andropogon glomeratus 73
Cenchrus incertus 71C 73
Eragrostis prolifera 71C 73
Eustachys petraea 71C 73
Paspalum distichum 71C 73
Paspalum laxum 73

Spartina patens 71C
Sporobolus sp. 61S

Cyperus ligularis 73

Cyperus planifolius 73
Cyperus sp. 61S

Fimbristylis cymosa 73
Fimbristylis sp. 61S

*Cocos nucifera 61S 71S 73
Hymenocallis littoralis 71S 73
Myrica cerifera 73

Coccoloba uvifera 61S 71S 73
Batis maritima 71C 73
Tresine diffusa,61C 72 73
Philoxerus vermicularis 73
Portulaca oleracea 718
Sesuvium portulacastrum 61S 73
Suriana maritima 61S 71S 73

Euphorbia blodgettii 72
Euphorbia mesembrianthemifolia
hes
Euphorbia trichotoma 61C
Euphorbia sp. 61S 73
Passiflora suberosa 71C 73
Rhizophora mangle 61S 71S 73
Conocarpus erectus 61S 71S 73
Laguncularia racemosa 71C
Rhabdadenia biflora 73
Ipomoea pes-caprae 61S 73
Cordia sebestena 61S 71C 73
Tournefortia gnaphalodes 61S
TAC oH 3
Avicennia germinans 61S 71C 73
Stachytarpheta jamaicensis
(ere 7 is}
Capraria biflora 71C 73
Erithalis fruticosa 71C 73
Ageratum littorale 71C
Ageratum maritimum 73
Borrichia arborescens 71C 73
Eclipta alba 73
Wedelia trilobata 71S 73

For further notes on this island, see Stoddart (1962, pp. 93-95, figs.
46-47).

Figures 14-15, Plates 6,
16 and 18

Southwest Cay II (16°41%5'N, 87°50'W)

Southwest Cay II is the most southerly island on Glover's Reef.
Although immediately adjacent to Southwest Cay I it is of very different
character. It is aligned at an angle of 45° to the reef edge, and
shingle is only found along the southem shore close to the reef. The
rest of the island is sandy, and there is no interior depression or
mangrove Swamp. When mapped in 1961 the island was 560 m long and
had a maximum width of 235 m; its area was 6.66 ha. The narrow
southerm shingle ridge reached about 1.5 m above sea level. Much of
the eastern shore was low and sandy, with many Rhizophora seedlings,
in spite of being on the seaward side of the island; it faced a broad

i

shallow reef flat connecting with Southwest Cay I. The western shore
is also sandy, but narrow and undercut: in its northernpart it was
lined with coconut boles and fallen trunks. There were several
exposures of relict beachrock on this western shore, indicating both
beach retreat and re-alignment as the shoreline moved in a clockwise
direction. There was further evidence of shore erosion at the
southwest point, though masked in 1961 by a fresh sandspit. Beach
retreat was likewise evident on the east shore, as shown by undercut
coconuts and small patches of beachrock which unusually showed landward
dip. Physiographic changes had been minor by 1971. There had been
erosion round the southern part of the cay, and the southerm shingle
ridge was more extensive; there had also been erosion at the northern
end. As a result the overall length of the cay had decreased to 540 m
and the area to 6.23 ha. The previous beachrock outcrops were still
visible, together with another on the south shore.

The island is entirely covered with coconut woodland. Except in
the southern part this has an undergrowth of herbs and grasses. Apart
from occasional Conocarpus and Suriana bushes the only coastal
vegetation is a narrow belt of Rhizophora on part of the east shore.

40 species of plants are recorded from the island (12 in 1961, 37 in
1971, and 31 by Linhart in 1973), a reflection of the diversity of
the ground cover under the coconut woodland. They are:

Andropogon glomeratus 61S 71C Canavalia rosea 71C 73

Cenchrus incertus 71C 73 Suriana maritima 71C 73
Eragrostis prolifera 71C Euphorbia blodgettii 61C 71C
Eustachys petraea 71C 73 Euphorbia mesembrianthemifolia
Paspalum distichum 71C Teas}
Paspalum laxum 71C Euphorbia sp. 61S 73
Spartina patens 71C Passiflora suberosa 71C 73
Spartina spartinae 73 Rhizophora mangle 61S 71C 73
Sporobolus virginicus 71C 73 Conocarpus erectus 61S 71C 73
Sporobolus sp. 61S Terminalia catappa 71C
Cyperus ligularis 73 Ipomoea macrantha 71C 73
Cyperus planifolius 71C 73 Ipomoea pes-caprae 71C 73
Fimbristylis cymosa 71C 73 Ipomoea stolonifera 71C 73
*Cocos nucifera 61S 71S 73 Ipomoea sp. 61S
Hymenocallis littoralis 61S 71C 73 Tournefortia gnaphalodes 71C 73
Coccoloba uvifera 71C Stachytarpheta jamaicensis
Batis maritima 73 Cle iG Hs
Iresine diffusa 71C 73 Capraria biflora 71C 73
Rivina humilis 71C 73 Erithalis fruticosa 71C 73
Portulaca oleracea 71C 73 Ernodea littoralis 71C 73
Sesuvium portulacastrum Morinda citrifolia 71C

GS Ears Ageratum littorale 71C 73
Cakile lanceolata 61S 71C Wedelia trilobata 71C 73

Salvin visited this island on 12 May 1862. He found "noddies
everywhere ... many thousands in all", and speculated "what must the
numbers have been when the Sooty Terns flocked to the same island"
(Salvin 1864, p. 383). Coues (1864) listed Anous stolidus (on coconuts)

14

and Anous tenuirostris (on mangroves). The brown noddies still inhabit
the cay but in small numbers; there are no records of Sooty Terns
breeding there. For further details, see Stoddart (1962, pp. 95-98,
Lilgee48) a

SUMMARY AND CONCLUSIONS

These surveys have established the variability of reef islands
even over so short a period as a decade. All of the larger islands
have decreased in area (Table 2), with the possible exception of Middle
Cay, and these changes have been mainly effected by erosion and
remodelling of windward coasts during major storms. Change has been
most severe at the eastern end of Long Cay, but can be traced on all
the islands in changed shore morphology, distribution of windward
shingle ridges, and exposure of beachrock. Leeward or lagoon shores,
on the other hand, have been more stable, and in some cases leeward
sand spits have increased in area (Long Cay, Middle Cay). The islet
of Long Cay North is the only one to have increased markedly in size,
but it is so small that the change could readily be reversed.

Our data do not allow inferences about colonisation of the cays by
new plant species between the surveys, nor do we wish to discuss the
floristic biogeography of the islands in this paper. We can, however,
record the following local extinctions: Brassavola nodosa and Suriana
maritima on Northeast Cay; Hibiscus tiliaceus on Middle Cay; and
Sophora tomentosa on Long Cay and Long Cay North. All were represented
in 1961 by single specimens. Brassavola is an uncommon epiphyte; the
Hibiscus was a Single lagoon-shore tree; Sophora is a beach legume
liable to substrate disturbance. This last effect is also seen in
the widespread disappearance of Tournefortia and Coccoloba on the
windward shingle ridges of Long Cay between 1961 and 1971, as a result
of topographic change; they did not become locally extinct only because
they existed in other habitats on the cay.

The total recorded terrestrial flora of Glover's Reef now stands
at 70 species. Of these, 30 percent (21 species) occur on only one
island. Taking the species recorded from all the Belizean atolls
(Lighthouse Reef, Turneffe, Glover's Reef), 28 species or 40 per cent of
the Glover's Reef flora are recorded only from that atoll: this
undoubtedly reflects the state of collecting on the other atolls rather
than any local peculiarity on Glover's Reef. There is a simple
relationship between number of plant species and area of the cays,
though the anomalously low figure of species for Long Cay indicates the
effect of coconut planting and clearing on that island. The species
diversity figures for the Glover's Reef cays are markedly higher (by a
factor of two) than those of other Caribbean and Gulf of Mexico reef
island groups, such as the Dry Tortugas, Alacran, the Pedro Cays, and
the Morant Cays. This probably results from the much wetter
environment of Glover's Reef rather than from differing proximity of
islands to source areas. The floristic biogeography of western
Atlantic reef islands will form a separate paper; that of the Belize
cays in general has been examined by Stoddart and Fosberg (in press).

15

Table 2. Change on the cays 1961-1971

Antonius, A. 1972. Hurricane Laura, witnessed in British Honduras.
AVEOIIL IRIS 5 MoulIs U6AS) Ikelas

Boone, L. 1927. Scientific results of the First Oceanographic
Expedition of the "Pawnee" 1925. Crustacea from tropical east
American seas. Bull. Bingham oceanogr. Coll. 1(2): 1-147.

Island Area in hectares Percentage Species of Total number
1961 ALS) 7/ aL change land plants of species of
according land plants

to) Milman te recorded Bs
(1980) \-
Northeast 4.73 Ake 552 3 32 40 .
Long North 0.44 Oo 53} T2ORS Ig) 22 ms:
Long 6.69 Sy ots5) —12 6 Dad 28 Ral
Middle 6.03 D697 IL 5© B2 42 13
Southwest I 52 5 HO 2365 35 40 pee
Southwest II 6.66 6523 -7.1 SL 40 HD
Total 33.57 31.76 -5.39 59 70 34
REFERENCES ul

Boone, L. 1928a. Scientific results of the First Oceanographic
Expedition of the "Pawnee" 1925, Mollusca from tropical east
American seas. Bull. Bingham oceanogr. Coll. 1(3): 1-20.

Boone, L. 1928b. Scientific results of the First Oceanographic
Expedition of the "Pawnee" 1925. Echinodermata from tropical
east American seas. Bulg Bingham oceanogr. Coll. 1(4)i:, 1-22. g

Boone, L. 1928c. Scientific results of the First Oceanographic
Expedition of the "Pawnee" 1925. Coelenterata from tropical east
American seas. istulil , dellinepaevas Ceeeiaocpas (Goulils Ik (S)) 39 yale

Breder, C. M. Jr. 1927. Scientific results of the First Oceanographic
Expedition of the "Pawnee" 1925. Fishes. Bull. Bingham
oceanogr. Coll. (ih) = 1-90.

Coues, E. 1864. Notes on certain Central-American Laridae collected
by Mr Osbert Salvin and Mr F. Godman. Mong, (i), Os SI7os) sie

Cran A Com IOI) 1 GSOGraphy OE LShing) im Eaistes sh HOongunas) and
adjacent coastal areas. Louisiana State University, Coastal
Studies Institute, Technical Report 28, i-xv, 1-143.

16

Dahl, A. L., Macintyre, I. G. and Antonius, A. 1974. A comparative
study of coral reef research sites. Atoll Res. (Bits Ma] 25 Si
T2ON

Devaney, D. M. 1974. Shallow-water echinoderms from British Honduras,
with a description of a new species of Ophiocoma (Ophiuroidea).
Bull. mar. Sci. 243. 922-64

Dillon, W. P. and Vedder, J. G. 1972. Structure and development of
the British Honduras continental margin. Earth plan. Sci. Lett.
AB eel S/S SIO)e

Dillon, W. P. and Vedder, J. G. 1973. Structure and development of
the continental margin of British Honduras. Geol. Soc. Am. Bull.
sion. DHSS AW SF «

Doubleday, C. W. 1886. Reminiscences of the "Filibuster" War in
Nicaragua. New York and London: G. P. Putnam's Sons. ix,
22> E's

Fosberg, F. R., Stoddart, D. R., Sachet, M.—H.,.and. Spellman) Deyeae
1981. Plants of the Belize cays. Atoll Res. Bull. 258.

Haas, F. 1941. Marine shells from the Mandel Caribbean Expedition.
Publis Field Muse nabs HtSt. , ZOOM. Sens 2a tsi yiar

Hughes, S. M. 1973. Foraminifera and sediments of patch reefs,
Glover's Reef, British Honduras. University of Montana, senior
thesis. 36pp. [not seen].

Jefferys, T. 1775. The Bay of Honduras. In: The West India Atlas
or a general description of the West Indies. London.

Linhart, Y. B. 1980. Local biogeography of plants on a Caribbean
atoll. TesBLOgeogra, H2= E59—N 7 iN

Maury, H. 1859. Loss of the Susan. The Mobile Daily Register, 32
(L329) ep 1s:

Sachet, M.-H. and Dahl, A. L., eds. 1974. Comparative investigations
of Tropical Reef Ecosystems: background for an integrated coral
reef program. AGO] LD eRESR a Bilal ope] Zn liGOr.

Salvin, O. 1864. A fortnight amongst the sea-birds of British
Honduras. PDUs (1) "6s Si 2-367).

Schafersman, S D. 1972. Carbonate sediments and Foraminifera of
patch reefs, Glover's Reef, British Honduras. Northern Illinois
University, Master's thesis. 99 pp. [not seen].

Schmidt, K. P. 1941. The amphibians and reptiles of British Honduras.
Rield= Mus. nat. HiSt.p, ZOOL. Sere 228) 2" 475-510.

Scroggs, W. O. 1916. Filibusters and financiers: the story of
William Walker and his associates. New York: Macmillan. xii,
408 pp. Reprinted (1969) by Russell and Russell.

IY)

Speer, J. S. 1771. The West-India Pilot. Containing piloting
directions ... from Jamaica to Black River on the Mosquito Shore,
and from thence to every Bay, Harbour, River, &c. in the Bay of
Honduras. ... By an Officer who has served upwards of Twenty Years
in the West Indies. . Second Edition. London.

Stoddart, D. R. 1962. Three Caribbean atolls: Turneffe Islands,
Lighthouse Reef, and Glover's Reef, British Honduras. Atoll Res.
xo {37/94 ISIS AL 6

Stoddart, D. R. and Fosberg, F. R. In press. Species-area
relationships on small islands: floristic data from Belize sand
cays, western Caribbean. Smiehson.| Contr. mar. Sci. oe 1980)

Todd, W. E. C. 1942. List of the hummingbirds in the collection of
the Carnegie Museum. Annis Carnegie Mus. 29: 271-370.

Tsuda, R. T. and Dawes, C. J. 1974. Preliminary checklist of the
marine benthic plants from Glover's Reef, British Honduras.
ACOs Res, Bull. 17S als

Uchupi, E. 1973. Eastern Yucatan continental margin and western
Caribbean tectonics. Am. ASS. petrol: Geol. Bull. 57: 1O075=
1085.

Vindel, F. 1955. Mapas de America en los libros Espanoles de los
siglos XVI al XVIII (1503-1798). Madrid.

Wallace, R. J. 1974. A reconnaissance of the sedimentology and
ecology of Glover's Reef atoll, Belize (British Honduras).
Princeton University, Ph.D. thesis. 140 pp. [not seen].

Wallace, R. J. and Schafersman, S. D. 1977. Patch-reef ecology and
sedimentology of Glover's Reef Atoll, Belize. Am. Assoc. petrol.
Geol. Stud. Geol. 4: 37-52.

York, M. E. 1971. Patch reef coral communities of southern British
Honduras and illustrated catalogue of common British Honduras
corals. In K. F. Wantland and W. C. Pusey III, eds.: A guidebook
for the field trip to the southern shelf of British Honduras,
October 10-13, 1971 (New Orleans: New Orleans Geological Society),
Appendix 1.

nears

Glovers//
Reef /’

) ae
9 6 5 :
rt B lame

ay
=

Figure l. Location and submarine topography of Glover's Reef. After
Dillon and Vedder (1973)

87°45’

87° 50'

kilometres

ss

16°50’

16°45)

16°45'

Peripheral reef

MIDDLE CAY

Inner edge of low platform

Larger reef knolls

Patch reef

°
Oo
Qo

87°45'

3Y/)

IEaL Oye

From Stoddart (1962,

Topography of Glover's Reef.

2s

Figure

Pie GolGAle

Oo mata Uibbile
Shingle

Sand

Fresh sandspit
Undercut shoreline
Beach ridge-crest

Beachrock

NAN NANNY

SAQAQQ0a0y»y, Phosphate rock

NANNNAANAN

Figure 3. Legend to the detailed physical and vegetation maps of the

cays

VEGETATION

limees
c ||| Coconut; Coconut woodland (dense, open)

Q Rhizophora

-O- Avicennia

Laguncularia

A Coccoloba

fo} Cordia

M Morinda

F Riles

T Terminalia

8 Bursera

Shrubs

Tournefortia
Conocarpus

> Suriana
Borrichia
Sophora

x Indeterminate

H

ii) Hymenocallis
Sesuvium

=a Heérpssahnd grasses

ae: -—Recent undercutting

1961

Large stranded
boulder

6. A
7,
° : Se 0 i oS 3 . x é phe ORE te DP aaa ; \
NEN hae BAe . t pee Bee c 7 File 5 eg aoa bahay os Oe 3) °
1971 Coo : : Ett ys eeu ere Hes ; 0 a 5 Pai 5 Rp ee Sle
° . . . 5 . AS 2 o ond Uc 5 4
© .
° ake
° .
ope S
° °
° ° 3 Z,
; °
G °
© °
°
oO oN é A |
9 ie)
° . i . . °
NS . . fe) °
New Beachrock ~ = Oe 2
Pe) ° ° °
) ° ° °

° °

SSIS
<I

O metres 100 IS
[Siar tal See ae ew SS]

Figure 4. Northeast Cay 1961 and 1971: topography

S
RK

OP
dy SY
RIAA
SRRRPH

1961

B)

C5) Var TTT

1971

os
Ss
HI
so

OO

1

Northeast Cay 1961 and 1971: vegetation

Figure 5.

1961

1971
O metres SO)
oS ele ee
Figure 6. Long Cay North 1961 and 1971:

topography

Figure 7. Long Cay North 1961 and 1971: vegetation

Figure 8.

Longe Cayz IG yand: OVAL:

topography

Pool

47

YOIAA7
4/474
°

4
\

O metres 100
|

S SS
MESSI SS

O
me
|

Figur
en)
5 ‘ae
gC
ay 1961 and 1
971:
: vegeta
tion

INV EIVWOUTIAIIIYD

Ar See oe a

—

Figure 10.

aime ane Soo oO
( Shaliow\: .* Re
Nas pit \ 5 *o
> .
~
~ i
Se o
NOOSE
Q\

Mitcdilies Cay gtIG1 “ang e197 1h:

: Sie

Be 9 fie: S a
eg ic 6 :
9G) ° 2
s Og ~
; : ° 5
ate h c

: : of
ACN is fs 5
Brora ets a ° S Se
N -
ee. 6 a fs 5
2 Oo 0 5 P
5 Oy ee ° z
: y/eraorie ia
: Jo ° D>)
: 5 )
: y 2
; OK. E
v 6 S -
5
)

topography

O metres 100
| |

eo ee el et

wo owe wes

eh ha

eevwwoesewoewst

ezruek eb 6 Outer

a a aS |

3
Q=
=a
S)
ce”
“,
3

/

O metres 100

Middle Cay 1961 and 1971: vegetation

Higure) J...

Figure 12.

Southwest Cay I

UO GH eranceslOy/As:

topography

O metres 100
[Sebesoaestsuriat aca |

ww

INV EINWOTILITUD

ee ee ee

tTWOTTTUTIUIN

QIVEDDTIOUWIVIFIN

ew eb tr wrt tek tO uD wo tee LIVANNIOYD

ae Nowe

O metres 100
{ee

Figure 13. Southwest Cay I 1961 and 1971: vegetation

= > ee
enmiuvudain 1Innmranien ePaarrriaAar AAD omemwowier amas Ai7vAh a ae aoe Revere iervcrn st a a tAres Cag tM ee ws es Ge

— co CAAITUICAATIAAI PAIPewerseawrsmas AIMIDIMPtLI_ean API RIVCH DD PATE

Aydez6odo} :TL6T pue TI6T II ACD JAsemyANoS “Ppl omznbty

fF
OOL seujaw CO

uoTRzeReHSeA 6FTLET pue T96T IT ASD FsemuQNOS "SGT eanbdta

[ots ee
OOL Se4}jeW O

sow ws rere ww

Plate 1. Northeast Cay: aerial view from the east 1961

Plate 2. Northeast Cay: storm block and shore erosion on the east
side 1961

Plate 3. Long Cay North: Suriana scrub on the north shore 1961

Plate 4. Long Cay North: the south shore in 1971

ZI6T BSeSeUuaNosS su WOAF MSTA

Tetzee :jprT AeD AsemuQnos

WOTF META TeTrSe

Z96T FSeEUINOS 9A

:Kea Huot

"9 e2eTd

“G 93eTd

Pilate, 7: Long Cay: Tournefortia on the northeast spit 1961

Plate 8. Long Cay: upper surface of the windward shingle ridge 1961

L Be 2 : epee z Y 3 a *, = ete * _ t . Tee y Lay ; a = % ag = s

= : Search Bete a es Se Kt ecponatt : - =
Plate 9. Long Cay: sandy island surface with backslope of the
windward shingle ridge 1971

Plate 10. Long Cay: mud hole and standing water at the east end,
seen from the windward shingle ridge 1961

windward shingle ridge with hedge of

Middle Cay

Plate ll.

SUS) 7A

Tournefortia

low lagoon shore with Conocarpus 1971

°

Middle Cay

Plate 12.

Plate 13. Middle Cay: Rhizophora at the east end of the lagoon
shore 1971

Plate 14. Southwest Cay I: seaward rubble shore 1971

mangrove pool at the southern end of the

Southwest Cay I:

Plate 15

island 1961

on the retreating

exposed 'rootrock'

.

Southwest Cay II

Plate 16.

lagoon shore 1971

PAeMPULT UTM Yyooryoreq

T96T SAOUS pAeMedS 9YyQ UO dtp
?T~ AeDd qsemyqnoS “gT e2eTd

T96T eAoczhuew usdo

>~T AeDd Asemyanos

‘Pe

ee

“LT 938Td

ATOLL RESEARCH BULLETIN |
No. 258 !

PLANTS OF THE BELIZE CAYS
by

F.R. Fosberg, D. R. Stoddart, Marie-Héléne Sachet, and
David L. Spellman

Issued by
THE SMITHSONIAN INSTITUTION
Washington, D. C., U.S.A.
April 1982

CONTENTS

Abstract

Introduction

Systematic list of plants
Literature cited

Index

List of cays cited

page

73

76

PLANTS OF THE BELIZE CAYS

lONi7 IP SIE Fosberg, ! D.R. Stoddart, 2 Marie-Héléne Sachet, | and
David L. Spellman 3

ABSTRACT

All of the vascular plants known to us from the sand and mangrove
cays off the coast of Belize, Central America, are listed, with
citations of specimens, sight records, and literature records. 178

species of land plants in 145 genera are listed, together with 4 species

and 4 genera of sea-grasses. Citations are by individual islets or
cays. These are classified into atoll cays, barrier reef cays, and
lagoon cays.

INTRODUCTION

This is an annotated checklist of the vascular plants of the small
islands off the coast of Belize, formerly British Honduras, Central
America, aS known to date. It is based principally on specimens and
observations collected during a series of investigations on these
Po Wandsmby mocoddarcein 959 I6O; i IOl el OG2 mlIGS,u LO7d andy 1972-8 by,
Sachet inidlo7- = bys hosbexg iin, 1971) and) 1972." and by Spellman in 19/2:
In addition specimens are cited collected by Dr John Dwyer and his
students in 1967 and 1980, and by R.W. Pippen in 1972, James Pringle in

1979-80, and by several other visitors to the area. Citations tof
literature records are also included, notably those listed by Linhart
(1980) based on a visit to Glover's Reef in 1973. When we have not

1 smithsonian Insti tucizon, Washingten, D.C., U.S.A;
2 Department of Geography, Cambridge University, England
Missouri Botanical Garden, St Louis, Missouri, U.S.A.
IMSWE (Investigations on Marine Shallow Water Ecosystems) Contribution
No. 76

Manuscript received January 1981 -- Eds.

Beta wiwi WW tINUOMLIFVOD V2

swt tas

crew

seen the specimens we have adjusted the nomenclature as best we could
to that based on actual specimens that we have seen. We cannot, of
course, vouch for the correctness of these identifications, or of our
interpretations of them. Nor can we vouch for the correctness of the
numerous sight records included, but since the flora is a small and
simple one, they are probably mostly reliable. The records cited from
literature refer to the papers listed at the end of this paper.

Entries cited with a generic name and sp. are mostly for literature
citations by genus only. In a few instances, e.g., "Ficus sp. probably
new", no binomial is available for the species, or as in the case of
Eugenia sp. our material is inadequate for determination. When we are
reasonably certain that species are of human introduction, their names
are preceded by an asterisk.

The herbarium symbols in parentheses after the specimen citations,
indicating where the specimens are deposited, are those in Index
Herbariorum, ed. 6, by Holmgren and Keuken, Regnum Vegetabile 92 (1974),
except for the symbol Fo, which indicates specimens still in possession
of the authors. These latter will mostly be deposited eventually in
the Pomona College Herbarium (POM).

In listing the species we have made every effort to use the names
that seem to us correct according to our ideas of the taxonomy of the
plants and following the International Code of Botanical Nomenclature.
We have cited few synonyms, only those that we know to have been used
in records of the plants of these cays or that are necessary to clarify
difficult applications of names. We hope that we have caught and
corrected all inconsistencies resulting from the many revisions of this
US E

We gratefully acknowledge the courtesy of R.W. Pippen and James
Pringle in showing us and permitting us to cite their collections from
the cays, and that of John Dwyer in giving us a list of his collections
from the area. We appreciate the hospitality of the Missouri
Botanical Garden and the privilege of examining specimens housed there.
Special thanks must go to Miss Dulcie A. Powell for her patient and
meticulous work in assembling and arranging the specimen citations.
Stoddart's initial investigations were supported by the British
Department of Scientific and Industrial Research, the Royal Society of
London, the Office of Naval Research, and the Coastal Studies Institute
of Louisiana State University. We also acknowledge the financial
support of the Smithsonian CITRE (Comparative Investigations of Tropical
Reef Ecosystems) Programme in 1971, and of the Smithsonian IMSWE
(Investigations of Marine Shallow Water Ecosystems) Programme in 1972,
of which this investigation is a part.

PSILOTACEAE
Psilotum Sw.
Psilotum nudum (L.) Beauv.
Atoll Cays. -- Glover's Reef: Long Cay, S. end, Pippen 907 (WMU),

"25 July '72, rare, 9 depauperate plants, sporulating on 2 burned out
decaying coconut stumps."

POLYPODIACEAE
Acrostichum L.

Acrostichum aureum L. (Mangrove fern, swamp fern)
Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg & Sachet
53849 (US), Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1631
CUS) sp asauavaine (GlS64) vine] 862/77 lamharrtm( L980), 1p scl'62)). Lighthouse Reef:

Half Moon Cay, Stoddart 48 (Fo).

Barrier Reef Cays. -- Northeast Sapodilla Cay, Stoddart sight
record in 1960, Spellman & Stoddart 2299 (US, MO, BH, Fo); Frank's
East Cay, Spellman & Stoddart 2420 (US, MO); Hunting Cay, Spellman &
Stoddart 2477 (US, MO).

Lagoon Cays. -- Cary Cay, Stoddart sight record in 1962 (1963,
PEO) aWeSicy Snake) Cayy, Stoddart 8) (Fo), (1965, pe 137).

Nephrolepis Schott

Nephrolepis multiflora (Roxb.) Jarrett ex Morton (Boston fern)
Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2330 (US,
MO, BH, Fo); Lime Cay, Spellman & Stoddart 2363 (US, MO, BH, Fo).

Polypodium L.
Polypodium lycopodioides L.

Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2332 (US,
MO, BH, Fo).

Polypodium polypodioides (L.) Watt var. polypodioides (Hard fern,
Resurrection fern)

Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2336 (US,
MO, BH, Fo).

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Pityrogramma Link

Pityrogramma calomelanos (L.) Link (Silver-backed fern)
Lagoon Cays. -- Wild Cane Cay, Stoddart 20 (Fo).
ZOSTERACEAE

Halodule Endl.
Halodule wrightii Aschers. (Small sea-grass)
Diplanthera Wrightii (Aschers.) Aschers.
Atoll Cays. -- Glover's Reef: Long Cay, Fosberg 53802. (US, MO,
BH, Fo, BM); southeast sector of lagoon, Tsuda 4256 (US) (Tsuda and
Dawes 1974).
Barrier Reef Cays. -- Tom Owen's East Cay, Spellman & Stoddart 2279

(US, MO); Frank's Cay, Spellman & Stoddart 2389 (US, MO); Nicolas Cay,
Spellman & Stoddart 2323 (US, MO).

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54454 (US);
Buttonwood Cay, Fosberg & Spellman 54427 (US); Hatchet Cay, Fosberg &
Spellman 54397 (US); Little Water Cay, Fosberg & Spellman 54355 (US).

Syringodium Kiitz.
Syringodium filiforme Kitz. (Eel grass, Manatee grass)

Cymodocea manatorum Aschers.

Atoll Cays. -- Glover's Reef: Long Cay, Fosberg & Sachet 53850
(US), Tsuda 4165 (US) (Tsuda and Dawes 1974); southeast sector of

lagoon, Tsuda 4257 (US) (Tsuda and Dawes 1974).

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2512 (US,
MO, BH); South Water Cay, Pringle 1920 (HAM, UNA), 1923 (HAM, MO,
UNA); Carrie Bow Cay, Fosberg & Spellman 54443 (US), Riitzler 3
(US) .

Lagoon Cays. -- Little Water Cay, Fosberg & Stoddart 54358 (US).

HYDROCHARITACEAE

Halophila Thouars
Halophila engelmanni Ascherson

Atoll Cays. -- Glover's Reef: southeast sector of lagoon, Tsuda
4258 (US) (Tsuda and Dawes 1974).

Thalassia K6nig
Thalassia testudinum Banks ex K6nig (Turtle-grass)

Atoll Cays. -- Turneffe Atoll: s.1. Burdon s.n. (K); Indian Cay,
Colman in 1937 (BM) (citations from den Hartog 1970, p. 226);
Crickozeen Creek, Stoddart sight record in 1962 (1963, p. 87); Harry
Jones, Stoddart sight record in 1960 (1962a, p. 48); Big Calabash Cay,
Stoddart sight record in 1960 (1962a, p. 42); Little Calabash Cay,
Stoddane sight record in 1960 and 1962) (1962a, p-) 41) 1963, pe 76)
Lighthouse Reef: Northern Cay, Stoddart sight records in 1960 and 1962
GiSo2a, jpraSo, 1968, pxe92);2 Sandbore Cay, Stoddart=sight record in
1962 (1963, p. 90); Saddle Cay, Stoddart sight record in 1960 (1962a,
Demos) Longe Cay), otoddart sight record in, 1960 (1962a;, pp. 79); “Halt
Moon Cay, Stoddart sight record in 1960. Glover's Reef: Northeast
Gay, Stoddart sight record in 1961 (1962a, p. 88); Long Cay, Stoddart
sight record in 1961 (1962a, p. 91), Tsuda 4164 (US) (Tsuda and Dawes
1974); Middle Cay, Stoddart sight record in 1961 (1962a, p. 93);
Southwest Cay I, Stoddart sight record in 1961 (1962a, p. 95); West
Channel, Fosberg 53907 (US); southeast sector of lagoon, Tsuda 4255
(US) (Tsuda and Dawes 1974).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight records in
1960 and 1962 (1963, p. 42); English Cay, Stoddart sight record in
1960 (1963, p. 45); Rendezvous Cay, Stoddart sight records in 1960 and
IIG2e ClIGSip 49) 77 Cay, Glory, Stoddart) sight record an 1960 (1963),
p. 53); South Water Cay, Stoddart sight records in 1960 and 1962 (1963,
p. 56), Pringle 1918 (HAM); Carrie Bow Cay, Stoddart sight record in
1960 (1962a, p. 164, 1963, p. 57), Fosberg & Spellman 54441 (US), 54435
(US); Curlew Cay, Stoddart sight record in 1960 (1963, p. 58); South
Water Cay, Stoddart sight records in 1960 and 1962 (1963, p. 56).

Lagoon Cays. -- Cay Chapel, Stoddart sight record in 1962 (1969,
p. 7); Little Water Cay, Fosberg & Spellman 54359 (US).

92

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GRAMINEAE

Andropogon L.

Andropogon glomeratus (Walt.) Britton, Sterns &Poggenburg (Broom-sedge)

Atoll Cays. -- Lighthouse Reef: Northern Cay, Stoddart sight
record in 1960 (1962a, p. 62); Sandbore Cay, Stoddart 553 (Fo) (1962a,
Dist Die IGS), spear Oe Glover's Reef: Middle Cay, Sachet & Stoddart
1621. (US; MO, BH, Fo), Linhart (1980, p. 162); Southwest Cay 1, amalmhbare

(1980, p. 162); Southwest Cay If, Steddart sight record ingle
(1962a, p. 98), Fosberg & Stoddart 53860 (US, MO, BH, Fo, BM).

Barrier Reef Cays. -- Goff's Cay, Spellman & Stoddart 2525 (US, MO,
BH, Fo); .South Water Cay,,-Stoddart. 92 (Fo) (1963;,.p.7 56) ,. Spel lmang&
Stoddart 2210 (US, MO, BH), Pringle 1919 (HAM); Frank's Cay, Spellman
& Stoddart 2391 (US, MO, BH); Nicolas Cay, Spellman & Stoddart 2342
(US, MO, BH, Fo); Lime Cay, Spellman & Stoddart 2376 (US, MO, BH, Fo).
Andropogon sp.

Barrier Reef Cays. -- Frank's East Cay, Stoddart sight record in
1961; Frank's West Cay, Stoddart sight record in 1961.

Lagoon Cays. -- South Snake Cay, Stoddart sight record in 1961
(1965; peeS8)<
Anthephora Schreb.
Anthephora hermaphrodita (L.) Kuntze
Barrier Reef Cays. -- South Water Cay, Pringle 1926 (HAM, MO);
Lime Cay, Spellman & Stoddart 2368 (US, MO BH).
Cenchrus 1).
Cenchrus incertus M. A. Curtis (Sand-burr)
Cenchrus pauciflorus Benth.
Atoll Cays. -- Lighthouse Reef: Northern Cay, Stoddart sight

record in 1962 (1963, p. 93,as Cenchrus); Sandbore Cay, Stoddart sight
record in 1962 (19637 p..91)) and 1965 "G@i9697 p. l7,as °C. tribuidordes)y

Half MoonsCay,. Stoddart 134 (Fo) (1963, sD. 98). Glover's Reef:
Northeast Cay, Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart
1625 (US); Southwest Cay I, Linhart (1980, p. 162); Southwest Cay II,

Fosberg & Sachet 53891 (US), Linhart (1980, p. 162); Southwest Cay,
Pippen 896 (US).

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2149
(US, MO, BH, Fo), Pringle 1906 (HAM); Lime Cay, Spellman & Stoddart
2364 (US, MO, BH, Fo).

Lagoon Cays. -- Cay Chapel, Stoddart sight records in 1962 (1963,
DiS )and 965.
*Cymbopogon Spreng.
*Cymbopogon citratus (DC) Stapf (Fever grass, Lemon grass)

Andropogon citratus DC.

Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54361 (US).

Digitaria Heist. ex Fabr.
Digitaria horizontalis Willd. (Crab grass)

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 133 (Fo).

Distichlis Raf.
Distichlis spicata (L.) Greene (Seashore saltgrass)
Barrier Reef Cays. -- South Water Cay, Pringle 1894 (HAM).

lagoon, Cays\. —— Ambergris) Cay,, West, (1977, p-. 198, Ege 9k 3)

*Eleusine Gaertn.

*Eleusine indica (L.) Gaertn. (Goose grass)
Barrier Reef Cays. -- Tobacco Cay, Stoddart 112 (Fo) (1963, p. 55),

Fosberg & Spellman 54222 (US); Hunting Cay, Spellman & Stoddart 2444

(US, MO, BH, Fo).

Eragrostis Wolf

EGAGLOS ET Si Ci naieiise a(liie)) kien (Bax. (Love grass)

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 547 (Fo),
136 U(Fo)miG9I62a,, sp. 76):

Barrier Reef Cays. -- Tobacco Cay, Stoddart 113 (Fo) (1963, p. 55);
South Water Cay, Spellman & Stoddart 2197 (US, MO, BH), Pringle 1895
(HAM); Hunting Cay, Spellman & Stoddart 2446 (US, MO, BH).

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Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54364 (US).
Eragrostis prolifera (Sw.) Steud. (Love grass)
Eragrostis domingensis (Pers.) Steud.

Atoll Cays. -- Lighthouse Reef: Long Cay, Stoddart 456 (Fo)
(1969, p. 18 as E. domingensis). Glover's Reef: Northeast Cay,
Stoddart 62 (Fo) (1962a, p. 88), Fosberg & Sachet 53818 (US), Linhart
(1980, p. 162); Middle Cay, Sachet & Stoddart 1619 (US), Linhart (1980,
p. 162); Southwest Cay I, Fosberg & Stoddart 53863 (US), Linhart (1980,
p. 162); Southwest Cay II, Fosberg & Sachet 53871 (US).

Barrier Reef Cays. -- Goff's Cay, Spellman & Stoddart 2534 (US,
MO); Middle Silk Cay, Fosberg & Spellman 54285 (US, MO), 54277 (US,
MO); South Silk Cay, Fosberg & Spellman 54259 (US, MO, BH, Fo, BM);
Ranguana Cay, Stoddart sight in 1960, Spellman & Stoddart 2252 (US, MO);
Northeast Sapodilla Cay, Spellman & Stoddart 2300 (US, MO, BH, Fo);
Frank's Cay, Spellman & Stoddart 2405 (US, MO); Nicolas Cay, Spellman
& Stoddart 2325 (US, MO, BH); Hunting Cay, Spellman & Stoddart 2454
(US, MO); Lime Cay, Spellman & Stoddart 2378 (US, MO).

Lagoon Cays. -- Cay Caulker, Stoddart 417 (Fo) (1969, p. 6);
Tobacco Range, Fosberg & Spellman 54459 (US, MO); Buttonwood Cay,
Stoddart 90 (Fo) (1963, p. 64), Fosberg & Spellman 54415 (US, MO, BH,
Fo); Hatchet Cay, Fosberg & Spellman 54383 (US, MO, BH, Fo); Little
Water Cay, Fosberg & Spellman 54329 (US, MO, BH, Fo, BM); Moho Cay,
Stoddart 4 (Fo).

Eustachys Desv.
Eustachys petraea (Sw.) Desv.
Chloris petraea Sw.

Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg & Sachet
53812 (US), Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1620
(US), Linhart (1980, p. 162); Southwest Cay I, Fosberg & Stoddart
53862 (US), Linhart (1980, p. 162); Southwest Cay II, Fosberg &

Sachet 53870. (US) , 53902) (US), linhare (1980, ps We2)r:

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2175
(US, MO, BH), Pringle 1913 (HAM); Ranguana Cay, Spellman & Stoddart
2250 (US, MO, BH, Fo, BM); Northeast Sapodilla Cay, Spellman & Stoddart
2312 (US, MO, BH); Frank's Cay, Spellman & Stoddart 2410 (US, MO, BH,
Fo, BM), Stoddart 504 (Fo); Frank's Cay East, Stoddart sight record in
1960; Nicolas Cay, Spellman & Stoddart 2348 (US, MO, BH, Fo, BM);
Hunting Cay, Spellman & Stoddart 2437 (US, MO, BH, Fo, BM); Lime Cay,
Spellman & Stoddart 2362 (US, MO, BH, Fo).

Lagoon Cays. -- Cay Caulker, Stoddart 416 (1969, p. 6, as Chloris
petraea), St George's Cay, Spellman 1457 (MO, US); Tobacco Range,
Fosberg & Spellman 54462 (US); Water Range, Spellman & Stoddart 2236
(US, MO, BH, Fo, BM); Weewee Cay, Stoddart 123 (Fo) (1963, p. 61, as
Chloris petraea); Buttonwood Cay, Stoddart 86 (Fo), Fosberg & Spellman
54403 (US); Hatchet Cay, Fosberg & Spellman 54370 (US); Little Water
Cay, Fosberg & Spellman 54248 (US); Moho Cay, Stoddart 10 (Fo), 117
(Fo).

Panicum L.
Panicum pilosum Sw.

Barrier Reef Cays. -- Lime Cay, Spellman & Stoddart 2374 (US, MO,
BH).

Panicum virgatum L.

Barrier Reef Cays. -- South Water Cay, Pringle 1925 (HAM, MO).

Paspalum L.
Paspalum blodgettii Chapman

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2213
(US), 2176 (US, MO, BH, Fo, BM); Northeast Sapodilla Cay, Spellman &
Stoddart 2321 (US, MO); Frank's Cay, Spellman & Stoddart 2401 (US, MO,
BH, Fo), 2407 (US, MO, BH, Fo); Nicolas Cay, Spellman & Stoddart 2335
(US, MO, BH); Hunting Cay, Spellman & Stoddart 2464 (US, MO, BH, Fo);
Lime Cay, Spellman & Stoddart 2380 (US, MO, BH, Fo, BM).

Lagoon Cays. -- Little Water Cay, Fosberg & Spellman 54349 (US, MO,
BH, Fo, BM).
Paspalum distichum L. (Salt grass)

Paspalum vaginatum Sw.

Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg & Sachet
5S oso (US, emo} SH) ;ebinhart 980; p. 162); Small Cay (Long Cay North)’;
Fosberg & Sachet 53776 (US), Linhart (1980, p. 162); Middle Cay,
Sachet & Stoddart 1627 (US), Linhart (1980, p. 162); Southwest Cay I,
Fosberg & Stoddart 53859 (US), Linhart (1980, p. 162); Southwest CayII,
Fosberg & Sachet 53873 (US) (all Linhart citations as P. vaginatum).

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2531 (US);
Rendezvous Cay, Spellman & Stoddart 2508 (US), 2509 (US, MO), 2510 (US,

MO); StoddartyllOs (Fo); CaynGlony, Stoddart Sill (Fo)s (19637) pe 52);
Tobacco Cay, Fosberg & Spellman 54244 (US); South Water Cay, Spellman
& Stoddart 2214 (US, MO, BH, Fo, TI); Carrie Bow Cay, Spellman &

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Stoddart 2218 (US, MO), 2219 (US), 2574 (US, MO), Fosberg & Spellman
54437 (US), 54438 (US), Ferraris 3 (US); North Silk Cay, Fosberg &
Spellman 54269 (US); Middle Silk Cay, Fosberg & Spellman 54290 (US);
Round Cay, Fosberg & Spellman 54302 (US); Ranguana Cay, Spellman &
Stoddart 2251 (US, MO); Tom Owen's Cays (West), Spellman & Stoddart
2278 (US, MO, BH, Fo); ‘Tom Owen's Cays (East), Spellman & Stoddart 2285
(US, MO, BH); Northeast Sapodilla Cay, Spellman & Stoddart 2317 (US,
MO); Nicolas Cay, Spellman & Stoddart 2350 (US, MO); Hunting Cay,
Spellman & Stoddart 2465 (US, MO, BH); Seal Cay, Spellman & Stoddart
2487 (US, MO).

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54448 (US);
Coco Plum Cay, Spellman & Stoddart 2564 (US, MO, BH); Water Range,
Spellman & Stoddart 2230 (US); Little Water Cay, Fosberg & Spellman
54342 (US).

Paspalum laxum Lam.

Atoll Cays. -- Glover's Reef: Northeast Cay, Stoddart 63 (Fo),
Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1616 (US);
Southwest Cay I, Linhart (1980, p. 162); Southwest Cay II, Fosberg &

Sachet 53872 (US):

Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54386 (US, MO, BH),
54391 (US).

Paspalum paniculatum L.

Lagoon Cays. -- Moho Cay, Stoddart 118 (Fo).
Paspalum sp.

Barrier Reef Cays. -- Ranguana Cay, Stoddart sight in 1960; Round
Cay, Stoddart sight in 1960; Tom Owen's Cays (West), Stoddart sight in
1960; Frank's Cay (West), Stoddart sight in 1960; Frank's Cay (East),
Stoddart sight in 1960.

Lagoon Cays. -- St George's Cay, Stoddart sight in 1962 (1963,
Deo OAs
Phragmites Trin.

Phragmites cf. australis (Cav.) Trin. ex Steud. (Reed)

Phragmites communis Trin.

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54211 (US,
MO); South Water Cay, Spellman & Stoddart 2181 (US, MO, BH).
Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54467 (US, MO,

BH):

*Saccharum L.

*Saccharum officinarum L. (Sugar-cane)

Lagoon Cays. -- Frenchman's Cay: Stoddart sight in 1961.

Spartina Schreb.

Pm Weds fit |

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Spartina patens (Ait.) Muhl.

Atoll Cays. -- Glover's Reef: Long Cay, Fosberg & Sachet 53916
(US, MO, BH); Southwest Cay I, Fosberg & Stoddart 53858 (US);
Southwest Cay II, Fosberg & Sachet 53875 (US).

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54226 (US);
South Water Cay, Spellman & Stoddart 2206 (US), Pringle 1929 (HAM);
Frank's Cay, Spellman & Stoddart 2418 (US, MO).

Spartina spartinae (Trin.) Merr.

Atoll Cays. -- Glover's Reef: Southwest Cay II, Linhart (1980,
p. 162) (could this be Spartina patens?).

Barrier Reef Cays. -- Sergeant's Cay, Spellman & Stoddart 2553
(US, MO); South Water Cay, Spellman & Stoddart 2191 (US, MO, BH, Fo,
BBE)

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54447 (US);

Coco Plum Cay, Spellman & Stoddart 2562 (US, MO, BH, Fo); Man-o'-War
Cay, Fosberg & Spellman 54208 (US, MO); Buttonwood Cay, Fosberg &
Spellman 54423 (US); Hatchet Cay, Fosberg & Spellman 54375 (US);
Little Water Cay, Fosberg & Spellman 54335 (US).

Sporobolus R. Br.

Sporobolus jacquemontii Kunth

Sporobolus indicus of authors, non (L.) R. Br.

Barrier Reef Cays. -- Northeast Sapodilla Cay, Spellman & Stoddart
2304 (US, MO, BH, Fo).

Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54360 (US); Wild
Cane Cay, Stoddart 23 (Fo).

Sporobolus virginicus (L.) Kunth (Salt grass)
Atoll Cays. -- Lighthouse Reef: Sandbore Cay, Stoddart 145 (Fo);

Halt SMoon: Cay;,wStoddartn4 ©) (Ho), 937 (Fo); Long (Cay) Steddart)455, (Fo).

Glover's Reef: Northeast Cay, Fosberg & Sachet 53825 (US); Small Cay

(Long Cay North), Fosberg & Sachet 53779 (US), Linhart (1980, p 162);

12

Long Cay, Pippen 910 (US), Fosberg & Sachet 53805 (US), Linhart (1980,
p. 162); Middle Cay, Sachet & Stoddart 1628 (US); Southwest Cay II,
Fosberg & Sachet 53874 (US), Linhart (1980, p. 162).

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2526 (US,
MO, BH, Fo, TI); Tobacco Cay, Fosberg & Spellman 54213 (US); South
Water Cay, Sampson 27 (US), Spellman & Stoddart 2182 (US, MO, BH, Fo),
2215 (US, MO), Pringle 1911 (HAM); Pompion Cay, Fosberg & Spellman
54311 (US); Ranguana Cay, Spellman & Stoddart 2253 (US, MO, BH, Fo);
Tom Owen's East Cay, Spellman & Stoddart 2286 (US, MO, BH, Fo, TI);
Tom Owen's West Cay, Spellman & Stoddart 2275 (US, MO, BH, Fo, TI);
Frank's Cay, Spellman & Stoddart 2414 (US, MO, BH, Fo); Nicolas Cay,
Spellman & Stoddart 2354 (US, MO, BH, Fo); Hunting Cay, Spellman &
Stoddart 2462 (US, MO, BH, Fo); Lime Cay, Spellman & Stoddart 2382
(US, MO, BH, Fo).

Lagoon Cays. -- Coco Plum Cay, Spellman & Stoddart 2573 (US, MO,
BH, Fo); Water Range, Spellman & Stoddart 2234 (US, MO); Jack's Cay,
Fosberg & Spellman 54429 (sterile) (US); Little Water Cay, Fosberg &
Spellman 54328 (US, MO, BH, Fo, BM); Moho Cay, Stoddart 7 (Fo).

Sporobolus sp.

Atoll Cays. -- Turneffe Atoll: Soldier Cay, Stoddart sight records
in 1960 and 41962 -(1962a,, p= 44, 1963), pis 75921969 , (p< 13) eat Big icallabash
Cay, Stoddart sight record in 1965 (1969, p. 14); Deadman Cay II,
Stoddart sight records in 1960 and 1962 (1962a, p. 38, 1963, p. 80);
Deadman Cay III, Stoddart sight records in 1960 and 1962 (1962a, p. 39,
1963, p. 81); Deadman Cay IV, Stoddart sight records in 1961 and 1965
(1962a, p. 39, 1963), pe 81;: 1969, p. 15); » Deadman -Cay V:; Stoddart’ signi
records in 1960. and 1962 (1962a, p. 40, 1963, p. 82)); | Big Cay: Bokel,
Stoddart sight record in 1965. Lighthouse Reef: Sandbore Cay,
Stoddart sight records in 1961 and: 1965: ((1962a\, +p 3-57- 1963), (pagao;
1969, p. 17); Northern Cay, Stoddart sight records in 1960 and 1965
(1962a, p. 62, 1969, p. 17); Hat Cay, Steddart sight record--anelgyco
(1962a, p. 81, 1963, p. 100); Half Moon Cay, sight records in «i960;
1961 and 1965 (1962a, ». 75,. 1963,. p.198),) 1969) ip. 7) = -Longicaya
Stoddart ‘sight record in 1960° (1962a, pe" 79 P1969 “pees Glover's
Reef: Northeast Cay, Stoddart sight record in 1961 (1962a, p. 88);
Southwest Cay I, Stoddart sight record in 1961 (1962a, p. 95);

Southwest Cay IL, Stoddart sight record in 196 (G962a,2p.) 97):

Barrier Reef Cays. -- Rendezvous Cay, Stoddart sight records in
1960," 1962 ‘and 1965"(1963), ‘p.- 485 1969F* p>. 9s) Tobacco Cay, Stoddart
sight record in 1965 (1969, p. 10); South Water Cay, Stoddart sight
records “in/1960"-and 1965 (1963, p. 56, 1969, p.(clal!) 2s Ranguanay Cays,
Stoddart sight record in 1960; Tom Owen's West Cay, Stoddart sight
record in 1960; Frank's Cay, Stoddart sight record in 1960; Frank's
West Cay, Stoddart sight record in 1960; Hunting Cay, Stoddart sight
record)jin 1960..(1962b,, .p.) 164).;

153

Lagoon Cays. -- Ambergris Cay, Stoddart sight records in 1962 and1965
GI969P pro) musi GeorgeusmCays) oteddantastghitanrecond 1m 1965 (19697,
Penel2) i; eCollsont Cay, stoddarnt ssigh te record: in 1965e@l969)7 sp). 1:2);
SeipiloyCay,, Steddart sight record in, 1965) (969) pm. 22); West Snake
Gay, (Stoddart esight; necond in 961 C965) pre 37) 7 West Snake, Cay.,

West Islet, Stoddart sight record in 1961.

Stenotaphrum Trin.

Stenotaphrum secundatum (Walt.) O. Ktze. (Buffalo grass, pimento
grass, crab grass)

Barrier Reef Cays. -- Northeast Sapodilla Cay, Spellman & Stoddart
2313 (US, MO, BH, Fo); Frank's Cay, Spellman & Stoddart 2409 (US, MO,
BH, Fo); Lime Cay, Spellman & Stoddart 2370 (US, MO, BH, Fo).

Lagoon Cays. -- Little Water Cay, Fosberg & Spellman 54346 (US);
Moho) Cay, Stoddart 2a (Fo), 15 (Fo);

CYPERACEAE

Cladium P. Browne
Cladium jamaicense Crantz (Saw-grass)
Mariscus jamaicensis (Crantz) Britt.

Lagoon, Gays). .—- Harvest. Cay, Stoddart 120 (Fo) (1963, p. 68);
Water Range, Spellman & Stoddart 2239 (US, MO).

Cyperus L.
Cyperus ligularis L.

Atoll Cays. -- Glover's Reef: Northeast Cay, Linhart (1980,
DPelG2) i Small (Cay e(hong tay, .Nowth)s, Linharte @l980), p47 162))- 5 Longi.Gay,;
Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1622 (US, MO,
BH), Ganhant GOS80;, palo2); = Southwest Cay 1; \Linhart ,(1980,- p. 62);
Southwesie Caywit, sianhawzey (l9S8SO;.p. 62.

Barrier Reef Cays. -- Rendezvous Cay, Stoddart 103 (Fo); Tobacco
Cay, Fosberg & Spellman 54227 (US); South Water Cay, Spellman &
Stoddart 2211 (US, MO); Hunting Cay, Spellman & Stoddart 2450 (US, MO,
al, INO, BIE).

Lagoon Cays. -- Cay Caulker, Stoddart 402 (Fo) (1969, p.6);
Buttonwood Cay, Fosberg & Spellman 54413 (US), 54414 (US), Stoddart 87

(Fo)), 89 (Fo) (19637, p. 64ras Cs planifolius);, Gitte Water (Cay;
Fosberg & Spellman 54338 (US); Moho Cay, Stoddart 8 (Fo).

Po eo i

EUW ECLIILILcOivi INVIIWNUOMLIOVSD

riwoce dF ity € tlie

soWU IW

14

Cyperus peruvianus (Lam.) Williams
Kyllinga peruviana Lam.

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2171
(US, MO, BH,Fo,TI), Pringle 1893 (HAM); Frank's Cay, Stoddart 505 (Fo),
Spellman & Stoddart 2400 (US, MO, BH); Nicolas Cay, Spellman & Stoddart
2343 (US, MO, BH, Fo); Hunting Cay, Spellman & Stoddart 2469 (US, MO,
BH); Lime Cay, Spellman & Stoddart 2381 (US, MO, BH).

Lagoon Cays. -- Weewee Cay, Stoddart 121 (Fo) (1963, p. 61);
Hatchet Cay, Fosberg & Spellman 54376 (US); Moho Cay, Stoddart 14
(Fo).

Cyperus planifolius L. C. Rich

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 41 (Fo)
(19627 pp: 76)E: Glover's Reef: Northeast Cay, Stoddart 64 (Fo),
Fosberg & Sachet 53820 (US, MO, BH, Fo, BM), Linhart (1980, p. 162);
Small Cay (Long Cay North), Fosberg & Sachet 53780 (US, MO, BH, Fo),
53781 (US, MO), Linhart (1980, p. 162); Long Cay, Fosberg & Sachet
53913 (US, MO, BH, Fo, BM), Pippen 909 (US), Linhart (1980, p. 126);
Middle Cay, Sachet & Stoddart 1614 (US, MO, BH, Fo, BM), Linhart (1980,
p. 126); Southwest Cay I, Linhart (1980, p. 126); Southwest Cay II,
Fosberg & Sachet 53884 (US, MO, BH, Fo), 53899 (US, MO, BH, Fo, “BM)’,
Linhace (980m Dp el26)=

Barrier Reef Cays. -- Goff's Cay, Spellman & Stoddart 2537 (US,
MO, BH, Fo); Rendezvous Cay, Spellman & Stoddart 2503 (US, MO, BH,
Fo); Tobacco Cay, Fosberg & Spellman 54216 (US), 54248 (US); South
Water Cay, Spellman & Stoddart 2170 (US, MO, BH, Fo), 2196 (US, MO),
2202 (US, MO, BH, Fo), Pringle 1892 (HAM, MO); Carrie Bow Cay,
Spellman & Stoddart 2216 (US); North Silk Cay, Fosberg & Spellman
54270 (US); Middle Silk Cay, Fosberg & Spellman 54282 (US); South
Silk Cay, Fosberg & Spellman 54262 (US, MO); Round Cay, Fosberg &
Spellman 54296 (US), 54299 (US), 54304 (US); Pompion Cay, Fosberg &
Spellman 54326 (US); Ranguana Cay, Spellman & Stoddart 2257 (US, MO,
BH, Fo); Tom Owen's West Cay, Spellman & Stoddart 2272 (US, MO, BH,
Fo); Tom Owen's East Cay, Spellman & Stoddart 2289 (US, MO, BH, Fo);
Northeast Sapodilla Cay, Spellman & Stoddart 2310 (US, MO, BH, Fo);
Frank's Cay, Spellman & Stoddart 2408 (US, MO, BH); Nicolas Cay,
Spelman & Stoddart 23244 (US; Mo), “BH) Fo)", 2347 "(US ;) MO} ‘BH, -Fo; TP);
Hunting Cay, Spellman & Stoddart 2422 (US, MO, BH); Lime Cay, Spellman
& Stoddart 2372 (US, MO), 2379 (US, MO, BH, Fo); Ragged Cay, Spellman
& Stoddart 2385 (US, MO, BH, Fo); Seal Cay, Spellman & Stoddart 2488
(US; MO, BH; Fo).

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54469 (US);
Coco Plum Cay, Spellman & Stoddart 2560 (US, MO, BH, Fo); Water Range,
Spellman & Stoddart 2237 (US, MO, BH, Fo); Buttonwood Cay, Fosberg &
Spellman 54413 (US); Hatchet Cay, Fosberg & Spellman 54382 (US);
Little Water Cay, Fosberg & Spellman 54334 (US).

LS)

Cyperus sp.

Atoll Cays. -- Turneffe Atoll: Mauger Cay, Stoddart sight record
Le OCOm(lIo2ay inp. Ae LICS pk li, saSnC planishod7S) ie COCKLOach
Cave Vie ocoddanniysaghit second Gin! 9654 G9 69) p13), | Cockroach) (Cay,
Stoddart saghita records) an 1962 and) 1965 "969; ps3) 5_ Solider (Cays,
Sitoddarsts signe: recordsean 1962" and 1965 ¢G.963), pe 75,6 LOCI ap), ols);
Deadman Cay I, Stoddart sight records in 1960 and 1962 (1963, p. 80,
IIGI toe 45); Deadman) Cay, Vi;7) Stoddart isighit) record jan 96O) (“L96Zai,
Pea On sasyG. plan trol TUsy)y Lighthouse Reef: Sandbore Cay, Stoddart
sight records in 1961 and 1962; Northern Cay, Stoddart sight records
ine GA aCVCSre pin 9S)i-on Hal taMoonyGay, .stoddank sighitrecord) iny 1965
GISCIE pe LG); =LoOngn Cay) oLoddantr sight wecord in 962 “963i. pe, 995,
asmGe. planwrol 1s). Glover's Reef: Small Cay (Long Cay North),
Stoddart sight record in 1961; Long Cay, Stoddart sight record in
1961; Southwest Cay I, Stoddart sight record in 1971.

Barrier Reef Cays. -- Rendezvous Cay, Stoddart sight record in
ICD eIChIGSi, pis 49), ase GC. plant folius, 1969) sp) 9) 7) se Tobacco Cay,
Stoddart) sight cLecords) in 1961 and 1965 \Gi969),, p. TO); South Water
Cay) Stoddart "sight second an 1960!) (i963), p. 56, ‘as (GC. “plans follius));;
North Silk Cay, Stoddart sight record in 1961; Round Cay, Stoddart
Sight record in 1960; Ranguana Cay, Stoddart sight record in 1960;
Tom Owen's East Cay, Stoddart sight record in 1960; Tom Owen's West
Cay, Stoddart sight record in 1960.

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1962;
Cay Chapel, Stoddart sight record in 1962; St George's Cay, Stoddart
sight =cecords in N962 Vand =1965 (963), p. 40, 1969, p: 7); ~Robiinson
PointuGay,,) Stoddart Ssighterecord in) 1961 (1963, p. 46,,as Cx
planiroOlius)i; LLapp Si Cay, Stoddart’ sight record in 1962 (1963, p. 64);
Cat Cay, Stoddart sight recordin 1962 (1963, p. 62); Bugle Cay, Stoddart
Sight recordin 1965 (1969, p. 11); East Snake Cay, Stoddart sight record
in 1961 (1965, p. 138, asC. ligularis); Middle Snake Cay North Islet,Stoddari
Sight recordin1961; South Snake Cay, Stoddart sight recordin 1961; West
Snake Cay, Stoddart sight recordin 1961 (1965, p. 137).

Fimbristylis Vahl
Fimbristylis cymosa R. Br. var. spathacea (Roth) Koyama
Fimbristylis spathacea Roth

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 135
(Fo). Glover's Reef: Northeast Cay, Fosberg & Sachet 53815 (US),
Linhawt (Go 8O; ap.l62)sne Long ‘Cay, Hosberg & Sachet 53792, (US;* MO; BH,
Fo;, .BM)),) 2tppeny 908: (US). Linharti(1980, sp. 162) 9 Middle: Cay ,, Stoddart
80 (Fo), Sachet & Stoddart 1615 (US, MO, BH, Fo), Linhart (1980, p. 162);
Southwest Cay I, Linhart (1980, p. 162); Southwest Cay II, Fosberg &
Sachety5s869 US — MO BH» Fo); Linhactwi(1980, pie 162) -

Po es i

BUCCI LEI Ltiacoinve INVIIWUOMLIEYS

ritceaaeE Peay bg ehusiv

Su yIWrraiy

16

Barrier Reef Cays. -- Rendezvous Cay, Stoddart 106 (Fo) (1963,
p- 49), Spellman & Stoddart 2511 (US, MO, BH); South Water Cay,
Spellman & Stoddart 2177 (US, MO, BH, Fo), Pringle 1888, 1892 (HAM);
Frank's Cay, Spellman & Stoddart 2415 (US, MO, BH, Fo); Nicolas Cay,
Spellman & Stoddart 2345 (US, MO, BH, Fo, TI); Hunting Cay, Spellman &
Stoddart 2457 (US, MO, BH, Fo, TI); Lime Cay, Spellman & Stoddart 2377
(US, MO, BH, Fo).

Lagoon Cays. -- Cay Caulker, Stoddart 414 (Fo) (1969, p. 6);
Weewee Cay, Stoddart 122 (Fo) (1963, p. 61); Buttonwood Cay, Fosberg &
Spellman 54401 (US); Hatchet Cay, Fosberg & Spellman 54371 (US);
Little Water Cay, Fosberg & Spellman 54344 (US); Moho Cay, Stoddart 11
(Fo).

Fimbristylis spadicea (L.) Vahl

Lagoon Cays. -- Coco Plum Cay, Spellman & Stoddart 2565 (US, MO);
Hatchet Cay, Fosberg & Spellman 54374 (US); Wild Cane Cay, Stoddart 22
(Fo); Moho Cay, Stoddart 3 (Fo).

Fimbristylis sp.

Atoll Cays. -- Glover's Reef: Southwest Cay I, Stoddart sight

record in 1971.

PALMAE

Acoelorrhaphe H. Wendl.
Acoelorrhaphe wrightii (Griseb. & Wendl.) Wendl. ex Becc.

Paurotis wrightii (Griseb. & Wendl.) Britt. ex Britt. & Schafer

Barrier Reef Cays. -- Lime Cay, Spellman & Stoddart 2356 (US, MO).
* Cocos iin
*Cocos nucifera L. (Coconut palm)
Atoll Cays. -- Turneffe Atoll: Vermeer (1959, p. 93); Dixon

(1956, p. 26); Romney et al. (1959, p. 59); Mauger Cay, Stoddart

sight record an-'1960 (1962a7, “p.. 47" 1963)" p 2 71); > Dog£lea ‘Cay, stoddart
sight record in 1960 (1962a, p. 46); Three Corner Cay, Stoddart sight
record in ‘1960, (1963, p. 71); Pelican Cay (Cockroach Group)’, Stoddart
sight record in 1962 (1963, p. 72); Cockroach Cay, Stoddart sight
records: ‘in 1960 and*1962 (1962a, p.#46,,71963;- p.| 73)4°2 Cockroach ‘Cay? V,
Stoddart’ sight xrecord’ in: ‘1962 -(1963,,-p. 73) ;-'Pelican Cay,. Stoddart
sight record in 1960 (1962a, 45, 1963, p. 74); Blackbird Cay, Stoddart
sight record in 1960 (1962a, p. 44); Soldier Cay, Stoddart sight

17

records in 1960 and 1962 (1962a, p. 44, 1963, p. 75); Harry Jones
Point, Stoddart sight record in 1960 (1962a, p. 48); Big Calabash
Cay, Stoddart sight records in 1960, 1962 and 1965 (1962a, p. 42, 1963,
Die lo Ole wpe 4) eb. Calabash basic, Cays ya scoddartysignit) records
in 1960 and 1962 (1963, p. 78); Big Calabash East Cay II, Stoddart
sight record in 1960 (1962a, p. 42,-1963, p. 78); Little Calabash Cay,
Stoddart sight record in 1960 (1962a, p. 41, 1963, p. 76); Grand: Bogue
Point, Stoddart sight record in 1962 (1963, p. 84); Deadman Cay I,
Stoddartusigatyrecordssing| 9607 l962zandylI6Sa(LI6Za,. p. 38,0 1965),
DeLee 1969; ep. Lo). | Deadman Cay, 11, Stoddart sight recoxds in; 1960),
IIG2 Tanda l965s, CICS so. cO) 19697 peak) >. Deadman Cay Dil,-Stoddart
sight records in 1960 and 1962 (1962a, p. 39, 1963, p. 81); Deadman
Cay IV, Stoddart sight records in 1960, 1962 and 1965 (1962a, p. 39,
LI6S% pe Ol I69F mp. 15)» Deadman, Cay V,- 9 coddart sight records: in
1960 and 1962 (1962a, p. 40, 1963, p. 82); Big Cay Bokel, Stoddart
sight record in 1962 (1963, p. 83); Cay Bokel, Glennie sight record in
192i Stoddart sight record a nydlI9oOl e(1962a,) pe 9S? ,71963,- paz)
Lighthouse Reef: Sandbore Cay, Stoddart sight records in 1960, 1961
and 1962 (1962a, p. 57, 1963, p. 89); Northern Cay, Stoddart sight
HOCOLdSy 1 nw IOO;mlIOly eIG25 sand 1965) @962a; vp. 6196S), pe 193);
Saddle Cay, Stoddart sight record in 1960 (1962a, p. 63, 1963, p.94);
Half Moon Cay, Uring in 1720, Owen in 1828 (1962a, p. 73), Honduras
Aulmanacks (L830; pe 150), Salvin sight record in862 (Salvin 1864,
pesos), Vermeer’ (1959, figs. 923 and 25), Stoddart sight records in
ICO) andelIGIs (LI6Za- penw>)) 7. LOo2a (L968 pee 95) pe and=1965)(19695 5.17) ;
long Cay, Vermeer) (1959, 5p, 97), vgs 21) Stoddart sight records iin
ICG MUIo LA IIG2Z Mand 1I65—q(1I96Za\jao. 78, 1963), (pe 199), 819697) pe. 18))r
Glover's Reef: Northeast Cay, Stoddart sight record in 1961 (1962a,

p. 88), Fosberg and Sachet sight record in 1971, Linhart (1980, p. 162);
Small Cay (Long Cay North), Stoddart sight record in 1961 (1962a,

p. 89), Fosberg and Sachet sight record in 1971, Linhart (1980, p. 162);
Long Cay, Stoddart sight record in 1961 (1962a, p. 91), Fosberg and
Sachet sight record in 1971, Linhart (1980, p. 162); Middle Cay,
Stoddart sight record in 1961 (1962a, p. 93), Fosberg and Sachet sight
record in 1971, Linhart (1980, p. 162); Southwest Cay I, Stoddart
Sight record in 1961 (1962a, p. 94), Fosberg and Sachet sight record
ingle swanhart (980; sp. 162); Southwest Cay 1, ‘Stoddartsight
record in 1961 (1962a, p. 98), Fosberg and Sachet sight record in 1971,
Lanhart MGEI8O0, sp. 4/62)

Barrier Reef Cays. -- Vermeer (1959, p. 30); Paunch Cay, H.M.S.
Rambler sight record in 1896-7; Sergeant's Cay, H.M.S. Rambler sight
record in 1896-7, Vermeer (1959, figs. 1, 11, 13), Stoddart sight record
imMpelIGOL(LI63), Da142) +) GOLE_S Cay, .H.Mic. Rambler Sight’ record an
1896-7, Vermeer (1959, fig. 15), Stoddart sight records in 1960 and
LIS SEACotanced). (969 aoe no, STA pe Avo); English Cay, ctoddart sight
records in 1960, 1962 and 1965 (1963, p. 44, 1969, p. 9); Rendezvous
Cay, H.M.S. Mutine sight record in 1921-2, Stoddart sight records in
19607" 1965" (planted) and "1972 (1969,*p: 9), 1974, p.” 480); ~ Cay Glory,

oa

PUVICLIILILIOING INVINUSOMLIEVS

HIWOTIEETULIVIN

eJIVIL BTEC IW EPRI

18

Stoddart sight*record in: 1960F (19672b; +p." 16454196877 pa S2) = Tobacco
Gay; Stoddart sights records any 1960; 71961 A962) A965) ands 1:97 2e4l96sF
Dp. 54));)+South: Water Cay, Stoddart™sight? records™in 196077 1961) aoG5
and 1972 (1969, p. 11), Pringle, sight record in 1980; Carrie Bow Cay,
Stoddart Sight’ records®in 1960,>1965;) and W9725 (4962b;, ps 164-57 19687,
pe S75 21969) peti); North» Silky Cay, sSteddar sight records tans 19 Gl
and 1972; Middle Silk Cay, Stoddart sight records in 1961 and 1972;
South Silk Cay, Stoddart sight records in 1961 and 1972; Round Cay,
Stoddart sight recordsin 1960 and 1972; Pompion Cay, Stoddart sight
records in 1960 and 1972; Ranguana Cay, Vermeer (1959, p. 91), Stoddart
sight records in 1960 and 1972; North Spot, Stoddart sight records in
1960 and 1972; Tom Owen's East Cay, Stoddart sight records in 1960 and
1972; Tom Owen's West Cay, Stoddart sight records in 1960 and 1972;
Northeast Sapodilla Cay, Stoddart sight records in 1960 (1962b, p. 164)
and 1972; Frank's Cay, Stoddart sight records in 1960 and 1972;
Frank's Cay East Islet, Stoddart sight records in 1960 and 1972;
Frank's Cay West Islet, Stoddart sight records in 1960 and 1972;
Nicolas Cay, Stoddart sight records in 1960 and 1972; Hunting Cay,
Stoddart sight records in 1960 and 1972 (1962b, p. 164); Lime Cay,
Stoddart sight recordsin 1960 and 1972; Ragged Cay, Owen sight record
in 1835, Stoddart sight records in 1960 and 1972; Seal Cay, Owen sight
record in 1835, Vermeer (1959, p. 90), Stoddart sight records in 1960
and 1972.

Lagoon Cays. -- Ambergris Cay, Vermeer (1959, p. 30, fig. 6),
Stoddart «sight records 7in 1960 and+19657 (1963, %p. “33, /1969%i p= 6)
Cay Caulker, Vermeer (1959, fig. 7), Stoddart sight records in 1960,
1961, “1962 ) and 1965 (19637 pir34) 7 '*Cay°Chapel’, ~yetferys’ in=27/757
Stoddart sight) records: ‘in 1962 ‘and=1965 (1963, -p.935, "19697 %pF°6))-
St George's Cay, Stoddart sight records in 1960, 1962, 1965 (1969,
p. 7 and 1972); Robinson Point Cay, Vermeer (1959, p. 92) Stoddart
Sight record in 1961 (1963, p. 46); Robinson Cay, Stoddart sight
record in-1962°(1963, p-..47); Spanish Cay, Stoddart siight *records=in
1961 and 1962 (1963, p. 47); Tobacco Range, Stoddart sight record in
1972; Weewee Cay, Stoddart sight record in 1965; Buttonwood Cay,
Stoddart "sight. records -in1961° (1963, ps 64) .and’ 19723" Cary Cay?
Stoddart tsight =records: in y1962°(1963, ‘p. 63)’; -Cat-Cay;, Stoddart*sighe
record in 1962°(1963, "=p /°62)'; “Laughing Bird"Cay, Stoddart“sightsrecord
in 1962 (1963, p. 65); Colson Cay, Stoddart sight records :in’ 1962¥*and
1965 (1963, p. 67); Scipio Cay, Stoddart sight records in 1962 and
1965. (1963, .,p. 66);. Owen..Cay, Stoddart sight record. in« 1962.,(1963;,
p. 66); Bugle Cay, Stoddart sight records in 1962 and 1965; Harvest
Cay, Stoddart sight record ,in, 1962 (1963;,,p..468);.. .Trappts.Cayy,
Stoddart sight records in 1962 (1963, p. 64).and 1972; Moho Cay,
Stoddart sight record in 1961; Frenchman's Cay, Stoddart sight record
in 1961; Wild Cane Cay, Stoddart sight record in 1961; East Snake
Cay,. Vermeer (1959), p.. 92), Stoddart sight record jine1961. Gl965;.-peq3 8);
Middle Snake Cay, Stoddart sight record in 1961; South Snake Cay,
Stoddart sight record in 1961 (1965, p. 138); West Snake Cay, Stoddart
SiqhiyreconrdenalI Gin MIG Sys LS 7i)r

iL)
Thrinax Sw.
Thrinax radiata Lodd. ex Schultes & Schultes. (Sea thatch)
Thrinax parviflora sensu auct.
Thrinax floridana Sarg.

Thrinax multiflora sensu Read in Adams, Flowering Plants of
Wamancari(l97 2 spi. W/O).

Atoll Cays. -- Turneffe Atoll: Pelican Cay (Cockroach Group),
Stoddart sight record in 1962 (1963, p. 72); Deadman Cay.I, Stoddart
Saghit record, in) 1965 (1969, p. 15) Deadman Cay IL, Stoddart sight

record in 1965 (1969, p. 15); Deadman Cay IV, Stoddart sight records
in t96OWand 1962 {(963,/ p. 81); Big Cay Bokel, Stoddart sight record
iin IDSS6 Lighthouse Reef: Northern Cay, Stoddart sight record in

1965 (1969, p. 17); Half Moon Cay, Honduras Almanack (1830, p. 150).
Glover's Reef: Northeast Cay, Stoddart 77 (Fo), Linhart (1980, p. 162);
Long Cay, Fosberg & Sachet 53791 (US, MO, BH, Fo, BM), Linhart (1980,

p. 162); Middle Cay, Fosberg & Stoddart 53919 (US, MO).

Barrier Reef Cays. -- South Water Cay, Stoddart sight record in

1965 (1969, p. 11), Spellman & Stoddart 2145 (US); Northeast Sapodilla
Cay, Stoddart sight record in 1960 (1962b, p. 164), Spellman & Stoddart
2298 (US, MO); Frank's Cay, Stoddart sight record in 1960, Spellman &
Stoddart 2417 (US, MO); Frank's Cay East Islet, Stoddart sight record
in 1960; Nicolas Cay, Spellman & Stoddart 2333 (US, MO); Hunting Cay,
Spellman & Stoddart 2476 (US, MO); Lime Cay, Spellman & Stoddart 2357
(US, MO).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1962
GSC peasy Gaye Chapel stoddart) sight ecord sin 1962 4 (19637 pi1935));
Robinson Point Cay, Stoddart sight record in 1961 (1963, p. 46);

Tobacco Range, Fosberg & Spellman 54445 (US), 54446 (US); Coco Plum
Cay, Spellman & Stoddart 2557 (US, MO); Water Range, Spellman &
Stoddataz222) (USP IMO) ia 22405(US;,) MO; BH, SEO, Tl); Hatchet™ Cay,

Fosberg & Spellman 54372 (US); Cat Cay, Stoddart sight record in 1962
(ises;, “pia, GZ) >= sCollsoniiCay, Stoddart sight records in 1962 and 1965
GIGS pi. sO wel I60 Ps pewl2) I SCiploO@Cay, Stoddart sight records: an’ 1962
ancmlI6o7 (19637, pe CO pulLI6O; pe a)e sOwen: Cay, Stoddart isucght record
nll IOS) a loon nenCany (Cay), Stoddart. Sight, record! in, 1962; Moho
Cay,Stoddart sight record in 1961; Trapp's Cay, Stoddart sight record
nel I62 | @lIGS ssp.) OS) a harvesituCay, sctoddart—si ght recordsin 1962
(1963, p. 68); East Snake Cay, Stoddart sight record in 1961 (1965,

p. 138); Middle Snake Cay North Islet, Stoddart sight record in 1961;
Southmonake Cay, stoddagtesighturecord in 1961) (1965p. 138):

Po i

eUUECLEILILVIVI INVIINUSOMLIVVS

PICBWOprrik Frill

QrRoOVUJIWIPIy

— ww

20

PONTEDERIACEAE
Eichhornia Kunth
Eichhornia crassipes (Mart.) Solms (Water hyacinth)
Barrier Reef Cays. -- Carrie Bow Cay, Ferraris 17 (US) (seedling).

LILIACEAE sensu lato

Agave L.

Agave sp. (Century plant, maypole)
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2189

(US).

Crinum, ji.
*Crinum amabile Donn ex Ker-Gawl. (Crinum lily)

Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg & Sachet
53842 (US); Small Cay (Long Cay North), Fosberg & Sachet 53779 (US).

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54209 (US);
South Water Cay, Spellman & Stoddart 2193 (US, MO).

Lagoon Cays. -- Tobacco Range, south end of principal cay,
Fosberg & Spellman 54450 (US).
*Dracaena Vandelli ex L.
*Dracaena sp.

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2159
(US, MO, BH, Fo, BM), Pringle sight record in 1980.
Hippeastrum Herb.
Hippeastrum puniceum (Lam.) Urb.

Atoll Cays. -- Glover's Reef: Small Cay (Long Cay North), Linhart
(IOS) pr L62)e.

Dad

Hymenocallis Salisb.
Hymenocallis littoralis (Jacq.) Salisb. (Spider-lily)
Pancratium littorale Jacq.

Atoll Cays. -- Turneffe Atoll: Eastern sand ridge, Stoddart sight
record in 1960 (1962a, p. 47); Harry Jones Point, Stoddart 512 (Fo)
(1962a, p. 48); Deadman Cay I, Stoddart sight records in 1962 and
IGS (1963, p. 79, 1969) p. 5); Deadman Cay IE, Stoddart sight records
immo o2 and 1965 eCl969), p. 15); Cay, Bokel), “Stoddart sight record, 2n
IMG (IMOZas Oe Sp MWIOSA jos SZ) S Lighthouse Reef: Northern Cay,
Stoddart sight record in 1962 (1963, p. 93); Half Moon Cay, Stoddart
BIO), ocoddare Ssagnt records) in) W6O- and 1965 (1962a, p. 757) 1963),
p. 98, IGT pial S)s) song uCay,, Stoddart: sught: record in 1960) (1l962Zai,
DemscO) re Hat) Cay), otoddant, sight record in 1961 (1l962a, p. 8), 1963,

Dis HOO) Glover's Reef: Northeast Cay, Fosberg & Sachet 53841 (US);
Small Cay (Long Cay North), Linhart (1980, p. 162); Long Cay, Fosberg
emSachet 5 s95) (US), Stoddart sight urecord in 1961 (1962a, pi Oil),
Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1634 (US),
Stoddarts1 ght m-ecord ini 96I 962ay sp. 93), binhart, (1980), \p. 162);
Southwest, Cay £, Stoddart sight record in 1971, Linhart, (1980, p. 162);
Southwest Cay II, Fosberg & Sachet 53905 (US), Stoddart sight record in
IIo 6Za, pO), Lana (l9SO; pe enlG2)) :

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54228 (US),
Stoddart sight records in 1961, 1962 and 1965 (1963, p. 54, 1969,
p. 10); South Water Cay, Spellman & Stoddart 2168 (Fo), Stoddart sight
record in 1965 (1969, p. 11), Pringle 1930 (HAM); Carrie Bow Cay,
Ferraris 16 (?) (US) (Seedling), Ferraris, photograph, Pringle, single
plant observed in 1979; North Silk Cay, Fosberg & Spellman 54265 (US);
South Silk Cay, Fosberg & Spellman 54258 (US), Stoddart sight record in
1961; Round Cay, Fosberg & Spellman 54308 (US), Stoddart sight record
in 1960; Pompion Cay, Fosberg & Spellman 54327 (US), Stoddart sight
record in 1960; Ranguana Cay, Spellman & Stoddart 2244 (US, MO),

Stoddart sight record in 1960; Tom Owen's East Cay, Spellman & Stoddart

2288 (US, MO), Stoddart sight record in 1960; Hunting Cay, Spellman
& Stoddart 2449 (US, MO, BH, Fo), Stoddart sight record in 1960 (1962b,
p. 164); Lime Cay, Spellman & Stoddart 2360 (US, MO); Seal Cay,
Spellman & Stoddart 2478 (US, MO).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965
(1969, p. 6); St George's Cay, Stoddart sight records in 1962 and 1965
GOGsi press, 1969), pa i Robinson Point Cay, Stoddart sight, record
ine I6i (l963; pi. 46) > Trappis Cay, Stoddart sight record in, 1962
(1963, p. 64); Hatchet Cay, Fosberg & Spellman 54389 (US); Moho Cay,
Stoddart sight record in 1961; Cary Cay,Stoddart sight record in 1962
GI963);, 26S) te Owen=Cays, (S Ctoddare sight record in 1962 (19637, p-. 66);
Laughing, Bird) Cay, Stoddart sight) record in 1962, (1963, p..65));) Bast
Snake Cay, Stoddart sight record in 1961 (1965, p. 138); Middle Snake
Cay North Islet, Stoddart sight record in 1961; South Snake Cay,
Stoddantysignitarecordein ew I6iea(lI965. 1. 138)).

Pees i

EUV ILIILILOQIVI INVIINUSOMLIYYS

TINO, tty bt ga

CUM DPE RPOULJIWIPALY

22

MUSACEAE
*Musa L.
*Musa Ssapientum L. (Banana)
Barrier Reef Cays. -- Northeast Sapodilla Cay, Spellman & Stoddart
2326 (US, MO).
*Musa paradisiaca L. (Plantain)
Lagoon Cays. -- Cay Caulker, Vermeer (1959, p. 58), Stoddart sight

record in 1960 (1963, p. 34); Frenchman's Cay, Stoddart sight record
in L9Giy.

ORCHIDACEAE
Brassavola R. Br.
Brassavola nodosa (L.) Lindl. (Orchid)
Atoll Cays. -- Glover's Reef: Northeast Cay, Stoddart 65 (Fo).
Barrier Reef Cays. -- Frank's East Cay, Spellman & Stoddart 2396

(US, MO); Hunting Cay, Spellman & Stoddart 2423 (US, MO).

Lagoon Cays. -- Jack's Cay, Fosberg & Spellman 54428 (US, MO, BH,
Fo); East Snake Cay, Stoddart 94 (Fo); Middle Snake Cay South Islet,
Stoddart sight<record=in- 196.
Laelia Lindl.
Laelia tibicinis (Batem. ex Lindl.) L. Wms. (Orchid)

Schomburgkia tibicinis Batem. ex Lindl.

Lagoon Cay. -- Wild Cane Cay, Stoddart 21 (Fo) (on prone tree trunk
on sand-mangrove cay).

23

CASUARINACEAE
*Casuarina Adans.
*Casuarina equisetifolia L. (Iron-wood)
Atoll Cays. -- Glover's Reef: Small Cay (Long Cay North), Fosberg

and Sachet sight record in 1971, 1 seedling only,2.5 dm tall, Linhart
(IDEO) 4 joe INGA) 6

Barrier Reef Cays. -- RendeZvous Cay, Stoddart sight record in
1965 (1969, p. 9); South Water Cay, Spellman & Stoddart 2199 (US, MO),
Stoddart sight record in 1965 (1969, p. 11), Pringle sight record in
1980; Carrie Bow Cay, Ferraris 14 (US); Hunting Cay, Spellman &
Stoddart 2472 (US, MO, BH), Stoddart sight record in 1960; Ragged Cay,
Spellman & Stoddart 2384 (US, MO, BH, Fo), Stoddart sight record in
1960.

Lagoon Cays. -- Cay Caulker, Stoddart 410 (Fo), Stoddart sight
record in 1962 (1969, p. 6); Water Range, Spellman & Stoddart 2224
(US, MO). Na RET Ret ere, eae

MYRICACEAE
Myrica L.
Myrica cerifera L.
Aeoliln Cays.. —- Glover's Reef: ;Southwesit: Cay 1, Linhart, s.n-.
(IMO) A Wakinvehae (AUOR jo, Wash) e
MORACEAE

Ficus L.
Ficus sp. probably new

Ficus cf. hemsleyana Standley of Linhart (1980)

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Verner (1959,
95 I-38, Moi, jog S74, Sweoeleics (Seen ide V4, IOs, jon Saye Glover's
Reef: Northeast Cay, Stoddart 71 (Fo) (1962a, p. 88, as F. ovalis),
Fosberg & Sachet 53845 (US, MO, BH, Fo, BM), Linhart (1980, p. 163);
Long Cay, Linhart (1980, p. 163); Middle Cay, Sachet & Stoddart 1630
US WO), sist SM into) Fh iningkater (II EO)> fog GS) G

> 4

INVIMULUDPRILIVVS

OW t LIILILoimwl

BR DIW

gsiwoy Pee ig

WIWELE BP CUSULU JIM IRIS

I Iirmroreamiieogosty

24

Barrier Reef Cays. -- Tobacco Cay, Smith (1842, p. 732); Water
Cay, Dwyer et al. 672 (MO) (figs pedunculate); Round Cay, Fosberg &
Spellman 54298 (US); Pompion Cay, Fosberg & Spellman 54310 (US);
Northeast Sapodilla Cay, Speliman & Stoddart 2307 (US, MO, BH); Frank's
Cay, Spellman & Stoddart 2397 (MO, Fo); Nicolas Cay, Spellman &
Stoddart 2328 (US, MO, BH); Hunting Cay, Spellman & Stoddart 2434
(US, MO).

Lagoon Cays. -- East Snake Cay, Stoddart 95 (Fo) (1965, p. 138, as
Ee Ovals).

OLACACEAE
Ximenia L.
Ximenia americana L.
Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Verner (1959,

pp. 3=8,.. 19610) pis 2582) Stoddart e(1962a7"p. 74)¥.

POLY GONACEAE
Coccoloba P. Browne
Coccoloba uvifera L. (Sea-grape)
Atoll Cays. -- Turneffe Atoll: Cockroach Cay V, Stoddart sight

records: in’ 1962Z2andi 1965" (1963) pis 73, 1969" “pe -L3) “Cockroach Cays,
Stoddart sight record in 1962; Soldier Cay, Stoddart sight records in
1962 and 1965 (1963, p. 75, 1969, p. 14); Harry Jones Point, Stoddare
sight record in 1960 (1962a, p. 48); Deadman Cay IV, Stoddart sight
records in 1960 and 1962 (1963, p. 81); Big Cay Bokel, Stoddart sight record
aly ol, ALEX Sy Sr Lighthouse Reef: Sandbore Cay, Stoddart sight records in 1960,
1961, 1962°and 1965 (1962a, p. 57, 1963, p..89, 1969 ;,:p. 16); Northern Cay,
Stoddart sight records in 1960, 1961 and 1965 (1962a, p. 62, 1969, p. 17); Halt
Moon Cay Honduras Almanack(1830, p.150). Glover's Reef; Northeast Cay,
Fosberg & Sachet 53831 (US), Stoddart sight in 1961 (1962a, p. 88), Linhart
(1980, p. 163) ; Small Cay (Long Cay North) , Linhart (1980, p. 163); Long Gay,
Fosberg & Sachet 53926 (US), Stoddart sight recordin 1961 (1962a, p. 91),
Linhart (1980, p. 163); Middle Cay, Sachet & Stoddart 1633 (US), Linhart
(1980, p. 163); Southwest Cay I, Stoddart sight records in 1961 and
1971 (1962a, p. 94), Linhart (1980, p. 163)3 Southwest Cay II, Fosberg
& Sachet 53894 (US).

Barrier Reef Cays. -- Goff's Cay, Stoddart sight records in 1960
and 1962 (1963, p. 43, 1969, p. 9); Sergeant's Cay, Spellman & Stoddart
2542 (US, MO), Stoddart sight records in 1965 (seedling) and 1972

(1969, p. 8, 1974, p. 478); English Cay, Stoddart sight record in 1960

935i

25

(1963, p. 45); Rendezvous Cay, Spellman & Stoddart 2495 (US, MO),
Stoddart sightarecords: ane lI6O, e962, 1965 and 1972" (19631-7487 1969,
pe 9, 1974, p. 480); “Lobacco) Cay, ) Hosberg -& Spelimany54214.— (US) ,
Stoddart sugnitarecordse in I61), e962) andy 1965). (l963 7p. 54), 1969,

p- 10); South Water Cay, Spellman & Stoddart 2151 (US, MO, BH),
Stoddawmtesught records pind 96On ands 1965. (19637 0p. 756;64969,, =p." I),
Pringle 1889 (HAM); Carrie Bow Cay, Spellman & Stoddart 2220 (US, MO),
Ferraris 11 (US); South Silk Cay, Fosberg & Spellman 54261 (US),
Stoddart sight record in 1961; Pompion Cay, Fosberg & Spellman 54322
(US), Stoddart sight record in 1960; Ranguana Cay, Spellman & Stoddart
2242 (US, MO, BH), Stoddart sight record in 1960; Tom Owen's West Cay,
Spellman & Stoddart 2265 (US, MO), Stoddart sight record in 1960; Tom
Owen's East Cay, Spellman & Stoddart 2290 (US, MO, BH), Stoddart sight
record in 1960; Northeast Sapodilla Cay, Spellman & Stoddart 2306 (US,
MO)F pc toddastas ight, record in L960 (1962b), 9p... 164) 5 “Frank's Cay,
Spellman & Stoddart 2390 (US, MO), Stoddart sight record in 1960;
Frank's Cay East Islet, Stoddart sight record in 1960; Nicolas Cay,
Stoddart sight record in 1960; Hunting Cay, Spellman & Stoddart 2448
(US MO) Ee, . Sitoddarta sa ght record in) 1960) (19625 ps, 64) “iilme (Cay,
Spellman & Stoddart 2358 (US, MO, BH, Fo), Stoddart sight record in
1960; Seal Cay, Spellman & Stoddart 2479 (US, MO).

INVIINUOMLIYVD

OWWELIILILOIVI

CIVIL E TIOUINIATY IIWODGrLU ERrUWIN

oi. a. oF ee ee Gee |

Lagoon Cays. -- Cay Caulker, Stoddart 430 (Fo), Stoddart sight
record in 1962 (1963, p. 35); Cay Chapel, Stoddart sight records in
IIG2Z and LI6S (Gl96Sy ap. S5));, mSit Georges ‘Cay, Stoddaresignt records
ew OOMAaniG el 96> (IGS, pe 1S6),, L969, pi. 7));) sRobimson Point Cay,
Stoddart sight record in 1961 (1963, p. 46); Buttonwood Cay, Fosberg &
Spellman 54416 (US), Stoddart sight record in 1961 (1963, p. 64);
Trapp"s Cay, Stoddart sight record in 1962 (1963, p. 64); Moho Cay,
Stoddart sight record in 1961; Cary Cay, Stoddart sight record in 1962
(1963, p. 63); East Snake Cay, Stoddart sight record in 1961 (1965,
p.- 138); Middle Snake Cay North Islet, Stoddart sight record in 1961;
Middle Snake Cay South Islet, Stoddart sight record in 1961; West
Snake Cay, Stoddart sight record in 1961 (1965, p. 137); West Snake :
Cay West Islet, Stoddart sight record in 1961; Wild Cane Cay, Stoddart :
sight record in 1961.

)
BATIDACEAE |
Batis P. Browne
Batis maritima L. (Samphire, Pickle-weed)
Atoll Cays. -- Turneffe Atoll: Mauger Cay, Stoddart 542 (Fo)

GiSGs 7 ep; a eG Calabash) Cay, stoddart, sr1ght record lainnd96 5969),
p. 14); Little Calabash Cay, Stoddart sight record in 1965 (1969,
Dremel) a Lighthouse Reef: Long Cay, Stoddart sight record in 1962

(USS Si. 105, SIS) 4 Glover's Reef: Small Cay (Long Cay North), Linhart
(1980, p. 162); Southwest Cay I, Fosberg & Stoddart 53853 (US), Linhart
GiSs0;. pial62) i Southwest iCaywil, slinhart (L980; ip. 162)%

26

Barrier Reef Cays. -- Sergeant's Cay, Stoddart 440 (Fo) (1969,
p-. 8); South Water Cay, Spellman & Stoddart 2144 (US), Stoddart sight
record in 1965 (1969, p. 11), Pringle 1904 (HAM).

Lagoon Cays. -- St George's Cay, Spellman 1476 (MO), Stoddart sight
record in 1965 (1969, p. 7); Tobacco Range, Fosberg & Spellman 54458
(US); Coco Plum Cay, Spellman & Stoddart 2568 (US, MO, BH, Fo); Water
Range, Spellman & Stoddart 2232 (US, MO); Weewee Cay, Stoddart 124 (Fo)
(1963, p. 61); Man-o'-War Cay, Fosberg & Spellman 54206 (US); Jack's
Cay, Fosberg & Spellman 54431 (US); Buttonwood Cay, Fosberg & Spellman
54419 (US); Hatchet Cay, Fosberg & Spellman 54379 (US); Little Water
Cay, Fosberg & Spellman 54336 (US); Owen Cay, Stoddart sight record in
1962 (1963, p. 66); Bugle Cay, Stoddart sight record in 1965 (1969,
p..6)); ~Harvesit Cay; Stoddartysight) record int 1962e(1968, pl7es)e

CHENOPODI ACE AE
Salicornia L.
Salicornia perennis Miller (Samphire, glasswort)
Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54455 (US);

Water Range, Spellman & Stoddart 2221 (US, MO, BH, Fo).
Suaeda Forsk. ex Scop.
Suaeda linearis (Ell.) Moq.

Barrier Reef Cays. -- Carrie Bow Cay, Ferraris 18 (US).
AMARANTHACEAE

Alternanthera Forsk.
Alternanthera ramosissima (Mart.) Chod.
Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 42 (Fo),
140 XE) 7; Sol (Bo) (196247, ple 7G).
Iresine P. Browne
Iresine diffusa H. B. ex Willd.
Iresine celosia L.

Atoll Cays. -- Turmmeffe Atoll: Harry Jones Point, Stoddart 28 (Fo)
(1962a, p. 48, as I. celosia); Deadman Cay II, Stoddart 129 (Fo)

27

(IQS3 > jo {8}0)} c Lighthouse Reef: Half Moon Cay, Stoddart 452 (Fo)
(1969, p. 18). Glover's Reef: Southwest Cay I, Stoddart 82 (Fo),
Pippen 899 (US), Linhart (1980, p. 162); Southwest Cay II, Fosberg &
Sachet 53888 (US), Linhart (1980, p. 162).

Barrier Reef Cays. -- Tom Owen's West Cay, Spellman & Stoddart 2274
(US, MO, BH, Fo).

Philoxerus R. Br.
Philoxerus vermicularis (L.) Beauv.

Atoll Cays. -- Glover's Reef: Small Cay (Long Cay North), Fosberg
& Sachet 53785 (US); Southwest Cay I, Linhart (1980, p. 162).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight record in 1965
(1969, p. 8), Spellman & Stoddart 2556 (US, MO); English Cay, Spellman
& Stoddart 2523 (US, MO); Rendezvous Cay, Spellman & Stoddart 2506
(USta MO), Stoddart 107 (Fo) (1963, p. 49); Cay Glory, Stoddart 508) (Fo)
GiSGS pa D2) ie TobaccoCay, Fosberg (& Spellman 54236 (US)*,;,- 54256" (US) ;

South Water Cay, Spellman & Stoddart 2146 (US, MO, BH, Fo, TI), Pringle

1905 (HAM); Carrie Bow Cay, Spellman & Stoddart 2141 (US, MO),
Ferraris 6 (US); Pompion Cay, Fosberg & Spellman 54313 (US); Ranguana
Cay, Spellman & Stoddart 2241 (US); North Spot, Spellman & Stoddart
2292 (US); Tom Owen's East Cay, Spellman & Stoddart 2284 (US); Seal
Cay, Spellman & Stoddart 2492 (US, MO).

Lagoon Cays. -- Cay Caulker, Stoddart 415 (Fo), 440a (Fo) (1969,
p. 6); St George's Cay, Spellman 1475 (US, MO); Hatchet Cay, Fosberg &
Spellman 54398 (US); Little Water Cay, Fosberg 54357 (US).
NYCTAGINACEAE

Boerhavia L.
Boerhavia coccinea Mill.?

Barrier Reef Cays. -- South Water Cay, Pringle 1866 (HAM).

Neea Ruiz & Pavon

Neea choriophylla Standl.

The Glover's Reef specimens fit this well enough in leaf shape,
though the leaves of some of the specimens are larger than any so named
in the US. The leaves are comparably thick, the panicle is similarly
large and complex. The staminodia on the one pistillate collection
are 5, 6, 8, 5, 7, 7, 7, in the anthocarps examined on Fosberg 53846

ee

and two staminate collections Sachet & Stoddart 1600 and 1636 have 6-7

Pe yes fe |

SOIVIL LE DIOUIINIRIY IIVWQILTIOU TL ruin OWE LIILILOINVI INVIINUSMLIYYSD

Ne a a ee came |

28

and 6-8 stamens respectively. Whether this species is more than
varietally distinct from Neea psychotrioides Donn.-Sm. of the Central
American mainland is debatable. The latter has 8-10 staminoida

according to Lundell.

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Verner (1959,
pp. 3-8, 1961, Dp. 582)y,, stoddant s¢l9G2a pag/4 pel 9 OSs too) Glover's
Reef: Northeast Cay, Stoddart 68 (Fo), Fosberg & Sachet 53832 (US, MO,
BH), 53846 (US, MO, BH, Fo,- BM) .(1962a, p.- 88)y, Linhart (1980, p-l62)i-
Middle Cay, Sachet & Stoddart 1636 (US, MO, BH, Fo), 1600 (US, MO, BH,
Fo, BM).

PHYTOLACCACEAE
Rivina L.
Rivina humilis L. (Bloodberry, Dog's blood)
Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 44 (Fo),
545) (Ho), (Gig6 2a ps adie IOS; Panes) s Glover's Reef: Southwest Cay

II, Fosberg & Sachet 53889 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -- Round Cay, Fosberg & Spellman 54307 (US);
Pompion Cay, Fosberg & Spellman 54320 (US); Tom Owen's West Cay,
Spellman & Stoddart 2270 (US, MO, BH); Ragged Cay, Stoddart 503 (Fo),

Spellman & Stoddart 2387 (US, MO); Seal Cay, Spellman & Stoddart 2494
(US, MO).
Lagoon Cays. -- Cay Chapel, Stoddart 442 (Fo).

PORTULACACEAE

Portulaca L.

Portulaca oleracea L. (Purslane, Pusly)
Atoll Cays. -- Turneffe Atoll: Soldier Cay, Stoddart sight record
in, UI62 {CUICS p< fD)ie Lighthouse Reef: Hat Cay, Stoddart 56 (Fo).

Glover's Reef: Southwest Cay II, Fosberg & Sachet 53868a (US), Linhart
(SSO; ps dies).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart 110 (Fo) (1963,
p. 42), Stoddart sight record in 1972 (1974, p. 478), Spellman & Stoddart
2554 (US, MO); Goff's Cay, Spellman & Stoddart 2540 (US, MO); English
Cay, Spellman & Stoddart 2513 (US, MO), Dwyer et al. 669 (MO);
Rendezvous Cay, Stoddart 108 (Fo) (1963, p. 49), Stoddart sight record
in 1965 .(1969,,. p.,,9);,..Spellman .& .Stoddart, -2504 —(US;. MO) >... \Tobacco).Cay,,
Stoddart .510. (Fo), Stoddart sight records in, 196] = 1962 6(1963% p.-7 55)
and 1965, Fosberg & Spellman 54254 (US); South Water Cay, Spellman &

29

Stoddart 2147 (US, MO), 2166 (US, MO), Pringle 1896 (HAM, MO); Carrie
Bow Cay, Fosberg & Spellman 54436 (US), Spellman & Stoddart 2137 (US,
MO), Ferraris 8 (US); North Silk Cay, Fosberg & Spellman 54271 (US);
South Silk Cay, Fosberg & Spellman 54264 (US); Round Cay, Fosberg &
Spellman 54300 (US), 54305 (US); Pompion Cay, Fosberg & Spellman
54314 (US); Ranguana Cay, Spellman & Stoddart 2246 (US, MO); Tom
Owen's Cay, Spellman & Stoddart 2276 (US, MO); Tom Owen's West Cay,
Spellman & Stoddart 2281 (US, MO); Hunting Cay, Spellman & Stoddart
2442 (US) (seedling); Ragged Cay, Spellman & Stoddart 2388 (US, MO);
Seal Cay, Spellman & Stoddart 2491 (US, MO).

Lagoon Cays. -- St George's Cay, Spellman 1480 (US, MO);
Buttonwood Cay, Stoddart 91 (Fo) (1963, p. 64), Fosberg & Spellman 54424
(US); Hatchet Cay, Fosberg & Spellman 54363 (US); Little Water Cay,
Fosberg & Spellman 54350 (US).

Portulaca sp.

Atoll Cays. -- Glover's Reef: Southwest Cay I, Stoddart sight
igexeronctel akigy ALS)7/ IL

Barrier Reef Cays. -- Goff's Cay, Stoddart sight record in 1962
@isGsinup. 447, VWI69 Pop. 9, ,aAswe. oleracea =); “English iCay, Sitoddart sight
record vin LI65= (9697p. Q)i- saNorth Spot, Stoddart sight record in
1960; Tom Owen's East Cay, Stoddart sight record in 1960; Tom Owen's
West Cay, Stoddart sight record in 1960.

AIZOACEAE
Sesuvium L.
Sesuvium portulacastrum (L.) L. (Seaside Purslane)
Atoll Cays. -- Turneffe Atoll: Mauger Cay, Stoddart sight record

im LICO RdGoZavinp = 47 1963)" Pp. ol); ie PelvcaniCay, Stoddart isight record
in 1960 (1962a, p. 45); Cockroach Cay V, Stoddart sight record in 1965
(1969, p. 13); Cockroach Cay, Stoddart sight records in 1962 and 1965
(1969), p. 13); Pelican Cay (Cockroach Group), Stoddart sight record in
IGS) (i969 op... 13))-) BlackbirdiCay, Stoddart. sighterecordain 960" (1962),
p. 44); Soldier Cay, Stoddart sight records in 1960 and 1965 (1962a,

pee 44,, e963), op. Dp alOGO ip. 2lS)p OsHarry, Jones! SPoimt 7 “Stoddartasight
record in 1960 (1962a, p. 48); Big Calabash East Cay I, Stoddart sight
record in 1960 and 1962 (1969, p. 14); Little Calabash Cay, Stoddart
Sight record in 1965 (1969, p. 14); Deadman Cay I, Stoddart sight
TEGO. alin UYGO) ial” IHS (UGS Zey OG BiSin ISIS, og. WS MoS) 7 fod eabsyis
Deadman Cay II, Stoddart sight records in 1960, 1962, and 1965 (1962a,
De Soy peLIGsy Deco), p LICE woe; = DeadnanmCay, HilEhy ¥S coddantesi gin
records in W96OcandelI62 e Gig62ia, apis 39), L963), pe ob) DeadmanaCay LV;
Stoddart sight neconds! anel96O), \V962 vand- 1965" (ilseWa, 44743897 =963 7 4p 5 Sil,
1969, p. 15); Deadman Cay V, Stoddart sight record in 1960 (1962a,

Do SI p- NGOS, jo. SZ) % Lighthouse Reef: Sandbore Cay, Stoddart sight

05!

Swill CIOUINTAIN TINO TEIPULIUIN eur arinicoINt INVIINUSMLIVVD

ae oe oe eo a eo eee ee Gee |

ea "wea eae 8

30

record in 1965 (1969, p. 16); Northern Cay, Stoddart sight records in
1960 and 1961 (1962a, p. 62); Saddle Cay, Stoddart sight record in 1960
(1962a, pi (63; 1963, <p. 94) 7° Half Moon Cay, Stoddart (46 (Fo)l, 7449" (kop
(19620; p.875, L969) .p-517) 7 ae LongsCay jsStoddant siaghtenecords Fine! I6Or
1961, andi 1962: (1962a,;\-p4' 80, L963 mips IO) Hak) Cay , (Stoddart co 7mitlyova,
De Cm I6 3, spe LOO)t Glover's Reef: Northeast Cay, Stoddart 75 (Fo),
Fosberg '& Sachet 53835) (US) ((1962a, ps 188); sbinhart (i98Opap. 1162)F-

Small Cay (Long Cay North) , Stoddart sight record in 1961 (1962a, p: 89),
Fosberg & Sachet 53790 (US), Linhart (1980, p. 162); Long Cay, Fosberg
& Sachet 53910 (US), Linhart (1980, p. 162); Middle Cay, Fosberg &
Stoddart 53920 (US); Southwest Cay I, Stoddart sight record in 1971,
Linhart (1980,p.%l62)%)> {Southwest Cay oll, «Stoddart sightvnecordpin lg 6m
(1962a;:p. 97), *Fosberg=é .Sachet 53878) (US)I, Linharto(l98o7 ip. Sle2)r

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in I960' (1963, p. 38, 1969, ps 7); Sexgeant’s Cay, Stoddarak Miia (Ho))

(1963, p. 42), Spellman & Stoddart 2547 (US, MO, BH); Goff'sCay;
Stoddart sight records’ in 1960 and 11965 (1963, ip. 43, 1974, pea)

Spellman & Stoddart 2538 (US); English Cay, Stoddart sight record in
1965 (1969, p. 9), Spellman & Stoddart 2516 (US; MO, BH), Dwyerretraie
667a (MO); Rendezvous Cay, Stoddart 101 (Fo), Stoddart sight records

in’ 1960 and 1965 (1963, -p. .48,/1969, p. 9), Spellman & Stoddart 72505
(US: { 2MO) ; “Cay Glory, iStoddart (51/35 (Ho)i=(1962b, ip. 164751963 hip es2)u
Tobacce Cay, Stoddart sight records in 1961 and 1965 (1963, ps 54, 1969)
p. 10), Fosberg & Spellman 54234 (US); South Water Cay, Stoddart sight
records in 1960 and 1965 (1963, p. 56, 2969; p- 21')), Speliimanis Stoddart
2148 (US) Pringle 1867 (HAM); Carrie Bow Cay, Stoddart sight record in
1965 (1969, pp. 11), Spellman &*Stoddart: 2136) US), Ferraris 7: \(uSii-

North Silk Cay, Stoddart sight record in 1961, Fosberg & Spellman 54268
(US); Middle Silk Cay, Stoddart sight record in 1961, Fosberg &
Spellman 54288 (US); South Silk Cay, Stoddart sight record in 1961,
Fosberg & Spellman 54260 (US); Round Cay, Stoddart sight record in
1960, Fosberg & Spellman 54306 (US); Pompion Cay, Stoddart sight record
in 1960, Fosberg & Spellman 54318 (US); Ranguana Cay, Spellman &
Stoddart 2258 (US);" \Nomth Spot, Stoddart sight record gin’ 1960;

Spellman & Stoddart 2291 (US); Northeast Sapodilla Cay, Stoddart sight
record in 1960 (1962b, p. 164); Tom Owen's East Cay, Stoddart sight
record in 1960, Spellman & Stoddart 2280 (US); Tom Owen's West Cay,
Stoddart sight record in 1960, Spellman & Stoddart 2277 (US); Lime Cay,
Stoddart sight record in 1960 and 1972; Ragged Cay, Spellman & Stoddart
2386 (US, MO, BH), Stoddart sight record in 1960.

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1962;
St George's Cay, Stoddart sight records in 1960 and 1965 (1963, p. 38,
1969, p. 7), Spellman 1458 (US, MO); ‘Tobacco Range, Fosberg & Spellman

54465 (US); Man-o'-War Cay, Fosberg & Spellman 54207 (US); Jack's Cay,
Fosberg & Spellman 54430 (US); Buttonwood Cay, Fosberg & Spellman 54426
(US); Hatchet Cay, Fosberg & Spellman 54399 (US); Little Water Cay,
Fosberg & Spellman 54356 (US); Trapp's Cay, Stoddart sight record in

1962; Golson Cay, Stoddart. sight record n<71965 “(W9697pe 12): > ‘Scipio
Cay, Stoddart sight record in 1965 (1969, p.'12); Maughing Bird=Cay,

stoddart sight record in 1962 (19637. p: 65) +V BuglesCay,,7:Stoddart ssight
record in 1965. /(1969)"p2, 1h)s71)7 West «Snake Cay); Stoddart sight recordin

Shi

1961 (1965, p. 137); West Snake Cay West Islet, Stoddart sight record
in 1961; Middle Snake Cay North Islet, Stoddart sight record in 1961:
Middle Snake Cay South Islet, Stoddart sight record in 1961.

LAURACEAE
Cassytha L.
Cassytha filiformis L. (Love vine)
Atoll Cays. -- Lighthouse Reef: Sandbore Cay, Stoddart sight

meconrd aniw965) (1969), p. 16)';) Northern) Cay, Stoddart sight record’ in
1960; Half Moon Cay, Stoddart 32 (Fo); Long Cay, Stoddart sight
records in 1961 and 1965 (1969, p. 18).

Barrier Reef Cays. -- South Water Cay, Stoddart sight records in

1960 and 1965, Spellman & Stoddart 2163 (US, MO), Pringle 1861 (HAM) ;
Ranguana Cay, Spellman & Stoddart 2256 (US, MO).

Lagoon Cays. -- Cay Chapel, Stoddart sight record in 1965 (1969,
Demi Collison Cay. Stoddart sightimecord in 1965) (1969), p- 12).

CRUCIFERAE
Cakile Mill.
Cakile lanceolata (Willd.) O. E. Schulz (Sea rocket)
Atoll Cays. -- Turneffe Atoll: Cockroach Cay, Stoddart sight

record in 1962; Little Calabash Cay, Stoddart sight record in 1962
CSCS pi) 7) Deadman, Caya my, Stoddart 127 (Fo) (19637, pp. 80).
Lighthouse Reef: Sandbore Cay, Stoddart sight record in 1962 (1963,
p-. 91); Northern Cay, Stoddart sight records in 1960 and 1962 (1963,
Dey 23). Half Moone Cay, Stoddart 132 (Fo), Stoddart sight records in
IQ (UGeZery ids Wp Wass fos So Glover's Reef: Southwest Cay II,

Stoddart sight record in 1961 (1962a, p. 98), Fosberg & Sachet 53883
(US).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart 439 (Fo) (1969,
p- 8), Spellman & Stoddart 2548 (US); Goff's Cay, Spellman & Stoddart
2536 (US, MO) (1974, p. 479); Rendezvous Cay, Stoddart 104 (Fo),
Stoddart. sightmecord anil 965 (1963); pp.) 497,.119697 p29); Cay Glory,
Stoddart 509 (Fo) (1963, p. 52); South Water Cay, Stoddart sight record
in 1960 (1969, p. 11), Spellman & Stoddart 2200 (US, MO), Pringle 1884
(HAM); Carrie Bow Cay, Spellman & Stoddart 2575 (US) (seedling),
Ferraris 2; Round Cay, Stoddart sight record in 1960, (US); Ranguana
Cay, Spellman & Stoddart 2261 (US, MO); Pompion Cay, Stoddart sight
record in 1960; Northeast Sapodilla Cay, Stoddart 507 (Fo); Tom Owen's

roe

IWYCLIILIZON! INVIINUSALIVYS

RA UnrmAnmniCocwd QIVILITIOVIVIATY TIWOJVITUOULIVUIN |

vw eG Vw Gwe 8

32

West Cay, Stoddart sight record in 1960; Frank's West Cay, Stoddart
sight record in 1960; Frank's East Cay, Stoddart sight record in 1960;
Hunting Cay, Spellman & Stoddart 2473 (US, MO).

Lagoon Cays. -- Cay Chapel, Stoddart sight record in 1965 (1969,
p. 7); St George's Cay, Stoddart sightyrecord in 1962 (1963), p...40;
1969, p. 7); Hatchet Cay, Fosberg & Spellman 54385 (US); Middle Snake
Cay North Islet, Stoddart sight record in 1961; West Snake Cay,
Stoddart 19° (Fo)- (1965), p. 137).

CAPPARIDACEAE
Capparis L.
Capparis flexuosa (L.) L. (Bottle-cod root)
Atoll Cays. -- Lighthouse Reef: Half Moon Cay,Verner (1959, 3-8),
Stoddart TOlLS62Zai, ip. 5)
CRASSULACEAE
*Kalanchoe Adans.
*Kalanchoe pinnata (Lam.) Pers. (Air plant, leaf of life)

Bryophyllum pinnatum (Lam.) Oken
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2460 (US,
MO).

ROSACEAE

Chrysobalanus L.

Chrysobalanus icaco L. (Coco plum, Icaco plum)
Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg & Sachet

53830. (US) ,, Linhart! (1980;<p .= 163):

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2207
(US, MO); Northeast Sapodilla Cay, Spellman & Stoddart 2308 (US, MO,
BH); Frank's Cay, Spellman & Stoddart 2412 (US, MO, BH); Nicolas Cay,
Spellman & Stoddart 2329 (US, MO, BH); Hunting Cay, Spellman & Stoddart
2456 (US, MO).

33
LEGUMINOSAE
Caesalpinia L.
Caesalpinia bonduc (L.) Roxb. (Nicker nuts, ticksie, nickol)
Caesalpinia crista L.
Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54237 (US)
(seedling); Tom Owen's West Cay, Spellman & Stoddart 2262 (US, MO,
BH) .
Canavalia DC.
Canavalia rosea (Sw.) DC. (Seaside bean)

Canavalia maritima (Aubl.) Urb.

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 59 (Fo),
AVEvils (Gate). AOUSYOS A roc ske)))4 Glover's Reef: Southwest Cay II, Fosberg &
Sachetyose79" (US), binhante (198075. 163) as Cy maritima);

Barrier Reef Cays. -- Sergeant's Cay, Spellman & Stoddart 2552
(US) MO; BH, Fo); Tobacco Cay, Stoddarty515) (Fo)), S16 (Fo), (1963,
p. 54, 1969, p. 10), Fosberg & Spellman 54250 (US); South Water Cay,
Spellman & Stoddart 2184 (US, MO), Pringle 1863 (HAM); Tom Owen's West
Cay, Stoddart sight record in 1960; Hunting Cay, Spellman & Stoddart
ZA38) (US; MO) <

Lagoon Cays. -- St George's Cay, Spellman 1466 (US, 2 sheets, MO).
Canavalia sp.

Atoll Cays. -- Turneffe Atoll: Cockroach Cay, Stoddart sight
BECOnd yin 1965 S69 R ip. 13) Lighthouse Reef: Sandbore Cay,
Stoddart sight record in 1965 (1969, p. 17).

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 40); Goff's Cay, Stoddart sight record in 1960
(MQSSE MoS HSA
Crotalaria L.

Crotalaria retusa L. (Rattle-pod, rattleweed)

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2164
(US, MO), Pringle 1883 (HAM).

re |

INVINUOMLIVVO

UWI LIP ILILOIVI

—

HWOTIIUTLIUIN

Vivid VEIOWIVIFAIN
Sey lh a a re ae ee ee

eieUVIVerivniungd

34

Crotalaria verrucosa L. (Rattle-pod, blue rattleweed)

Lagoon Cays. -- Cay Caulker, Stoddart 407 (Fo) (1969, p. 6).

Dalbergia L. f.
Dalbergia ecastophyllum (L.) Taub.

Lagoon Cays. -- South Snake Cay, Stoddart 93 (Fo) (1965, p. 138).

Desmodium Desv.
Desmodium incanum DC. (Beggar's lice)
Desmodium canum (Gmel.) Schinz & Thell.

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2165
(US): MO},) BH)E 2202) (US)... prangllesto07) (HAM):

Desmodium tortuosum (Sw.) DC.

Barrier Reef Cays. -- South Water Cay, Pringle 1922 (HAM).

Erythrina L.

Erythrina sp. (Coral tree)
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2205
(Fo)F
Lagoon Cays. -- East Snake Cay, Stoddart 98 (Fo).

Gliricidia Kunth ex Endl.
Gliricidia sepium (Jacq.) Kunth ex Griseb. (Quickstick, St Vincent plum)
Barrier Reef Cays. -- Frank's Cay, Spellman & Stoddart 2402 (US);
Hunting Cay, Spellman & Stoddart 2439 (Fo, MO).
Leucaena Benth.
Leucaena leucocephala (Lam.) de Wit (Lead tree)
Leucaena latisiliqua sensu Gillis & Stearn non (L.) Gillis

Leucaena glauca sensu auct. non (L.) Benth.

bole ie |

2
35 5
E
5
Atoll Cays. -- Turneffe Atoll: Grand Bogue Point, Stoddart 131 3
(HO) (AGE Sn- 1s MIS))- Z
;
Mucuna Adans. i
5
Mucuna sp. (Horse eye bean, sea bean) >
I
Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg sight record =
in 1971; Long Cay, Fosberg & Sachet 53796 (US). is
ois ees ag yeu 2)
Pithecellobium Mart. :
Pithecellobium keyense Britt. :
>
Pithecolobium keyense Britt. ex Coker 12
Atoll Cays. -- Lighthouse Reef: Northern Cay, Stoddart 450 (Fo) uy
GiSGI es Dawlw))-aeHalbteMoon.Cay,—verner (19617 4p. 582) 7 Stoddart (1962a,
Dee) Glover's Reef: Northeast Cay, Linhart (1980, p. 163); Middle
Cay, Fosberg & Stoddart 53917 (US), Linhart (1980, p. 163).

Barrier Reef Cays. —-- Water Cay, Dwyer et al. 673 (MO); Frank”s
Cay, Stoddart sight record in 1972; Frank's Cay East, Stoddart sight
record in 1972; Nicolas Cay, Spellman & Stoddart 2339 (US, MO, BH);
Hunting Cay, Spellman & Stoddart 2453 (US, MO); Lime Cay, Spellman &
Stoddart 2367 (US, MO).

[er Oe oe |

Lagoon Cays. -- Cay Caulker, Stoddart 424 (Fo) (1969, p. 6); ;
Tobacco Range, Fosberg & Spellman 54474 (US), 54475 (US).

Sophora L. )
Sophora tomentosa L.

Atoll Cays. -- Turneffe Atoll: Pelican Cay (Cockroach Group),
Stoddart, sight necord in 1965 (1969) p. 13); Cockroach Cay, Stoddart
sight record in 1965 (seedling) (1969, p. 13); Soldier Cay, Stoddart
sight record in 1965 (1969, p. 14); Deadman Cay I, Stoddart sight record
in 19659 (seeding) (1969), ps-i5) = Lighthouse Reef: Sandbore Cay, I
Stoddart sight records in 1961 and 1965 (1969, p. 16); Long Cay,
Stoddart sight records in 1960, 1961 and 1965 (1962a, p. 80, 1969, p. 18).
Glover's Reef: Small Cay (Long Cay North), Stoddart sight record in
LOGIE (UI6Zay pec); sLong. Cay, stoddanty/9) (Fo) 7s) Middle Cay), "Sachet &
Stoddart) 635m (US), sbinhare «W980; =p: 163).

Barrier Reef Cays. -- Tobacco Cay, Stoddart sight record in 1965 :
(1969, p. 10); South Water Cay, Stoddart sight record in 1965, Spellman 4
& Stoddart 2153 (US, MO, BH, Fo), Pringle 1886 (HAM); Ranguana Cay,
Stoddart 501 (Fo); Northeast Sapodilla Cay, Spellman & Stoddart 2314 3
(US, MO); Frank's Cay, Spellman & Stoddart 2393 (US, MO, BH); Nicolas

36

Cay, Spellman & Stoddart 2352 (US, MO, BH, Fo, BM); Hunting Cay,

Spellman & Stoddart 2459 (US); Lime Cay, Spellman & Stoddart 2371
(US, MO, BH).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965 (1969,
p. 6); Tobacco Range, Fosberg & Spellman 54453 (US), 54470 (US); Peter
Douglas Cay, Stoddart sight record in 1961 (1963, p. 61); Little Peter

Cay, Stoddart sight record in 1961 (1963, p. 61); Colson Cay, Stoddart
sight record in 1965; Scipio Cay, Stoddart sight record in 1965; East
Snake Cay, Stoddart 97 (Fo) (1965, p. 138); Middle Snake Cay North
Islet, Stoddart sight record in 1961; South Snake Cay, Stoddart sight
record in 1961: (1965,°p. 138); Moho Cay, Stoddart 1 “(ho);-

*Tamarindus L.

*Tamarindus indica L.? (Tamarind)
Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2529

(seedling) (US).

Vigna Savi
Vigna luteola (Jacq.) Benth. (Beach pea)
Vigna repens (L.) Kuntze

Barrier Reef Cays. -- Tobacco Cay, Stoddart 115 (Fo), 518 (Fo),
Stoddart sight in 1965 (1963, p. 54), Fosberg & Spellman 54240 (US);
South Water Cay, Spellman & Stoddart 2173 (US, MO, BH), 2217 (US)
(seedling), Pringle 1874 (HAM); Ranguana Cay, Spellman & Stoddart 2260
(US, MO); Hunting Cay, Spellman & Stoddart 2435 (US, MO).

Lagoon Cays. -- Cay Caulker, Stoddart 405 (Fo); Buttonwood Cay,
Stoddart 88 (Fo), Fosberg & Spellman 54407 (US); Hatchet Cay, Fosberg &
Spellman 54394 (US); Little Water Cay, Fosberg & Spellman 54353 (US);
Moho Cay, Stoddart 2 (Fo).

RUTACEAE
*Citrus. L.
*Citrus aurantiifolia (Christm.) Swingle (Lime)
Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2327 (US,

MO, BH); Hunting Cay, Spellman & Stoddart 2440 (US, MO).

Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54367 (US).

Sh7/

*Citrus sp.

Lagoon Cays. -- Wild Cane Cay, Stoddart sight record in 1961.

SURIANACEAE

Suriana L.
Suriana maritima L. (Bay Cedar)

Atoll Cays. -- Turneffe Atoll: Dogflea Cay, Stoddart sight record
in 1960 (1962a, p. 46); Cockroach Cay, Stoddart sight record in 1965
ICO pe 2S); Cockroach Cay V7, Stoddare sight record in 1965 (19697,
p. 13); Soldier Cay, Stoddart sight records in 1960 and 1965 (1962a,
PEt GOS), Pe 1/5), 1969), Dp. 13); Harry vones Point, Stoddart sight
record in 1960 (1962a, p. 48); Big Calabash East Cay I, Stoddart sight
mecora in 1960 (1962a, p. 42, 1963, p. 78); Big Calabash East Cay II,
Stoddart sight record in 1960 (1962a, p. 42, 1963, p. 78); Little
Calabash Cay, Stoddart sight record in 1965 (seedlings) (1969, p. 14);
Deadman Cay II, Stoddart sight record in 1960 (1962a, p. 38, 1963,
p. 80); Deadman Cay III, Stoddart sight record in 1960; Deadman Cay
IV, Stoddart sight recordsin 1960 and 1965 (1969, p. 15); Deadman Cay
V, Stoddart sight record in 1965; Big Cay Bokel, Stoddart sight record
TnetO65* Lighthouse Reef: Sandbore Cay, Stoddart sight records in
Soi 962s ande1965),, Stoddantso4 (Fo), (1962a, p.. 57, 1963, p. 89, 1969,
p-. 16); Northern Cay, Stoddart sight records in 1960, 1961 and 1965
(ioGZay-p. 62, 1963), Dp. 93, 1969, p- 17); Halli Moon Cay, Stoddart. 47
(Hoje (Steddart 1962a,ep. 75, 1963;-—p.—98)— Long. Cay, Stoddart sight
HeCOras) in e960 Vand 1965 (196Z2a, 7p. 80), 1969) sp. 18); Hat Cay, Stoddart
sight record in 1961 (1962a, p. 81, 1963, p. 100). Glover's Reef:
Northeast Cay, Stoddart sight record in 1961 (1962a, p. 88); Small Cay
(Long Cay North), Stoddart sight record in 1961 (1962a, p. 89).
Fosberg & Sachet 53775 (US, MO, BH, Fo), Linhart (1980, p. 163); Long
Cay, Stoddart sight record in 1961 (1962a, p. 91), Fosberg & Sachet
53909 (US, Fo), Linhart (1980, p. 163); Middle Cay, Sachet & Stoddart
1623 (US), Linhart (1980, p. 163); Southwest Cay I, Stoddart sight
records in 1961 and 1971, Linhart (1980, p. 163); Southwest Cay II,
Fosberg & Sachet 53893 (US, Fo), Linhart (1980, p. 163).

Barrier Reef Cays. -- Sergeant's Cay, Spellman & Stoddart 2551
(US; MO; BH); Stoddart sight rwecord in 1965 (1969; p-"8,-1974, p. 4278) ;
English Cay, Spellman & Stoddart 2530 (US, MO); Rendezvous Cay,
Spellman & Stoddart 2496 (US, MO), Stoddart sight record in 1965 (1969,
p. 9, 1974, p. 480); Tobacco Cay, Fosberg & Spellman 54217 (US); South
Water Cay, Stoddart sight record in 1960 (1963, p. 56), Pringle 1869
(HAM); North Silk Cay, Stoddart sight record in 1961, Fosberg &
Spellman 54266 (US); Middle Silk Cay, Stoddart sight record in 1961,
Fosberg & Spellman 54287 (US); South Silk Cay, Stoddart sight record in
1961, Fosberg & Spellman 54263 (US); Round Cay, Stoddart sight record in
1960, Fosberg & Spellman 54292 (US); Pompion Cay, Stoddart sight record

EUV CLIILZILIIVI INVIINUOMLIIVGS Pe eee i |
—

HWOTTIUTIIUIN

VIVE DT EIOUWITIFAIY

a oe

38

in 1960, Fosberg & Spellman 54324 (US); Ranguana Cay, Stoddart sight
record in 1960; Northeast Sapodilla Cay, Spellman & Stoddart 2315
(US, MO).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965 (1969,
p. 6); Cay Caulker, Stoddart 421)\(Fo) “(1969;,.p2"6) 7 “Cay Chapels;
Stoddart ‘sight records an e962 and 2965" (1963, p. 357 19697 pas);
Robinson Point Cay, Stoddart sight record in 1961 (1963, p. 46);
Tobacco Range, Fosberg & Spellman 54472 (US); Coco Plum Cay, Spellman
& Stoddart 2566 (US, MO); Water Range, Spellman & Stoddart 2233 (US,
MO, BH, Fo); Little Water Cay, Fosberg & Spellman 54352 (US); Middle
Snake Cay North Islet, Stoddart sight record in 1961; Middle Snake Cay
South Islet, Stoddart sight record in 1961; West Snake Cay West Islet,
Stoddart sight record in 1961; South Snake Cay, Stoddart sight record
im, LOGI L965) 42D.)

BURSERACEAE
Bursera Jacq. ex L.
Bursera simaruba (L.) Sarg. (Red birch, Gumbo-limbo)
Atoll Cays. -- Turneffe Atoll: Pelican Cay (Cockroach Group),

Stoddart sight records in 1962 and 1965 (1963, p. 72, 1969, .p. 13):
Lighthouse Reef: Half Moon Cay, Verner (1959, pp. 3-8, 1961, p. 582),
Stoddart sight records in 1960, 1961, and 1965 (1962a, p. 74, 1963, p. 95,
HI69 jes el Opa Glover's Reef: Northeast Cay, Stoddart sight record in
1961 (1962a, p. 88), Fosberg & Sachet 53834 (US), 53847 (US), Linhart
(P9807, ps 2162):

Barrier Reef Cays. -- Carrie Bow Cay, Fosberg & Spellman 54440
(US) (seedling); Pompion Cay, Stoddart sight record in 1960, Fosberg
& Spellman 54323 (US); Tom Owen's East Cay, Stoddart 506 (Fo).

MELIACEAE
Swietenia Jacq.
Swietenia macrophylla King (Honduras mahogany, mahogany)
Atoll Cays. -- Turneffe Atoll: Soldier Cay, Romney et al. (1959,

p. 254), Stoddart (1962a, p. 44: not seen), Calabash Cays, Romney et
al. (1959, p. 254), Rope Walk, Romney et al. (1959, p. 254).

roe a |

33

NYIINUSMLIVVS

EUPHORBIACEAE

*Codiaeum Rumph. ex Juss. :
*Codiaeum variegatum ee) Blume (Croton) :
>
Barrier Reef Cays. -- Hunting Cay, cult., Spellman & Stoddart 2475 i
(US, MO). z
Drypetes Vahl =
Drypetes brownii Standl. i

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 60 (Fo).

Sivid VETOUWIVIPAIN

Euphorbia L.
Euphorbia blodgettii Engelm. ex Hitchc.

Chamaesyce blodgettii (Engelm. ex Hitchc.) Small

weiewVInNnNtud

Atoll Cays. -- Glover's Reef: Northeast Cay, Stoddart 70 (Fo),
Fosberg & Sachet 53843 (US, MO, BH, Fo, BM), 53822 (US, MO, BH, Fo), E
§3823 (US), 53828 (?identification) (US); Small Cay (Long Cay North), ,
Fosberg & Sachet 53782 (US, MO, BH, Fo, BM), 53786 (US, MO, BH, Fo, BM);

Long Cay, Linhart (1980, p. 163, ?as Euphorbia sp.); Middle Cay,
Sachet & Stoddart 1611 (US, MO, BH, Fo); Southwest Cay I, Pippen 902 ;
(US); Southwest Cay II, Fosberg & Sachet 53887 (US, MO, Fo, BH),

53876 (US, MO, BH, Fo, BM), Stoddart 84 (Fo).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight record in )

1965 (1969, p. 8); English Cay, Spellman & Stoddart 2515 (US, MO, BH, 2

Fo), 2518 (US, MO, BH, Fo, BM); Tobacco Cay, Fosberg & Spellman 54235
(US;eMO}, BHAeHO) 165423202 (US) 7554247 (US, MO)), 54255. (US, #MO, SBH) SEo,

BM), Stoddart 114a (Fo) (1963, p. 55); South Water Cay, Spellman &

Stoddart 2142a (US, MO, BH, Fo), 2178 (US, MO, BH), Stoddart sight

j

=

record in 1960 (1963, p. 55), Pringle 1899 (HAM); Carrie Bow Cay,
Spellman & Stoddart 2138 (US, MO, BH), Ferraris 5 (US); Ranguana Cay,
Spellman & Stoddart 2243 (US, MO, BH, Fo); Northeast Sapodilla Cay,
Spellman & Stoddart 2320 (US, MO); Frank's Cay, Spellman & Stoddart
2411 (US, MO, BH, Fo, BM); Hunting Cay, Spellman & Stoddart 2431 (US).

Lagoon Cays. -- Cay Caulker, Stoddart 418 (Fo) (1969, p. 6); 5
Tobacco Range, Fosberg & Spellman 54466 (US, MO, BH, Fo, BM); Buttonwood :
Cay, Fosberg & Spellman 54404 (US, MO, BH, Fo, BM), 54406 (US, MO, BH),

Stoddart 85 (Fo); Hatchet Cay, Fosberg & Spellman 54380 (US, MO, BH,
Fo, BM); Little Water Cay, Fosberg & Spellman 54345 (US, MO, BH, Fo, )
BM), 54330. (US, MO, BH, Fo, BM); Moho Cay; Stoddart 6 (Fo).

40

Euphorbia glomerifera (Millsp.) Wheeler

Chamaesyce glomerifera Millsp.

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54225 (US);
South Water Cay, Stoddart 522 (Fo).

(Stearn also determines Stoddart 6, from Moho Cay, as this, but
our specimen is E. blodgettii. The same is true of no. 85 from
Buttonwood Cay.

Euphorbia mesembrianthemifolia Jacq.
Euphorbia buxifolia Lam.
Chamaesyce buxifolia (Lam.) Small

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 33 (Fo),
D8 (Fo)” (196375 ps5 98). Glover's Reef: Northeast Cay, Stoddart 74
(Fo), Fosberg & Sachet 53811 (US, MO, BH, Fo, MO), 53837 (US, MO, BH,
Fo, BM), Linhart (1980, p. 163); Small Cay (Long Cay North), Fosberg
& Sachet 53788 (US, MO, BH), 53787 (US, MO, BH, Fo) (prostrate form),
Linhart (1980, p. 163); Long Cay, Fosberg & Sachet 53797 (US, MO, BH),
Linhart (1980, p. 163); Middle Cay, Fosberg, Sachet & Stoddart 53925
(US, MO, BH, Fo, BM), Sachet & Stoddart 1610 (US, MO, BH, Fo), Linhart
(1980, p. 163); Southwest Cay, Pippen 901 (US); Southwest Cay I,
Linhart (1980, p. 163); Southwest Cay II, Fosberg & Sachet 53885 (US,
MO,'°BH)*;' Linhart: (19807)<p..-163)i:

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 40); Sergeant's Cay, Stoddart 109 (Fo), Spellman &
Stoddart 2546 (US, MO, BH, Fo), 2550 (US, MO, BH), Stoddart sight
record in 1965 (1963, p. 42, 1969, p. 8); Goff's Cay, Spellman &
Stoddart 2533 (US, MO, BH, Fo, BM); English Cay, Dwyer et al. 670
(MO), Spellman & Stoddart 2514 (US, MO, BH), 2517 (US, MO, BH, Fo, BM);
Rendezvous Cay, Stoddart 102 (Fo) (1963, p. 49), Spellman & Stoddart
2500 (US, MO, BH, Fo, BM); Cay Glory, Stoddart 502 (Fo) (1963, p. 52);
Tobacco Cay, Stoddart 114 (Fo), Fosberg & Spellman 54239 (US, MO),
54249 (US, MO); South Water Cay, Spellman & Stoddart 2180 (US, MO, BH,
Fo), Pringle 1876, 1880 (HAM); Carrie Bow Cay, Spellman & Stoddart
2139 (US, MO, BH), Ferraris 4 (US); Middle Silk Cay, Fosberg & Spellman
54278, 54281, 54283 (all US, MO); South Silk Cay, Fosberg & Spellman
54257 (US, MO); Round Cay, Fosberg & Spellman 54301 (US, MO); Pompion
Cay, Fosberg & Spellman 54319 (US, MO), 54321 (US, MO); Ranguana Cay,
Spellman & Stoddart 2245 (US, MO, BH); Tom Owen's East Cay, Spellman
& Stoddart 2287 (US, MO, BH, Fo, BM); Frank's Cay, Spellman &

Stoddart 2395 (US, MO, BH, Fo, BM); Nicolas Cay, Spellman & Stoddart
2344 (US, MO, BH, Fo, BM); Hunting Cay, Spellman & Stoddart 2424 (US,
MO); Lime Cay, Spellman & Stoddart 2361 (US, MO, BH).

4

2)

&

41 z

n

=

lagoon ‘ays. —— Cay, Caulker, Stoddart 427 “(Fo) \(t969), p. 6) ; >

St George's Cay, Spellman 1478 (US, MO) (ordinary more or less erect E
form), 1479 (US, MO) (habit diffuse, leaves large, ovate, oblique at z
base); Water Cay, Dwyer et al. 677 (MO); Tobacco Range, Fosberg & 4
Spellman 54460 (US, MO); Coco Plum Cay, Spellman & Stoddart 2558 (US, zg
MO, BH, Fo, BM); Water Range, Spellman & Stoddart 2229 (US, MO, BH, 4
Fo); Buttonwood Cay, Fosberg & Spellman 54400 (US, MO), 54402 (US, "
MO); Hatchet Cay, Fosberg & Spellman 54388 (US, MO); Little Water Wy
Cay, Fosberg & Spellman 54337 (US, MO). >
3

*Euphorbia prostrata Ait. E
2)

Chamaesyce prostrata (Ait.) Small z
Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54219 (US). :

>

9

*Euphorbia thymifolia L. - :
Chamaesyce thymifolia (L.) Millsp. F

)

J

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2519 (US, E

MO, BH, Fo, BM). c
3

Euphorbia trichotoma Kunth }
Atoll Cays. -- Glover's Reef: Southwest Cay I, Stoddart 81 (Fo). :

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2203 }
(US, MO, BH, Fo), Pringle 1900, 1901 (HAM).

Euphorbia sp. 4

Atoll Cays. -- Turneffe Atoll: Three Corner Cay, Stoddart sight
record in 1962 (1963, p. 71); Cockroach Cay, Stoddart sight records ;
inet 960) andl962°0G1 962a, p.-'46, 1963), p.> 7377 19690) pr als) 7 2 Cockroach ,
Cay V, Stoddart sight record in 1965 (1969, p. 13); Soldier Cay, I
Stoddart sight records in 1960 and 1965 (1962a, p. 44, 1963, p. 75, 1969,
p. 13); Harry Jones Point, Stoddart sight record in 1960 (1962a, !
p. 48); Big Calabash Cay, Stoddart sight record in 1965 (1969, p. 14);
Big Calabash East Cay I, Stoddart sight record in 1960 (1969, p. 14);
Little Calabash Cay, Stoddart sight record in 1965 (1969, p. 14);

Deadmany Cay, ls Stoddart ‘sighterecord in 1962, (1962a, p= 38, 1963),.p. 79, D
1969, p. 15); Deadman Cay II, Stoddart sight record in 1962 (1962a, |
Desc Les, Dp. 60), 19695 p. 15); Deadman Cay IV, Stoddart: sight )
Becordsming 1960nand 1965 (d962a, pe 3972196387 p.m8l, 19697 p= 15)5
Deadman Cay V, Stoddart sight record in 1962 (1963, p. 82); Big Cay |
Bokel (Stoddart sight records in 1961 and 1965 (1963, p. 83).

Lighthouse Reef: Sandbore Cay, Stoddart sight records in 1960, 1961,
iIGZ Randel IGomGG62Zal—p. 157), 1963),op-. 69), 1 2609,-.p.. £6));, Northern) Cay,
Stoddart sight records in 1960, 1961, 1962 and 1965 (1962a, p. 62, |
1963, p. 92), 1969), p. 17); Half Moon Cay, Stoddart sight record in
1965 (1969, p. 17); Long Cay, Stoddart sight records in 1960, 1961,

42

1962VandmiGE5(962a, pe SO0;e 963,827 99) ~969,5 Dal). Glover's
Reef: Northeast Cay, Stoddart sight record in 1961 (1962a, p. 88),
Linhart (1980, p. 163); Small Cay (Long Cay North), Stoddart sight
record, in 1961 (1962a, p._ 89), Linhart +1980, p. 163);;" LongyCay;
Stoddart sight: record in 1961.) (1962a7 \p.) 91), .binhare (OS0 ipa l6ésnr
Middle Cay, Stoddart sight record in 1961, Linhart (1980, p. 163);
Southest Cay I, Stoddart sight record in 1961 (1962a, p. 95), Linhart
(1980, p. 163); Southwest Cay II, Stoddart sight record in 1961
(1962a;, -p. “97) 7 Linhartey (19807 p. Wes)

Barrier Reef Cays. -- Goff's Cay, Stoddart sight records in 1960
and 1965 (1963, p. 43, 1969, p. 9, as E. mesembrianthemifolia); English
Cay, Stoddart sight record in 1965 (1969, p. 9); Rendezvous Cay,

Stoddart sight records in 1960 and 1965 (1963, p. 48); Tobacco Cay,
Stoddart sight records in 1960 and 1965 (1963, p. 54, 1969, p. 10);
South Water ‘Cay, Stoddart sight records in 1960 and 1965 (1963, p.56,
1969, p. 11); Carrie Bow Cay, Stoddart sight record, in.-1960: (1963,

p. 57); Round Cay, Stoddart sight record in 1960; Pompion Cay,
Stoddart sight record in 1960; Tom Owen's East Cay, Stoddart sight
record in 1960; Frank's Cay, Stoddart sight record in 1960; Frank's
East Cay, Stoddart sight record in 1960; Frank's West Cay, Stoddart
sight record in 1960.

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965 (1969,
p. 6); Cay Chapel, Stoddart sight records in 1962 and 1965 (1963,
p.- 35); St George's Cay, Stoddart sight records in 1962 and 1965 (1963,
p. 38, 1969, p.. 7); Robinson Point Cay, Stoddart sight record in get
(1963), .p. 46); Cat, Cay, Stoddart sight; record in 1962, (1963 ,.p-y 62);
Colson Cay, Stoddart sight recordyin W965 (1969). p.. 12) ; pScipis0 Cay,
Stoddart sight recordin 1965 (19697 p. 12); “Laughing’Bird Cay,
Stoddart sight record in 1962 (1963, p. 65); Bugle Cay, Stoddart sight
record in 1965 (1969, p. 11); Buttonwood Cay, Stoddart sight record in
1961 (1963, pr 64); Spanish Cay, Stoddart, sight record in: 1961;; \West
Snake Cay, Stoddart sight record in 1961 (1965, p. 137); Middle Snake
Cay North Islet, Stoddart sight record in 1961; East Snake Cay,
Stoddart isight. nécord in, 1961 (965, pF 138):

Pedilanthus Poit.
*Pedilanthus tithymaloides (L.) Poit. (Monkey fiddle, shoe flower)

Lagoon Cays. -- Wild Cane Cay, Stoddart 27(Fo).

Phyllanthus L.

Phyllanthus amarus Schum. & Thonn. (Carry-me-seed, Egg woman)
Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2522 (US,
MO, BH); Tobacco Cay, Fosberg & Spellman 54221 (US); South Water Cay,

Spellman & Stoddart 2190 (US, MO, BH, Fo, BM), Pringle 1875 (HAM) ;
Hunting Cay, Spellman & Stoddart 2445 (US, MO, BH).

43
hagoon) CayS..-—-— lGbapps Cay, Stoddart milo (HO) Gl963, pe. 64).
ANACARDIACEAE
*Mangifera L.
*Mangifera indica L.
Lagoon Cays. -- Cay Caulker, Vermeer (1959, p. 58), Stoddart sight

record in 1960 (1963, p. 34); Frenchman's Cay, Stoddart sight record
in 1961; Wild Cane Cay, Stoddart sight record in 1961.

HIPPOCRATEACEAE
Hippocratea L.
Hippocratea volubilis L.
Barrier Reef Cays. -- Northeast Sapodilla Cay, Spellman & Stoddart

2303 (US); Hunting Cay, Spellman & Stoddart 2425 (US).

MALVACEAE
Hibiscus L.
*Hibiscus rosa-sinensis L. (Hibiscus, shoe-black)
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2158
(US).
Hibiscus tiliaceus L. (Seaside mahoe)
Atoll Cays. -- Glover's Reef; Middle Cay, Fosberg & Stoddart
53921 (US).
Barrier Reef Cays. -- Northeast Sapodilla Cay, Spellman & Stoddart

2309 (US, MO); Hunting Cay, Spellman & Stoddart 2429 (US, MO).

Lagoon Cays. -- Cay Caulker, Stoddart 435 (Fo); Buttonwood Cay,
Fosberg & Spellman 54422 (US); Hatchet Cay, Fosberg & Spellman 54384
(US); Little Water Cay, Fosberg & Spellman 54340 (US, MO).

Oivil I CIOUINIATY

te |

NVINUSALIWNS

OWN ILIILILOINI

——

HWOTTIUELIUN

BIVUNMNIOYD

44

Sida L.
Sida acuta Burm. f. (Broom-weed)
Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Vermeer (1959,

Pps, 3-8), steddart: (19624), prew/5) <

Lagoon Cays, -- Cay Caulker, Stoddart 411 (Fo) (sensu lato) (1969,
p. 6); Wild Cane Cay, Stoddart 26 (Fo).

Thespesia Solander ex Correa

Thespesia populnea (L.) Solander ex Correa (Seaside mahoe)
Barrier Reef Cays. -- Tom Owen's West Cay, Spellman & Stoddart

2264 (US, MO); Seal Cay, Spellman & Stoddart 2484 (US, MO).

Lagoon Cays. -- Cat Cay, Stoddart 119 (Fo) (1963, p. 62); Trapp's
Cay, Stoddart sight record in 1962 (1963, p. 64).

STERCULIACEAE
Waltheria L.
Waltheria indica L. (Raichie)
Waltheria americana L.
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2195

(US, MO), Pringle 1877 (HAM), Backman s.n. (MO).

TURNERACEAE
Turnera L.
Turnera ulmifolia L. (Ramgoat dashalong)
Lagoon Cays. -- Cay Caulker, Stoddart 404 (Fo), 419 (Fo) (1969,
iq (6)
PASSIFLORACEAE

Passiflora L.

Passiflora suberosa L. (Passion flower)

45

Atoll Cays. -- Glover's Reef: Northeast Cay, Stoddart 66 (Fo),
Fosberg & Sachet 53819 (US), Linhart (1980, p. 163); Middle Cay,
Middle Cay, Sachet & Stoddart 1618 (US), Linhart (1980, p. 163);
Southwest Cay I, Fosberg & Stoddart 53866 (US), Linhart (1980, p. 163);
Southwest Cay II, Fosberg & Sachet 53886 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2142
(US, MO, BH), Pringle 1924 (HAM, MO); Northeast Sapodilla Cay,
Spellman & Stoddart 2322 (US, MO); Frank's Cay, Spellman & Stoddart
2404 (US, MO); Nicolas Cay, Spellman & Stoddart 2337 (US, MO, BH, Fo,
BM); Hunting Cay, Spellman & Stoddart 2436 (US, MO); Lime Cay,
Spellman & Stoddart 2366 (US, MO).

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54461 (US).

CARICACEAE

*Carica L.

*Carica papaya L. (Papaya, Paw-paw)

Barrier Reef Cays. -- English Cay, Stoddart sight record in 1965
(U9G9); p29); erobaccomCay,,, Stoddart asaght necord ain 1965 s(969) p70),
Fosberg & Spellman 54218 (US).

CACTACEAE
Opuntia Mill.
Opuntia sp.
Lagoon Cays. -- Wild Cane Cay, Stoddart sight record in 1961.
CUCURBITACEAE

*Citrullus Schrad. ex Eckl. & Zeyh.
*Citrullus lanatus (Thunb.) Matsum. & Nakai (Water melon)
Citrullus vulgaris Schrad.

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2527 (US).

ro ie |

INVINUSFILIUNS

IWVILIILILOINI
—

HWOTETUTIUN

VIVILIMOUINIAIN

LIUNNNtOCSD

46

*Cucumis L.

*Cucumis melo L. (Melon)
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2447
(US, MO).
RHIZOPHORACEAE

Rhizophora L.
Rhizophora mangle L. (Red mangrove)

Atoll Cays. -- Turneffe Atoll: Mauger Cay, Stoddart sight record
in 1960 (1962a, p. 47, 1963), p.49/1)3)Pellacan: Cay. (Cockroach Group),
Stoddart sight record in 1962 (1963, p:. 73)); Cockroach Cay, Stoddart
sightrecords in 1960 and 1962 (1962a, p. -46, 1963, p. 73, 1969, pe ws)
Cockroach Cay V, Stoddart sight record in 1965 (1969, p. 13); Pelican
Cay, Stoddart sight record in 1960 (1962a, p. 45, 1963, p: 74)" “Crawl
Cay, Stoddart sight record in 1960 (1962a, p. 46, 1963, p. 72); Three
Corner Cay, Stoddart sight record in 1960 (1962a, p. 46, 1963, p. 71);
Blackbird Cay, Stoddart sight record in 1960 (1962a, p. 44, 1963,

p-. #5); Soldier Cay, Stoddart ‘sight record “in yl960!-(1962a, ps 44), sil96si,
p. 75); Harry Jones Point, Stoddart sight record in 1960 (19624), tpeev4s);
Big Calabash Cay, Stoddart sight record in 1960 (1962a, p. 42, 1963,

p. 77, 19609, p. 14); Big Calabash East Cay I, Stoddart sight record™im
1960 (1962a, p. 42, 1963, p. 78)i;* Big*’Calabash East Cay If, Stoddant
Sight record in 1960 (1962a, p. 42, 1963, p. 78); Deadman Cay I,
Stoddart sight record in 1960 (1962a, p. 38, 1963, p. 79); Deadman Cay
Il, Stoddart sight record in 1960 (1962a, p.. 38, 1963, p- 80)i;3;Deadman
Cay III, Stoddart sight record in 1960 (1962a, p. 39); Deadman Cay IV,
Stoddart sight records in 1960 and 1965 (1962a, p. 39, 1963, p8iF;
1969, p. 15); Deadman Cay V, Stoddart sight record'in 1960 (1962a,

p. 40, 1963, p. 42); Big Cay Bokel, Stoddart sight record in 1962
(1963, p. 83); Cay Bokel, Stoddart sight record in’ 1960 (1962a, Woes,
ISS Si Ils | A))ic Lighthouse Reef: Sandbore Cay, Stoddart sight records
in 1960 and 1961 (1962a, fig. 28); Northern Cay, Stoddart sight records
in 1960 and 1961 (1962a;, p. 59,1963; p. 93); Saddle Cay, Stoddant
sight record in 1960 (1962a, p. 63, 1963, p. 94); Half Moon Cay,
Stoddart sight record in 1962 (seedlings); Long Cay, Stoddart sight
records in 1960, 1961 and 1962 (1962a, p---78,- 1963; ‘p.--99); Hat Cay7,
Stoddart ‘sight record in 1961 ~(1962a, p2-Sl),--196377*-p. =100)% Glover's
Reef: Long Cay, Linhart (1980, p. 163); Middle Cay, Stoddart sight
record in 1961 (1962a, p. 92), Sachet & Stoddart 1603 (US, MO), Linhart
(1980, p. 163); Southwest Cay I, Stoddart sight records in 1961 and
1971 =(1962a, p. 94)? ‘Linhare (1980) *p.. “163); Southwest ‘Cay Ir;

Stoddart sight record in 1961 (1962a, p. 96), Fosberg & Sachet 53904
(Os); Linhart: (T960;2pis 163):

47
Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 48); Sergeant's Cay, Stoddart sight record in 1962
(seedlings); English Cay, Stoddart sight records in 1960 and 1962

(1963, p. 45); Rendezvous Cay, Stoddart sight records in 1960, 1962,
LIGS Wand sl 972 VI63i) p46, e1969 pis Sy, L974 pe 480), “Spellman &

Stoddart 2497 (US, MO, BH); Tobacco Cay, Stoddart sight records in
1962 and 1965 (1963, p. 54, 1969, p. 10), Fosberg & Spellman 54229
(US); South Water Cay, Stoddart sight records in 1960 and 1965 (1963,

Pema o, 1 I69 pp. 1), spellman ee Stoddartszls> (US, MO; BH), Pringle
1862 (HAM); North Silk Cay, Stoddart sight record in 1961, Fosberg &
Spellman 54273 (US); Pompion Cay, Fosberg & Spellman 54325 (US)
(seedling); Ranguana Cay, Spellman & Stoddart 2247 (US) (seedling) ;
North Spot Cay, Spellman & Stoddart 2295 (US, MO, BH); Tom Owen's
West Cay, Stoddart sight record in 1960, Spellman & Stoddart 2263 (US);
Frank's Cay, Spellman & Stoddart 2406 (US, MO, BH, Fo, TI); Frank's
East Cay, Stoddart sight record in 1960; Frank's West Cay, Stoddart
Sight record in 1960; Lime Cay, Stoddart sight record in 1960; Seal
Cay, Spellman & Stoddart 2480 (US, MO).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1960
GiSGs) spews 2) CayniCaulkernsesStoddact sight) cecordrim 1960) i963), pia 34,
1969, p. 6); Cay Cangrejo, Stoddart sight records in 1960 and 1962
(1963, p. 62); Cay Chapel, Stoddart sight records in 1962 and 1965 (1963,
p. 35); St George's Cay, Stoddart sight records in 1962 and 1965 (1963,
ESS mel IOS) ep) same CksauSmCaySisstoddaraen Saghit mecordyain: a962-sGl963;,
p. 36); Rider's Cays, Stoddart sight record in 1962 (1963, p. 36);
Drowned Cays, Stoddart sight record in 1962 (1963, p. 36); Robinson
Point Cay, Stoddartsight record in 1961 °.(1963,,p. 46); Robinson Cay,
Sicoddazs& saiqht secordeine L961 N(iG6s; ip. 47) ;= Spanish Cay, Stoddart
sight) cecords ain 1961 and) 1962 (1963),) p-2 47));) Drappis Cay, Stoddart
sight record in 1962 (1963, p. 64); Simmonds Cay, Stoddart sight record
in 1962 (1963, p. 47); Southern Long Cay, Stoddart sight record in
UI SZenChIGS pea Dive pStewaKgenCay,, Stoddart isightimecordwin’ 1962 (1 9638;,
p. 61); Peter Douglas Cay, Stoddart sight record in 1962 (1963, p. 61);
Weewee Cay, Stoddart sight record in 1962 (1963, p. 61); Baker's
Rendezvous, Stoddart sight record in 1962 (1963, p. 62); Quamino Cay,
Stoddart sight record in 1962 (1963, p. 62); Tobacco Range, Fosberg &
Spellman 54457 (US); Coco Plum Cay, Spellman & Stoddart 2570 (US);
Water Range, Spellman & Stoddart 2226 (US, MO, BH, Fo); Man-o'-War
Cay, Fosberg & Spellman 54203 (US); Jack's Cay, Fosberg & Spellman
54434 (US); Buttonwood Cay, Stoddart sight record in 1961, Fosberg &
Spellman 54418 (US); Hatchet Cay, Fosberg & Spellman 54396 (US);
Little Water Cay, Fosberg & Spellman 54341 (US); Cary Cay, Stoddart
Sight record in 1962 (1963, p. 62); Owen Cay, Stoddart sight record in
TO C2ZM (ICS 7a p'-"66));) Collison) Cay7,) Stoddanesicght records anpl962) and
1965) (1963iekige 42)% (Scipio: Cay, Stoddart. saghtirecords ini1962: and
1965: (1963) Eig: 42); Bugle Cay, Stoddart sight records in 1962 and
965 (hI 6S piano) \; = piltacenciianCayy, Stoddart isightrecord an ul962
(1963, p. 68); Laughing Bird Cay, Stoddart sight record in 1962 (1963,
p. 65); East Snake Cay, Stoddart sight record in 1961 (1965, p. 138);
Middle Snake Cay South Islet, Stoddart sight record in 1961; West Snake
Cay, Stoddart sight record in 1961 (1965, p. 137); South Snake Cay,

ro i |

— NY ILNILILONI INYINUSFALIUNDS

(IWOTIEETULIVUIN

Vivid I TIOUINIAIN

LIVUNNNtLgd

48

Stoddart sight record in 1961 (1965, p. 137); Frenchman's Cay,
Stoddart sight record in 1961; Wild Cane Cay, Stoddart sight record
siiay sl Srel

COMBRETACEAE

Bucida L.
Bucida spinosa (Northrop) Jennings

Lagoon Cays. -- Ambergris Cay, Pelzl (US); Cay Caulker, Stoddart
431 (Fo) (1969, p. 6, as Reynosia sp.).

Conocarpus L.
Conocarpus erectus L. (Buttonwood mangrove)

Atoll Cays. -- Turneffe Atoll: Mauger Cay, Stoddart 549 (Fo);
Cockroach Cay, Stoddart sight record in 1962 (1963, p. 73); Cockroach
Cay V,, Stoddart: sight mecordiein 1965) (1969p. 43)GaPelaicanyCay,
Stoddart sight» record? in} 1960)141962a),p.. 45741963, 4p. «7 aa Soldier
Cay, Stoddart sight record in 1965 (1969, p. 14); Blackbird Cay,
Stoddart sight recordin 1960 ((1962a,%p;: 44)7 11963),> p.9 75) ;4 Big
Calabash Cay, Stoddart sight record in 1965 (1969, p. 14); Big
Calabash East Cay I, Stoddart sight record in 1960 (1962a, p. 42,

1963, p. 78)3°° Little Calabash-Cay, “Stoddant-sight+recordrinti965

(1969, p. 14); Deadman Cay II, Stoddart sight records in 1960 and 1965
(1962a;, p.- 38, 1963; p. 80i,> 1969)¢pe. 15) ';> "Deadman Cay ELL," Stoddart
sight records in 1960 and 1962 (1962a, p. 39, 1963, p. 81); Deadman
Cay IV, Stoddart sight record in 1960. Lighthouse Reef; Sandbore Cay,
Stoddart 538 (Fo), Stoddart sight records in 1961, 1962 and 1965,
(1962a, p. 57, 1963, p. 89, 1969, p. 16); “Northern Cay,” Stoddart isight
records iniige0; 1961 and A965 (1l962a,i pi627,9 19638 ripe 927. L969 aioe ela ee
Saddle Cay, Stoddart sight record in 1960 (1962a, p. 63, 1963, p. 94);
Half Moon Cay, Stoddart sight record in 1961 (1962a, p. 76, 1963,

p. 98); Long Cay, Stoddart sight record in/960) (1962a;,- p=. 80) 4 senate
Cay, Stoddart sight record in 1961°(1962a, p. 81, 1963, p. 100).
Glover's Reef: Northeast Cay, Stoddart 67 (Fo) (1962a, p. 88),

Fosberg & Sachet 53826 (US); Small Cay (Long Cay North), Stoddart sight
record in 1961 (1962a, p. 89), Fosberg & Sachet 53777 (US), Linhart
(1980 ,p.-162)";: “Long Cay’, Stoddart/sight: record ‘in’ 1961) 1(196 2ayepem 9
Fosberg & Sachet 53803 (US), Pippen 911 (US), Linhart (1980, p. 162);
Middle Cay, Stoddart sight record in 1961 (1962a, p. 93), Sachet &
Stoddart 1632 (US), Linhart (1980, p. 162); Southwest Cay I, Stoddart
sight records in 1961 and #1971 (1962a, p2°94). > :Linhart: (1980 4p. 162);
Southwest Cay II, Stoddart sight record in 1961 (1962a, p. 97), Fosberg

& Sachet 53890! (US) ,- hanhare <(1.980 ,ip<. 162))F.

49

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 41); Sergeant's Cay, Stoddart sight records in 1965
and 1972 (1969, p. 8, 1974, p. 478), Spellman & Stoddart 2543 (US, MO,
BH); Rendezvous Cay, Stgddart sight records in 1965 and 1972 (1974,
p. 480), Spellman & Stoddart 2502 (US); Tobacco Cay, Stoddart sight
records in 1961 and 1965 (1963, p. 54, 1969, p. 10), Fosberg &
Spellman 54230 (US); South Water Cay, Spellman & Stoddart 2194 (US,
MO), Pringle 1891 (HAM); North Silk Cay, Stoddart sight record in
1961, Fosberg & Spellman 54274 (US); Middle Silk Cay, Fosberg &
Spellman 54276 (US); Round Cay, Fosberg & Spellman 54297 (US);
Pompion Cay, Fosberg & Spellman 54316 (US); Northeast Sapodilla Cay,
Spellman & Stoddart 2305 (US, MO); Tom Owen's East Cay, Spellman &
Stoddart 2283 (US, MO); Tom Owen's West Cay, Stoddart sight record in
1960, Spellman & Stoddart 2267 (US, MO); Frank's Cay, Stoddart sight
record in 1960, Spellman & Stoddart 2399 (US, MO); Frank's East Cay,
Stoddart sight record in 1960; Frank's West Cay, Stoddart sight record
in 1960; Nicolas Cay, Stoddart sight record in 1960, Spellman &
Stoddart 2351 (US, BH, MO, Fo); Hunting Cay, Stoddart sight record in
1960 (1962b, p. 164), Spellman & Stoddart 2426 (US, MO); Lime Cay,
Stoddart sight record in 1960, Spellman & Stoddart 2365 (US, MO) ;
Ragged Cay, Stoddart sight record in 1960; Seal Cay, Spellman &
Stoddart 2485 (US, MO, BH, Fo).

Lagoon Cays. -- Cay Chapel, Stoddart sight record in 1965 (1969,
Ds io pSteGeorges Cay, Sstoddartrsight necord vin 19652(1969),, p:'s7).,
Spellman 1464 (US, MO); Water Cay, Dwyer et al. 676 (MO); Robinson
Point Cay, Stoddart sight record in 1961 (1963, p. 46); Spanish Cay,
Stoddart sightrecords in 1961 and 1962 (1963, p. 47); Cat Cay,
Stoddart sight record in 1962 (1963, p. 62); Trapp's Cay, Stoddart
Sight record in 1962 (1963, p. 63); Tobacco Range, Fosberg & Spellman
54449 (US); Coco Plum Cay, Spellman & Stoddart 2559 (US, MO, BH, Fo);
Water Range, Spellman & Stoddart 2227 (US, MO, BH); Buttonwood Cay,
Stoddart sight record in 1961 (1963, p. 64), Fosberg & Spellman 54417
(US); Hatchet Cay, Fosberg & Spellman 54369 (US); Little Water Cay,
Fosberg & Spellman 54331 (US); East Snake Cay, Stoddart sight record
in 1961; Middle Snake Cay North Islet, Stoddart sight record in 1961;
Middle Snake Cay South Islet, Stoddart sight record in 1961; West
Snake Cay West Islet, Stoddart sight record in.1961; South Snake Cay,
Stoddart sight record in 1961 (1965, p. 138); Wild Cane Cay, Stoddart
sight record in 1961.

Laguncularia Gaertn. f.
Laguncularia racemosa (L.) Gaertn. f. (White mangrove)

Atoll Cays. -- Turneffe Atoll: Harry Jones Point, Stoddart sight
record in 1960; Deadman Cay II, Stoddart sight records in 1960 and
1ICZAGISS Za; wees 6,7 L968), -p. SO0)i; | Deadman) Cay ELL, Stoddart Sight
Becora nw 1960 N(19637 1p. S15; Deadman Cay LV, Stoddart sight) record
ab AGO (USA, i956 Sp) es} 5 js Sl). Glover's Reef: Northeast Cay,
Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1602 (US),
Linhart (1980, p. 162); Southwest Cay I, Fosberg & Stoddart 53855 (US).

oe |

NVINUSHLIIVS ©

IWUILIILILONI

INSTITUTION _

OWI IMOUINTAIN

LIDNANICS

ouvuwe ot

=)

+e ews ce em

50

Barrier Reef Cays. -- Rendezvous Cay, Spellman & Stoddart 2501
(US, MO); Tobacco Cay, Fosberg & Spellman 54245 (US); South Water Cay,
Spellman & Stoddart 2156 (US, MO, BH, Fo), Pringle 1891 (HAM); Frank's
Cay, Stoddart sight record in 1960, Spellman & Stoddart 2392 (US, MO);
Frank's Cay East, Stoddart sight record in 1960; Seal Cay, Spellman &
Stoddart 2486 (US, MO, BH, Fo).

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54476 (US);
Coco Plum Cay, Spellman & Stoddart 2561 (US, MO); Water Range,
Spellman & Stoddart 2228 (US, MO); Man-o'-War Cay, Fosberg & Spellman
54205 (US); Jack's Cay, Fosberg & Spellman 54432 (US, MO); Buttonwood
Cay, Fosberg & Spellman 54420 (US); Hatchet Cay, Fosberg & Spellman
54390 (US); Little Water Cay, Fosberg & Spellman 54333 (US).

*Terminalia L.

*Terminalia catappa lL. (Almond, Indian Almond, Tropical Almond)
Atoll Cays. -- Glover's Reef: Northeast Cay, Fosberg & Sachet
53809 (US) , Linhart (1980), p. 162); Middile"Cay, Sachets Stoddart el629

(US); Southwest Cay II, Fosberg & Sachet 53882 (US).

Barrier Reef Cays. -- Tobacco Cay, Stoddart sight records in 1961,
1962 and 1965 (1963, p. 54, 1969, p. 10), Fosberg & Spellman 54215
(US); South Water Cay, Spellman & Stoddart 2204 (US, MO); Carrie Bow
Cay, Fosberg & Spellman 54439 (US) (seedling); Ranguana Cay, Spellman

& Stoddart 2259 (US); Northeast Sapodilla Cay, Spellman & Stoddart
2301 (US, MO); Frank's Cay, Spellman & Stoddart 2394 (US, MO, BH, Fo);
Nicolas Cay, Spellman & Stoddart 2334 (US, MO, BH, Fo, BM); Hunting
Cay, Spellman & Stoddart 2421 (US, MO); Lime Cay, Spellman & Stoddart
23859 (US; *MOy 78H, #0):

Lagoon Cays. -- Ambergris Cay, Stoddart sight records in 1962 and

1965 (1963, p- 33); “Cay Caulker, Stoddart ‘sight records in @ig6, andi

1965 (1963, p. 34); Hatchet Cay, Fosberg & Spellman 54387 (US);
Little Water Cay, Fosberg & Spellman 54332 (US); East Snake Cay,
Stoddart 96 (Fo)- (1965, p. 138).

MYRTACEAE

Eugenia L.
Eugenia sp. (Rodwood)

Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2427 (US).

Syl
*Psidium L.
*Psidium guajava L. (Guava)
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2161
(US, MO).
Lagoon Cays. - Hatchet Cay, Fosberg & Spellman 54366 (US); Wild

Cane Cay, Stoddart 24 (det. W. T. Stearn, not seen).
SAPOTACEAE

Bumelia Sw.
Bumelia retusa Sw.
Bumelia americana (Mill.) Stearn
Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Verner (1959,

PPE SO oOl = De) SoZ), stoddane (dg962a, pi. 74): Glover's Reef:
Northeast Cay, Fosberg & Sachet 53824 (US), Linhart (1980, p. 163);

Long Cay, Fosberg & Sachet 53796 (US), Pippen 912 (US), Linhart (1980,
p. 163); Middle Cay, Sachet & Stoddart 1601 (US), Linhart (1980,
Dee 163).

BarEter Reet "Gays. '——" Tobacco Cay, "Stoddart 4374(Fo)!

Lagoon Cays. -- Tobacco Range, Fosberg & Spellman 54444 (US).

Manilkara Adans.

Manilkara zapota (L.) P. van Royen (Naseberry, Sapodilla, Nispero,
Chiclé)
Atoll Cays. -- Turneffe Atoll: Soldier Cay, Romney et al. (1959,

p. 254), Stoddart (1962a, p. 44: not seen); Calabash Cays, Romney et
alu(t9S9ap.. 254), Rope Walk, Romney et all. (1959, p: 254).

Barrier Reef Cays. -- Hunting Cay, Vermeer (1959, p. 90).

Pouteria Aubl.
Pouteria rivicoa (Gaertn.f.) Ducke (Egg fruit)

Lucuma rivicoa Gaertn.f.

Pouteria campechiana (H.B.K.) Baehni

INDSETTUTION NWUILMLILONI NVINUSHLING 3d]

OWI MNOUINTAIN

LIDNANICS

cuvuws yp

52

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Verner (1959,
PP. 3-87, LISI pp. DS2)), Stoddart (d962Zay pa 74)re Glover's Reef:
Northeast Cay, Fosberg & Sachet 53833 (US), Linhart (1980, p. 163);
Middle Cay, Sachet & Stoddart 1604 (US), Linhart (1980, p. 163).
Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2338 (US,
MO; BH)r.
LOGANIACEAE

Polypremum L.
Polypremum procumbens L.

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2209
(US, MO, BH) Pringle 1903 (HAM, NY).

GENTIANACEAE
Eustoma Salisb.
Eustoma exaltatum (L.) G. Don
Barrier Reef Cays. -- Sergeant's Cay, Stoddart 438 (Fo) (1969,
Depo)
Lagoon Cays. -- St George's Cay, Spellman 1472 (US, MO).
APOCYNACEAE
*Catharanthus G. Don
*Catharanthus roseus (L.) G. Don (Ramgoat rose, Madagascar
periwinkle)
Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54224 (US,
MO).
Lagoon Cays. -- Cay Caulker, Stoddart 409 (Fo).
Echites Browne
Echites umbellata Jacq.
Atoll Cays. -- Turneffe Atoll, Deadman Cay I, Stoddart 458 (Fo)

(1969), tps. 15)i5

53
Lagoon Cays. -- St George's Cay, Spellman 1461 (MO); Tobacco
Range, Fosberg & Spellman 54473 (US).
*Nerium L.
*Nerium oleander L. (South Sea rose, Oleander)

Barrier Reef Cays. -- South Water Cay, Pringle sight record in
1980; Nicolas Cay, Spellman & Stoddart 2331 (US, MO, BH); Hunting Cay,
Spellman & Stoddart 2452 (US, MO).

Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54365 (US, MO).

Plumeria L.

*Plumeria rubra L. (Frangipani)
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2150

(US, MO); Hunting Cay, Spellman & Stoddart 2474 (US, MO).

Rhabdadenia MU11.-Arg.

Rhabdadenia biflora (Jacq.) MUll.-Arg.

Rhabdadenia paludosa (Mill.) Miers

Atoll Cays. -- Glover's Reef: Southwest Cay I, Linhart (1980,
as EQ Ve
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2432 (US,
MO, BH).
CONVOLVULACEAE

Ipomoea L.
Ipomoea aSarifolia (Desr.) R. & S.
BarnieruRcer Caysi., =~ Cay Glory ,, Stoddart 516 i(Fo)ei (196372 pee 52) .
Ipomoea macrantha R. & S. (Moon flower)
Ipomoea tuba (Schlect.) Don
Atoll Cays. -- Turneffe Atoll: Pelican Cay (Cockroach Group),

Stoddart 146 (Fo) (1963, p. 72 as I. tuba). Lighthouse Reef: Half
Moon Cay, Stoddart 34 (det. W. T. Stearn, not seen), Stoddart 448 (Fo).

WUITLIILILONI

—

INSTTIULION

LIDNANICS

oumvumwe ii

vo

NVINUSHLINS S41

EROS Kite TT RO wea

OWL IMOUINIAIN

a eeR hs A ed

54

Glover's Reef: Small Cay (Long Cay North), Fosberg & Sachet 53784
(US), Linhart (1980, p. 163) (as I. tuba); Southwest Cay II, Fosberg
& Sachet 53896 (US), Linhart (1980, p. 163) (as I. tuba).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight record in
1965 (1969, p. 8), Spellman & Stoddart 2549 (US, MO); South Water Cay,
Stoddart sight record in 1965, Pringle 1928 (HAM); North Silk. Cay,
Fosberg & Spellman 54267 (US); Middle Silk Cay, Fosberg & Spellman

54280 (US); Round Cay, Fosberg & Spellman 54303 (US); Ranguana Cay,
Spellman & Stoddart 2249 (US, MO); North Spot Cay, Spellman & Stoddart
2293 (US, MO); Tom Owen's West Cay, Spellman & Stoddart 2273 (US, MO),

2271 (US) (seedling); Seal Cay, Spellman & Stoddart 2482 (US, MO).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965
(1969, p. 6, as I. tuba); West Snake Cay, Stoddart 17 (Fo) (1965,
ido LEW Ne

Ipomoea pes-caprae ssp. brasiliensis (L.) v. Ooststr. (Beach morning-
glory)

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

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 31 (det.
W. T. Stearn, not seen). Glover's Reef: Small Cay (Long Cay North),
Fosberg & Sachet 53789 (US), Linhart (1980, p. 163); Southwest Cay I,

Stoddart sight record in 1971, Linhart (1980, p. 163); Southwest Cay
It, Hosberg & Sachet 53892 (US), Linhart (1980), Ip 163):

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight record in 1965
(1969, p. 8), Spellman & Stoddart 2555 (US, MO); Goff's Cay, Spellman &
Stoddart 2541 (US, MO, BH); English Cay, Spellman & Stoddart 2532 (US,
MO, BH, Fo); Rendezvous Cay, Stoddart sight record in 1962 (1963,

p. 49), Speldman &, Stoddart, 2507. (US, MO); Tobacco Cay, Stoddart; sight
records in 1962 and 1965 (1963, p. 54), Fosberg & Spellman 54253 (US);
South Water Cay, Spellman & Stoddart 2187 (US, MO), Pringle 1881 (HAM,
MO); Carrie Bow Cay, Spellman & Stoddart 2576 (US) (seedling); Middle
Silk Cay, Fosberg & Spellman 54284 (US); Round Cay, Fosberg %&

Spellman 54294 (US); Pompion Cay, Fosberg & Spellman 54315 (US); North
Spot Cay, Spellman & Stoddart 2294 (US, MO); Tom Owen's West Cay,
Spellman & Stoddart 2269 (US, MO).

Lagoon Cays. -- St George's Cay, Stoddart sight record in 1962
(1963, p. 38); Hatchet Cay, Fosberg & Spellman 54381 (US); West Snake
Cay, Stoddarity dict i(Fo)gi(l965 2p Rais):

Ipomoea stolonifera (Cyr.) Gmel.

Atoll Cays. -- Glover's Reef, Southwest Cay II, Fosberg & Sachet
>SIOtP (US) i, la nivartee((9 SO pre Os)) ee

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2524 (US,
MO), .2528:..(US) (seedling) +s- Tobacco: Cay,:.Stoddart,526\4 (Fo)a. (1963,

55

p- 54), Fosberg & Spellman 54238 (US); South Water Cay, Stoddart 436
(Fo), Spellman & Stoddart 2174 (US, MO, BH, Fo), Pringle 1913 (HAM);
Carrie Bow Cay, Ferraris 10 (US).

NVYINUSHLINGS Odi

Lagoon Cays. -- St George's Cay, Spellman 1460 (US, MO).

Ipomoea sp.

IWUILIILILONI

—

Atoll Cays. -- Turneffe Atoll: Cockroach Cay, Stoddart sight
BeCcoOradmin NICS uI6IF sp. visi Soldier) Cay, Stoddane sight second) in
1965 (1969, p. 13); Deadman Cay III, Stoddart sight records in 1960
ancy Io2mCI6Zay, pi S97 UI637, 3p. Si); eDeadman, Cay, TV, ‘Stoddart ‘sight
record in 1965; Eastern Sand Ridge, Stoddart sight records in 1960
andro Gia n(loGZay spr) «47/9. Lighthouse Reef: Sandbore Cay, Stoddart
Sughiaaseconds ine ICG; I6laand 1965. (1962a, pa a, 19637 p., 89, 1969,
p- 16); Half Moon Cay, Stoddart 37 (Fo), Stoddart sight records in
ILQS6O. etal AlIel” (ALoZe.5 6 Pao WIGS > jo5 (Seis Glover's Reef: Southwest
Cayeil mocoddarntasigh te necocdminy 196) (lr962Zaypi 97):

INSTITEULIUON

OWL LMOUINIAIN

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 40); Goff's Cay, Stoddart sight records in 1960 and
1965 (1963, p. 43, 1969, p. 9); Rendezvous Cay, Stoddart sight record
inl I65);) (elobacco.Cay),-Stoddaneasight record any l965 (1969), pe -10));
South Water Cay, Stoddart sight records in 1960 and 1965 (1963, p. 56,
USGI pap) CacaresBow. Cay, Stoddart sight secord=in- 1965-—(1969,
Dee) SeNortheSilkiGay,, Stoddart isight. record an 19615" Middle Sidik
Cay, Stoddart sight record in 1961.

LIDNARICS

wvyuwei s

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965 (1969,
Pp. 6); Cay Caulker, Stoddart sight record in 1962; St George's Cay,
Sroddantmsrght, rcecordman: 1965) (1969p. 8). Scipio ay, Stoddart sight H
record in 1962; Laughing Bird Cay, Stoddart sight record in 1962
(1963, p. 65); West Snake Cay West Islet, Stoddart sight record in ‘y
1961; Middle Snake Cay North Islet, Stoddart sight record in 1961;

wot

Middle Snake Cay South Islet, Stoddart sight record in 1961; East
Snake Cay, Stoddart sight record in 1961 (1965, p. 138).
)
J
Jacquemontia Choisy iy
i
Jacquemontia havanensis (Jacq.) Urb. ,
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2167 7
(US, MO), Pringle 1865 (HAM). f
Merremia Dennst. ex Endl.

Merremia dissecta (Jacq.) Hall. f. (Noyo)

Lagoon Cays. -- Cay Chapel, Stoddart 441 (Fo) (1969, p. 7).

56

BORAGINACEAE
Cordia L.
Cordia sebestena L. (Geiger Tree, Scarlet Cordia)
Atoll Cays. -- Turneffe Atoll: Pelican Cay (Cockroach Group),

Stoddart, sight’ records. in; 1962 andul965 .A(19637iep -.572),1. 1969) apere2 ee
Cockroach Cay, Stoddart sight record in 1962 (1963, p. 73); Harry Jones
Point, Stoddart sight: record in, 1962 3(1963;, p.2 86). Lighthouse Reef:
Half Moon Cay, Verner (1959, pp. 3-8), Stoddart sight records in 1960,
1962 and, 1965 (19624), -p.. 75), 1.963) .2p 1.95), 2! 969). Div lic) in Stoudantasse
(Fo); Long Cay, Stoddart sight record in 1965 (1969, p. 18). Glover's
Reef: Northeast Cay, Stoddart 76 (Fo) (1962a, p. 88), Fosberg & Sachet
53839 (US), Linhart (1980, p. 162); Long Cay, Fosbert & Sachet 53914
(US), Linhart (1980, p. 162); Middle Cay, Sachet & Stoddart 1605 (US),
Linhart (1980, p. 162); Southwest Cay I, Fosberg & Stoddart 53856 (US),
ELinhart (1980; p; 162).-

Barrier Reef Cays. -- Tobacco Cay, Stoddart 517 (Fo), Stoddart
Sight records in 1961 and 1965 (1963, p. 54, 1969, p. 10), Fosberg &
Spellman 54241 (US); South Water Cay, Spellman & Stoddart 2152 (US,
MO, BH); Pringle 1921 (HAM); Pompion Cay, Stoddart sight record in
1960, Fosberg x Spellman 54309 (US); Seal Cay, Spellman & Stoddart 2483
(USF MO} BH tok

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965 (1969,
p- 6); Cay Caulker, Stoddart 403 (Fo) (1969, p. 6); St George's Cay,
Spellman 1459 (MO); Robinson Point Cay, Stoddart sight record in 1961
(1963, p. 46); Cary Cay, Stoddart: sight, record:in 1962 (1963), peaGS)r
Scipio Cay, Stoddart sight record in 1962 (1963, p. 66); Owen Cay,
Stoddart sight record in 1962 (1963, p. 66); Colson Cay, Stoddart sight
records .in 1962 and. 1965 .(1963),..p...67, 1969),p-..12));. East Snakevcayy
Stoddart sight record in 1961 (1965, p. 138).

Tournefortia L.
Tournefortia gnaphalodes (L.) Kunth (Seaside Lavender)
Mallotonia gnaphalodes (L.) Britt.

Atoll Cays. -- Turneffe Atoll: Pelican Cay (Cockroach Group),
Stoddart sight record in 1965 (1969, p. 12); Cockroach Cay, Stoddart
sight records in 1962 and 1965 (1969, p. 13);. Cockroach Cay V,
Stoddart sight record in 1965 (1969, p. 13); Soldier Cay, Stoddart
sight records in 1960 and 1965 (1962a, p. 44, 1963, p. 75, 1969,

p. 13); Harry Jones Point, Stoddart sight record in 1960 (1962a,

p- 48); Little Calabash Cay, Stoddart sight record in 1965 (1969,

p. 14) (seedlings); Deadman Cay I, Stoddart sight records in 1960 and
1965 (1962a;- p. 38; 1963. p:-79," 1969,7 p.. 14)i;; - Deadman’ Cay 1m,

Sd

Stoddart siighty records in: 1960 Jandeli96Se@962a,, pi. 38, 1963, p. 80,
1969, p. 15); Deadman Cay IL, Stoddart sight records in 1960 and 1962
(1962a, p. 39, 1963, p. 81); Deadman Cay IV, Stoddart sight record in
NIGOM(AG62Zay pi 9), 1963), pi. Sl); sBig Cay, sBokell, “Steddazt sights record
inl 965). Lighthouse Reef: Sandbore Cay, Stoddart sight records in
HIG etIoL setooZz and 1965. (1 962a, pr=5/), 1963, =p. 39), 1969), mpee V6);
Northern Cay, Stoddart sight records in 1960, 1961 and 1965 (1962a,
DEO LOS; abs 9S nn) SOO, Ds wl) ny Hal feMoon Cay motoddartteuA Sea(ho)s
Stoddart. srghtarecord sine! S65en(hI6Za\, spam dion, ol IOS), ap. IS L969 pe 1)!
Long Cay, Stoddart sight record in 1965 (1969, p. 18). Glover's Reef:
Northeast Cay, Stoddart sight record in 1961 (1962a, p. 88), Fosberg &
Sachet 53836 (US), Linhart (1980, p. 162); Small Cay (Long Cay North),
Stoddart sight record in 1961 (1962a, p. 89), Fosberg & Sachet 53783
(US) ie binhant “9860; ips 62); 2hongeCay,, Stoddart sight record in’ 196%,
(1962a, p. 91), Fosberg & Sachet 53908 (US), Linhart (1980, p. 162);
Middle Cay, Stoddart sight record in 1961 (1962a, p. 93), Fosberg &
Stoddart 53924 (US), Sachet & Stoddart 1606 (US), Linhart (1980,
Pewlo2)ie Southwest Cayo, Stoddart sight record an 19619 (l962a, pz 95),
Fosberg & Stoddart 53861 (US), Linhart (1980, p. 162); Southwest Cay
IL, Fosberg & Sachet 53900 (US), Linhart (1980, p. 162).

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 40); Sergeant's Cay, Stoddart sight records in 1965
and 1972 (1969, p. 8, 1974, p. 478), Spellman & Stoddart 2544 (US, MO);
English Cay, Spellman & Stoddart 2525 (US); Rendezvous Cay, Stoddart
SaighitzenecordsminelI6>vanded 97/2EK(1 9697 oe See t9V4 = p.* 47/8) ;o Tobacco
Cay, Stoddart sight record in 1965 (1969, p. 10), Fosberg & Spellman
54231 (US); South Water Cay, Stoddart 520 (Fo), Stoddart sight records
ine ol Tande 1965) (96s) sips oO elo pawl) Speliimanee. Stoddart 243
(US, MO), Pringle 1914, 1927 (HAM) ; Carrie Bow Cay, Stoddart sight recordin
NIGS (1969, pe Il), a Speliimanws Stoddart 240) (US, SMO) Rerraris 9° "(US)>;
North Silk Cay, Stoddart sight record in 1961, Fosberg & Spellman
54275 (US); Middle Silk Cay, Stoddart sight record in 1961, Fosberg &
Spellman 54289 (US); Round Cay, Stoddart sight record in 1960,
Fosberg & Spellman 54295 (US); Pompion Cay, Stoddart sight record in
1960, Fosberg & Spellman 54312 (US); Ranguana Cay, Stoddart sight
record in 1960, Spellman & Stoddart 2255 (US, MO); North Spot Cay,
Spellman & Stoddart 2296 (US, MO); Tom Owen's West Cay, Stoddart sight
record in 1960, Spellman & Stoddart 2266 (US, MO); Tom Owen's East Cay,
Stoddart sight record in 1960; Nicolas Cay, Stoddart sight record in
1960; Hunting Cay, Stoddart sight record in 1960.

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965 (1969,
Paue);) 1Cay~ Caulker,, Stoddart 422, (Fo) (969, p-* 6));3 Cay ‘Chapel’,
Stoddart sight, wecordsyin L962 andi1965.5(1963),>p- 35,7 1969), pe 7);

St George's Cay, Spellman 1465 (MO); Tobacco Range, Fosberg & Spellman
54463 (US); Coco Plum Cay, Spellman & Stoddart 2569 (US, MO); Colson
Cay, Stoddart sight record in 1965" (1969)," p. 12));>> Scipio Cay, Stoddart
sight record in 1965; East Snake Cay, Stoddart sight record in 1961;
Middle Snake Cay North Islet, Stoddart sight record in 1961; West Snake
Cay, Stoddart. sight record: in 1961 (1965, p. 137).

_NVINUSHLINS 331

WUTLIILILONI

—

OWIINSUNIAN TINSTITEULIUON

LIDNANICS

oevunaouesd b

ve

ow w COUN Cee
aa

re Cem ew

58

VERBENACEAE
Avicennia L.
Avicennia germinans (L.) L- (Black mangrove)
Avicennia nitida Jacq.
Atoll Cays. -- Turneffe Atoll: Cockroach Cay, Stoddart sight

record. ine 1962) (1963, p--73); "PeliicanyCay, Stoddartssiight recordwam
1960 (1962a, p. 45, 1963, p. 74); Blackbird Cay, Stoddart sight record
in 1960 (1962,-p. 44, 19637, p-275)%* Solldiler! Cay, .Stoddart 4617 ao)
(1969, p. 14); Harry Jones Point, Stoddart sight record in 1960 (1962a,
p. 48); Big Calabash; Cay, Stoddart sight, record in) 1960) (1962a,ap 427
1963, p. 77); Big Calabash East Cay I, Stoddart sight records in 1960
and 1962 (1962a,;p-.4~42, 1963), p:> 78), p Big) Calabash) East) Cay, iii,
Stoddart sight record in 1960 (1962a, p. 42, 1963, p. 78); Deadman Cay
I, Stoddart, sight records in 1960) and: 1962 (1962a%,cp-5 38).11963;, P-nwoF
1969, p. 15); Deadman Cay II, Stoddart sight records in 1960 and 1962
(1962a, p. 38, 1963, p. 80); Deadman Cay III, Stoddart sight records in
1960 and 1961 (1962a, p. 39, 1963, p. 81); Deadman Cay IV, Stoddart
sight record in- 1960 (1962a,°p. 39, 1963), ps. 81)%- Deadman’ Cay. Vi;
Stoddart sight record in 1960 (1963, p. 82); Big Cay Bokel, Stoddart
sight record in 1962. Lighthouse Reef: Sandbore Cay, Stoddart sight
records in 1960 and: 1961, s(4962a;, p. 5776 1963)" p.5 89) 92 NortEhernacay,
Stoddart. sight records in 1960 and 1961 (1962a, p. 62); Saddle Cay,
Stoddart sight. record in 1960; (1962a;/p263,. 19635) p.. 94)i;4e LongaiCay,
Stoddart sight records in 1960 and 1961 (1962a, p. 80); Hat Cay,
Stoddart. 55.) (Fo)/a(1962a,“p-= 1815, a1963'7 p-(wl00): Glover's Reef:
Southwest Cay I; Stoddart sight record in 1961 (1962a, p. 94),

Fosberg & Stoddart 53854 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight record in
1960 (1963, p. 42); Goff's Cay, Stoddart sight record in 1960 (1963,
p. 43); Rendezvous Cay, Stoddart sight records in 1960 and 1972 (1963,
p. 48, 1974, p. 480), Spellman’ & Stoddart 2498 (US, MO,. BH, Fo);
Tobacco Cay, Stoddart sight records in 1961 and 1965 (1969, p. 10),
Fosberg & Spellman 54243 (US); South Water Cay, Stoddart sight record
in 1960 (1965, p. 57), Spellman & Stoddart 2157 (US, MO), Pringle 1908,
1909 (HAM); North Silk Cay, Stoddart sight record in 1960, Fosberg &
Spellman 54272 (US); Frank's Cay, Stoddart sight record in 1960,
Spellman & Stoddart 2419 (US, MO, BH, Fo); Frank's Cay East, Stoddart
Sight record in 1960; Frank's Cay West, Stoddart sight record in 1960;
Seal Cay, Spellman & Stoddart 2481 (US, MO, BH, Fo).

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1960 (1963,
p., 33); +>Spanish Cay,: Stoddart sight; records: in’ 19617 andy 19627219637,
p-. 47); Robinson Point Cay, Stoddart sight record in 1961 (1963, p. 46);
Cat Cay, Stoddart sight record in 1962)5(1963," 62) 5. sCaxny. Cay,
Stoddart sight record in 1962 (1963, p. 63); Tobacco Range, Fosberg &
Spellman 54456 (US); Coco Plum Cay, Spellman & Stoddart 2567 (US, MO);

Sy)

Water Range, Spellman & Stoddart 2225 (US, MO); Man-o'-War Cay (west
of Tobacco Cay), Fosberg & Spellman 54204 (US); Jack's Cay, Fosberg &
Spellman 54433 (S); Buttonwood Cay, Stoddart sight record in 1961
(1963, p. 64), Fosberg & Spellman 54421 (US); Hatchet Cay, Fosberg &
Spellman 54395 (US); Little Water Cay, Fosberg & Spellman 54343 (US);
OwenmCay7,, Sitoddageasrght secord: imi 1962 "(196s),, p. 66)"; Colson Cay,
Stoddart, srghtrnecords) an? 1962mandel965 (1963), ip. 67), 1969, pe 2))i;
Scipio Cay, Stoddart sight records in 1962 and 1965 (1963, p 66, 1969,
Dewitt); laughing = Bird Cay, Stoddart sight record “in” 1962; Bugle ‘Cay,
Stoddart sight record in 1962; East Snake Cay, Stoddart sight record
in 1961; Middle Snake Cay South Islet, Stoddart sight record in 1961;
West Snake Cay, Stoddart sight record in 1961 (1965, p. 137).

Citharexylum L.
Citharexylum caudatum L.

Barrier Reef Cays. -- Northeast Sapodilla Cay, Spellman & Stoddart
2302 (US, MO, BH, Fo, BM); Hunting Cay, Spellman & Stoddart 2455 (US,
MO, BH, Fo).
Lantana L.
Lantana involucrata L.

Atoll Cays. -- Lighthouse Reef: Northern Cay, Stoddart 53 (Fo);
MOnGmCay,,, os toddamt 455) (Ho)r* (i969) apa. 8).

Lagoon Cays. -- Cay Caulker, Stoddart 429 (Fo), 434 (Fo) (1969,
p. 6); Water Cay, Dwyer et al. 675 (MO); Buttonwood Cay, Fosberg &
Spellman 54405 (US, MO); Hatchet Cay, Fosberg & Spellman 54378 (US,
MO).
Lippia L.

Lippia nodiflora (L.) Michx. (Lippia)

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2183
(US, MO, BH, Fo, BM), Pringle 1885 (HAM); Tom Owen's West Cay,

Spellman & Stoddart 2268 (US, MO, BH); Frank's Cay, Spellman & Stoddart

2398 (US); Nicolas Cay, Spellman & Stoddart 2349 (US, MO, BH);
Hunting Cay, Spellman & Stoddart 2471 (US, MO, BH, Fo); Lime Cay,
Spellman & Stoddart 2375 (US, MO, BH, Fo); Seal Cay, Spellman &
Stoddart 2489 (US, MO).

Lippia strigulosa Martens & Gal.

Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54377 (US); Little

Water Cay, Fosberg & Spellman 54354 (US).

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Stachytarpheta Vahl

Stachytarpheta jamaicensis (L.) Vahl (Vervine, false verbena)
Atoll Cays. -- Turneffe Atoll: Big Cay Bokel, Stoddart sight
record in 1965. Lighthouse Reef: Half Moon Cay, Stoddart 35 (Fo),

Stoddart sight record in 1965.(1962a),, p:. 753 1963,, sp..-98i,, 1969). Deselone
Long Cay, Stoddart sight records in 1962 and 1965 (1963, p. 99, 1969),
Da 8) a Glover's Reef: Northeast Cay, Linhart (1980, p. 163);

Middle Cay, Stoddart sight record in 1961 (1962a, p. 93), Sachet &
Stoddart 1609 (US), Linhart (1980, p. 163); Southwest Cay I, Fosberg

& Stoddart 53864 (US), Linhart (1980, p. 163); Southwest Cay II,
Stoddart 83 (Fo), Fosberg & Sachet 53880 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -- Tobacco Cay, Stoddart 525 (Fo), Stoddart
sight records in 1961 and 1965 (1963, p. 54, 1969, p. 10), Fosberg &

Spellman 54251 (US); South Water Cay, Stoddart sight record in 1965

(1969, p.’ 11), Spellman & ‘Steddart 2188 (US), MO; BH), Pringle 882
(HAM); Ranguana Cay, Spellman & Stoddart 2248 (US); Northeast
Sapodilla Cay, Spellman & Stoddart 2319 (US, MO, BH); Frank's Cay,

Spellman & Stoddart 2416 (US, MO, BH, Fo); Hunting Cay, Spellman &

Stoddart 2458 (US, MO, BH, Fo); Lime Cay, Spellman & Stoddart 2369

(US MO;e BH):

Lagoon Cays. -- St George's Cay, Stoddart sight record in 1965
(1969, p. 8), Spellman 1467 (US, MO); Robinson Point Cay, Stoddart
Sight record in 1961: (1963, p. 46); Cary Cay, Stoddart sight record an
1962 (1963, p. 63); East Snake Cay, Stoddart sight record in 1961
(1965, p. 138); > Wild Cane’ Cay, Stoddart 25 /(Fo)i,

Stachytarpheta mutabilis (Jacq.) Vahl var. maxonii Mold. ?

Lagoon Cays. -- Cay Caulker, Stoddart 408 (Fo) (1969, p. 6).

SOLANACEAE

Solanum L.
Solanum blodgettii Chapm.

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 139
(US)

Lagoon Cays. -- Ambergris Cay, Stoddart 443 (US, MO, Fo, BM)
(1969, p. 6); Cay Caulker, Stoddart 412 (US, MO, Fo, BM), Stoddart 423
(US)= (L969) pe 6):

Solanum campechiense L.

Barrier Reef Cays. -- Rendezvous Cay, Stoddart 126 (Fo).

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*Solanum lycopersicum L. (Tomato)

Barrier Reef Cays. -- Rendezvous Cay, Stoddart 125 (Fo) (1963,
p. 49); South Water Cay, Pringle sight record in 1980.

SCROPHULARIACEAE

Capraria L.
Capraria biflora L.

Atoll Cays. -- Glover's Reef: Southwest Cay I, Fosberg & Stoddart

53867 (US), Linhart (1980, p. 163); Southwest Cay II, Fosberg & Sachet
53877 (US), Linhart (1980, p. 163); Southwest Cay, Pippen 905 (US).

*Russelia Jacq.

*Russelia equisetiformis Cham. & Schlecht. (Mary's tears, Fire-cracker
flower)
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2468 (US,

MO) BHO!)

Stemodia L.
Stemodia maritima L.
Atoll Cays. -- Turneffe Atoll: Big Cay Bokel, Stoddart 459 (Fo).

Lighthouse Reef: Half Moon Cay, Stoddart 49 (Fo); Long Cay, Stoddart
143) (Fo):

BIGNONIACEAE
Enallagma (Miers.) Baill.
Enallagma latifolia (Mill.) Small (Wild Calabash)
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2441 (US,

MO, BH, Fo).

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RUBIACEAE
Diodia L.
Diodia serrulata (Beauv.) Tayl.
Diodia maritima Thonn.
Atoll Cays. -- Glover's Reef: Middle Cay, Fosberg & Stoddart
93922 (US):
Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2346 (US,

MO, BH, Fo, BM); Hunting Cay, Spellman & Stoddart 2428 (US, MO, BH,
Fo); Lime Cay, Spellman & Stoddart 2373 (US, MO, BH, Fo).

Erithalis P. Browne
Erithalis fruticosa L.

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 30 (Fo),
Stoddart 546 (Fo) (1962a, p. 76). Glover's Reef: Northeast Cay,
Stoddart 73 (Fo) (1962a, p. 88), Fosberg & Sachet 53814 (US), Linhart
(1980, p. 163); Long Cay, Fosberg & Sachet 53806 (US), Linhart (1980,
p- 163): Middle Cay, Fosberg & Stoddart 53918 (US), 53923 (US),

Sachet & Stoddart 1607 (US), Linhart (1980, p. 163); Southwest Cay I,
Fosberg & Schreiber 53868 (US), Linhart (1980, p. 163); Southwest Cay
II, Fosberg & Sachet 53895 (US), Linhart (1980, p. 163); Southwest Cay,

Pappen/897 (US).

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2160
(US, MO, BH), Pringle 1864 (HAM); North Silk Cay, Stoddart sight record
in 1961; Middle Silk Cay, Fosberg & Spellman 54286 (US); Pompion Cay,
Stoddart sight record in 1960; Northeast Sapodilla Cay, Spellman &
Stoddart 2316 (US, MO); Tom Owen's West Cay, Stoddart sight record in
1960; Tom Owen's East Cay, Stoddart sight record in 1960; Frank's
Cay, Spellman & Stoddart 2413 (US, MO, BH, Fo); Nicolas Cay, Spellman &
Stoddart 2340 (US, MO, BH); Hunting Cay, Spellman & Stoddart 2463
(US, MO, BH); Lime Cay, Spellman & Stoddart 2383 (US, MO).

Lagoon Cays. -- St George's Cay, Spellman 1463 (US, MO); Tobacco
Range, Fosberg & Spellman 54452 (US); Coco Plum Cay, Spellman &
Stoddart 2572 (US, MO, BH, Fo); Water Range, Spellman & Stoddart 2238
(US, MO, BH); East Snake Cay, Stoddart sight record in 1961.

Ernodea Sw.

Ernodea littoralis Sw.

Atoll Cays. -- Turneffe Atoll: Deadman Cay I, Stoddart sight
BECOrd ein 1 I65. 11969 so el 5)k. Lighthouse Reef: Sandbore Cay,

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Sitoddartesightrecond: anelI65 (1969, p. 17); “Northern Cay, Stoddaritz051
(Ho) oteddart sight wecond: ine 1965) (1969/7 spa, f7);) - Halli Moon’ Cay,
Stoddasty552 (Homo eZaymon 76), 19687 p. 98); long Cay, sStoddacte sii
(Ho) ocoddart 457) (Ho) (19637 pp. 99!,, 19697 4p. 18): Glover's Reef:
Northeast Cay, Fosberg & Sachet 53827 (US); Middle Cay, Sachet &
Stoddarelols) (US), Linhact 1980); sp. 163); Southwesie Cay Tl,

Fosberg & Sachet 53898 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -— Goff"s Cay, Stoddart sight record in 1965
ChIGS sp. oS, L974 Pp. 479), Spellman & Stoddart 2539 (US, MO); English
Cay, Dwyer et al. 667 (MO); South Water Cay, Spellman & Stoddart 2162
(US, MO), Pringle 1898 (HAM); Nicolas Cay, Spellman & Stoddart 2353
(US, MO); Hunting Cay, Spellman & Stoddart 2461 (US, MO, BH).

Lagoon Cays. -- Cay Caulker, Stoddart 432 (Fo) (1969, p. 6); Cay
Chapel, Stoddart sight record in 1965 (1969, p. 7); St George's Cay,
Spellman 1473 (US, MO); Tobacco Range, Fosberg & Spellman 54451
(OSDir
Hamelia Jacq.

Hamelia patens Jacq.

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Stoddart 36 (Fo),
Siroddant ise, (Ho) W(962a7, spa 76)i:

Lagoon Cays. -- Ambergris Cay, Stoddart 444 (Fo) (1969, p. 6).

Hedyotis L.
Hedyotis corymbosa (L.) Lam.

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2521 (US,
MO); South Water Cay, Spellman & Stoddart 2208 (US, MO).

Lagoon Cays. -- Buttonwood Cay, Fosberg & Spellman 54410 (US).

*Hedyotis lancifolia Schum.
Oldenlandia lancifolia (Schum.) DC.

Oldenlandia herbasea sensu auct. non (L.) Roxb.

Barrier Reef Cays. -- South Water Cay, Pringle 1902 (HAM).

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*Ixora L.
*Ixora coccinea L.
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2451

(US, MO).

Morinda L.

Morinda citrifolia L. (Hog apple)
Atoll Cays. -- Glover's Reef: Southwest Cay II, Fosberg & Sachet
53881 (US).
Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2198

(US, MO); Round Cay, Fosberg & Spellman 54293 (US); Pompion Cay,
Fosberg & Spellman 54317 (US) (seedling).

Psychotria L.
Psychotria nervosa Sw. sensu lato
Psychotria undata Jacq.
Psychotria granadensis Benth. ?
Barrier Reef Cays. -- Nicolas Cay, Spellman & Stoddart 2341 (US,
MOF ABH ero).
Spermacoce L.
Spermacoce assurgens R. & P. (Peeny bush, button weed)
Spermacoce suffrutescens Jacq.
Borreria laevis sensu auct., non Lam.
Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54223 (US,
MO); South Water Cay, Spellman & Stoddart 2179 (US, MO, BH, Fo, BM),
Pringle 1868 (HAM); Hunting Cay, Spellman & Stoddart 2470 (US, MO, BH,
FO)

Spermacoce prostrata Aublet

Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2430 (US,
MO, BH, Fo, BM).

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Spermacoce suaveolens (Mey.) O. Ktze.
Borreria suaveolens Meyer
Lagoon Cays. -- Cay Caulker, Stoddart 4265 (Fo) (l9697" pe):
Spermacoce verticillata L.
Borreria verticillata (L.) G. F. W. Meyer

Lagoon Cays. -- St George's Cay, Spellman 1477 (US, MO).
COMPOSITAE

Ageratum L.
Ageratum littorale Gray

Atoll Cays. -- Turneffe Atoll: Mauger Cay, Stoddart 543 (Fo)
(1962a, p. 47, 1963, p. 71, as A. maritimum); Deadman Cay II,
Stoddart 128 (Fo), Stoddart 130 (Fo), Stoddart sight record in 1965
CSGsh pa C0, 19695" pe ao) Lighthouse Reef: Half Moon Cay, Stoddart
43 (Fo), Stoddart 48 (Fo) (1962a, p. 76, as A. maritimum). Glover's
Reef: Long Cay, Fosberg & Sachet 53793 (US) (abnormal), 53911 (US,
MO, BH, Fo, BM), 53912 (US, MO), Stoddart 78 (Fo), Linhart (1980,

p. 163); Middle Cay, Sachet & Stoddart 1612 (US, MO, BH, Fo, BM);
Southwest Cay I, Fosberg & Stoddart 53865 (US); Southwest Cay II,
Fosberg & Sachet 53897 (US, MO, BH, Fo, BM), Linhart (1980, p. 163);
Southwest Cay, Pippen 9 (WMU).

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54212 (US,
MO); South Water Cay, Spellman & Stoddart 2185 (US, MO, BH, Fo, BM),

Pringle 1870 (HAM): Hunting Cay, Spellman & Stoddart 2438 (US, MO,
BH, Fo, BM).

Lagoon Cays. -- Cay Caulker, Stoddart 413 (Fo), 425 (Fo), 428 (Fo)
(1969, p. 6, inc. A. maritimum); Tobacco Range, Fosberg & Spellman
54468 (US); Water Range, Spellman & Stoddart 2223 (US); Hatchet Cay,

Fosberg & Spellman 54392 (US).

Ageratum maritimum Kunth

Atoll Cays. -- Lighthouse Reef: Half Moon Cay, Verner (1959,
pp. 3-8). Glover's Reef: Middle Cay, Linhart (1980, p. 163);
Southwest Cay I, Linhart (1980, p. 163) (probably all A. littorale
Gray).

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Ageratum maritimum Kunth f. calvum Robinson

"Keys off the coast, N. S. Stevenson 152". Stanley & Record,
Flora of British Honduras, Fieldiana, Botany 12: 394, 1936 (probably
A. littorale Gray).

Ageratum sp.

Atoll Cays. -- Turneffe Atoll: Three Corner Cay, Stoddart sight
record in 1962 (1963, p. 71); Cockroach Cay, Stoddart sight records in
1962 and 1965)(1969,, p-.. 13)4... Cockroach Cay. V,,..\Steddart sightyrecord
in 1962 (1963, p. 73); Soldier Cay, Stoddart sight records in 1962 and
1965 *(1963, p. 75, 1969) p. U3). Big CalabashuGay,, Stoddart sights
record in 1965 (1969, p. 14); Little Calabash Cay, Stoddart sight
records in 1960, 1962 and 1965 (1963, p. 76, 1969, p. 14); ‘Deadmangeay
I, Stoddart sight record in 1962 “(1963 p. 80, 1969), ips "1'5)i- Deadman:
Cay IV, Stoddart ‘sight record an 962 5(1963i p. i8ith)r. Lighthouse Reef:
Sandbore Cay, Stoddart sight record in 1965 (1969, p. 17, as A.
maritimum); Long Cay, Stoddart sight record in 1965 (1969, p. 18).

Barrier Reef Cays. -- Rendezvous Cay, Stoddart sight record in
1965) (CLI69 apa)

Lagoon Cays. -- Cay Chapel, Stoddart sight record in 1965 (1969,
p. 7); St George's Cay, Stoddart sight records in 1962 and 1965 (1963,
p. 40, 1969; p. 7)% Cary Cay, Stoddart sight mecord, an. 1962 7GL963),
iDe, L6yS)))

Ambrosia L.

Ambrosia hispida Pursh (Bay tansy)
Atoll Cays. -- Turneffe Atoll: Harry Jones Point, Stoddart sight

record: in.1960;, Cay Bokel, Stoddart sight record, in 1961 (1962a,, passi,

(USES 1G tevA)lee Lighthouse Reef: Sandbore Cay, Stoddart 50 (Fo), 554

(Fo)>,) Stoddart «sight xecondstan; 1962 and’ 1965 (1962a,. p. 577, 19687
p.~89 ,° 1969, p. 16); Northern, Cay, Stoddart sight.records,<'n -1-960Rand
1961 (1962a,) pi .62.9 1963 ,,1p.--92)-;,.. Long Cay:,, Steddar sight. recondain
1965: “CL969: ap.” WSi)r. Glover's Reef: Northeast Cay, Stoddart 72 (Fo)
(1962a, p. 88), Fosberg & Sachet 53840 (US), Linhart (1980, p. 163);
Middle Cay, Stoddart sight record in 1961 (1962a, p. 93), Sachet &
Stoddart 1608 S(US), Linhart (1980; 7p. 163):

Barrier Reef Cays. -- South Water Cay, Stoddart sight record in
1965. (1963i;"p~ 56: 1969, %p.- 11;)), Spellman. i&, Stoddart, 12201, (US, Mops,
Pringle 1897 (HAM); Tom Owen's West Cay, Stoddart sight record in
19602

Lagoon Cays. -- Ambergris Cay, Stoddart sight record in 1965
(1969, p. 6).

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Bidens L.
Bidens cynapiifolia Kunth (Spanish needle, beggar's ticks)
Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54246 (US)
(seedling).
Lagoon Cays. -- Moho Cay, Stoddart 5 (Fo).
Bidens pilosa lL. (Spanish needle, beggar's ticks)
Barrier Reef Cays. -- Hunting Cay, Spellman & Stoddart 2443
(WS, WO).
Borrichia Adans.
Borrichia arborescens (L.) DC. (Seaside ox-eye)

Atoll Cays. -- Turneffe Atoll: Cockroach Cay, Stoddart sight
record in 1962; Cockroach Cay V, Stoddart sight record in 1962 (1963,
Peas) eb lackbird = Cay, Stoddart sight record in 1960; Soldier ‘Cay;
Stoddart sight record in 1960; Big Calabash Cay, Stoddart sight
records in 1960 and 1965 (1969, p. 14); Big Calabash East Cay I,
Stoddart sight records in 1960 and 1962 (1963, p. 78, 1969, p. 14);

Big Calabash East Cay II, Stoddart sight record in 1960; Little
Calabash Cay, Stoddart sight record in 1965 (1969, p. 14); Deadman Cay
I, Stoddart sight record in 1960 (1963, p. 80, 1969, p. 15); Deadman
Cay sil Stoddant sight recondsping1 96071962 and 19658 (1963, ps 80,
1969, p. 15); Deadman Cay III, Stoddart sight record in 1960; Deadman
Cay iV Stoddart isight second ian 1965 (969) ps WS). Deadman Gaye Vv,
Stoddart sight record in 1965. Lighthouse Reef: Sandbore Cay,
Stoddart sight’ “record in.1965..(1969), p.= 17); Northern Cay, Stoddart
sight record in 1965 (1969, p. 17); Half Moon Cay, Stoddart 548 (Fo);
mongmCay, stoddant, l420(Fo)y, stoddart Sight record in’ 1965 (1963, p 99),
IQYS9> Wo LS) 7 Isles Cenyp Sieosickeucie D4! (Galo) (iSozei, joe Billy WS. jos 1000).
Glover's Reef: Northeast Cay, Fosberg & Sachet 53810 (US); Small Cay
(Long Cay North), Stoddart sight record in 1961, Fosberg & Sachet 53778
(US), Linhart (1980, p. 163); Long Cay, Fosberg & Sachet 53804 (US),
Pippen 916 (US), Linhart (1980, p. 163); Middle Cay, Stoddart sight
Gecord=in 196l Sachets Stoddart 11624 (US); Linhant (1980), p. 163);
Southwest Cay I, Fosberg & Stoddart 53897 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -- St George's East Cay, Stoddart sight record
in 1960 (1963, p. 41); South Water Cay, Stoddart sight records in 1960
ance i9GSenC 96s pe 56, 1969), pe oli) ispelulimanseastoddantz 2154) (US), MO),
BH, Fo), Pringle 1871 (HAM).

Lagoon Cays. -- Ambergris Cay, Stoddart 445 (canescent form) (Fo),
446) (Fo) (1969p. 6); "Cay Caulker, Stoddart (420) (Fo) (1969, p. 6);
Cay Chapel, Stoddart sight record in 1965 (1969, p. 7); Tobacco Range,
Fosberg & Spellman 54464 (US); Water Range, Spellman & Stoddart 2231
(US, MO).

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Conyza L.

Conyza canadensis (L.) Cronquist var. pusilla (Nutt.) Cronq.
(Small horse-weed)

Erigeron canadensis L.

Atoll Cays. -- Lighthouse Reef: Long Cay, Stoddart 454 (Fo)
(1969p e 1S)k

Barrier Reef Cays. -- South Water Cay, Spellman & Stoddart 2192
(US, MO); Hunting Cay, Spellman & Stoddart 2467 (US, MO).

Lagoon Cays. -- Moho Cay, Stoddart 9 (Fo) (seedling).
Eclipta L.

Eclipta alba (L.) Hassk.

Atoll Cays. -- Glover's Reef: Southwest Cay I, Linhart (1980,
p. 163)”

Barrier Reef Cays. -- English Cay, Spellman & Stoddart 2520 (US,
MO); South Water Cay, Pringle 1879 (HAM); Carrie Bow Cay, Ferraris
12 (US).

Lagoon Cays. -- Buttonwood Cay, Fosberg & Spellman 54409 (US);

Little Water Cay, Fosberg & Spellman 54339 (US).
Eclipta prostrata (L.) L.

Barrier Reef Cays. -- English Cay, Dwyer et al. 666 (MO).

Emilia Cass.
Emilia sonchifolia (L.) DC. ex Wight

Barrier Reef Cays. -- South Water Cay, Pringle 1878 (HAM).

Flaveria Juss.
Flaveria linearis Lag.

Lagoon Cays. -- Ambergris Cay, Stoddart 447 (Fo) (1969, p. 6).

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Melanthera Rohr
Melanthera nivea (L.) Small

Atoll Cays. -- Turneffe Atoll: Big Cay Bokel, Stoddart 460 (Fo).
Lighthouse Reef: Sandbore Cay, Stoddart 144 (Fo). Glover's Reef:
Long Cay, Fosberg & Sachet 53794 (US) (sterile).

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54233 (US),
54252 (US, MO); South Water Cay, Spellman & Stoddart 2169 (US, MO).

Lagoon Cays. ~—- St George's Cay, Spellman 1468 (US, MO);

Buttonwood Cay, Fosberg & Spellman 54412 (US); Hatchet Cay, Fosberg &
Spellman 54393 (US); Little Water Cay, Fosberg & Spellman 54351 (US,
MO) Mono Cay, steddant 12 i(Fo))<
Pluchea Cass.
Pluchea symphytifolia (Mill.) Gillis (Wild tobacco)

Pluehea carolinensis (Jaéq.) G. Don

Pluchea odorata (L.) Cass.

Barrier Reef Cays. -- Tobacco Cay, Fosberg & Spellman 54210 (US).

Lagoon Cays. -- Tobacco Range, coconut grove in nothern part of
Cay, Fosberg & Spellman 54471 (US); Coco Plum Cay, Spellman & Stoddart
2571 (US, MO); Buttonwood Cay, Fosberg & Spellman 54408 (US).
Synedrella Gaertn.
Synedrella nodiflora (L.) Gaertn.

Lagoon Cays. -- St George's Cay, Spellman 1469 (MO), 1474 (US,

MO).

Vernonia Schreb.

*Vernonia cinerea (L.) Less. (Little ironweed)
Lagoon Cays. -- Hatchet Cay, Fosberg & Spellman 54362 (US).

Wedelia Jacq.

Wedelia trilobata (L.) Hitchc. (Seaside marigold)

Atoll Cays. -- Turneffe Atoll: Soldier Cay, Stoddart sight record

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in 1965 (1969, p. 13); Little Callabash Cay, Stoddart sight records in
1962 and 1965 (1963, p. 77, 1969, p. 14); Deadman Cay I, Stoddart sight
record in 1965 (1969, p. 15); Big Cay Bokel, Stoddart sight record in
1965, Lighthouse Reef: Sandbore Cay, Stoddart sight record in 1965
(1969,,. p..17);- Half Moon Cay,..Stoddart.61, (Fo), Stoddart ‘sight record
in 1965 “(1962a,-P7- 73, 1969), p. l7/);. Long Cay, Stoddart sighterecond

da LOGS CLI 69 > pr ees) Glover's Reef: Northeast Cay, Stoddart 69

(Fo) (1962a, p. 88), Fosberg & Sachet 53821 (US), Linhart (1980, p. 163);
Middle Cay, Sachet & Stoddart 1617 (US), Linhart (1980, p. 163);
Southwest Cay I, Stoddart sight record in 1971, Linhart (1980, p. 163);
Southwest Cay II, Fosberg & Sachet 53903 (US), Linhart (1980, p. 163).

Barrier Reef Cays. -- Sergeant's Cay, Stoddart sight record in
1965 (1969, p. 8), Spellman & Stoddart 2545 (US); Rendezvous Cay,
Stoddart “sight records*in’ 1962-and" 1965. (1969),) p. = 9)\-= | TobaccorGay,
Stoddart 527 (Fo), Stoddart sight records in 1961, 1962 and 1965 (1963,
p. 54, 1969, p. 10), Fosberg & Spellman 54242 (US); South Water Cay,
Stoddart sight records in, 1960) and 1965 (i963) pe 56, l969),) pawl,
Spellman & Stoddart 2172 (US, MO, BH, Fo), 2186 (US, MO), Pringle 1872
(HAM); Carrie Bow Cay, Spellman & Stoddart 2135 (US, MO); Middle Silk

Cay, Fosberg & Spellman 54291 (US); Ranguana Cay, Spellman & Stoddart
2254 (US, MO); Tom Owen's East Cay, Spellman & Stoddart 2282 (US, MO,
BH); Tom Owen's West Cay, Stoddart sight record in 1960; Northeast

Sapodilla Cay, Spellman & Stoddart 2318 (US, MO, BH, Fo); Frank's Cay,
Stoddart sight record in 1960, Spellman & Stoddart 2403 (US, MO, BH,
Fo, BM).

Lagoon Cays. -- Cay Caulker, Stoddart 401 (Fo), 406 (Fo) (1969,
Pp. 6) 7. Cay’ Chapel ;~ Stoddartysight: record) ing 1965) (1969) pie. ns eee
George's Cay, Stoddart sight records in 1962 and 1965 (1963, p. 8,
L969, p. 7), Speisimans471==(MO)—"“CatCay, Stoddartasrqht recordmam
1962. \S6S3 7 L.-62)); "Cany Cay, —Stoddart. (sight record) dn 1962 -msCcoce

Plum Cay, Spellman & Stoddart 2563 (US, MO, BH, Fo); Water Range,
Spellman & Stoddart 2235 (US, MO); Buttonwood Cay, Fosberg & Spellman
54425 (US); Hatchet Cay, Fosberg & Spellman 54368 (US); Little Water
Cay, Fosberg & Spellman 54347 (US); East Snake Cay, Stoddart sight

record in 1961 (1965, p. 138); Middle Snake Cay North Islet, Stoddart
sight record in 1961; South Snake Cay, Stoddart sight record in 1961
(1965), <p). -138))5) 7 MohomCay;, Stoddareel3) (Fo):

Wak
LITERATURE CITED

Adams, C. D. 1972. Flowering plants of Jamaica. 848 pp. Mona,
Jamaica.

Anonymous. 1830. The Honduras Almanack for the year of our Lord 1830.
Belize, for the Legislative Assembly. 185 pp.

den Hartog C. 1970. The sea-grasses of the world. Verh. Kon. Ned.
Akad. Wetens. Naturk. II, 59(1): 1-275.

Dixon, C. G. 1956. Geology of southern British Honduras, with notes on
adjacent areas. Belize, Government Printer. 92 pp.

Jefferys, T. 1775. The Bay of Honduras. In: The West India Atlas or
a general description of the West Indies. London.

Linhart, Y. B. 1980. Local biogeography of plants on a Caribbean
atollta J. Blegeogres 7 M5917,

Owen, R. 1838. A nautical memoir descriptive of the surveys made in
H.M. Ships Blossom and Thunder from 1829 to 1837. Dublin.

ROME ZN ND Hi W(eGistor)),) Wireilght, As (Gs (Sy, Arbuckle; Ra H=, and Valal,
V. E. 1959. Land in British Honduras: report of the British
Honduras Land Use Survey Team. London, H.M.S.O.

Salvin, O. 1864. A fortnight among the seabirds of British Honduras.
MoS pS MANOS SIZ=I37 3

stanley, P. €. and Record, S. J. 1936. The forests and flora of
British Honduras. Fieldiana, Botany, 12: 1-432.

Stoddart, D. R. 1962a. Three Caribbean atolls: Turneffe Islands,
Lighthouse Reef, and Glover's Reef, British Honduras. Atoll Res.
Beli, OF Wasi S

Stoddart, D. R. 1962b. Physiographic studies on the British Honduras
reefs and cays. Geogrl J. 128: 161-171.

Stoddart, D. R. 1963. Effects of Hurricane Hattie on the British
Honduras reefs and cays, October 30-31, 1961. Atoll Res. Bull.
953 dala2s

Stoddart, D. R. 1965. British Honduras cays and the low wooded island
js lem, URS Ss IGNsie5 Uieo (Eeoles Sor slay

Stoddart, D. R. 1969. Post-hurricane changes on the British Honduras
reefs and cays: re-survey of 1965. Atoll Res. Bull. 131: 1-25.

Stoddart, D. R. 1974. Post-hurricane changes on the British Honduras
1g OERESS “ViIcS=SLaveyy Cre UTA iehaoyle Aistol awayes (Ciel Wx-are Geohutoc
2: 473-483.

WUILIILILONG NVINOUSHLINS SAI

—>

INSTIIUIION

OMI LASUINTAIN

LIONARICS

wotuvuai ys

3

ceuuiiaroww

wet oe ee

72

Tsuda, R. T. and Dawes, C. J. 1974. Preliminary checklist of the
marine benthic plants from Glover's Reef, British Honduras.
Atoll Res. ABull . 01737 51—13.

Uring, N. 1726. History of the voyages and travels of Captain
Nathaniel Uring. London, first edition (fourth edition 1928).

Vermeer, D. E. 1959. The cays of British Honduras. Berkeley,
Department of Geography, 127 pp., mimeographed.

Verner, J. 1959. Behavior of the Red-footed Booby (Sula sula) in
British Honduras. Louisiana State University, Department of
Zoology, M.S. thesis.

Verner, J. 1961. Nesting activities of the Red-footed Booby in
British Honduras, Auk 78: 573-594.

West, R. C. 1977. Tidal salt-marsh and mangal formations of Middle
and South America, pp. 193-213, in Goodall, D. W. (editor-in-
chief), Ecosystems of the World, I. Wet coastal ecosystems
(ed. V. J. Chapman). New York.

Acoelorraphe wrightii 16
Acrostichum aureum 3
Agave sp. 20

Ageratum littorale 65
Ageratum maritimum 65
Ageratum sp. 66
AIZOACEAE 29
Alternanthera ramosissima 26
AMARANTHACEAE 26
Ambrosia hispida 66
ANACARDIACEAE 43
Andropogon citratus 7
Andropogon glomeratus 6
Andropogon sp. 6
Anthephora hermaphrodita 6
APOCYNACEAE 52

Avicennia germinans 58
Avicennia nitida 58
BATIDACEAE 25

Batis maritima 25

Bidens cynapiifolia 67
Bidens pilosa 67
BIGNONIACEAE 61
Boerhavia coccinea 27
BORAGINACEAE 56

Borreria laevis 64
Borreria suaveolens 65
Borreria verticillata 65
Borrichia arborescens 67
Brassavola nodosa 22
Bryophyllum pinnatum 32
Bucida spinosa 48
Bumelia americana 51
Bumelia retusa 51
Bursera simaruba 38
BURSERACEAE 38

CACTACEAE 45

Caesalpinia bonduc 33
Caesalpinia crista 33
Cakile lanceolata 31
Canavalia maritima 33
Canavalia rosea 33
Canavalia sp. 33
CAPPARIDACEAE 32
Capparis flexuosa 32
Capraria biflora 61
Carica papaya 45
CARICACEAE 45

Cassytha filiformis 31
Casuarina equisetifolia 23
CASUARINACEAE 23

V3)

INDEX

Catharanthus roseus 52
Cenchrus incertus 6
Cenchrus pauciflorus 6
Chamaesyce blodgettii 39
Chamaesyce buxifolia 40
Chamaesyce glomerifera 40
Chamaesyce prostrata 41
Chamaesyce thymifolia 41
CHENOPODIACEAE 26
Chloris petraea 8
Chrysobalanus icaco 32
Citharexylum caudatum 59
Citrullus lanatus 45
Citrullus vulgaris 45
Citrus aurantiifolia 36
CHiagns Sido S7/

Cladium jamaicense 13
Coccoloba uvifera 24
Cocos nucifera 16
Codiaeum variegatum 39
COMBRETACEAE 48
COMPOSITAE 65

Conocarpus erectus 48
CONVOLVULACEAE 53
Conyza canadensis 68
Cordia sebestena 56
CRASSULACEAE 32

Crinum amabile 20
Crotalaria retusa 33
Crotalaria verrucosa 34
CRUCIFERAE 31

Cucumis melo 46
CUCURBITACEAE 45
Cymbopogon citratus 7
Cymodocea manatorum 4
CYPERACEAE 13

Cyperus ligularis 13
Cyperus peruvianus 14
Cyperus planifolius 14
Cyperus sp. 15
Dalbergia ecastophyllum 34
Desmodium canum 34
Desmodium incanum 34
Desmodium tortuosum 34
Digitaria horizontalis 7
Diodia maritima 62
Diodia serrulata 62
Diplanthera wrightii 4
Distichlis spicata 7
Dracaena sp. 20
Drypetes brownii 39

— NYILMLILONI NVINUSHLINS Sal

INSTITIUTLION

LIDNANRICS OMIINOUNIAWN

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74

Echites umbellata 52
Eclipta alba 68

Eclipta prostrata 68
Eichhornia crassipes 20
Eleusine indica 7

Emilia sonchifolia 68
Enallagma latifolia 61
Eragrostis ciliaris 7
Eragrostis domingensis 8
Eragrostis prolifera 8
Erigeron canadensis 68
Erithalis fruticosa 62
Ernodea littoralis 62
Erythrina sp. 34

Eugenia sp. 50

Euphorbia blodgettii 39
Euphorbia buxifolia 40
Euphorbia glomerifera 40

Euphorbia mesembrianthemifolia 40

Euphorbia prostrata 41
Euphorbia thymifolia 41
Euphorbia trichotoma 41
Euphorbia sp. 41
EUPHORBIACEAE 39
Eustachys petraea 8
Eustoma exaltatum 52
Ficus cf. hemsleyana 23
FLCUS) ‘Sp. 23
Fimbristylis cymosa 15
Fimbristylis spadicea 16
Fimbristylis spathacea 15
Fimbristylis sp. 16
Flaveria linearis 68
GENTIANACEAE 52
Gliricidia sepium 34
GRAMINEAE 6

Halodule wrightii 4
Halophila engelmanni 5
Hamelia patens 63
Hedyotis corymbosa 63
Hedyotis lancifolia 63
Hibiscus rosa-sinensis 43
Hibiscus tiliaceus 43
Hippeastrum puniceum 20
Hippocratea volubilis 43
HIPPOCRATEACEAE 43
HYDROCHARITACEAE 5
Hymenocallis littoralis 21
Ipomoea asarifolia 53
Ipomoea macrantha 53
Ipomoea pes-caprae 54
Ipomoea stolonifera 54
Ipomoea tuba 53

Ipomoea sp. 55

Iresine celosia 26
Iresine diffusa 26

Ixora coccinea 64
Jacquemontia havanensis 55
Kalanchoe pinnata 32
Kyllinga peruviana 14
Laelia tibicinis 22
Laguncularia racemosa 49
Lantana involucrata 59
LAURACEAE 31

LEGUMINOSAE 33

Leucaena leucocephala 34
LILIACEAE 20

Lippia nodiflora 59
Lippia strigulosa 59
LOGANIACEAE 52

Lucuma rivicoa 51
Mallotonia gnaphalodes 56
MALVACEAE 43

Mangifera indica 43
Manilkara zapota 51
Mariscus jamaicensis 13
Melanthera nivea 69
MELIACEAE 38

Merremia dissecta 55
MORACEAE 23

Morinda citrifolia 64
Mucuna sp. 35

Musa paradisiaca 22

Musa sapientum 22
MUSACEAE 22

Myrica cerifera 23
MYRICACEAE 23

MYRTACEAE 50

Neea choriophylla 27
Nephrolepis multiflora 3
Nerium oleander 53
NYCTAGINACEAE 27
OLACACEAE 24

Oldenlandia herbasea 63
Oldenlandia lancifolia 63
Opuntia sp. 45
ORCHIDACEAE 22

PALMAE 16

Pancratium littorale 21
Panicum pilosum 9
Panicum virgatum 9
Paspalum blodgettii 9
Paspalum distichum 9
Paspalum laxum 10
Paspalum paniculatum 10
Paspalum vaginatum 9

Paspalum sp. 10

Passiflora suberosa 44
PASSIFLORACEAE 44

Paurotis wrightii 16
Pedilanthus tithymaloides 42
Philoxerus vermicularis 27
Phragmites cf. australis 10
Phragmites communis 10
Phyllanthus amarus 42
PHYTOLACCACEAE 28
Pithecellobium kenense 35
Pithecolobium keyense 35
Pityrogramma calomelanos 4
Pluchea carolinensis 69
Pluchea odorata 69

Pluchea symphytifolia 69
Plumeria rubra 53
POLYGONACEAE 24
POLYPODIACEAE 3
Polypodium lycopodioides 3
Polypodium polypodioides 3
Polypremum procumbens 52
PONTEDERIACEAE 20
Portulaca oleracea 28
Porcewliaca Spi. 29
PORTULACACEAE 28

Pouteria campechiana 51
Pouteria rivicoa 51
Psidium guajava 51
PSILOTACEAE 3

Psilotum nudum 3
Psychotria granadensis 64
Psychotria nervosa 64
Psychotria undata 64
Reynosia sp. 48
Rhabdadenia biflora 53
Rhabdadenia paludosa 53
Rhizophora mangle 46
RHIZOPHORACEAE 46

Rivina humilis 28

ROSACEAE 32

RUBIACEAE 62

Russelia equisetiformis 61
RUTACEAE 36

Saccharum officinarum 11
Salicornia perennis 26
SAPOTACEAE 51
Schomburgkia tibicinis 22
SCROPHULARIACEAE 61

75

Sesuvium portulacastrum 29
Sida acuta 44

SOLANACEAE 60

Solanum blodgettii 60
Solanum campechiense 60
Solanum lycopersicum 61
Sophora tomentosa 35
Spartina patens 11
Spartina spartinae 11
Spermacoce assurgens 64
Spermacoce prostrata 64
Spermacoce suaveolens 65
Spermacoce suffrutescens 64
Spermacoce verticillata 65
Sporobolus indicus 11
Sporobolus jacquemontii 11
Sporobolus virginicus 11
Sporobolus sp. 12
Stachytarpheta jamaicensis 60
Stachytarpheta mutabilis 60
Stemodia maritima 61
Stenotaphrum secundatum 13
STERCULIACEAE 44

Suaeda linearis 26

Suriana maritima 37
SURIANACEAE 37

Swietenia macrophylla 38
Synedrella nodiflora 69
Syringodium filiforme 4
Tamarindus indica 36
Terminalia catappa 50
Thalassia testudinum 5
Thespesia populnea 44
Thrinax floridana 19
Thrinax multiflora 19
Thrinax parviflora 19
Thrinax radiata 19
Tournefortia gnaphalodes 56
Turnera ulmifolia 44
TURNERACEAE 44

VERBENACEAE 58

Vernonia cinerea 69

Vigna luteola 36

Vigna repens 36

Waltheria americana 44
Waltheria indica 44
Wedelia trilobata 69
Ximenia americana 24
ZOSTERACEAE 4

NUITLIILILONI NVINOSHLINS Sil

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INSTITUTION

LIDNANICS OSOMIIASUNIAIWN

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Ter Ot Oe te

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)

LIST OF CAYS CITED

The islands in the systematic lists are grouped into Atoll Cays

(Turneffe Islands; Lighthouse Reef; Glover's Reef); Barrier Reef
Cays; and Lagoon Cays. Within each category the islands are cited
from north to south. For locations of cays on Turneffe Islands and
Lighthouse Reef, see Atoll Res. Bull. 87 (1962); for location of cays

on Glover's Reef, the Barrier Reef, and in the Lagoon, see Atoll Res.
Bull., Ehis rssues

TURNEFFE ISLANDS BARRIER REEF CAYS
Mauger Cay St George's East Cay
Dogflea Cay Paunch Cay
Three Corner Cay Sergeant's Cay
Crawl Cay Goff's Cay
Pelican Cay (Cockroach Group) English Cay
Cockroach Cay Rendezvous Cay
Cockroach Cay V Cay Glory
Pelican Cay Tobacco Cay
Blackbird Cay South Water Cay
Soldier Cay Carrie Bow Cay
Harry Jones Point North Silk Cay
Big Calabash Cay Middle Silk Cay
Big Calabash East Cay I South Silk Cay
Big Calabash East Cay II Round Cay
Little Calabash Cay Pompion Cay
Grand Bogue Point Ranguana Cay
Deadman Cay I North Spot
Deadman Cay II Tom Owen's East Cay
Deadman Cay III Tom Owen's West Cay
Deadman Cay IV Northeast Sapodilla Cay
Deadman Cay V Frank's Cay
Big Cay Bokel Frank's Cay East
Cay Bokel Frank's Cay West

Nicolas Cay
LIGHTHOUSE REEF Hunting Cay

Lime Cay
Sandbore Cay Ragged Cay
Northern Cay Seal Cay
Saddle Cay
Long Cay LAGOON CAYS
Half Moon Cay
Hat Cay Ambergris Cay

Cay Cangrejo
GLOVER'S REEF Cay Caulker

Cay Chapel
Northeast Cay St George's Cay
Long Cay North (Small Cay) Hick's Cays
Long Cay Rider's Cays
Middle Cay Drowned Cays
Southwest Cay I Water Cay

Southwest Cay II Robinson Point Cay

Robinson Cay
Spanish Cay
Tobacco Range
Stewart Cay

Weewee Cay

Coco Plum Cay
Water Range (Twin Cays)
Peter Douglas Cay
Cat Cay

Man-o'-War Cay
Crawl Cay

Baker's Rendezvous
Quamino Cay

Jack's Cay

Moho Cay (northern)
Buttonwood Cay
Simmond's Cay
Southern Long Cay
Trapp's Cay
Hatchet Cay

Ua

Little Water Cay

Cary Cay

Laughing Bird Cay

Owen Cay

Colson Cay

Scipio Cay

Bugle Cay

Placencia Cay

Harvest Cay

Moho Cay (southern)
Frenchman's Cay

Wild Cane Cay

East Snake Cay

Middle Snake Cay

Middle Snake Cay North Islet
Middle Snake Cay South Islet
South Snake Cay

West Snake Cay

West Snake Cay West Islet

_NVINUSHLINS Sil

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ATOLL RESEARCH BULLETIN
No. 259

FLORISTIC OBSERVATIONS ON SOUTH WATER AND CARRIE BOW
CAYS, STANN CREEK DISTRICT, BELIZE, in 1979-1980

NULLIILILONI NVINUSHLINS _ bee fa |

—_

by

James S. Pringle

1 LIDKRARICS OSOMIINSUNIAN INSITIULION

ww wVwaotuvudqi

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Issued by .

THE SMITHSONIAN INSITUTION

Washington; De Gay sUeseA.
April 1982

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-

FLORISTIC OBSERVATIONS ON SOUTH WATER AND
CARRIE BOW CAYS, STANN CREEK DISTRICT,
BELIZE, IN 1979-1980!

NVINUSHLIWS S31

James S. Pringle

South Water Cay is an island of ca. 5.17 nha in Stann Creek District, Belize, 16° 48’
21-42’’ N, 88° 04’ 50-58” W, 22.4 km SE of Dangriga (Stann Creek Town) and 20.9
km from the nearest mainland point. It is at the southern tip of Tobacco Reef (part of
the Barrier Reef), with the reef edge paralleling its eastern and southern coasts 150-250
m offshore. This cay is much used for teaching and research by biologists from Queen's
University, the Smithsonian Institution, and elsewhere. Therefore, records of its flora are
especially desirable, for use in ecological research and as a basis for observations of
floristic changes.

NUILIILILONI

—_

INSTIITULION

The cays of Belize have been described by Stoddart (1962a, 1965a). South Water
Cay is a sand cay. (In the more detailed classification of Stoddart 1965a, it is mapped,
from pre-1961 observations, as a vegetated sand cay with a windward shingle ridge. The
shingle ridge, nowever, is confined tu a very small area at the northeast tip, and in 1979-
1980 was poorly developed.) On sand cays, deposits of sand, derived from coral and
snell fragmenis, permit the development of diverse vegetation including many species
tolerant of the dry coral-sand habitats, in contrast to that of the mangrove cays. The
vegetation of the uncleared sand cays consists largely of thickets in which Bursera
simaruba (L.) Sarg., Coccoloba uvifera, Cordia sebestena, and Thrinax radiata Lodd. ex
J.A. & J.H. Schultes are among the dominant species (Stoddart, 1965b). This was pro-
bably the original vegetation of South Water Cay, although some of the usual dominant
species are now absent. Many of the sand cays, including South Water Cay, have been
cleared of much of this vegetation and planted to coconuts, Cocos nucifera. Clearing has
left such cays more susceptible to wind erosion, and extensive areas of largely bare sand
are now common (Stoddart, 1965b).

LIDNANICS. SMLIMSUNIAN

vaotuvudi jt

—o

On 30-31 October 1961, Hurricane Hattie, the most severe of several hurricanes
since coconut planting had become widespread, killed so many coconut palms and demo-
lished so many structures that many of the coconut plantations and associated facilities
on the Belizean cays were abandoned, and some recovery of the original thicket vegeta-
tion has been noted since then. The hurricane also caused extensive physiographic modifi-
cations and damage to natural vegetation. Mangroves -- Avicennia germinans, Conocarpus
erectus, Laguncularia racemosa (L.) Gaertn. fil:, and Rhizophora mangle -- were killed
through defoliation, and the littoral thickets of such species as Sophora tomentosa,
Borrichia arborescens, Suriana maritima, and Tournefortia gnaphalodes were extensively
eliminated by uprooting, to the extent that, shortly after the hurricane, the three last-
named species, although they had been extensively dominant in the cays, could not be
found within 20 mi (ca. 32 km) of the center of the storm-track (Stoddart, 1965b).
South Water Cay, ca 54 km from the storm-track’s center, was less severely affected -¢
than some of the cays studied by Stoddart. There was, however, extensive erosion at the 2
southern tip of the island, as shown in Stoddart’s (1963) maps of the 1960 and 1962 )
coastlines, and also significant erosion on the west-central coast, where a large resort 2
building was destroyed (Stoddart, 1963). Fresh exposure of beach-rock and some sand- a

eh ee

‘Contribution No. 42 from the Royal Botanical Gardens, Hamilton, Ontario, Canada.
Royal Botanical Gardens, Box 399, Hamilton, Ontario, Canada L8N 3H8.

5%
|

2

stripping and root exposure were noted at several places along the coast of the island
(Stoddart, 1962b). New plantings of coconuts were made on South Water and Carrie Bow
cays following the 1961 hurricane (Stoddart, 1969).

Human use of South Water Cay was originally focused around a well near the
center of the island, now surrounded by concrete pavement. Near this well are buildings
used as a field station by biologists. Elsewhere there are vacationers’ cottages, fishermen’s
residences, and a retreat belonging to a religious order. Except for coconuts, no food
crops or other economic plants are grown. Ornamental gardens were planted behind the
above-mentioned resort building near the center of the island, and a hard-surfaced tennis
court (not extant) was constructed just north of these gardens prior to 1960 (Stoddart,
1963, 1969). Some additional ornamentals were planted around several of the existing
buildings and along paths. Most of the introduced ornamentals remain confined to the
sites where they were planted, with no tendency to spread. Hymenocallis /ittoralis, in
contrast, has become extensively naturalized throughout the island, and some plants of
Casuarina equisetifolia \ikewise appear to be spontaneous. Whether all plants of these
species are derived from those planted on the island, or whether natural colonization has
also occurred, cannot be determined. Eritha/is fruticosa has been used in the ornamental
plantings, but can be assumed to be part of the native flora.

Much of South Water Cay is covered by coconut palms. Vegetation is sparse in their
shade, consisting of small, scattered plants of Fimbristylis cymosa, Cyperus peruvianus,
C. platifolius, and several species that are more abundant and luxuriant elsewhere, e.g.,
Euphorbia mesembryanthemifolia and Wedelia trilobata. Only two herb-subshrub com-
munities are at all extensive. A dry, sunny area of ca. 0.29 ha near the field station is
dominated by /pomoea sto/onifera, with numerous plants of Ageratum //ttorale and
Stachytarpheta jamaicensis, moderate numbers of Crota/aria retusa, Desmodium incanum,
and Wa/theria indica, and scattered individuals of many other species (Fig. 1). Just north
of this community, an evidently less xeric area of ca. 0.44 ha is densely covered by
approximately equal quantities of /oomoea pes-caprae, Vigna luteola, and Wedelia
trilobata (Fig. 2). There are some relatively high areas where dryness, probably in com-
bination with wind action and foot traffic, keeps the vegetation very sparse, with only a
few plants of such species as Euphorbia blodgettii, E. trichotoma, Phyllanthus amarus,
and Sporobolus virginicus being present (Fig. 3). Some areas near the spring are regularly
mowed as well as being much trampled; these are the principal habitats of such prostrate
or semi-prostrate species as Po/ypremum procumbens and Hedyotis lancifolia. Along the
eastern coast are occasional plants or groups of Avicennia germinans and Conocarpus
erectus at the water’s edge and Rhizophora mang/e in the shallows (Fig. 4), but there are
no extensive mangrove communities. |mmediately inshore from the mangroves are rem-
nants of a littoral-thicket community, with a few plants of Borrichia arborescens, Cordia
sebestena, and Sophora tomentosa, as well as the more abundant Cocco/oba uvifera.
Patches of Sesuvium portulacastrum (the most abundant of these succulents), Phi/oxerus
vermicularis, and Batis maritima grow in open areas near the water’s edge, especially
around exposed beach-rock and shingle near the northeast tip of the island. Panicum
virgatum grows in a dense stand just north of the field station and in smaller patches
elsewhere, and other graminoids, especially Sporobolus virginicus and Fimbristylis
cymosa, occasionally form dense patches. A few other species, including Lipp/a nod/flora,
Ernodea /ittoralis, and Ambrosia hispida, form patches up to several m in diameter, but
do not dominate more extensive communities. Cassytha filiformis infestations are con-
spicuous in parts of both /pomoea-dominated communities (on numerous hosts) and on
Ernodea littoralis. Large, dense beds of Thalassia testudinum, Syringodium filiforme, and
mixtures of both occur submerged between the cay and the reef edge to the east and
south and in a narrower zone along the west coast.

3

Sixty-eight vascular-plant species, listed in Table 1, were found on South Water Cay
and in the immediately adjacent shallow waters.

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Table 1. Vascular plant species observed on South Water Cay in 1979 and 1980. a
Hydrocharitaceae 4 ;
Thalassia testudinum K.D. Koenig 11 - 1918, HAM 5 i
Cymodoceaceae 2
Syringodium filiforme Kutzing (Cymodocea filiformis (Kutzing) Correll) im:
11 1920, HAM, UNA; 1923, HAM, MO, UNA 2E
Gramineae a
Andropogon glomeratus (Walt.) B.S.P. 7 - 1919, HAM, MO fs
<ieg
Anthephora hermaphrodita (L.) Kuntze 7 - 1926, HAM, MO |
Cenchrus incertus M.A. Curtis (C. pauciflorus Benth.) 5,3 - 1906, HAM, MO ce
Distichlis spicata (L.) Greene var. spicata 3 - 1894, HAM =i
Eragrostis ciliaris (L.) R.Br. 7 - 1895, HAM af
Eustachys petraea (Sw.) Desv. (Chloris petraea Sw.) 7 - 1913, HAM “B
Panicum virgatum L. var. virgatum 7 - 1925, HAM, MO a
Spartina patens (Ait.) Muhl. 3,7 - 1929, HAM of
Sporobolus virginicus (L.) Kunth 3 - 1911, HAM “e
Cyperaceae 3
Cyperus peruvianus (Lam.) F.N. Williams (Kyllinga peruviana Lam.) 7,6? - 5
1893, HAM 7
Cyperus planifolius L.C. Rich. 7,6? - 1892, HAM, MO , ¢
Fimbristylis cymosa R. Br. subsp. spathacea (Roth) Koyama (F. spathacea Roth) :f
1,7 - 1887, 1888, HAM ig

?
7
Palmae :
Cocos nucifera L. 9 7.
Liliaceae :
Dracaena sp. 9 i
Amaryllidaceae 4
Hymenocallis littoralis (Jacq.) Salisb. 10 - 1930, HAM : 4
Casuarinaceae ) i
Casuarina equisetifolia J.R. & G. Forst. 10 7.

Amaranthaceae
Philoxerus vermicularis (L.) Beauv. 3,1 - 1905, HAM

.

Polygonaceae

Coccoloba uvifera (L.) L. 1,6 - 1889, HAM

Bataceae

Batis maritima L. 3,2,1 - 1904, HAM

Nyctaginaceae

Boerhavia coccinea Mill. (B. diffusa L., s. lat.) 5 - 1866, HAM

Aizoaceae

Sesuvium portulacastrum (L.) L. var. portulacastrum 3,7 - 1867, HAM

Portulacaceae

Portulaca oleracea L. 1,7,2? - 1896, HAM, MO
Portulaca pilosa L. 7 - 1873, HAM

Lauraceae

Cassytha filiformis L. 1,6 - 1861, HAM

Cruciferae

Cakile lanceolata (Willd.) O.E. Schultz 1 - 1884, HAM

Leguminosae

Canavalia rosea (Sw.) DC. (C. maritima (Aubl.) Thouars; C. obtusifolia
(Lam.) DC.) 1 - 1863, HAM

Crotalaria retusa L. 1 - 1883, HAM

Desmodium incanum DC. var. incanum (D.canum Schinz & Thell.) 5 - 1907,

HAM
Desmodium tortuosum (Sw.) DC. 5 - 1922, HAM
Sophora tomentosa L. 1 - 1886, HAM
Vigna luteola (Jacq.) Benth. 1 - 1874, HAM

Surianaceae

Suriana maritima L. 1,3 - 1869, HAM

Euphorbiaceae

Euphorbia blodgettii Engelm. ex Hitchc. (Chamaesyce blodgettii (Engelm. ex
Hitchc.) Small) 7 - 1899, HAM, MO

Euphorbia mesembryanthemifolia Jacq. (Chamaesyce mesembryanthemifolia
(Jacq.) Dugand; E. buxifolia Lam.) 7,1? - 1876, 1880, HAM

Euphorbia trichotoma H.B.K. 7 - 1900, 1901, HAM
Phyllanthus amarus Schum. (P. niruri sensu auctt., non L.) 7 - 1875, HAM

Malvaceae

Hibiscus rosa-sinensis L. 9

Sterculiaceae
Waltheria indica L. (W. americana L.) 7,62 - 1877, HAM; Backman s.n., MO

NVINOSHLINS Sil

Passifloraceae
Passiflora suberosa L. 6,1? - 1924 HAM, MO

NUILIILILSONI

Rhizophoraceae
Rhizophora mangle L. 2 - 1962, HAM

Myrtaceae
Psidium guajava L. 9

INSTITULION

Combretaceae
Conocarpus erectus L. 1,3 - 1891, HAM
Terminalia catappa L. 9?,1?

Loganiaceae
Polypremum procumbens L. 7 - 1903, HAM, NY

Apocynaceae
Nerium oleander L. 9

LIORARICS OMIIMNSUNIAN

Plumeria rubra L. 9

Convolvulaceae
Ipomoea macrantha Roem. & Schult. 1 - 1928, HAM
|pomoea pes-caprae (L.) R. Br. ssp. brasiliensis (L.) Ooststr. 1 - 1881, HAM,
MO

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lpomoea stolonifera (Cyrillo) J.F. Gmel. 1 - 1913, HAM

Jacquemontia havanensis (Jacq.) Urb. (J. jamaicensis (Jacq.) Hallier fil.) 1 - 1965,
HAM

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Boraginaceae
Cordia sebestena L. 1 - 1921, HAM

Tournefortia gnaphalodes (L.) R.Br. ex Roem. & Schult. (Mallotonia gnaphalodes
(L.) Britt.) 1,7 - 1914, 1927, HAM

Verbenaceae
Avicennia germinans (L.) L. (A. nitida Jacq.) 2 - 1908, 1909, HAM 2
Lippia nodiflora (L.) Michx. (Phyla nodiflora (L.) Greene) 6,7 - 1885, HAM ;
Stachytarpheta jamaicensis (L.) Vahl 7,1? - 1882, HAM

Solanaceae '

Lycopersicon esculentum Mill. (Solanum lycopersicum L.) 8

Rubiaceae

Spermacoce assurgens Ruiz & Pavon (Borreria laevis sensu auctt., non (Lam.)
Griseb.) 7 - 1868, HAM

Erithalis fruticosa L. 6 - 1864, HAM
Ernodea littoralis Sw. 6 - 1898, HAM

Hedyotis lancifolia Schumach. (Oldenlandia lancifolia (Schumach.) DC.;
O. herbacea sensu auctt., non (L.) Roxb.) 7 - 1902, HAM

Compositae

Ageratum littorale A. Gray f. setigerum Robins. 5,4 - 1870, HAM
Ambrosia hispida Pursh (A. crithmifolia DC.) 1 - 1897, HAM
Borrichia arborescens (L.) DC. 1,3 - 1871, HAM

Eclipta alba (L.) Hassk. (E. prostrata (L.) L.) 5,7 - 1879, HAM, MO
Emilia sonchifolia (L.) DC. ex Wight 4,5 - 1878, HAM

Wedelia trilobata (L.) Hitchc. 1 - 1872, HAM

The first number following each name indicates the probable most important means

of inter-island dispersal, or status if introduced, according to the list below. The second
is my collection number for the voucher specimen(s), with abbreviations for herbaria
following Holmgren & Keuken (1974).

Probable Dispersal Adaptations

Adapted to dispersal by oceanic transport of seeds or fruits; seeds or fruits are
capable of floating, often for long periods.

Adapted to dispersal by oceanic transport of young seedlings; includes viviparous
species (seedlings transported with fruit).

Dispersal probably effected by oceanic transport of whole plants, stem fragments,
rhizomes, or bulbs; plants may be characterized by ample, buoyant storage tissue,
brittle stems, etc.

Adapted to dispersal by wind; fruits or seeds are plumed or winged.

Having definite adaptations to transport attached to feathers or feet of birds, hair
of mammals, and/or clothing; seeds, fruits, or associated structures are barbed or
sticky.

Adapted to internal transport by birds; seeds are protected by hard seed coats,
endocarps, etc., and in many cases are enclosed in fleshy fruits.

Probably dispersed largely as fruits or seeds on “rafts’’ of dislodged vegetation or
floating debris, in mud adhering to feet of birds, shoes of humans, etc.; in many
cases lacking obvious adaptations to other means of long-distance dispersal; seeds
often small and numerous.

Probably introduced via seeds discarded in garbage; not established.

9. Introduced species persisting where planted but not spreading.
10. Planted species now reproducing spontaneously and spreading.

11. Marine species dispersed largely through fragmentation.

The Dracaena sp. is a cultivated species seen only without flowers.

Carrie Bow Cay, a smaller islet 1.14 km S of South Water Cay, is likewise largely
vegetated by planted coconut palms. The herbaceous vegetation was sparse in 1979, com-
prising only small patches and a few scattered individuals of Sesuvium portulacastrum,
Portulaca oleracea, and Cakile lanceolata, plus one Hymenocal/is Iittoralis near one of the
buildings.

Most of the species listed above are widely distributed in the Caribbean area. Of the
60 native and naturalized species, all but two -- Panicum virgatum and Euphorbia
trichotoma -- are also present in the flora of Jamaica (1015 km EbN) and/or the
Cayman Islands (on the basis of being listed by Adams, 1972, with the addition of
Ageratum l/ittorale from Johnson, 1971), and all but three - Anthephora hermaphrodita,
Cyperus peruvianus, and Hedyotis /ancifolia -- are known from southern Florida (ca.
1080-1230 km NE) (Long & Lakela, 1971). Some of the widespread species, e.g.,
Canavalia rosea, Sophora tomentosa, Suriana maritima, and Waltheria indica, occur along
seacoasts throughout much of the tropics or at least in widely separated areas, presumably
as a result of long-distance dispersal by ocean currents. Of the 26 species listed by
Fosberg (1974) as being characteristic of coral-island vegetation throughout the tropics,
11 are present on South Water Cay. Other species, such as Desmodium incanum, Eclipta
alba, and Emilia sonchifolia, are weedy, commonly colonizing recently disturbed sites,
and in many localities are evidently introduced inadvertently through human activities
(see Johnston, 1949).

Adaptations to dispersal listed above are based on my observations, published des-
criptions of fruits and seeds, and information compiled by Guppy (1917), Ridley
(1930), and Gunn & Dennis (1976). A high proportion of the plants of this cay are
adapted to oceanic dispersal of seeds or fruits. Some of these are in families and genera in
which most species do not exhibit such adaptations, e.g., the Cyperaceae, Leguminosae,
and Compositae (cf. Ridley, 1930, p. 250). Because of the relatively short distances
among the cays and from the mainland, buoyancy for two or three days or even less
would probably sometimes suffice for dispersal.

Stoddart’s (1962b, 1965a) observations of shoreline erosion and the disappearance of
many populations of shrubs that characteristically grow close to the water’s edge in-
dicates the frequency with which large portions of plants, including storage organs, may
be uprooted during storms and presumably carried by ocean currents to other islands.
Many species, including most of the perennials, may sometimes be dispersed in this way.

Seeds of virtually all of these species could at times be dispersed via floating
vegetation, in mud adhering to the feet of birds, or in debris inadvertently transported by
humans. Guppy (1917), Ridley (1930, esp. p. 252), and authors cited by them considered
transport of fruits and seeds on floating logs to be particularly important in the inter-
island dispersal of many shoreline shrubs, specifically mentioning Borrichia arborescens
and Suriana maritima, among others, and also some of the annual psammophytes such as
Portulaca oleracea. According to F.R. Fosberg (personal communication in 1980), this

-NVINUSHLINS S31

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concept should be expanded to emphasize “'rafts’’ of dislodged vegetation more than
smaller fragmenis such as logs; such ‘‘rafts’’ may carry seeds and similar small propagules
as well as transporting the larger plants they comprise. In the above list, this mode of
transport is noted only when it seems likely to be of major importance.

Tne Great-tailed Grackle, Cassidix mexicanus, a common species on South Water
Cay, is probably by far the most important agent of endoornithochorous dispersal in the
Belizean cays. This species consumes a wide range of food items, including various berry-
like fruits (Skutch, 1958). Only those species appearing to be specifically adapted to
endoornithochorous dispersal are noted in the list, but a small percentage of seeds
normally destroyed when eaten by birds may occasionally be egested undamaged. Like-
wise, wind, during severe storms, may transport a wide range of relatively light seeds.

Stoddart’s (publ. 1963, 1969) observations of cays including South Water and
Carrie Bow cays in 1960, 1962, and 1965 provide a basis for assessing changes in
the vegetation over a 20-year period. Following the destruction of the major resort build-
ing on South Water Cay in 1961, the ornamental gardens have scarcely been maintained as
such, and much of the area is now more or less open coral sand or vegetated by the
/pomoea stolonifera-dominated community, although many of the ornamental shrubs re-
main. The effects of the weather and the spread of vegetation, plus new cottage con-
struction and coconut plantings on or near the site, have eliminated all traces of the
tennis court. The present field station and most of the cottages have been built since
1961. Early in this period, further clearing was carried out on the northern part of
the island, where several cottages were built. The vegetation there can no longer be
called ‘‘dense palm thickets’ as it was by Stoddart in 1963, and none of the Thrinax
radiata palms recorded (as 7. parviflora Sw.) as ‘thriving’ there as late as 1965 was
seen in 1979 or 1980. The swampy zone of standing water mapped by Stoddart
(1963) near the northern tip of the island apparently no longer forms, although inter-
mittently damp depressions remain. Most of the ground-cover species recorded for the
northern part of the island in 1962 were still fairly plentiful in 1979-1980. Along the
shingle ridge at the northern end, Sesuvium portulacastrum was still present although
perhaps less extensive, but the crest-zone of Tournefortia and Suriana on this part of
the island had disappeared, presumably because of further erosion.

Comparing the present vegetation with that recorded by Stoddart in 1965 (publ.
1969), the two /pomoea-dominated communities on South Water Cay, described above,
appear to have changed little; the dominants then present in both were present in similar
quantities in 1979-1980. With the exception of Thrinax radiata, all of the other species
mentioned by Stoddart (1969) in his description of the vegetation of South Water Cay
were still present.2 Rhizophora mangle has continued to recover since the 1961 hurricane;
Stoddart reported only “one or two” individuals surviving in 1965, whereas several size-
able colonies and numerous small seedlings were present in 1979-1980. Tournefortia
gnaphalodes has made less of a recovery, with only a few small plants being present in
1980. The coconuts planted on the southern part of the island during the 1960's are now
large trees.

2 According to F.R. Fosberg (personal communication in 1980), however, specimens collected on
South Water Cay by Stoddart & Spellman in 1972 or earlier by Stoddart included Erythrina sp.
(Leguminosae); Euphorbia glomerifera (Millsp.) L.C. Wheeler (Chamaesyce glomerifera Millsp.)
(Euphorbiaceae); Laguncularia racemosa (L.) Gaertn. fil. (Combretaceae); and Hedyotis corymbosa
(L.) Lam. (O/den/landia corymbosa .) (Rubiaceae), none of which were encountered in 1979 or 1980.

9

According to F.R. Fosberg (personal communication in 1979), recurrent coloniza-
tions and extinctions are characteristic of the flora of Carrie Bow Cay. Since 1965, its
vegetation has declined both in biomass and in number of species, although human use
of the island remains similar. Euphorbia (presumably £. b/odgettii) was said to form a
“patchy ground cover” in 1965, and /pomoea (probably /. sto/onifera), Tournefortia,
and grasses were also reported, but none of these were seen in 1979. Of the taxa men-
tioned by Stoddart, only Sesuvium portulacastrum and the planted coconuts remained.

_NVINUSHLINS S3l.

NUILIILILONI

—>

Acknowledgments

| am grateful to Dr. J. Allen Keast for suggesting this study and for reviewing the
manuscript; to Mr. W. Floyd Connor for his contributions to the organization of the
trips to Belize and for his suggestions of useful literature; to Dr. F. Raymond Fosberg
for his review of the manuscript and for information on nomenclatural matters; to Dr.
Fosberg and Dr. John D. Dwyer for the identification of some of the specimens; and to
Ms. Cathy Backman for some data used in this paper.

INSTIIULION

Literature Cited

Adams, C.D. 1972. Flowering Plants of Jamaica. Mona, Jamaica: University of the West
Indies. 848 pp.

Fosberg, F.R. 1974. Phytogeography of atolls and other coral islands. /n: Cameron, A.M.,
et al., eds. Proceedings of the Second International Symposium on Coral Reefs.
Brisbane: The Great Barrier Reef Committee. 1:389-396.

LIOKRARICS _OMIILAOUNIAN

Gunn, C.R., & J.V. Dennis. 1976. World Guide to Tropical Drift Seeds and Fruits.
New York: The New York Times Book Co. xiii + 240 pp.

Vatuvugdi s

Guppy, H.B. 1917. Plants, Seeds, and Currents in the West Indies and Azores: the
Results of Investigations Carried out in those Regions between 1906 and 1914.
London: Williams and Norgate. xi + 531 pp. + 1 pl. + 3 maps.

Holmgren, P.K., & W. Keuken. 1974. Index Herbariorum. Part |: The herbaria of the
world, ed. 6. Regnum Veg. 92. vii + 397 pp.

tev towwvuliiaiow

owe
a

Johnson, M.F. 1971. A monograph of the genus Ageratum (Compositae -- Eupatorieae).
Ann. Missouri Bot. Gard. 58:6-88.

Johnston, |.M. 1949. The botany of San Jose Island (Gulf of Panama). Sargentia 8.
ii + 806 pp. + 17 pl.

Long, R.W., & O. Lakela. 1971. A Flora of Tropical Florida: a Manual of the Seed
Plants and Ferns of Southern Peninsula Florida. Coral Gables: University of

Miami Press. xvii + 962 pp.

Ridley, H.N. 1930. The Dispersal of Plants Throughout the World. Ashford, U.K.:
L. Reeve & Co. xx + 744 pp. + 22 pl.

Skutch, A.F. 1958. Cassidix mexicanus mexicanus (Gmelin): Boat-tailed Grackle.
Bull. U.S. Natl. Mus. 211: 335-350.

Stoddart, D.R. 1962a. Cambridge Expedition to British Honduras. ||. Physiographic
studies on the British Honduras reefs and cays. Geogr. J. 128: 161-171 + 3 pl.

10

Stoddart, D.R. 1962b. Catastrophic storm effects on the British Honduras reefs and
cays. Nature 196:512-515.

Stoddart, D.R. 1963. Effects of Hurricane Hattie on the British Honduras reefs and cays,
October 30-31, 1961. Atoll Res. Bull. 95. iv + 142 pp. + 65 figs.

Stoddart, D.R. 1965a. British Honduras cays and the low wooded island problem.
Inst. Brit. Geogr. Trans. 36:131-147.

Stoddart, D.R. 1965b. Resurvey of hurricane effects on the British Honduras reefs
and cays. Nature 207: 589-592.

Stoddart, D.R. 1969. Post-hurricane changes on the British Honduras reefs and cays:
Resurvey of 1965. Atoll Res. Bull. 131. 25 pp. + 15 maps.

NOTE

As this paper was aoina to press, new information
was provided by Robert H. Sims, just returned from Carrie
Bow Cay. He reported that, for some years, the proprietors
of the island have, at intervals, systematically removed the
vegetation, excepting coconut trees, with the purpose of dis-
couraqing or eliminatina insects.

This would account for the naucity of the flora of this
tiny cay (see pages 7 and 9).

[Editors ]

Figs. 1-4. Plant communities on South Water Cay, photographed in 1979. Fig.
1. Community dominated by /pomoea stolonifera. Fig. 2. Community dominated
by /pomoea pes-caprae, Vigna luteola, and Wedelia trilobata, with Cocos nucifera
trees in the background.

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Fig. 3. Sparsely vegetated area with Euphorbia spp. and
graminoids, and planted row of Hymenocallis littoralis. Fig. 4. Mangroves --
Rhizophora mangle and Avicennia germinans -- along windward (east) side of island.

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