J . Linn. SOC.(Bot.)60, 382, p. 99
With 2 platea and 3 text-figurea
99
Printed i n areat Britain
July, 1967
A revision of the marine algae of SBo Tom6 and Principe
(Gulf of Guinea)
BY MARGARET STEENTOFT, F.L.S.
In-stitut for Sporeplanter, University of Copenhagen*
(Accepted for publication December, 1965)
In all 101 species and varieties of marine dgaa are mentioned for S. Tom6 and Principe. These
records arise from examination of the material in the herbarium of the Botanical Institute of
the University of Coimbra, together with material collected in 1956 by Dr C. A. Thorold.
From the latter material there are 17 new records for the island of 8. Tom&
No new species have been described, but Acanthophora ramuloaa Kutz. is emended, and
Bryopais denaa Pilger and Struvea multi-prtita Pilger are reduced to the synonymy of Byopaia
pennata var. aecunda and Struvea anaatomoaana respectively.
Information on the climate and tides of the islands is given, and a series of ‘seasonal
coincidences’ likely to influence coastal marine conditions is described for the Gulf of Guinea.
CONTENTS
.
.
Introduction
Localities.
Climate
Tides
.
Oceanography
Climatic and marine seasons, and the littoral environment
Systematic list
Chlorophyceae
Phaeophyceaa
Rhodophycem
Cyanophyceae
Acknowledgements
References
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PAQE
99
102
103
104
105
105
107
107
115
121
142
142
143
INTRODUCTION
While examining a collection of marine algae from SBo Tom6 made by Dr C. A. Thorold
in 1956, existing collections in the herbarium of the Botanical Institute of the University
of Coimbra were consulted, and it became apparent that some revision of this material
might prove interesting.
Through the kindness of Prof. Dr A. Fernandes, the Coimbra material was borrowed
on two occasions, in 1957 and in 1964-6. It proved to contain not only material listed in
Henriques (1886, 1917) and Hariot (1908), but also some undetermined material. Some
other material has also been traced and as full as possible an account is given of these
various collections.
Thorold’s collection contains 32 species, 23 of which constitute new records for SHo
Tom& Carpine (1959) lists 34 species, 13 of which are new records, and a further six are
*
Present address: Denmark8 Lcererhojskole Copenhagen.
7
ML~~GABET
STEENTOFT,
F.L.S.
100
new records in common with those in Thorold’s collection. With the rate of acquisition
of new records so high, the flora cannot be considered to be well known.
It can be claimed that identifying marine algae from the Gulf of Guinea is often a
difficult undertaking. Few species have been described from the area, so the solace of the
type specimen is denied, and endemic species are correspondingly few. The only noncalcareous one from Siio Tomb is &milaria henriqmiana Hariot.
The cauae of diEculty in identification often lies in the small size, even stunted form,
of many specimens, and hairs for example may be little developed or even absent. Unless
comparison has been possible with the Cameroun specimens collected by Lawson, those
of Lawson and of Foote in Ghana, or those collected by the present writer in Nigeria and
Ghana (Fox, 1957), there is also a general paucity of material from the Gulf. The northern
shore of the Gulf is relatively overrepresented a t present.
Usually, then, the Siio Tom6 and PrIncipe plants have had to be compared with material
from places with very different climatic and marine regimes, including (most often) West
Africa north of Sierra Leone, South Africa and western tropical Atlantic coasts. In order to
put into perspective, as it were, the Gulf of Guinea specimens discussed below, the main
features of the coastal climates of the Gulf are described, and a brief account of the
investigationsof Berrit (1962, etc.) of the oceanography of the Gulf is given. The climatic
and marine seasons are then related to each other to produce a series of seasonal coincidences, five within the Gulf alone, which somewhat enlarge the impression prevailing
of conditions round the Gulf. The better known northern shore (CapePalmas to Cameroun)
represents only one type of coincidence.
To these seasonal fluctuations must be added the effect of the periodicity of the lower
low waters. In the Gulf of Guinea these occur by day in the northern hemisphere winter,
by night in the northern hemisphere summer.
In all 101 species and varieties of marine algae are mentioned below.
*
Table 1. History of the marine and algal collectionsfrom S. Tom6 and Prlncipe
Principal collections
No. specimens traced/
Previously
species reoognized, in
listed (pro
Collector
Island
Date
present revision
parte only)
F. Welwitsch
Prhcipe
Sept. 1863
516
1 specimen mn. wid. }Barton, 1897
S. Tom6
Dec. 1860
F. Newton
8. Tom6
Mar. to Apr. 1881
Henriques, 1886
mn. uid. 6 spp.
reported
A. F. Moller
13. Tom6
June to Sept. 1886
Henriques, 1886
24117
F. J. Dias
S. Tom6
Sept. to ?Dec.
Foslie, 1897,
38/31
1886
. 1900 a, b, c,
Quintas
Ilheu das
1907, 1909, 1929
Rolas
Hariot, 1908
F. Newton*
S. Tom6 (IlhBu June 1887
daa Cabras)
Hariot, 1908
July to Aug. 1887
loll0
Prhcipe
Hariot, 1908
J. J. Ribeiro
8.Tom6
1894
18/17
S. Tom6
Apr. 1906
616
Hariot; 1908
C. Graviert
A. A. Praenpa-see under c4racilaria henriqueakna
H. Cardoso
8.Tom6
?
17/12
Rodrigues, 1906
A. W. Exell
9.Tom6
Nov. 1932
616
4 specimens
}Exell, 1944
non. vid.
C. Carpine
8.Tom6 and
June 1966
mn. vid. 34 species
Prfncipe
reported
Carpine, 1969
C.A. Thorold
8. Tom6
Aug. 1966
34/32
* Nylander (1888) confirms date of Newton’s visit as 1887.
t Gravier (1906, 1907) describes coastal conditions. Geology is described in Cotelo Neiva (1968a, b ) ,
and much helpful general information is to be found in Monod (1960).
Marine algae of SCo TmC and Principe
101
The specimens in this list are mainly to be found in the Coimbra Herbarium (COI),
and in the exsiccata lists following (except for Hb. Fox Nielsen specimens) this can be
assumed unless some other herbarium is named. Henriques (1917) is not given as a reference
in the list of principal collections, because it is based on Henriques (1885, 1886), Hariot
(1908) and Foslie (1900~).In addition it is full of minor errors. The numbering of the
Moller and Quintas specimens is very confused.
Dr Thorold's collection, made a t the request of the present writer, constitutes nos. 221
to 250, 356 and 370 in Hb. Fox Nielsen," part existing in a parallel collection in Hb. Br.
Mus.
There are three minor collectors to be mentioned. A. Franklin is given as the collector
of a specimen of Caulerp mexicum found by M. Denizot in Hb. Mus. Hist. nat., Paris.
Second, Griife is given as the collector of Bryothumnion spp. in De Toni (1903 : 974-975).
Third, J. D. Epffenhausen is the probable collector of two specimensof &milaria spp. and
two of Laurencia obtwa found in Hb. Binder, Hamburg.
O"26f
6'47'E
3'E
I
0'26"
Dias
3ia
elia
>-
S Jo60 Dos Angolare;
3.
SCry
,#'
Angra
Ponta de
DoS.
I6J a i a
Praia Grandee,--
,'
c
Canal Das Rolas
A
V I l h k u Dos Rolas
lOkm
0"
6'
Text-fig. 1. Map of S. Tom& Localities at which collections have been made are referred
to on p. 102. The broken line delimits the superhumid climatic area to the southwest (Lains e
Silva, 1968). m, mangrove; E, sand.
Mrs Margaret Nielsen, n6e Fox,now Steentoft.
I
MAEGARHITSTEENTOFT,
F.L.S.
102
Epffenhausen was a Hamburg merchant who collected for Binder in Gambia (1837)
and Angola (1839, 1842) as well as in 'Guinea' (Ghana). The specimens do not bear his
name, but only 'St Thomas' or 'St Thorn.' and the year, 1838 or 1839, but his ship, the
Herold, left Hamburg on Oct. 31st. 1838 and returned there on Oct. 25th. 1839 from 'St
Thomas, on the coast of Guinea' (seeP. E. Schramm, 1950).According to the same source,
Epffenhausen is presumed to have travelled regularly with his ship. It cannot however, be
absolutely excluded that these four plants came from St Thomas in the West Indies.
Two partial revisions of the Coimbra material have been attempted. Dr R. Lami
revised the Chlorophyceae, but did not publish his determinations, which the present
writer is grateful for permission to quote. More recently, Dr J. E. de Mesquita Rodrigues
has revised the Phaeophyceae (Rodrigues, 1960).
A study of the Cyanophyceae of the islands is being made by Dr J. Sampaio (1958,
1962a,b, 1963)and only the Cyanophyceaefrom Dr Thorold's collection will be mentioned
in the following list. Sampaio has so far noted only one larger alga, Cladostephw,
verticillatwr
.
LOUALITIES
In the exsiccata lists following, information about localities is quoted directly from
the labels, translations being given in inverted commas, the comments of the present
writer in square brackets.
It is not always clear where some of the localities lie. Moller collected only in the Bahia
de Ana Chaves; his second locality, Praia Lagarto, is part of the shore of this bay. Quintas
collected on the southern shore of the Canal das Rolas, on the Ilh6u das Rolas on the
equator ('Equador') and this is where his sublittoral specimens come from, at depths
7'20'E
1.45'
7'32'E
1'45'N
10km
I
lo30'
7
3'E
"
"
"
"
I
? 30"
7' 2"
Text-fig.2. Map of Prhcipe. For remarks see legend to Text-fig. 1.
Marine algae of S6o Tom6 and Principe
103
down to 10 ft, i.e. 3 m. The maximum depth of the Canal is 30 m. But some specimens
are labelled simply ‘Ilh6u das Rolas’ and may come from another place. His ‘Angolares’
specimens come from the coast of S. Tom6 itself, between Praia Inhame and S. Cruz.
Three of Gravier’s localities are clear, Praia Inhame, Ferniio Dias and Bella Vista, but
the fourth, Ribeiro Peixe, is doubtful. The river a t present known by this name seems a
little far from S. JoHo to qualify.
Thorold‘s material came from four localities which were not kept separate. They are :
‘Beach near S. Tom6 City [Baia de Ane Chaves] composed of silt or sand with some
exposed rocks’, ‘ Beach near airport with no exposed rocks’, Praia S. Jer6nymo and Agua
Iz6, the last being purely rocky.
CLIMATE
Both islands have a southern equatorial type of climate, the dry season and the lowest
air temperatures occurring in the northern hemisphere summer, and there being a ‘twopeak’ rainy season.
Lains e Silva (1958 :64) classifiesthe climate of the islands in terms of evapotranspiration,
following Thornthwaite (1948) (Table 2).
Table 2. Comparison of seasonal distribution of coastal rainfall
Ghana*
Takoradi
(semiarid)
Dry
season
Little
rains
Little
dry
season
Rains
S. Tom6 and Principe
(aridsemiarid*)
Gabon*
Libreville
October
Aug. to Sept.
Dec. to Jan.
Apr. to July
Feb. to May
Cameroun
Victoria
(subhumidhumidsuperhumidt) (superhumid) (superhumid) (superhumid)
Nov.toMarch June to Sept. June to Sept. June to Aug.
October
Kribi
January$
(Feb.)March March to May Feb. to May
to April
to June
Jan.t(Feb.) February
July
Oct. to Nov.
(December)5
Sept. to Jan.
Dec.toJan.
-
Aug. t o Nov. Feb. to May to
December June to Oct. to
November
Bold face indicates hot season.
* After Anon (1958).
t After Lains e Silva (1958). I n the arid-semiarid zones of S. Tom6 and Principe, Lains e Silva makes
the dry season longer (May to Oct.), fixes the little dry mason in November, and shortens the rains
(March to April). The subhumid climate differs from the humid and superhumid types in having the
shorter little rainy season and the longer little dry season.
$ Season of doubtful occurrence, somewhat less wet than preceding and following months.
3 Part of rainy season in humid type climate only.
SBo Tom6 and Principe have a climatic regime similar to that of the Gabon-Rkpublique
du Congo type. The drier parts of Siio Tom6 differ only from the wetter parts, and from
Principe, in the transposition of the main and little rainy seasons. The hot season is also
wet, while on the northern coast of the Gulf (Cape Palmas to Victoria), the hot season is
dry.
The dry season is conventionally recognized as a period of reduced rainfall (the ‘conventional dry season’ of Table 3), but rainfall can only have a minor direct effect on littoral
algae if, as seems probable, they grow low in the intertidal zone or under continuous
wave or surf action (as described for example in Lawson, 1955). Prolonged periods of low
relative humidity could, in certain circumstances, have more effect.
Figures are available for only the grosser aspects of climate on the shores of the Gulf,
but i t has been possible to redefine ‘dry season’ as ‘the period in which the mean monthly
MARGARET
STEENTOIW,
F.L.S.
104
relative humidity is less than the median minimum relative humidity’. I n Table 3, the
prolonged hot dry season of the northern shore of the Gulf is shown very clearly using
this definition. There are both practical and theoretical criticisms of such a definition,
but unfortunately no detailed studies of the shore environment in the Gulf exist. At least
the studies of Lawson (1955, 1957)’ showing the very restricted algal zone of sheltered
conditions (minimum wave and surf action) in Ghana, and the retreat of several species
into the lower part of the intertidal zone during the dry season, do not contradict the
definition immediately.
Table 3. Comparison of climatic semom on the shorefl of the Gulf of Guinea
Duration of dry
season (months) and Temperature Mean
range of minimum
season
annual
relative humidities
(h, hot;
rainfall
(% R.H., in brackets) c. cool)
(in.)
Cape Palm(Tabou)*
Takoradi
Accra
Victoria
Kribi
Libreville
S h Tom6
7+2
7
7
7
7
3
4
Loangot
Loande
MoqImedee
1 1 (73 to 74)
2
(74 to 76)
7
(72 t o 7 6 )
(77to80)
(87 to 74)
(81 t o 6 8 )
(74 to 80)
(77 t o 8 1 )
(89to71)
(70 to 74)
+
h
h
h
h
h
C
C
Rainfall in dry mason
(approx. % of total
rainfall), and minimum monthly rainfall
(in., in brackets)
92.6
48.6
28.6
168.8
122.3
98.8
38.0
48 (1.6)
64 (1.0)
64 (0.8)
24 (1.3)
47 (3.8)
1 (0.1)
8 ( < 0.1)
48.1
12.9
2.1
13 ( 0 )
34 ( < 0.1)
100 ( < 0.1)
c+h
h
h
Conventional
dry maaon
hot, humidity
low
Conventional
dry season
cool, humidity
low
Conventional
dry seaaon
cool, humidity
high
* Two months in the cool, rainy mason also have low minimum relative humidities.
t Climate intermediate, one month (July) of dry season cool and totally dry,the other (March)hot
and wet.
TIDES
Tidal ranges in the Gulf of Guinea are relatively small, though it must be remembered
that wave and surf action can be considerable.
Table 4. Mean tidal ranges in the aUlf of Guinea (ft)*
SBo Tom6
(City)
Prhcipe
(8. Antbnio)
Ghana
Nigerie
(Takoradi) (Lagos)
Bay
Mean tidal range
Mean tidal range
(wings)
Mean tidal range
(new4
3.4
4.8
643
3.9
3.0
4.2
2.6
3.4
2.3
2.3
1.9
1.6
* After Admiralty (1984).
Throughout the Gulf of Guinea the lowest low waters (those no higher than mean lower
low water, the average of the lower of the two daily low waters throughout the year)
occur only or mainly by day in the period October-November to March-April. Between
March-April and October-November they occur only or mainly by night. The result of
this seasonal incidence of extra low tides is that maximum littoral exposure occurs during
the hot, dry and least humid season a t places west of Kribi, during the hot, wet and most
Marine algae of S b Tom6 and Principe
105
humid seasons from Loanda southwards while i t occurs during the hot, wet and most
humid season a t places such as SBo Tom6 and in Gabon.
OCEANOQRAFHY
Berrit has described the oceanic conditions prevailing in the eastern tropical Atlantic,
between Cap Blanc and Cabo Frio, in a series of papers (listed in Berrit, 1962) to which
reference should be made for details. Only the characteristics of the surface waters will be
mentioned here.
Briefly it appears that, in the northern part of the Gulf of Guinea, between approximately the R. Sierra Leone and Cap Lopez, is a mass of ‘Guinea Water’, which is permanently warm (above 24°C) and of low salinity (below 35%,). There is only periodic
upwelling of cooler, saltier water t o the surface, occurring a t points between Cape Palmas
and Cape Formosa.
The northern and southern boundaries of the Guinea Water oscillate seasonally. The
northern boundary moves from the R. Sierra Leone as far north as Nouachott in the
northern hemisphere summer, while the southern boundary moves as far north as Cap
Lopez. I n the southern hemisphere summer the southern boundary moves south from
Cap Lopez to Loanda, the northern boundary moving south again to the R. Sierra Leone.
Similar and parallel oscillations of the other two types of water mass adjacent to the
Guinea Water also occur. These two types are the ‘eaux chaudes sal6es ’ immediately
adjacent to the north and south of the Guinea Water, and the ‘eaux froides salks’ of
Berrit lying still further north and south respectively. The former have a temperature of
a t least 24°C and a salinity of a t least 35%,, the latter a temperature lower than 24°C
and a salinity of a t least 35%,.
Both Principe and Sio Tom6 lie in the area occupied by Guinea Water where the
temperature lies between 24 and 30 to 31”C, falling to between 24 and 27°C only in the
cool season (June to August). SBo Tom6 lies further south than Principe and can be
expected to experience a somewhat longer period of reduced sea temperatures. It is
more difficult to generalize about salinities, but it seems probable that Sio Tom6 has the
higher values for a longer period of the cool season than Principe, though whether the
value of 35%, is passed, and if so, by how much and for long, is not clear.
Salinities in the areas immediately offshore to the mainland coast of the Gulf of Guinea
are very much influenced by freshwater coming from rivers. I n Fox (1957: 617) approximate values of 25 to 35Z0are given for high tides a t Lagos all the year round and for low
tides in the dry season, but a t low tides in the rains and just after a value of €i%
is given.
,
Watts (1960: 1162) has measured salinities just within the mouth of the R. Sierra Leone
and gives extreme values of 3 ~ 4 %
and~ 34.4%,. Thus the amount of fluctuation of salinity
in the offshore water a t the mouth of one of the larger rivers and of one of the larger lagoon
systems is known and it is probable that the Guinea Water as a whole undergoes fluctuations in the course of the year, though the fluctuations are probably not so large away
from the mainland coast.
There are thus marine seasons in the Gulf, i.e. fluctuations of temperature and salinity
just as there are climatic seasons, i.e. fluctuations of rainfall, temperature and minimum
relative humidity.
CLIMATIC AND MARINE SEASONS, AND THE LITTORAL ENVIRONMENT
Because of the climatic and oceanographic peculiarities of the eastern tropical Atlantic,
the littoral environments existing on the mainland coast of the Gulf of Guinea and on the
shores of the islands of the Gulf are probably very varied. I n Table 5 the nine possible
combinations of climatic and marine season are summarized. It should be remembered
that the lower low waters occur only or mainly by day in the period of the northern hemisphere winter throughout the Gulf.
MARGARETSTEENTOFT,
F.L.S.
106
Except for Sourie (1954) there are no detailed studies of the physical conditions of the
littoral habitat for the Atlantic coast of Africa south of Cap Blanc (excluding South
Africa). In the absence of such information only a very approximate idea of the nature
of the littoral environment can be arrived at.
But it would seem that a good deal of variety is possible and that S&o Tom6 and
Table 5 . Climatic and marine setmom O R the Atlantic coast of Africa between Cap Bhnc
and Cab0 Frfo
N. hemisphere summer
Lower low tides by
night
N. hemisphere winter
Lower low tides by
day
Climatic
mwon
Mauritanie
Maxine
aeason
Climatic
mason
Cap Blanc
cool, wet
humidity
low
1
hot. dry
humidity
high
Nouakchott
cool,
dry
2
hot,
wet
Port. Guinea
Cabo Verde Is.
Gambia
R. Cacheu
Guin6e
Cap Verga
Sierra Leone
Gulf of Guinea
Liberia
Cameroun
R. Sierra Leone
FS
Marine
aeason
1
CS
3
hot
dry
4
6
cool,
wet
Guinea
water
cool,
h Y
humidity
high
FS
Cap Palmas
Kribi
6
SBo Tom6 and Prlncipe
Gabon
Cap Lopez
Angola
Loanda
Mopamedes
S.W.Africa
Cab0 Frio
hot,
wet
7
hot,
wet,
humidity
low
cs
8
9
FS
S. hemisphere summer
9
.
S hemisphere winter
Oceanographic information summarized from Berrit (1961, Fig. 9) omitting details of zones of
upwelling. No scale of latitude is implied in the spacing of the places mentioned. The dry season is the
conventional one (p. 103).
FS-'eaux
froides sal6es' of Berrit. Temperature always under 24"C, salinity always over 36%.
CS-'eaux chaudes sal6es' of Berrit. Temperature always over 24"C, salinity always over 36%.
Guinea water temperature always over 24°C. salinity always under 36%.
1 to 9, various types of climatic and marine seasonal coincidences.
Except where specified, the dry aeason can be m u m e d t o have the lowest minimum relative
humidities, the wet semon the highest ones.
Marine algae of Stio Tomi and PrCncipe
107
Principe are probably unique in the particular coincidence of climatic and marine conditions they enjoy (type 6 in Table 5). While the mainland coast south of Kribi as far as
the north of Angola has a similar pattern of seasonal coincidence, it also has to be pointed
out that this part of the mainland coast has a more variable climate in which the extremes
of temperature and relative humidity are wider apart.
To what extent, and in what manner, the littoral environments of the shores of the
Gulf are affected by the seasonal coincidences demonstrated remains to be shown. But it
is reasonable to expect that, the grosser environmental features of the shore will be
biologically operative to some extent, a t any rate on the morphologically more plastic,
or physiologically more sensitive, species.
SYSTEMATIC LIST
Geographical distributions refer to each species as a whole, unless infraspecific taxa
are treated separately. In these distribution records, ‘Africa’ means the Atlantic coast of
Africa.
Herbarium abbreviations are those used in Index Herbariorum (Lanjouw t Stafleu,
1964).
Chlorophyceae
Ulvaceae
Enteromorpha Link 1820
Enteromorpha clathrata (Roth) Grev., Alg. Br.: 181 (1850),insensu Bliding, SOC.Bot,
Lund. 8 (3): 106 (1963).
Three collections of this species were found among Thorold’s material. I n no. 222 two
forms were present, one coarser and with a cell size more typical of the species as it is
seen in Europe, the other a finer form with cells smaller than is usual for the European
form. The latter is also known from Lagos (Fox, 1957 : 618) where i t is seen both a t the
landward limit of algal growth on the Moles, and relatively high on the outer faces of the
Moles in the surf zone (Steentoft Nielsen, 1959: 35). It is possible that similar conditions
exist in S. Tom&,but no information is available.
Exsiccata s. TO&: Fox Nielaen 222, 235 (epiphytic on Padina commersonii) and 238
(epiphytic on Sargassum vulgare var. foliosissimum), sin. loo., August 1956
( =Thorold 620 in BM spirit collection).
Distribution Africa: a new record for S. Tom&;see also Fox (1957: 618).
Enteromorpha flexuosa subsp. flexuosa Bliding, SOC.Bot. Lund., 8 : 73 (1963); E .
intermedia in Fox (1957 : 618) ; as E . prolifera in Hariot (1908 : 162) and Henriques (1917 :
165).
This would appear to be the most constant of the Enteromorpha species on the Atlantic
coast of Africa.
Exsiccata s. TOM$: Ribeiro 18, Ropa Praia da Nazareth, 1894. Fox Nielsen 225, sin. loc.,
August 1956 ( =Thorold 610 in BM spirit collection).
Distribution Africa (putative, pending examination of African material) : S. Tom6; as
E . flexuosa in South Africa, S&n&gal,
Cab0 Verde Is., Maroc; as E . compressa var. $exuosa
in Tanger ;as E . intermedia in Nigeria, Ghana and Sierra Leone ;as E . prolifera in ?Annobon
and Apores; as E . lingulata in ?Canary Is. Cosmopolitan.
108
MAR~ARET
STEENTOFT,
F.L.S.
TJlva Thuret 1864
Ulva faaciata Delile, PI. Egypte: 163 (1812).
This species was first reported in Exell (1944 : 386) and is also present in Cardoso’s and
Thorold‘s material. Only the last mentioned plants have the ribbon-shaped thallus
supposed to be characteristic of the species. U . fmchta seems to be the common Ulva
species of the warmer eastern Atlantic.
Exsiccata s.
TO&: Cardoeo a m , sin. loc. Exell 464, 472 and 473, Praia Pequeno,
30.ix.1932 (BM).Fox NieLen 221, sin. loc., August 1956 (duplicate BM).
Distribution Africa : South Africa, Angola, Annobon, S. Tomb, Ghana, ?Guide,
SBndgal, Cab0 Verde Is., Mauritanie, Maroc, Tanger. Pantropical.
Ulva lactuca L., Spec. pl. 2: 1163 (1753).
Both Thorold’s and Newton’s plants are small, with thalli up to 6 cm long. These are
new records for both islands.
Exsiccata
8 . TO& :Fox NieLen 356, sin. loc., August 1956.
P R ~ C I P E :Newton a. n., Praia de Sundy, 1887.
Distribution Africa: South Africa, Angola, St. Helena, RBpublique du Congo, S. Tom6,
Principe, Ghana, ?Guin&, S6n6ga1, Cab0 Verde Is., Mauritanie, Canary Is., Maroc,
Madeira, Tanger, Agores. Cosmopolitan.
Cladophoraceae
Chaetomorpha Kiitz. 1845
Chaetomorpha antennina (Bory) Kutz., Spec. a&. : 379 (1849).
Cardoso’s plants are relatively stunted, as are Nigerian plants (Fox,1957: 618) but
with the filaments slightly wider, up to 600 p, than in the latter. The species would appear
to be a constant and probably conspicuous one on the Atlantic coast of Africa. The
record for S. Tom6 is new.
Exsiccata s. TO& : Cardoso am., sin. loc.
Distribution Africa : Portuguese East Africa, S. Tom& See also Fox (1957 : 618).
Chaetomorpha clavata (C. Ag.) Kutz., Bot. Ztg, 5 : 166 (1874).
The plants referred to this species are of the same size and general appearance as those
of C. antennina mentioned above, but proved upon examination to have filaments
distinctly broader at their upper ends than at their lower ends. The top part of each
filament was 1 mm wide on average, while the lower part of the same filament is only
300 p. The plant is about 10 cm high.
Papenfuss (1940 : 200) differentiates between the West Indian species C. clavata and
the South African C . robwta (Aresch.) Papenfuss largely on grounds of size. He states
that the dimensions of the type specimen of C . clavata (2 cm tall with filaments 700 p
wide) show that it is a different species from C. robwta (20 cm tall or more, with filaments
1 to 1.5 mm wide). However, judging from Taylor (1960 : 73) C. clavata as found on the
warm Atlantic coasts of the Americas is a more variable species than hitherto recognized,
up to 30 cm high with filaments 600 to 760 p wide basally and 1.5 mm wide apically. The
S. Tom6 plant can reasonably be considered to belong in C. clccvatcc as thus defined, especially as reduction in size is quite common among Gulf of Guinea species. This is a new
record for S.Tom&
Marine algae of S& Tomi and Principe
109
The independance of C. robwta as a separate species should perhaps be reinvestigated.
Exsiccata s. T
O I :~Cardoso
s.n., sin. loc.
Distribution Africa : Portuguese East Africa, ?South Africa, S. Tom&,Pantropical.
Chaetomorpha crassa (C. Ag.) Kiitz., Spec. alg. : 379 (1849);Phyc. germ.: 204 (1845).
This occurs free floating, tangled with other algae. The filaments are 400 p wide on
average, with cells twice as long as wide. More material of this species is desirable. The
record for S. Tom6 is new.
Exsiccata
S. TOM^: Fox Nielsen
223, sin. loc., August 1956.
Distribution Africa : Portuguese East Africa, South Africa, S. Tom6, Agores. West
Indies, Mediterranean, Atlantic coasts of Europe, Indonesia. Perhaps cosmopolitan.
Cladophora Kiitz. 1843
Cladophora laetevirens (Dillw.) Kutz., Phyc. gen. : 267 (1843); Soderstrom, Bot. Goth.,
1 : 113 (1963).
Of Cardoso’s seven plants Soderstrom says (in litt.) ‘Probably detached CZudophora
laetevirens (Dillw.)Kiitz. The size of the cells and the end clusters are just like this species.
However, C1. laetevirena is not reported earlier in a detached form?’
Newton’s specimen on the other hand, Dr Soderstrom says (in litt.), ‘is quite like
English laetevirens’. These are new records for both islands.
It is difficult to give any idea of the distribution of this species, but following the
synonymy reported by Soderstrom (1963: 113) a putative distribution has been arrived
at, though it should be regarded with great caution.
Exsiccata s. TOM& : Cardoso s.n., sin. loc. [two sheets].
PRfNcIPE: Newton s.n., Praia de Sundy, 1887.
Distribution Africa: S. Tom6, Principe, Cab0 Verde Is., Maroc, Canary Is., Madeira,
Tanger . Perhaps most Atlantic and Mediterranean coasts. ?New Zealand.
Cladophora oblitterata Soderstrom, Bot. (20th.: 1,47 (1963).
According to Van den Hoek (1963 : 186) C. oblitterata is to be included in C . dalmatica
Kutz., but Soderstrom disagrees. A putative distribution has been arrived a t in the same
way as for the previous species.
Exsimta S. TOM&.: Fox Nielsen 229, sin. loc., August 1956 (=ThoroZd 618 in BM
spirit collection).
Distribution Africa: S . Tomb, Sbnbgal, Canary Is., perhaps most Atlantic and Mediterranean coasts.
Cladophora prolifera (Roth) Kiitz., Phyc. germ., 207 (1843); Hariot (1908: 162);
Henriques (1917: 165); as C. catenata (f. robustior) in Henriques (1886: 217; 1917: 165).
Hariot originally revised the name of Quintas’s plant, and Lami (unpub.) and Soderstrom have since agreed with him. It seems probable that Norstedt or J. Agardh created f.
robustior in order to accommodate this plant, whereas its larger size and different structure
should have placed i t in C. prolifera.
Some records of C. catenata in semu Kiitz. may relate to C .prolifera. Hamel (1931 : 172)
also considered that C. multiJida Kiitz. (1849: 390) was possibly part of this species too.
C. multi$da is known from Gabon, Cameroun and SBn6gal.
MARGARETSTEENTOJTT,
F.L.S.
110
Exsiccata s. TOM&: Quintarr sm., Equador, Canal das Rolas (alga do fundo, basalto)
1885.
Distribution Africa : Portuguese East Africa, South Africa, S. TomB, ?Gabon,
?Cameroun,SBnbgal, Cab0 Verde Is., Canary Is.,Maroc, Madeira, Tanger, Aqores. Widely
spread in warmer waters.
Lola A. & G. Hamel 1929
Lola gracilis (Kutz.) Chapman, Bull. I m t . Jamaica Sci. Ser., 12 (1): 73 (1961).
The filaments are 45 to 65 p wide with cells two to six times as long as wide. Thorold’s
plants occurred in small floating masses.
Exsiccata s. TOMI%: Quintas 29, Equador, Canal das Rolas. Alga a 10 pBs de fundo
calcario. 1885. Fox Nielsen 227, sin. loc., August 1956 (=ThoroZd 614 in BM
spirit collection).
Distribution Africa : S . Tom& Probably pantropical.
Rhizoclonium Kiitz. 1843
Rhizoclonium implexum (Dillw.)Kutz., Phyc. germ.: 206 (1845);Koster, Pubbl. Staz.
zool. Napoli, 27: 343 (1955).
The cells are mostly 1 2 6 p wide by three to five times longer. The filaments grew
entangled with Bostychh rdicans and Murrayella pericludos, as might be expected of a
species which can occur in brackish water as well as salt water.
Exsiccata s. TOMI%:Fox Nielsen 232, sin. loo., August 1956 (=Thorold 609 and 611 (as
Bostychia moritziana) in BM spirit collection).
Distribution Africa : S . Tomb, Nigeria, Sierra Leone, Maroc, Tanger. Cosmopolitan.
Anodyomenaceae
Microdictyon Decne. 1839
Microdictyon calodictyon (Mont.) Decne., P1. Arab. in A r c h mus. nutn. Hwt. nut.,
Paris. 2, 115 (1841) : Bergesen, R. a’unske Videmk. Selsk., Biol., 5 (3): 32 (1925).
Cardoso’s plant forms an irregular black net, up to 5 cm across and not adhering to
paper. The main filaments are somewhat obscure, composed of cells (100)-120-(150) p
wide by up to 260 p long. As a rule the cells are two to three times as long as wide and
somewhat swollen. The interstices in the thallus are relatively small compared with the
cell diameters.
This species is known with certainty only from the type locality, the Canary Is. Taylor
(1960: 121) gives two records for Brazil as ‘uncertain’. The record for South Africa is
from Barton (1896 : 458).
Exsiccata s. TO&
: Cardoeo s.n., sin. loc.
Distribution Africa : South Africa, S. TomB, Canary Is.
Bryopsidaceae
Bryopsis Lamour. 1809
Bryopsis pennata var. secunda (Harp.) Coll. & Herv., Proc. Am. A d . Arts Sci., 53:
62 (1917);as B . plumosa in Hariot (1908: 162) and Henriques (1917 : 165). Bryopsie demu
Pilger, Bot. Jb., 5 7 , l (1920).
Marine algae of 850 T m t and Principe
111
Lami (unpub.) renamed Ribeiro’s plant B . rumulosa Mont., but the present writer is
unable to agree with this. It seems more likely that it is a small plant of B. pennata var.
secunda.
The plant is dark green and about 3 cm high. The main axes are 300 to 350 p wide,
many of them arcuate in form, naked below and the pinnules thus apparently rather
closely set towards their tips. However, the two rows of scars running up each main axis
converge, and the pinnules themselves are set in two very closely approaching rows, or
even lie over together to one side. Occasionally three scars are found a t one level on a main
axis. The pinnules are (47)-60-( 125) p wide and all of approximately the same length until
the tip of the frond is reached.
B. pennata has not been reported in the Gulf to the west and south of Nigeria, though
the B. balbisianu var. distich listed in Henriques (1885: 132) for Angola should be
re-examined.
An isotype of B. dema Pilger from Annobon, found in Hb. Bargesen, has proved to be
a small, perhaps younger, form of B. pennata var. secunda, with which it agrees particularly
in the arrangement of the pinnules.
A specimen from Ghana, Lawson A 1009 in Hb. Fox Nielsen, also belongs to the latter
species. There are thus new varietal records for Annobon, S. Tom6 and Ghana.
Exsiccata s. TOA
: Ribeiro 19, Ropa Praia da Nazareth, 1894.
Distribution Africa : ?Angola, Annobon, S. Tom6, Cameroun, Nigeria, Ghana, Sierra
Leone, SBnBgal, Maroc. Pantropical.
Caulerpaceae
Caulerpa Lamour. 1809
Caulkrpa cupressoides (Vahl)C.Ag. emend. W. v.Bosse, Monogr Caulerpes. : 323 (1898) ;
Henriques (1917: 165) ; as Chauvinia cupressoides in Henriques (1886: 218).
The species is, as a whole, pantropical and known in several varieties and forms from
many places, for example, from the warmer Atlantic coast of the Americas. It is recorded
from Cameroun (Engler, 1895: 24) but no specimen has been traced to substantiate this
record.
There is also a record for Annobon of ‘Caulerpa cf. lessonii Bory’ in Pilger (1920: 2))
where Weber v. Bosse’s somewhat inconclusive remarks are also quoted. C . lessonii is
usually considered an Indian and Pacific Ocean species but the distinction between
certain forms of this species and of C. cupressoides is certainly difficult to make.
Exsiccuta s. TOM&: Quintas s.n., Equador, Canal das Roles, Alga do fundo (basalto),
1885.
Distribution Africa : Portuguese East Africa, S. Tom6, ?Cameroun. Pantropical.
Caulerpa mexicana Sonder ex Kutz. in Papenfuss, S. Afr. J . Bot., 22 (2) :65 (1956); as
C. taxifolia var. crmsifolia in Henriques (1885: 133).
Newton’s plant from Principe has never been found, but if correctly determined would
now be known as C . mexiwna. Askenasy’s record (Askenasy, 1896: 157) is probably
based on this specimen, or on Henriques record of it (Henriques, 1885: 133), though he
actually refers to C. crassifolia var. mexicana, another synonym.
The Franklin specimen found by M. Denizot in Paris is included because the locality
is given as ‘ Insel St. Thoma’, but no corroboration for Franklin’s visit to S. Tom6 has
been found. Carpine has however dredged C. mexicuna in 11 m. of water (Carpine, 1959:
77).
Both varieties of C . mexicuna appear to be recognized on the Atlantic coast of Africa.
112
MARGARET
STEENTOIW,
F.L.S.
Exsiccuta s. TOMI%:Franklin s.n.*sin. loc. 16 Mars 1904 (PC).
Distribution Africa : Portuguese East Africa, South Africa, S. TomB, Ghana, SBnBgal,
Cab0 Verde Is., Canary Is. Probably in most warmer waters.
Caulerpa racemosa var. occidentalis (J.Ag.) B~rgesen,
K. danske Vidensk.Selek. Skr.,
7 (4): 5379 (1907); as Chauvinia clavifera in Henriques (1886: 218) ;as Caulerpa racernosa
var. clavifera in Hariot (1908: 162); as C. racemsa in Henriques (1917: 165).
Lami (unpub.) revised Quintas’s Ilhdu das Rolas plant and the Equador plant is very
similar.
C. racemosa is a pantropical species and in the western Atlantic. Taylor (1960: 52)
recognizes eight varieties and three forms. Of these var. occitlentulis seems to be limited
to the Atlantic. A Jadin specimen from Mauritius (in Hb. Bsrgesen) can only be included
with considerable doubt.
Var. occidentalis is easily confused with var. clavifera on the one side and with C. peltata
on the other. Welwitach 215 from Cab0 Verde Is., published in Barton (1897 : 370) as
C. clavifera Ag., is more likely composed of young and growing plants of var. midentalia,
since many branch tips are relatively longer than in var. clavifera, and therefore are
probably unswollen and immature.
Dangeard (1952, Fig. 3) shows, at least in part, var. occidentalis, from SBnBgal, and
Lawson (1954:64 and 1957 : 172) for Sierra Leone, reports plants ‘nearest to var. clavifera’.
Exsiccata s. TO&: Quintas 15, I l h h das Rolas, 1885. Quintas s.n., Equador, Canal das
Rolas (alga do fundo calc8reo-c8r verde) 1885.
Distribution Africa: var. occidentalis S. TomB, Ghana, Cab0 Verde Is., SQndgal.Warm
Atlantic coasts. Var. clavifera Portuguese East Africa, South Africa, Annobon, ?Sierra
Leone, SBnBgal. Atlantic and Indian Oceans.
Caulerpa scalpelliformis (R. Br.) C. Ag., Spec. a&., 1: 437 (1823); Barton (1897: 370);
Exell (1944: 386); Henriques (1917: 165); as var. intermedia in Hariot (1908: 162); as
C. denticulatu in Henriques (1886: 218; 1917: 165).
Moller collected numerous plants of both var. intermedia and var. denticulata. The
duplicates in Hb. Agardh have rather better developed stolons than the Coimbra plants,
and one of them is number ‘ 18’, whereas in both Coimbra and Paris specimens, no. 18 is
C. taxifolia.
Exell’s material also consists of both varieties and grew attached in the littoral zone.
Thorold’s plants are of var. intermedia only, and were washed up. The most recent record
is that of Carpine (1959: 78), who collected the species once in 5 m of water, and again in
5 to 7 m of water off Praia Lagarto.
The main area occupied by C. scdpelliformis is in the western Pacific, and Indian
Ocean and the Red Sea. It is also known with certainty from South Africa, Angola and
SBo Tom& Engler’s record (Engler 1895: 24) for Senegambia is somewhat doubtful,
though a specimen labelled Senegambia exists in S.
There are specimens from Brazil, Mexico and the West Indies in HBG, and two from
Brazil (one leg. Salzman)in Hb. Agardh, but neither Taylor (1960) nor Bsrgesen 1913)
refers to these. Thus the furthest certain extension of C. scalpelliformis into the Atlantic is Siio Tom&
Exsiccata s. TOMI%:Exell 470, Praia Pequena. Near the town [Siio Tom6 City] 3 0 . i .1932
(BM).Fox Nielsen 228, sin. loc., August 1956 (duplicate in BM). Moller 11,
Bahia de Ana Chaves et Praia Lagarto, 9/85 (in Hb. Agardh no. 16432, LD).
Moller 18, Bahia de Ana Chaves, 5/85 (in Hb. Agardh no. 16430, LD). Moller
19, sin. loc., 1886. MoZZer 30, Bafa de Ana Chaves, 1885 [2 sheets]. Welwitsck
Marine algae of S& Tom6 and Principe
113
262, ‘Ad litt. insulae S. Thorn6 (sub aequatore) frequentissime ejecta unacum
Berkeleyana e t Codio tomentoso. Decbr. 1860.’ [‘sub aequatore ’ probably
means ‘almost on the equator’; there is no evidence to show that Welwitsch
went to the Ilh6u das Rolas, where he could have been, at least in a sense,
‘under’ or ‘below’ the equator. Berkeleya and Codium have not been found.]
(BM).
Distribution Africa: Portuguese East Africa, Angola, S. Tom& Pacific and Indian
Oceans, Red Sea.
Caulerpa sertularioides (Gmel.) Howe, Bull. Torrey bot. Club, 32: 576 (1905); as C.
plumaris in Henriques (1886 : 218, 1917 : 165) ; Hariot (1908 : 162).
Lami (unpub.) has examined this material and revised the name. The species appears
to be a sublittoral but shallow water one here, as in other parts of the tropics.
Specimens from Angola and Senegambia exist in S and in Hb. Binder (HBG), while
De Toni (1889: 453) refers to a specimen from ‘Guinea’. This could mean Jardin’s Bissagos
plant (Jardin, 1850-51 : 10) in Hb. Lenormand.
Exsiccata s. TO^: Quintas 24, Ilh6u das Rolas, 1885. Quintas s.n., Equador, Canal das
Rolas (alga de 5 p6s de fundo-basalto) 1885. Ribeiro s.n., Roga Praia da
Nazareth, 1894.
Distribution Africa : South Africa (Barton), Angola, S. Tom6, Cameroun, Ghana,
Guinke, Cab0 Verde Is., Senegambia. Pantropical.
Caulerpa taxifolia (Vahl) C. Ag., Spec. alg.: 435 (1823); Henriques (1886: 218; 1917:
165); Hariot (1908: 162).
Newton’s plant from Principe, listed as C . taxifolia var. crassifolia in Henriques (1885 :
33) has not been traced, but presumably belongs rather in C . mexicum.
Carpine (1959: 77) has recently confirmed the occurrence of the species on S. Tom&,
which has hitherto rested on Moller 18 only. S. Tom6 is the only locality known for the
species on the Atlantic coast of Africa, since the other two records (Trochain, 1940: 108;
Jardin, 1850-51 : 10) are for plants washed up.
Ezsiccuta s. TO^: Moller 18, sin. loc., 1885 (2 sheets COI+duplicate PC).
Distribution Africa : S. Tom6, ?S6n6gal. Probably pantropical.
Codiaceae
Codium Stackhouse 1797
Codium tomentosum Stackh. in Welwitsch ms.
No specimen has been traced. The specimen was said to have been cast up with Caulerpa
scalpelliformis q.v. but it is very unlikely that it is Codium tomentosum.
Boodleaceae
Struvea Sonder 1845
Struvea anastomosans (Harv.) Piccone, Croc. Cors. Madera e Canur.: 20 (1884); as
S . delicatula in Hariot (1908 : 162) and Henriques (1917 : 165). Struvea multipartita Pilger,
Bot. Jb., 57: 2 (1920).
Thorold’s and Ribeiro’s plants both belong t o var. anastornosam, which has some
irregularly branched nets and shows some attachment of the end cells. The species is
MAEWARETSTEENTOFT,
F.L.S.
114
recorded for Guinde, Ghana and western Cameroun without mention of variety, though
by implication the greater part of such material may belong to var. anastornosans.
On the southern Cameroun coast Pilger (1911: 296) reports var. caracasana Grun.,
distinguished by its regularly bipinnate nets and the absence of attachments between
the end cells of the branches. Pilger states that not all the nets showed these features.
According to Egerod (1952:361) each net, in the course of its development, goes through
a Atage when it is bipinnate and lacks terminal attachments of the branches. Since the
plants Pilger examined (Lederman 262) were collected in the rainy season when the
maximal development of Struvea takes place in the minimal conditions of exposure, it is
possible that a young population of var. anastornosans was developing, and the existence
of var. curacasana in the Gulf of Guinea is thus somewhat doubtful.
In Table 6 S. anastornosans and S . multipartita are compared. The plant examined of
the latter species (Mildbrued 6659 in Hb. Bsrgesen) was collected in an intertidal pool (on
Annobon), a type of habitat which, as Sourie (1954: 273) has shown, is a very special one
on a tropical shore. Pilger separated the two species on the grounds of the small nets and
irregular branching of the latter, but it seems more likely that it is only a growth form of
S. anaatmnosans and is therefore reduced t o the synonymy of S. anastornosans.
Table 6. Comparison between Struvea anastornosans and S. multipartita
S . anaatomoeana
Western Atlantic
(Bargesen, 1913: 54,
and Hb. Bergeaen)
(Taylor, 1960 : 122)
Cameroun
(Pilger, 1912: 296)
Szlo Tom6
(Ribeiro no. 1(?)
Fox Nielsen no. 248)
S. multipclrtita
Annobon
(Mildbrmdno. 6659)
Exsiccata
Height of plant
(mm)
Nets
length x width
(mm)
Main axes
length (mm) x width ( p )
20 to 50
(up to) 10 x 5
(up to) 40 x 200 to 900
often branched
26
1 5 x 15
10 x 750
usually branched
15 to 30
5X3to5
10to25x750
unbranched
25
3x3c.
c. 22 x 500
irregularly branched in
several degrees
(proliferated?)
9. TOM$: Fox Nieben 248, sin. loc., August 1956. Ribeiro [l?],Ropa Praia da
Nazareth, 1894.
Distribution Africa : Portuguese East Africa, Annobon, S. Tom6, Cameroun, Ghana,
Guinde, SBnBgal, Cabo Verde Is., ?Mauritanie. Apparently pantropical between about
30"N and 30"s wherever there is a warm current, though not known from the South
American coast south of Tobago, where the warm Brazilian current is present.
Siphonocladaceae
Ernodesmis Bmgesen 1912
Ernodesmis verticillata (Kiitz.) Bsrgesen, Bot. Tidsskr., 32 : 259 (1912).
The Principe plant is rather small, only 2 cm high, but the S. Tome one is 3 cm high and
well branohed. These are new records for both islands.
Ezsiccata s. TOP& : Cardoso sm., sin. loc.
PRfNomE: Newton cia., Praia de Sundy, 1887.
Marine algae of
Tom6 and Principe
115
Distribution Africa: ?St. Helena, S. Tomb, Prhcipe, Ghana, Cab0 Verde Is., ?S6nbgal,
Canary Is. Atlantic and Indian Oceans.
Siphonocladus Schmitz 1878
Siphonocladus tropicus (Crouan) J. Ag., Alg. syst.: 5, 105 (1887); Bmgesen, Dansk
Bot. Ark., 1 (4): 61 (1913).
The largest plant is 4 cm high and sterile, the other two are smaller but more branched
and show zoospore production.
Exsiccata 9. TO~W?:Cardoso s.n., sin. loc.
Distribution Africa: S. Tomb, Canary Is., Maroc. Also Portuguese East Africa and
Mauritius. Pantropical.
Phaeophyceae
Ectocarpaceae
Bachelotia (Bornet) Kuckuck ex Hamel 1939
Bachelotia antillarum (Griin.) Gerloff, Nov. Hedwigia, 1 (1): 38 (1959) ; as B. fulvescem
in Fox (1957: 621).
The filaments are 28 to 36 p wide and there are numerous unilocular sporangia from
32 to 40 p wide by 20 to 28 p high. The filaments are thus somewhat wider, and of more
regular width than those found a t Lagos (Fox, 1957: 621) where the filament width was
from 16 to 36 p.
These plants are intermediate in dimensions between B . fulvescens as reported in the
western Atlantic (Taylor, 1960: 198, filament width 30 to 47 p, unilocular sporangia
57 to 65 p wide by 26 to 31 p high) and the former Pylaiella antillarum (Griin.) De Toni
(1805: 535) which Gerloff (1959: 38) has transferred to Bachelotia, with B. fulvescena
(Schousb. ex Bornet) Kuckuck ex Hamel as a synonym of it. P . antillarum is reported with
filament widths of 11 to 25 p, with unilocular sporangia 18 to 15 p wide.
Although Lindauer & Chapman (1961 : 142) prefer to call their New Zealand material
B . fulvescens, on grounds of its greater dimensions, the Gulf of Guinea plants clearly
support Gerloff's view.
Exsiccata s. TOM&: Fox Nielsen 230 and 231, sin. loc., August 1956 (with Murray&
pericludos etc.).
Distribution Africa : S . Tomb; see also FOX(1957 : 621). Probably widespread in all but
the coldest waters.
Giffordia (Butt.)Hamel 1939
Giffordia mitchellae (Ham.) Hamel, Phueophyckes de France, 29 : (1931-39).
Fertile, with meiosporangia. This is a new record for S. Tom6 and the species is now
known from many parts of the Guinea coast.
Exsiccuta s. TOM&: Fox NieLen 234, sin. loc., August 1956.
Distribution Africa: S. Tom6; see also FOX(1957: 622). Apparently pantropical.
Sphacelariaceae
Sphacelaria Lyngbye 1819
Sphacelaria sp.
Sterile fragments of this species have been found, particularly with Chaetomorpb
crmsa. This is a new record for the genus from S. Tomb, though previous records for the
8
116
MARGARET
STEENTOFT,
F.L.S.
Gulf of Guinea are to be found in Dickinson (1953 : 42) for Ghana, Lawson (1955 :86) for
Cameroun, and Pilger (1920 : 4) for Annobon.
Cladostephaceae
CladostephusC. Ag. 1817
Cladostephus verticillatus (Lightf.) C. Ag., Symp. [email protected] c u d . , xvi (1817).
In Sampaio (19623:24) this speciesisgiven as the substrate of Hydrocoleumlyngbyaceum,
as in Kiitzing (1843 : 196) Kiitzing actually refers to C. spongwsus, which is now included
in C. verticillatus by Taylor (1937: 135).
C. verticillatus is recorded only as far south as the Canaries, on the Atlantic coast of
Africa.
Cutleriaceae
Cutleria Greville 1830
Cutleria multifida (Smith) Grev., Alg. Br.: 60 (1830); Henriques (1917: 165); as
Aglaozonia reptans in Henriques ( 1886 : 219).
This determination must be regarded as uncertain since Quintas’s plant consists of a
small black crust on rock, and is not in a state permitting examination. Aglaozonia sp. is
reported by Carpine (1959: 87) from Principe, collected in a depth of between 3 and 12 m
but the species is otherwise confined to Maroc on the Atlantic coast of Africa.
Exsiccata s . TOM&: Quintas 35, Ilh6u das Rolas, 1885 (Hb. Agardh no. 47945, LD.).
Dictyotaceae
Dictyota Lamour. 1809
Dictyota barteyresiana Lamour., J. Bot., Paris, 2: 43 (1809); Rodrigues (1960: 585
and fig. 1 (a)); Hariot (1908 : 162);Henriques (1917 : 166) p.p. ; as D. ciliata in Henriques
(1886 : 219) p.p.
Quintas’s plant, referred to in Henriques (1886: 219), has been renamed by Hariot,
and Rodrigues (1960 : 585) has made new determinations for a Cardoso and a Newton
specimen.
A thallus tip from Newton’splant is shown by Rodrigues (1960: 586, fig. l(a))a t what is
said to be twice natural size, whereas c. x 11 would be a more correct figure. The same
illustration shows ‘sori of tetrasporangia’ in rows across the thallus, but in fact this
effect is produced by the coagulation of the oontents of the medullary cells, and these dark
brown granular masses happen to lie more or less in rows across the thallus, showing
through the superficial layer of cells. They give the thallus a darker colour than seems
usual in D. barteyresiam and although Taylor (1928: 117 and P1. 16, fig. 11) shows a
similar form (as pointed out by Rodrigues) more material is desirable.
A Ribeiro specimen found by M. Denizot probably alao belongs in this species, and
Carpine (1959: 80) reports D. barteyresiam from three places on S. TomB, two sublittoral
stations and one littoral.
Three forms are included here in D. barteyruiana, and further collections are obviously
necessary.
Exsiccuta s. TOM&: Cardoso sm., sin. loc. Fox Nieben 237, sin. loc. August 1956.
Quintas 10, Ilh6u das Rolas,1885 (duplicate PC). Ribeiro s.u., Ropa Praia
da Nazareth, 1894 (PC).
PB~NNOIPE
: Newton rm.,Bahia de Sto Antonio, Julho de 1887.
Marine algae of S& Tom6 and Principe
117
Distribution Africa: Annobon, S. Tom6, Principe, Cameroun, Ghana, Cab0 Verde Is.
Probably pantropical.
Dictyota cervicornis f. pseudobarteyresiana Taylor, Mar. a&. Florida : 118 andpl. 16,
fig. 15 (1928); Mar. alg. . Americas: 222 and pl. 31, fig. 2 (1960); Rodrigues (1960:
585 and P1. 1); as D . Barteyresiana in Hariot (1908: 162) and Henriques (1917: 166)
both p.p.
..
Taylor (1960: 222) now doubts the value of the three forms of D. cervicornis established
by him. There is a single S. Tom6 plant, 20 cm high, which Rodrigues has identified as f.
pseudobarteyreaiana.
Exsiccuta s. TO& : Ribeiro 5, Ropa Praia da Nazareth, 1894.
Distribution Africa : S. Tom& Otherwise on the tropical and subtropical east coast of
the Americas.
Dictyota ciliata J. Ag., Linnea, 15: 5 (1841); Henriques (1886: 219; 1917: 166); Hariot
(1908: 162); Rodrigues (1960: 587 and pl. 2 and fig. l(b)).
Three forms seem to be present. Moller’s and Quintas’s plants look like the usually
accepted forms of the West Indies, as represented in Hb. Bsrgesen and illustrated in
Taylor (1960 : 223). Ribeiro 6 is a large luxuriously branched female plant which corresponds with the description of some Ghana forms in Dickinson (1953 : 41). Thorold’s
plants on the other hand are small, about 6 cm high and lack the marginal ciliations
supposed to be characteristic of the species. Similar forms have been collected in Ghana
by the present writer, both from pools and from rocks covered by sand.
Emiccata s. TO&: Fox Nieken 236, sin. loc., August 1956 (duplicate BM). Moller s.n.,
sin. loc., 1885. Quintas 25, Equador, Canal das Rolas. Alga do fundo, 10 p6s.
1885 (duplicate PC). Ribeiro 6, Ropa Praia da Nazareth, 1894.
Di8tribution Africa : S . Tom6, Nigeria, Ghana, SBn6gal. Probably pantropical.
Dictyota crenulata J. Ag., Ofwera. Vetensk.Akad.Forh. Stockh.: 1847,7 (1848); Taylor,
Pacific mar. alg. . . . Galapagos, Alhn Hancock Paci$c Expeditim, 12: 90 and pl. 10,
fig, 1 (1945); as D. ciliata p.p. in Rodrigues (1960: 587), Hariot (1908: 162) and Henriques
(1917: 166); as D. dichotoma in Barton (1897: 371).
Ribeiro 3, included by Rodrigues in D . ciliata, consists of at least three plants, one
tetrasporangial, one female and the third probably male, though in a bad state of preservation. While the two sexual plants seem to agree quite well with an isotype of D . crenuhta
in Hb. Bsrgesen, the tetrasporangial plant causes some doubt, and Rodrigues was
probably correct in placing it in D . ciliata.
Taylor (1960: 223) doubts the occurrence of D . crenulata in the Atlantic, and leaves it
among the ‘uncertain records’ for the genus. Instead D . jamaicensis Taylor is described
to receive plants of this form, largely it seems on the grounds that the Atlantic plants are
larger (and more robust?) than Liebmann’s plant. Having seen very little material of
D. crenulata (what appears to be an isotype is present in Hb. Bmgesen) and none of D.
jamaicensis the present writer prefers Agardh’s name for the present.
Welwitsoh’s plant is very young and sterile, but on grounds of habit, branching and
thallus tip form, is thought to belong here. But typically developed marginal teeth are
absent.
Exsiccata s. TOM^ : Ribeiro 3 p.p., Ropa Praia da Nazareth, 1894.
P R f N c n w : Welwitsch 258, Bahia de s. Antbnio, 20 Sept. 1863 [between midand low-tide levels, ‘not infrequent’] (BM).
118
WGARET
STEENTOFT,
F.L.S.
Distribution Africa: Annobon, 8. Tom6, Principe, SBndgal, Cab0 Verde Is. Warm
Atlantic and Pacific Oceans.
Dictyota dentata Lamour., J. Bot., Paris,2 : 42(1809);Henriques (1886: 219; 1917: 166
Hariot (1908: 162); Rodrigues (1960: 588, fig. l(c) and pl. 3); as D. Mertensii (Martensii)
in Hariot (1908: 162) and Henriques (1917 : 166).
D . mertensii was the name originally given to alternately branched forms of Dictyota
in which the tips of the thallus lobes were rounded, not toothed as in D. dentata. Hauck
(1888 : 466) placed D. mertensii among the synonyms of D. dentuta, presumably because
he could find no sharp and constant differences between the two among the material from
Puerto Rico which he examined.
Taylor (1960 : 224 and P1.30, fig. 5 ) remarks, of D . dentata, ‘branch tips broadly rounded
on young shoots’ and this could well apply to Ribeiro’s plant. There is however the
possibility that it is a Dilophw species, since there are two layers of medullary cells in
one margin of the thallus in the basal part of the plant.
The remark of Murray (1888: 36) that the type of Dilophus guineensis (Kiitz.) J. Ag.
originates in S l o Tom6 and not in St Thomas, West Indies is unfounded if, as Kutzing
(1843: 339) says, it was collected by Ehrenberg (who collected in the West Indies in
1828-9).
Dickie’s record for Cab0 Verde Is. remains the only one.
Exsiccata s. TO&: Cardoso s.n., sin. loc. Quintas [6?], Ilh6u das Rolas, 1885. Ribeiro
2, RoOa Praia da Nazareth, 1894.
Distribution Africa : S . Tom6, ?Cab0Verde Is. Warmer Atlantic coasts of the Americas.
Dictyota dichotoma (Huds.) Lamour., J. Bot., Paris, 2: 42 (1809); ?Henriques (1885:
133; 1917: 166).
The current record depends on Carpine (1959: 77, 88) and on a plant collected by
Gravier, and not mentioned in Hariot (1908).
Exsiccata s. TO&: Gravier s.n., Praia da Bella Vista, a mar6e basse sur la basalte.
29 Avrill906 (sub.m.D . barteyreaiana in PC).
Distribution Africa: South Africa, S. Tom6, Principe, Canary Is., Maroc, Tanger.
Widely spread in all but the coldest waters.
Dictyota sp.
Reported in Carpine (1959 : 83) from S. Tom&
Padina Adanson 1763
Padina commersonii Bory, Voy. Coquille, Bot.: 144 (1828); Weber v. Bosse, Liste . . .
Siboga, 1: 178 (1913).
Thorold‘s plants are small but nevertheless have the distinctive characteristics of a
thallus only three cells thick, with sori of oogonia standing immediately apical to rows of
paraphyses. The sori lack indusia. This is a new record for S l o Tom6 and represents a
considerable extension into the Atlantic Ocean of a species hitherto known from the
Pacific and Indian Oceans.
Exsiccata s. TOM& :Fox Nielsen 235, sin. loc., August 1956.
Distribution Africa: Portuguese East Africa, South Africa, S. Tom&Pacific and Indian
Oceans.
Marine akae of 8& Tome‘and Principe
119
Padina vickersiae Hoyt ex Howe, in Britton and Millspaugh, Bahama Flora: 595
(1920);Hoyt, Bull. Bur. Fish., Wash. (1920):36 0921); Rodrigues (1960: 558 and fig. l ( d )
and pl. 4) ; as Zonaria pavonia in Henriques (1886: 219) ; as Padina variegata in Hariot
(1908: 162); as P d i n a pavonia and Zonuria variegata in Henriques (1917: 166); as
Pocockiella variegata in Rodrigues (1960 : 588 and pl. 5 ) .
The plant referred to Zonaria pavonia Ag. in Henriques (1886: 219) was collected by
Moller, part being retained in Coimbra and part being sent to Lund. These plants are quite
different from P . pavonicu (L.) Thivy (Taylor, 1960: 234). The plant in Hb. Agardh
matches Moller 7 among the Coimbra material determined by Hariot as P . variegata
(Kiitz.) Vickers (‘leg. Moller, 1885, SLo TomB’).
Ribeiro’s plant was also determined by Hariot as P . variegata (Kiitz.) Vickers and
with these identifications the present writer agrees. This combination is now a synonym
of P . vickersiae Hoyt ex Howe, and perhaps because of the maze of synonymy woven
around the epithet variegata as applied to Padina, Rodrigues (1960: 589) places both
Ribeiro’s and Moller’s plants in Powckiella variegata (Lamour.) Papenf. But the plants
differ from Pocockiella in having a rolled thallus apex and in lacking a distinct medulla.
Rodrigues (1960 : 589) also places a Newton specimen in P . vickersiae, but i t is somewhat
different from the rest of the material in having a three-layered thallus 80 to 105 p thick
for some distance behind the apex. All the cells are about a third larger in both length
and height than in the other plants, but the proportions and arrangement of the cells is
similar.
All plants, including Thorold’s, had one or two broken bands of sori, with indusia, per
fertile zone. There were no sterile zones.
A curious feature was the absence of paraphyses. Instead of a band of epidermal hairs
forming a piliferous zone, a band of epidermal cells all showed subdivision into small
cells, square in surface view. Their smaller size and dense pigmentation gave the effect
of a piliferous zones without any free-standing hairs being present. The same feature is
commented upon by Dangeard (1952: 249) in plants from Cap Vert. Dangeard favours
P . durvillei or P . tetrastromaticu as possible names for his material, but the latter is certainly impossible for the S. Tom6 plants.
Ifthe absence of hairs isnot a character sufficient to demarcate a new genus, as suggested
by Feldmann (in Dangeard 1952 : 249), then the S. Tom6 and Principe plants may possibly
belong in P . durvillei or P . mexicunu.
P . vickersiae is reported (in litt.) by Lawson for Cameroun.
Exsiccata s. TOME: Fox Nielsen 244, sin. loc., August 1956; Moller 7, sin. loc., 1885
[+duplicate? s.n.1; Moller 7 , Bahia de Ana Chaves et Praia Lagarto, 9/85
(in Herb. Agardh no. 48348, LD); Newton s.n., I l h h das Cabras, 6-1889
[1887]; Ribeiro s.n., Ropa Praia da Nazareth, 1894.
PR~NCIPE:Newton s.n., Bahia de S. Antbnio, 7-1887.
Distribution Africa : S. TomB, Principe, Cameronn, ?SBnBgal, Canary Is. Warmer
Atlantic coasts of America.
Punctariaceae
Chnoospora J. dg. 1847
Chnoospora minima Hering in Papenfuss, S. Afr. J . Bot., 22: 69 (1956).
The single specimen available is smaller, more delicate and narrower in the thallus than
is usual among the Lagos material with which the present writer is familiar (Fox, 1957 :
623) and no tufts of hairs could be found. It was most reminiscent of plants growing high
in the surf zones a t Lagos. The present writer has also collected the species in Ghana.
MARGARETSTEENTOFT,
F.L.S.
120
The record for the Cab0 Verde Is. appears to rest only on Bolle’s specimens mentioned
in hkenasy (1896: 160) as ‘Ch. fmtigiata J. Ag.?’
Exsiccata
8 . TO&
: Cardoso s.n., sin. loc.
Distribution Africa : South Africa, Annobon, S. Tom6, Nigeria, Ghana, Cab0 Verde Is.
Widespread in warmer waters.
Colpomenia Derb. & Sol. 1866
Colpomenia sinuosa (Roth) Derb. et Sol., Hem.
. . .physiol. Algues: 11 (1866).
This is one of Carpine’s new records for S. Tom6 (Carpine, 1959 : 77). It appears to be a
widespread species on the Atlantic coast of Africa. Distribution South Africa, Angola,
Ghana, SBnBgal, Cab0 Verde Is., Mauritanie, Canary Is., Maroc, Tanger, Madeira and
Agores, also Portuguese East Africa.
Sargassaceae
Sargassum C. Ag. 1821
Sargassum vulgare C. Ag., Sp. alg., 1: 3 (1821); Hariot (1908: 162); Henriques (1917:
165); ?Exell (1944: 386) ; Rodrigues (1960: 590 and pl. 6 and fig. 2(a));as S. boryanum
in Henriques (1886: 219) p.p.
Var. vulgare
MoZZer 9a appears to be well developed in the sense of Taylor (1960 : 268) and resembles
his description and figures (Taylor 1960: 272 and pl. 38, fig. 1; pl. 40, fig. 6).
Moller’s plants also closely resemble 8.CheiriifoZium Kutz., based on Lewmand 214
from the mouth of the Gabon River, and this name should probably be added to the
synonymy of S. vulgccre, rather than to that of 5.cyntosuln Ag. as was done by De Toni
(1895: 87). However there seems to be a certain amount of confusion about the type
specimen of S. cheirifolium for Kiitzing (1861 : 7 and pl. 21.21, figs a to d) gives the type
locality as Senegambia.
ExsicccctcG
TO^: ?Exell 465-8, Praia Pequena, on lava rocks, near low-tide level
(?in BM, non. vU.).Holler 9 sin. lac., 1885 [2 sheets]. MoZZer 9a, Bahia de
Anna Chaves, 9-1885. Holler s.n., sin. loc. 1885.
8.
Distribution Africa: South Africa (Barton), S. Tomb, lcameroun, Ghana, ?Sierra
Leone, S6n6ga1, Mauritanie, Cabo Verde Is., Maroc, Aqores. Atlantic coasts of Portugal,
tropical and subtropical Americas, Mediterranean.
var. foliosissimum (Lamour.) J. Ag., Spec. Sargass. Austral. : 108 (1889); Hariot
(1908 : 162) ; Rodrigues (1960 : 592 and fig. 2(b)); as S. boyanum in Henriques (1886 :
133; 1886: 219) p.p.; as Marginaria boryana in Henriques (1917: 165); as Sargassum sp.
in Barton (1897 : 374).
In view of the variability of Sargmsum spp., and the limited amount of material
available in the present case, it seems better not to distinguish forms, though Rodrigues
(1960: 692) considers that Quintas’s plants belong in fa. ‘pteropus (Kutz.) Grunow’.
Similar plants have been collected by the present writer in drift at Cotonou (Dahomey)
and Lagos (Nigeria).
The record for Cab0 Verde Is. appears to rest on Newton’s plant from 1881 (Henriques,
1885: 133).
Marine algae of SaO TmC and Principe
121
Exsiccata s. TOME: Fox Niehen 238, sin. loc., August 1956 (duplicate BM). Quintus
23, Ilh6u das Rolas, 1885. Quintas31, Angolares, 1885. Quintas s.n.,Equador,
Canal das Rolas. Alga a 10 p6s de fundo. C8r verde. 1885.
PR~NCIPE: Welwitsch 223, Sept. 1853 (BM).
Distribution Africa : S. Tom6, Prhcipe, Cameroun, ?Nigeria, ?Dahomey, SBn6ga1,
?Cab0 Verde Is., Tropical and subtropical Atlantic coasts of the Americas, Australia.
Rhodophyceae
Helminthocladaceae
Liagora Lamour. 1812
Liagora megagyna Brargesen, Dansk Bot. Ark., 3: 77 and figs 82-84 (1915-20).
The single plant available is in poor condition, but most resembles L. megagyna, known
only from the female plant and from the type locality, St Croix. The present determination
must be regarded as uncertain.
Exsiccata
PRiNcrPE : Newton
s.n., Praia de Sundy, Agosto 1887.
Distribution Africa: S. Tom& Otherwise only from St Croix.
Chaetangiaceae
Galaxaura Lamour. 1812
The complexities of this genus are well known. It was monographed by Kjellman (1900)
and this was presumably available to Hariot in 1908. Kjellman must have known of the
existence of specimens of marine algae from S. Tom6 (see p. 128) but the only West
African Q a h a u r a mentioned is Q. ventricosa Kjellm. described from what is stated to be
one of Jardin’s Gabon specimens.
Hariot lists three species of Gakaura, but one of them, G. rugosa, is not fully dealt
with by Kjellman, and another, a. murginuta, is very much narrowed, a profusion of new
species being created on rather trivial characters.
Since 1900 the works of Chou (1945,1947), Bmgesen (1915-20,1927, 1942 etc.), Howe
(1917, 1918), Svedelius (especially 1946, 1953) and Taylor (1928, 1960) have appeared.
Much new information is available and re-examination of the S. Tom6 and Principe
material shows that these islands probably have the richest Galaxaura flora in the Gulf of
Guinea.
The distribution of each species given below is regarded as putative, and the authority
for each record is quoted. It has not yet been possible to examine all the West African
material available, and it is hoped to make this the subject of a separate investigation.
Galaxaura cylindrica (Ell. et Sol.) Lamour., Expos. mbthod. : 2 (1821) ; Henriques
(1886: 218) and, p.p. (1917: 166); ?Hariot (1908: 162).
Both Quintas’s plants belong here, but the plant collected by Newton, and presumably
named by Hariot, is more doubtful. The small piece remaining intact is coarser and has
more divergent branching, and seems to belong in Q. oblongata. Moller’s material (Henriques, 1886: 218) has not been seen.
Q. cylindrica is perhaps the sexual phase corresponding to the asexual phase known as
Q. lupidescem. The latter is not knownfrom anywhere nearer to S. Tom6 than the Canary Is.
Exsiccata s. TOM&: Quintas 13, Inter ins. St. Thorn6 et Rolas (no. 32449 in Hb. Agardh,
LD). Quintas 20, Ilh6u das Rolas, 1885 [with immature conceptacles].
Quintas s.n., Equador, Canal das Rolas. Alga a 5 p6s de fundo calchreo.
1885.
122
MARGARET
STEINTOFT,
F.L.S.
Distribution Africa: S. Tom6, Cabo Verde Is. (Dickie), Canary Is. (Bmgesen). Pantropical.
Galaxaura Alamentosa Chou, Pap. Mich. Acad. Sci., 30 (1) : 39 (1944); Svedelius, Nova
Acta R. SOC.Scient. upsal., ser. 4, 15 (9):33 (1953); as Q. hpidescens in Henriques (1886:
218; 1917: 166).
This is a highly distinctive ‘species’possessing only long assimilators which lack both
swollen (tumid) basal cells and distinct supporting cells. There is hardly any difference
between medulla and cortex. The sexual generation corresponding to Q. filamentosa has
yet to be suggested.
The two S. Tom6 plants have been compared with an isotype of Q. filamentosa in Hb.
Bergesen, and agree well with it except that, in habit, the S. Tom6 plants are more
fastigiately branched and, in structure, the assimilators are somewhat shorter (about
1 mm on average).
A similar form from Ceylon is commented upon by Svedelius (1945 : 66) under the name
Q. tomentosa Kutz.
Exsiccata s . TOM&: Quintas 17, Inter ins. St. Thorn6 et Rolas (no. 32498 in Hb. Agardh,
LD, S u b m m . Q. h p i d e m m ) .Quintas s.n., Equador, Canal das Rolas. Alga
do fondo. C6r roxo terra. 1885.
Distribution Africa : S. Tom& Northern Pacific Ocean (Chou).
Galaxaura marginata (Ell. et Sol.) Lamour., Hist. Polyp.: 264 (1816); Kjellman, Om
Ploridd-Sligtet Galaxaura etc. : 77 (1900); Taylor, Mar. alg. . Americas: 343 and pl. 44,
fig. 2, pl. 45, figs 7 to 8 (1960; Henriques (1886: 218; 1917: 166); Exell (1944: 386); as
Brachytrichia [ =Brachycladia?] murginata in Hariot (1908 : 162).
. .
The above name is given with reservations pending enquiry into the Atlantic and
Indian Ocean species of the sections Arboreae Chou and Veprecuhe Kjellm.
Three forms are present among the S. Tom6 and Principe plants, one of them accompanied by a sexual specimen showing characteristics intermediate between a. ventricosa
Kjellm. (describedfrom Gabon)and Q. uepreculaKjellm. (describedfrom the Indian Ocean
and since reported from the Pacific by Chou (1945: 16).
The asexual forms show much in common with a. stupimula, a. frutescens and Q.
murginata. The two former species were split from the latter by Kjellman, and the whole
complex should be re-examined together.
Exsiccata s. TO&: Exell 471, Praia Pequena. Near the town [SBo Tom6 City]. On
lava rocks at low tide level. 30.xi.1932. (BM). Moller 20, Bahia de Anna
Chaves, 6/85 (no. 32516 in Hb. Agardh, LD). Moller 20, sin. loc., 1885 [asex.
and sex. plants] (duplicate PC). MoZZer 21, Bahia de Anna Chaves et Praia
Lagarto, 7/85 (no. 32514 in Hb. Agardh, LD). Quintas 3, Inter ins. St. Thomd
et Rolas (no. 32522 in Hb. Agardh, LD). Quintm 22, Ilh6u das Rolas, 1886.
(duplicate PC) Q u i W s.n., Equador, Canal das Rolas. Alga a, 5 p6s do
fundo calcireo. 1885. Quintm s. n., Equador, Canal das Rolas. Alga do fundo.
C6r roxo terra. 1885.
PafNcmc: Newton s.n., Praia de Sundy, Agosto de 1887.
Distribution Africa : S. Tom&,Principe, Ghana (Dickinson).Warm Atlantic?
Galaxaura oblongata (Ell. et Sol.) Lamour., Hist. Polyp.: 262 (1816): as 0.fragilis in
Barton (1897: 371) and (?) Askenasy (1896: 165); as a. cylindrim in Hariot (1908: 162)
and Henriques (1917 : 166) p.p.
The specimen is commented upon under Q. cylindrica above. The corresponding tetrasporic form is reputed to be Q. c o m n a , but this is not so far known in the eastern Atlantic.
Marine algae of Sizo Tom6 and Prtncipe
Exsiccata
PR~NCIPE: Newton
123
s.n., Praia [de] Sundy, Agosto de 1887.
Distribution Africa :Principe, Ghana (Dickinson),Cab0 Verde Is. (Bartonand Askenasy),
Canary Is. (Bclrgesen). Pantropical and subtropical, also Mediterranean.
Galaxaura rugosa (Ell. et Sol.) Lamour., Hist. Polyp.: 263 (1816); Svedelius, Nova
Acta R. SOC.
Scient. upsal., ser. 4, 15 (9): 18 (1953);p.p. in Henriques (1886: 218; 1917:
166))also in Hariot (1908: 162).
Three species, G. rugosa, G. squalida and G. subfruticulosa, have previously been confused among the s. Tom6 material. The thalli of clr. rugosa are quite glabrous, annulate
rugose and cylindrical (apart from the thallus tips which are not fully calcified and tend
to collapse). Thallus structure agrees with that of Chou’s section Squulidae, with three
layers of cortical cells forming a subparenchymatous cortex, the large lobed cells innermost. The ‘segments’ are up to 1 cm long and regularly 2 mm wide. The lower nodes are
distinctly constricted. There is no stipose region a t the base of the plant.
According to Bclrgesen (1927 : 70), the clr. rugosa reported for the Canary Is. in Piccone
(1884: 35) is G. squalida.
The tetrasporic generation corresponding to G . rugosa is said to be a. verticillata, which
is not known in the ea,stern Atlantic.
It is not known from the way the material is mounted which of the three species mentioned first above, grew together. Moller’s material (Henriques, 1886: 218) has not been
seen.
Exsiccata s. TO& : Quintas 4a, Inter ins. St. Thorn6 et Rolas (no. 32472 in Hb. Agardh,
LD). Quintas [21?], pep.,Ilh6u das Rolas, 1885.
Distribution Africa : Ascension (Askenasy),S. Tomk, Cameroun (Pilger), Cab0 Verde Is.
(Dickie and Askenasy). Pantropical.
Galaxaura squalida Kjellm., Om Floridk-Sliigtet Galaxaura etc. : 57 (1900) ; Taylor,
Mar. alq. . . . Americas: 339 and pl. 44, fig. 3 and pl. 45, fig. 9 (1960); as clr. rugosa p.p. in
Hariot (1908: 162) and Henriques (1917: 166).
The plants here described were found mixed with clr. rugosa and (3. subfruticulosa q.v.
They are densely branched and much proliferated with dense whorls of long assimilators
a t each node where proliferation has taken place, as well as groups and patches of long
assimilators on other part of the thalli where a general covering of rather shorter
assimilators also occur. The younger parts are rugose, the calcification setting in unevenly
as described by Svedelius (1945 : 4) and complete whorls of assimilators are also seen on
some of the younger branches. I n general appearance the plants are like the one illustrated
by Bclrgesen (1915-20: 102) but more divergently branched and very much more articulate
because of the proliferation.
The cell arrangement and measurements agree well with those given by Chou (1947 : 9).
Two forms of G . squalida are present. The plants Quintas collected sublittorially
together with some of those from Ilhku das Rolas bear many young proliferations. The
basal parts are quite without long assimilators, while the proliferations are glabrous and
only slightly calcified. These plants may have recently been in a reproductive phase, while
the remaining plants were in a pre-reproductive phase,
This is a new record for s. Tom&
Bsrgesen (1927 : 65) considers that G. $qelliformis is the asexual form corresponding
to 0. squalida, but it is not known from anywhere nearer to S. Tom6 than the Canary Is.
E&cmta s. TOMI? : Quintas 4, Inter ins. St. Thorn6 et Rolas (no. 32471 in Hb. Agardh,
LD, sub nom. G. rugosa). Quintas 16 and 21 [two unlabelled packets on same
sheet] p.p., Ilhku das Rolas, 1885. Quintas s.n., p.p., Equador, Canal das
Rolas. Alga do fundo (calchreo) c6r verde. 1885.
124
MARGARET
STEENTOFT,
F.L.S.
Distribution Africa : S . Tomk, Canary Is. (Bsrgesen). Pantropical.
Galaxaura subfruticulosa Chou, Pap. MicA. A d . Sci., 30 (1944) (1) : 41 and pl. 2,
fig. 6 and pl. 8, fig. 2 (1945); Svedelius, Nova Acta R. SOC.Scient. upsal., ser. 4, 15 (9):51
and figs 43-7 (1953); as Q. nqosa p.p. in Henriques (1886: 218; 1917: 166), and Hariot
(1908 : 62).
This is the third of the species found in Quintas 21 and was also found with G. sqwclida
in an unnumbered Quintas collection. The latter is labelled as growing sublittorally
(‘alga do fundo’) but presumably Quintas 21 was also found sublittorally.
C. subfruticulosa is very nearly related to Q. eubverticillata Kjellm. and hitherto only
the latter of the two species has been recognized in the Atlantic, the former species being
apparently confirmed to the Pacific.
Both Quintas’s plants are 5 to 5.5 cm high (the holdfasts are missing). Branching is
almost regularly dichotomous fastigiate. Most of the segments are about 1 cm long and
have continuous internodes. In the basal parts of the main branches the segments are very
short, from 1 to 4 mm long. This presumably where regeneration has taken place.
The branches are terete, slightly collapsed in the younger parts and mostly 1 mm wide,
though in the older parts the width is somewhat uneven, up t o 1.5 mm, and these segments
are rather heavily calcified.
Long assimilators occur all over the thallus, being especially prominent round the
bases of the proliferations. I n the dried specimen they appear as blackish spots (sometimes
arranged verticillately) because they adhere together in groups. Examination has shown
that clear bands of short assimilators alternating with bands of long assimilators are not
present, except in the very youngest parts and the resemblance to a. subverticillata is
quite superficial.
Another difference from a. subverticillata, is the development of secondary short
filaments from the basal parts of the plant. These are stiff and thick-walled (Svedelius,
1953, fig. 45(e) and account for the uneven width and furry appearance of the older parts
of the plant. Such filaments are absent in Q. subverticillata.
Ezsiccuta s. TO^: Quintas [21?] p.p., Ilhku das Rolas, 1885. Quintas s.n., Equador,
Canal das Rolas. Alga do fundo (calchreo) cBr verde. 1885.
Distribution Africa: S. Tom6. Pacific Ocean (Chou).
Gelidiaceae
Gelidiella FeMrn. & Hamel 1934
Gelidiella acerosa (Forssk.) Feldm. et Hamel, Revue gkn. Bot., 46: 528 (1934): as
Qeldidium claviferum fa. in Henriques (1886: 220; 1917: 166).
Qelidiella differs from Qelidiopsie in possessing a single apical cell to the thallus, but the
two genera resemble each other in lacking rhizines and in possessing tetrahedral
tetrasporangia.
The Sio Tom6 specimens have formed part of a turf, 2 to 3 cm high with intricately
woven creeping basal axes from which erect branches arise. The thalli are black and wiry
when dry, cartilaginous and light brown with markedly darker thallus tips when soaked
out. The branches are mainly cylindrical but the widest ones show slight flattening. Most
axes are 500 p to 800 p wide with 300 p and 1000 p as extreme values.
Branching is irregular, alternate as a rule, but one-sided, pinnate or even pseudodichotomous upon occasion. Regeneration is frequent and most of the shorter axes show
irregular rings a t intervals of 0.5 cm or so.
Fertile tips bearing tetrasporangia are slightly flattened. After the tetrasporangia
Marine algae of S& Tom6 and Principe
125
have been shed, the tip is for a time hollow, then regenerates from within, giving the
irregular rings mentioned above.
Anatomically the thallus possesses a medulla of relatively large elongate cells, 20 to 30
(40) p in transverse diameter with walls about 3 p thick. The cells become progressively
smaller through the cortex though the walls remain the same thickness. The superficial
layer consists of cells 4 p wide and 12 p long (the longer axis lying radially in the thallus),
arranged very regularly in a single layer.
More material of this species is desirable, since it has proved impossible t o demonstrate
satisfactorily the uniaxial nature of the thallus tip, and since i t differs somewhat in gross
morphology from the illustration of western Atlantic material in Taylor (1960, pl. 46,
fig. 5). However, there are specimens of similar habit in Hb. Bergesen.
Oelidium rigidum (Vahl) Grev. is usually a synonym of Gelidiella acerosa (Forssk.)
Feldm. et Hamel, but according to Papenfuss (1951 : 175) this is not so with regard to
South African material. Isaac (1956: 182) gives no details of the material collected by him
in Portuguese East Africa and named Qelidiopsis rigida (Vahl) Web. v. Bosse so that it is
impossible to decide the further distribution of the species on the Atlantic coast of Africa.
Ezsiccuta s. TOME: : Quintas 14, I l h h das Rolas, 1885. Quintas s.n., Equador, Canal das
Rolas. Alga do fundo (calcario). 1885.
Distribution Africa : S. Tom& Widespread in warmer waters.
Gelidiopsis Schmitz 1895
Gelidiopsis variabilis (Grev.) Schmitz, Bot. Jb., 21: 148 (1895); as a. intricata in Exell
(1944: 386).
Exell’s plants are 5 to 6 cm high, very irregularly branched and somewhat coarse with
branches of rather variable width (225 to 450 p wide). They are sterile.
They agree closely in form with the illustrations of Gelidium variabile (Grev.) J. Ag. in
Kutzing (1869: 9 and pl. 23, figs (c) and (d) and in cross-section the thallus present a
similar appearance to that of the type shown in Feldmann (1931 : 166, fig. 2(b)).
G. variabilis was a t the time considered a mainly Indian Ocean species, though it has
since been reported from Annobon, Cameroon, Ghana and from G u i d e with doubt (see
Sourie 1954: 306).
Both G. variabilis and a. intricuta (C. Ag.) Vickers are probably in need of critical
re-examination. The latter name is probably invalid, since i t is based on Sphuerococcus
intricutw C . Ag., the ‘cotype’ of which Feldmann has examined and reported as a
Gelidium species, since intercellular rhizines are present (Feldmann, 1931 : 157).
Vickers’s species was in fact only described relatively recently in Taylor (1960: 353)
who reports it from various places along the warmer Atlantic shores of the Americas.
Exell’s plant appears to differ in habit from G. intricata as described by Taylor through
the range of thallus diameters is similar.
Vickers 127, Algues de la Barbade, consists of plants from a t least two places, since the
specimen in BM, presumably available to Tandy when identifying Exell’s plants, differs
in details of locality and date from those given in Vicker’s list (1905: 61). This plant has
been examined and is in many ways intermediate between G. variabilis and B. intricata
as generally understood with a n erect tufted habit and thalli of somewhat variable width.
Another specimen ex. BM, Hamel et Hamel-Joukov 88 from Guadeloupe, corresponds more
closely with Taylor’s description of Q. intricuta and with that given by Bergesen for a
Mauritius plant (Bmgesen, 1943 : 53).
Anatomically i t seems impossible to find any sharp and constant differences between
the two species.
On the Atlantic coast of Africa a. intricuta is reported in Trochain (1940: 109) washed
I
MARQARETSTEENTOET,
F.L.S.
126
upon the coast of SBndgal and from Cab0 Verde Is. (in Chevalier, 1935: 1069-1071).
Material from both places was identified by Feldmann.
After examining the Indian Ocean material of both species present in Hb. Brargesen,
i t seems to the present writer that there is some doubt as to the distinctness of G. variabilis
and Q. intricata. Larger tuft-forming plants seem to be placed in the former species, lowgrowing turf-forming ones in the latter. However a much greater acquaintance with the
American forms of the so-called Q. intricata is required before any conclusions can be
reached.
Exsiccuta s.
TO&:
ExelE 469, Praia Pequena, 30.xi.1932 (BM).
Distribution Africa : Annobon, S . TomB, Cameroun, Ghana, ?GuinBe, ?Canary Is.,
Indian Ocean.
Gelidium Lamour. 1813
Gelidium crinale (Turn.) Lamour. in Bory, Dict. class., 7 : 191 (1825); Hariot (1908:
162) and Henriques (1917: 166).
The condition of the single specimen makes comment on Hariot’s identification impossible. Morphologically there is good reaaon to believe it is correct. G. crinule is known
from Ghana, Mauritanie and Maroc.
Exsiccuta s. TOMB : Ribeiro !7, Roqa Praia da Nazareth, 1894.
Gelidium reptans (Suhr) Kylin, Acta Univ. Lund., 2 (34):6 (1938).
This is a tentative identification for aome fragments (with tetrasporangia) found
growing with Struvea amtomosans. The plants resemble the figures of Kylin (1938: 7,
fig. 2A to C), Dangeard (1949: 150, fig. 10) and Isaac (1956: 185, fig. 17). Carpine also
reports this species from the Ilh6u das Rolas (Carpine, 1959 : 83).
a. reptans has been considered a form of various other species; see, for example,
Bargesen (1943:6).
Exsiccuta s. TOME:Fox Nielsen 370, sin. loc., August 1956.
Distribution Africa : Portuguese East Africa, South Africa, S. Tom&,S&n&gal,Maroc.
Wurdemanniaceae
Wurdemannia Harvey 1853
Wurdemannia miniata (Drap.) Feldm. et Hamel, Revue g6n. Bot. 46: 544 (1934); as
setacea in Brargesen, Dansk Bot. Ark., 3 : 368 (1915-20); as Caulacanthus ustuhtua in
Henriques (1886 : 220) and Hariot (1908 : 162) ; as C. notuhtua in Henriques (1917 : 166).
w.
The multiaxial construction of the thallus clearly removes the S. Thomd plants from
&dacanthus, and the absence of thickened walls to the central cells of the thallus precludes
Qelidiopsis. Unfortunately there are no tetrasporangia, which could otherwise be used
as a deciding character. They are zonate in Wurdemannia but tetrahedral in Qelidiopsis.
I n Bergesen (1915-20, fig. 3 6 1 ( ~ )a)thallus apex is illustrated, and this aids in differentiating sterile specimens, for in Wurdemunniathe apex is narrow and made up few cells, in
Qelidiopsis it is broad and made up of many cells. Caulacanthus is widely reported on the
Atlantic coast of Africa and it is possible that some of these records refer to Wurdemannia.
The Moller specimens are undoubtedly wrongly attributed. There is no evidence that
Moiler ever visited the southern part of the coast of S. Tom&.Quintas probably collected
these plants, and only he is referred to in Henriques (1886: 220).
Marine algae of ScZo TomC and Principe
127
Exsiccuta s . TOM^: [Moller] s.n., Equador, Canal das Rolas. Alga a 5 p6s de fundo
(calchreo) [1885]. Quintas 8, Ilh6u das Rolas, 1885.
Distribution Africa: S. TomB, Canary Is. Atlantic coasts of tropical and subtropical
America, Mediterranean.
Squamariaceae
Hildenbrandia Nardo 1834
Hildenbrandia sp.; as H . rosea in Henriques (1886: 220; 1917: 166).
The specimen is sterile and takes the form of two small pink crusts growing on shells.
Since the species are separated on mode of division of tetrasporangia, further comment is
impossible, except to note that the name H . rosea Kiitz. is no longer valid.
Exsiccuta
8.TOMS
:Moller 22, Bahia de Ana Chaves, 9/85 (in Hb. Agardh, no. 27618) LD.
Peyssonnelia Dcne. 1841
Peyssonnelia rubra (Grev.) J . Ag.,Spec. alq., 2 (2):502 (1852);Barton (1897: 374).
The thalli are thin but apparently easily detached and relatively heavily calcified. The
material proved difficult to handle, but as far as could be seen the apical cells were
approximately of the same length as the height of the hypothallus cells, and there seems
therefore to be good reason to agree with Barton’s own determination.
In SBnBgal, Sourie (1954: 107) records only Peyssonnelia spp., and in Maroc, P. rubra
is mentioned only by Dangeard (1949 : 152), not by Gayral(l958).
Ezsiccafa PR~NCIPE: Welwitsch 256, 21 Sept. 1853 [just above low-tide level] (BM).
Distribution Africa : Angola, Principe, ?SBn&gal,Cab0 Verde Is., Canary Is., Maroc,
Madeira, Tanger. Probably in most warmer seas.
Corallinaceae
(Corallineae)
Amphiroa Lamour. 1812
Amphiroa sp., as A . capensis in Henriques (1886: 220; 1917: 166).
The specimen consists only of fragments, and showed little resemblance to the authentic
specimen of A . capensis Aresch. seen (in HBG). It seems best to leave this material without
specific determination.
Exsiccuta s. TOM^ : Quintas s.n., Equador, Canal das Rolas, 1885.
Jania Lamour. 1812
Jania adhaerens Lamour., Hist. Polyp.: 270 (1816); as Corallina adhuerans in Pilger
(1911: 312).
This is reported from the IlhBu das Rolas by Carpine (1959 : 83). Pilger’s seems to be
otherwise the first and only record for this species on the Atlantic coast of Africa. Distribution is more or less pantropical.
Jania rubens (L.) Lamour., Hist. Polyp., 272 (1816).
Small tufts of this species were found with Bryothamnion triquetrum. It is rather
more slender than is typical, the upper segments 60 p wide basally, tapering to 45
128
M ~ ~ G A RSTEINTOFT,
ET
F.L.S.
apically. The lower branahes are a t most 160 p wide. I n habit the erect branches are very
fastigiate, arising from a disc 375 p in diameter, to form tufts 0.5 to 1 cm high. The plants
are sterile.
J. natalensis is recorded from Gabon (leg. Soyeaux) by Schmidt (1930: 172) and
Corallinu (Junia)Eongifurca from R6publique du Congo (leg. Lecomte) by Hariot (1895 :
244), but it does not seem likely that either of these would be confused with J. rubem.
Exsiccatu s. TO&: [Quintas?]s.n., Ilh6u das Rolas [1885].
Distribution Africa : a new record for S. Tom6; see also Fox (1957 : 625).
Corallinaceae
(Melobesieae)
Lithophyllum Philippi 1837
Lithophyllum aequinoctiale (Foal.) Foslie, Alg. not. 6, K . norske Vidensk. Sebk. Skr.,
1909 (2):46 (1909) and Foslie (1929, pl. 70, figs 4 to 5).
This is another of the six species described by Foslie from S. Tom6 material and is one
of the two apparently endemic species.
Exsiccata s. TO&&: Quintas 33 p.p., Ilh6u das Rolas 1885 (with Lithophyllum simile)
( ? in Hb. Foslie, TRH, non. wid.)
Distribution Africa : S . Tom6.
Lithophyllum africanum f. intermedia Foslie, K. norske Videmk. Sebk. Skr., 1900
(3):3 (1900b) and 1900 (5): 18 (1900~);
Foslie (1929, P1.68, fig. 2); Lemoine, Annb Inst.
OCWTW~T., Mcnuzco, 2 (2): 146 (1911). As Liththamnium ponderosum in Henriques (1917:
166).
Moller 28 is the type of L . ponderosurn Foslie (1897 : 15), who comments that it ‘ seems
to have been dead when collected and the shape of the branches apparently in part to
have been caused by the in0uence of rapid tides’. I n Foslie (1909 : 42) it is stated that ‘the
only known specimen of L. ponderosum is an old and strongly waterworn form of Lithophyllum africanum with only the lower parts of the plant left . .’.
L. africanum is recorded by Foslie (1900b : 3) for Cap Vert, and in Foslie (1909 : 42) for
Cab0 Verde Is. Mme. Lemoine has seen these plants and also reports the species for Tanger
and the Canary Is. (Lemoine, 1911: 150).
.
Exsiccata
8 . TOM&: Moller 28, Bahia de Ana Chaves et Praia Lagarto, 7/1885. ‘Comm.
Kjellman ex Herb. Univ. Coimbra’ (in Hb. Foslie, TRH).
Distribution (of the species as a whole) Africa: Annobon, S. TomB, S6n6ga1, Cab0 Verde
Is., Canary Is., Maroc, Tanger.
Lithophyllum irregulare (Foslie)Huv6, Lithothmnium(?) irregulare Foslie, Alg. not. 3,
K. norske Vidensk. Selsk. Skr., 1906 (8): 6 (1907). Tenarea irregularis (Foal.) Lemoine,
K . danske Vidensk. Selsk., Biol., 8 (1): 56 (1929).
This species was described by Foslie from the S. Tom6 material, and is illustrated in
Foslie (1929, pl. 12, fig. 22). The type material has been examined. It consists of three
large pieces and a number of fragments, from which two slides have been made, accompanied by three lists of cell measurements in Foslie’s hand. One of the slides is labelled
‘ex Bot. Mus. Hamburg’, but the significance of this remark is not known.
The hypothallus cells are a little more variable in size than Foslie (1907 : 6) indicates.
and according to the lists the measurements should be 11 to 29 p long by 6 to 14 p wide,
Marine algae of Silo Tom6 and Principe
129
i.e. rather nearer to the 7 to 22 p long by 5 to 9 p wide given by Mme Lemoine for Brargesen’s
Canaries plants.
Exsiccata s. TO^: Moller 26, Bahia de Ana Chaves e t Praia Lagarto, 6/85 (in Hb.
Foslie, TRH).
Distribution Africa: S. Tom6 and Canary Is.
Lithophyllum retusum Foslie, K . norske Vidensk. Selsk. Skr., 1909 (2):33 (1909) and
1900 (5): 18 (1900b). Lithothamnium retusum Foslie, K . norske Videmk. Selsk. Skr., 1897
( l ) ,15 (1897). Non Lithophyllum retusum Foslie fa. in Hariot (1908: 164) and Herriques
(1917: 166).
This is another species described by Foslie from a Moller specimen, and illustreted in
Foslie (1929, pl. 64, figs 12 to 14).
Foslie refers to an additional locality, Fort Brandenburg in Ghana. This was one of the
two Brandenburger forts built near Cape Three Points and occupied up t o 1720. The
western fort (Gross Friedrichsburg) was known as Fort Brandenburg a t least until 1873,
though the town now there is known as Princestown. No specimen has been traced.
Gravier’s plant, referred to in Hariot (1908 : 164), has been examined by Mme Lemoine,
who states (in litt.) that i t is not a form of L . retusum, and it seems therefore that L .
retusum and another, undescribed species have been confused. Foslie has published no
comments on Gravier’s plant.
Exsiccata s. TOM$: Moller 24, Bahia de Ana Chaves et Praia Lagarto, 7/85 (in Hb.
Foslie, TRH ; also PC, ‘ex Chicago Museum ’).
Distribution Africa : S. Tom&,?Ghana.
Lithophyllum simile Foslie, Alg. not. 6 in K . norske Vidensk. Sebk. Skr., 1909 (2) : 30
(1909; 1929, pl. 63, fig. 22).
This is the fourth Lithophylhm species demribed by Foslie from a S. Tom6 specimen,
and one of the two apparently endemic species of Melobesieae.
Exsiccata s. TOM$: Quintas 33 p.p., Ilh6u das Rolas, 1885 [with Porolithon aequinoctiale]
(in Hb. Foslie, TRH).
Distribution Africa: S. Tom&.
Lithothamnium Philippi 1837
Lithothamnium brachycladum Foslie, K . norske Vidensk. Sebk. Skr., 1899 (5): 3
(19OOa). As L . crassuminDickie (1873: 179).
Foslie’s species is founded on a specimen in BM probably the same one referred by
Dickie to L . crassum Phil. (Dickie, 1873: 179). This Foslie first included in Lithophyllum
racemus (Lamour.) Foslie, later making it the type of Lithothamnium brachycladum,
to which he also referred Welwitsch’s Principe plant. The latter he did not however
examine.
Exsiccata PR~NCIPE: Welwitschnon. vid. (?BM).
Distribution Africa : St. Helena, ?Principe.
Lithothamnium floridanum Foslie, Alg. not. 2, K . norske Vidensk. Selsk. Skr., 1906
(2): 11 (1906).
Foslie states that he has seen two specimens from ‘St Thomas, West Coast of Africa’,
‘which both in habit and with regard to structure, stand very near to L.flbridanum, but
in that they are sterile, they cannot be definitely determined’.
130
MARQARETSTEINTOFT,
F.L.S.
L . JEoridanum is otherwise known only from the western Atlantic, the Florida type
specimen being illustrated in Foslie (1929, pl. 12, fig. 12).
Both Moller 25 and 27 are untraced and it seems likely these are the two Foslie is
reporting.
Exsiccata s. TOM&: ?Moller 25 and 27, non. vid.
Distribution Africa : ? S . Tom&.‘Florida, ?Cuba, Hispaniola’ (Taylor 1960 : 383).
Lithothamnium subtenellum Foslie in Hamel & Lemoine, A r c h Mus. natn Hist. nut.,
Paris, 7 (1) : 24 and 98 (1963). Lithophyllum subtenellum Foslie, Alg. not. 6, K . norske
Vidensk. Sebk. Skr., 1909 (2): 11 (1909); Hariot (1908: 164); Henriques (1917: 166).
Ooniolithon subtenellum Foslie, K . norske Vidensk. Selsk. Skr. 1898 (6): 11 (1098).
Foslie has published no comments on Gravier’s plants, and the illustration in Foslie
(1929, pl. 53, figs 1 to 2) is of material from the Atlantic coast of France.
Exsiccata s . TOM&: Oravier s.n., Bella Vista [1906] (in Hb. Foslie, TRH, also PC).
Distribution Africa : S . Tom6, Maroc. Atlantic coast of France, Mediterranean.
Lithothamnium tenuissimum Foslie, K . norske Vidensk. Selsk. Skr., 1899 ( 5 ): 20
(1900a and 1900 (5): 15 (1900~).
Foslie (1900a: 21) remarks that this species seems to be ‘of pretty common occurrence ’.
This is another of the six species described by Foslie from Quintas and Moller specimens.
I n this case it is Moller 23, sent t o Foslie as Henriques 23, and not to be confused with
Henriques 23 from St. Vincent (Lithophyllum ajricunum, Foslie 1909: 42; 1900b: 4).
Exsiccata s. TO&: Moller s.n., [i.e. 231 Bahia de Ana Chaves e t Praia Lagarto, 1885
(in Hb. Foslie, TRH).
Distribution Africa: S. Tom&,Canary Is., Maroc.
Mesophyllum Lemoine 1928
Mesophyllum canariense (Foal.) Lemoine, K . danske Vidensk. Sebk., Biol.,8 (1): 31
and pl. 1, figs 1 t o 2, pl. 2, figs 2 and 4 (1929).Lithothamnium cunariense Foslie, K . norske
Vidensk. Selsk. M w . Arsberetn, 1905: 1 ( 1 9 0 6 ~ )As
. Lithophyllummurlothii fa. in Hariot
(1908 : 164) and Henriques (1917: 166).
Two contradictory statements are found, in Foslie’s handwriting, on slips of paper
accompanying the two small fragments and slide of this species in Hb. Foslie. One reads
‘Usually [gj. =gjerne] the cells a little larger and wider, and smaller conceptacles than the
type’, while on a list of cell dimensions is written ‘Not usually with thicker or longer cells
than in Marlothii. Conceptacles 140-240 p’.
Mme Lemoine has examined the portion of Gravier’s material remaining in Paris, and
considers it identical with Memphyllum canariense (Lemoine in litt.).
Exsiccuta s . TOMI%:Gavier s.n., sin. loc. [1906] (in Hb. Foslie, TRH, and PC).
Distribution Africa: S . Tom&,Canary Is.
Neogoniolithon Setch. & Mason 1943
Neogoniolithon sp.
Reported by Carpine (1959 : 83) from I l h h das Rolas.
Marine algae of S& Tome‘and Principe
131
Porolithon Fmlie emend. Lemoine 191 1
Porolithon boergesenii (Fosl.) Lemoine, Damk Bot. Ark. 3 : 149 and 178 (1915-20).
Goniolithon Borgesenii fa. africuna Foslie, Agl. not. 3, K . norske Vidensk. Selsk. Xkr.,
1906 (8):20 (1907); Hariot (1908: 164); Henriques (1917: 166).
The species was described from material collected by Bsrgesen on St Croix, the fa. africuna being distinguished later to accommodate Gravier’s plant.
From Foslie’s slide i t can be seen that the thallus is extremely irregularly organized, no
rows of cells being traceable, but characteristically scattered heterocysts are not visible
and probably for this reason, Foslie created a separate form.
Exsiccuta s. TOMI$: Gravier s.n., sin. loc. [1906] (in Hb. Foslie, TRH].
Distribution Africa : S. Tom& Western tropical and subtropical Atlantic.
Gracilariaceae
Gracilaria Greville 1830
Gracilaria dentata J. Ag., Sp. alg., 2 (2): 603 (1852).
The single specimen found is labelled in Binder’s handwriting, and appears to be one
of the four possibly collected for him by Epffenhausen (see p. 101).
In form i t resembles some specimens from Ghana which have hitherto been known by
the name G. henriquesiana Hariot (in semu Dickinson and Foote, see below), and also
some forms collected a t Lagos by the present writer and known as G . dentata (Fox 1957 :
624).
Ezsiccuta S. TOMB: [Epflenhuusen?],St Thomas, 1838 (in Hb. Binder, HBG, sub. nom.
Sphuerococcus multipartitus nom. nud.).
Distribution Africa : RBpublique du Congo, S. TomB, Cameroun, Nigeria, Ghana,
Gambia, SBnBgal. Otherwise recorded, with doubt, for the warmer Atlantic coasts of the
Americas (Taylor, 1960: 451).
Gracilaria henriquesiana Hariot, J . Bot., Paris, 2e ser., 1 (7): 162 (1908). Non G .
henriquesiana in sensu Dickinson and Foote (1951: 270). As G . wrightii in Henriques
(1886 : 221 ; 1917 : 166) ; as G . Henriquesii in Henriques (1917 : 166).
The type material of Hariot’s species was collected partly by Moller, partly by Ribeiro.
Although Praenpa is mentioned as the collector by Hariot (1908 : 163), his name does not
accompany any of the specimens, and it has been impossible t o find out anything about
him. He was possibly a local assistant to either Moller or Ribeiro.
Moller’s material, now a t Coimbra, consists of four plants or parts of plants mounted
on two sheets. Three specimens are labelled G . wrightii, but this can hardly be considered
a correct identification. G. wrightii (now G. debilis (Forssk.) Borg.) is placed by Agardh in
the Pachycladia section of the genus, having a cortex composed of radial rows of cells.
Moller’s plants have a submonostromatic cortex and belong therefore in the section
Podeum.
Hariot was able to examine Ribeiro’s plants in addition to Moller’s. The type material
(in PC) consists of two gatherings, Ribeiro 14 and Moller 15, although the latter is now
represented by an empty packet. One may assume that Hariot had both plants in mind
when describing the species since he refers (1908: 163) to the type locality as ‘Praia
Lagarto (Rages Praia da Nazareth)’, a combination of two entirely different places. The
first relates to Moller 15 (COI), and the second t o Ribeiro 14 (PC) (sub. nom. clr. Henriquezianu). A third gathering, collector unknown, sub. nom. G . Henriquezii in PC is
apparently part of Ribeiro 14.
9
138
MARGARETSTEENTOFT,
F.L.S.
The only collector given by Hariot is ‘Dom. A. A. Praenva’. Which specimen Praenya
collected is not clear. There are thus two syntypes, perhaps three.
It is necessary to establish that Hariot used both collections in describing his species,
because there are differences between them and Hariot’s description applies more directly
to Ribeiro’s plant than to Moller’s. Colour, form and anatomy differ somewhat, and in
addition Hariot chose to illustrate only what appears to be part of Ribeiro 14 (Hariot,
1908: 163).
As regards colour, Moller’s plants are a faded shade of light brown, a common colour
for Qracilaria dentata, for example, in,the Gulf of Guinea, whereas Ribeiro 16 is a striking
red colour, such as one would expect to find only in plants growing in very shaded conditions in the Gulf.
I n habit and form, differences are not so great, though Moller’s plants are a little
broader and have less even margins than Ribeiro’s, and the branching is more open. These
differences are slight and, of course, purely qualitative.
Anatomically Moller 15 is indistinguishable from the bulk of Nigerian and Ghanaian
material of G. dentata examined by the present writer. The cortex is well developed, morc
than one cell thick and sometimes up to three cells thick, composed of very small cells
with well-defined walls. I n Ribeiro 14, the cortical cells are larger and rather irregularly
disposed, producing an uneven and ill-defined outer surface to the thallus, the cortex
never being the compact tissue usual for G . dentata.
There seems to the present writer some reason to believe that Moller 15 represents a
different speciesfrom Ribeiro 14, but it is proposed to examine the flat speciesof Gracilaria,
as they occur on the Atlantic coast of Africa, separately, and to retain Hariot’s delimitation of a. henriquesiana for the time being.
As regards the illustration of G. henriquesiana (Hariot, 1908: 163), this is in the first
place of poor quality. Secondly it appears to be composed of the two largest portions of
Ribeiro 14, though neither can be matched with certainty to particular parts of the
photograph.
It is also unfortunate that no scale is given, nor is size mentioned in the text of Hariot’s
description. If it is true that the photograph represents the assembled portions of Ribeiro
14, then i t has been reduced in reproduction by about 4 times (linear). The specimen must
have been approximately 40 cm across and 15 to 20 om high when growing.
If it were supposed that the photograph had been reduced by about twice linear, as is
commonly done today, then many of the Ghana forms of a. dentata J. Ag. would agree
reasonably well with the description and illustration of G . henriquesiana. This mistake
was made in Dickinson & Foote (1951 : 270) but it was equally a mistake, in Fox (1957 :
624), to reduce G. henriquesiana to the synonymy of G. dentata. Only the Ghana forms of
G . henriquesianasemu Dickinson and Foote should have been so treated. Having seen the
type of G. henriquesiana it is possible to state quite definitely that it is a form distinct from
G. dentata a8 known in Cameroun, Nigeria and Ghana. These latter plants have been
compared with the type of a. dentata in Hb. Agardh (leg. Epffenhausen, Gambia, 1837)as
well as with an isotype (in BM) and there is no doubt as to the identification is correct
(though there is doubt as to the distinction between G. dentata and G . foliifera, a point
which cannot be dealt with here).
I n Plates 1 and 2, the main portions of the syntypes and isotypes of 0.henriquesiana
are shown. For convenience, Hariot’s description is here reproduced :
‘Gracilaria Henriquesiana sp.n.
Fronde subcartilaginea, complanata, basi attenuata, sat regularite dichotomofastigiata, margine pro rite nuda ; segmentis linearibus, axillis rotundatis vel leviter
acutiusculis, apice furcatis multidentatis ve cystocarpis hemisphaericis (usque ad 1 mm
diam.) apice obtusis, per totam frondem sparsis, saepius marginalibus. Color intense et
nitenti roseus.’
Marine algae ojS6o Tomi and Principe
133
It can be seen that Moller’s plant has broader, more cuneate segments and less tendency
to pseudosecund branching. But all the specimens adhere well to paper and although
plants of G. dentata can be found with one or more of the characteristics of G. henriquesiana
none has been discovered which adheres well to paper.
Exsiccuta s. TOM^: Moller 15. Praia Lagarto, 8-1885 (isotype and syntype, COI);
Ribeiro 14, Roqa Praia da Nazareth, 1894 (isotypes COI, two syntypes PC).
Distribution Africa: S. Tom6 (endemic).
Gracilaria verrucosa (Huds.) Papenfuss, Hydrobiologia 2 : 195 (1950).
This is a widely distributed species in the Gulf of Guinea, previously reported from S.
Tom6 in 5 to 6 m of water, by Carpine (1959 : 77).
Exsiccata s. TOM^: [?Epffenhawen],St Thomas, 1838 (in Hb. Binder, HBG, sub nom.
Sphaerococcw confervoides v. caudatus Ag. ms.).
Distribution Africa : South Africa, Angola, R6publique du Congo, S. Tom&,Cameroun,
Ghana, SBn6ga1, Mauritanie, Maroc, Tanger. Widespread in all except the coldest waters.
H ypneaceae
Hypnea Lamour. 1813
Hypnea cervicornis J. Ag., S p . a@., 2 (2):451 (1852);Taylor, Mar. alg. . . . Americas:
466 and pl. 73, fig. 2 (1960); as H . mwcijormis in Henriques (1886: 220; 1917: 166), and
Hariot (1908: 163).
This is a new record for S. Tom&.Thorold’s plants are fertile, and the tetrasporangial
branches in Fox Nielsen 242 have a characteristically antlered form, but in Fox Nielsen
241 the tetrasporangial branches simply taper, as do the fertile zones of these branches.
The former plant is of the cushion type, the latter a softer and looser form.
Moller’s material, which gave rise to the original record of H . mwcijormis, is quite
lacking in hooked branches, and is almost all of the cushion-forming type.
Ribeiro’s plants are partly loosely branched, partly of the cushion-forming type. When
sterile the latter are difficult to distinguish from H . spinella but the cushions seem to be
softer and looser than those of H . spinella (as here distinguished) and there is a distinct
tendency for the main axes to be thicker than the side branches.
It seems therefore possible to distinguish H . cervicornis from H . spinella among the S.
Tom6 material, but not to take the problem further. Taylor (1960: 465) also distinguishes
the two species, as did Bsrgesen (1915-20: 385) though he thought they were possibly the
same, in which case the name H . spinella (C. Ag.) Kiitz. would have precedence.
The H . pannosa mentioned by Dangeard (1952 : 282 and pl. 19, fig. c) seems to be very
similar to H . cervicornis. Taylor (1945: 226) distinguishes H . pannosa as a Pacific and
Indian Ocean species, with H . cervicornis as the possible parallel Atlantic species.
Carpine (1959: 86) has the first record of H . cervicornis from Principe.
Exsiccata s. TOMI? :Fox Nielsen 241,242, sin. loc., August 1956 ( =Thorold 613 and 616
(the latter sub.nom. H . spinella) in BM spirit collection). Moller 13, sin. loc., 1885 [with
Acanthophora spicifera] (duplicate PC). Moller s.n., sin. loc., 1885. Ribeiro 11, Roqa
Praia da Nazareth, 1894.
Distribution Africa: S. Tom6 and Principe, SQnBgal,Canary Is., Maroc. Atlantic coasts
of tropical and subtropical America.
Hypnea musciformis (Wulf.) Lamour., Essai . . . Thalass.: 45 (1813); Henriques
?(1885:134), also (1886: 220; 1917: 166); non Hariot (1908: 163).
134
MARGARETSTEENTOFT,
F.L.S.
Henriques (1885: 134) refers to Newton’s plant, which has not been seen. Otherwise
previous records refer to H. cervicornis. Carpine (1959: 87) records H. musciformis for
Principe for the first time.
Barton (1893: 172) lists several collectors of this species in South Africa, but it is not
mentioned in the available papers of Papenfuss, nor in Stephenson’slist (1948 : 305-306).
Exsiccatu s. TOM$:Fox Nielson 239, sin. loo., Angust 1956 (=Thorold 617 in BM
spirit collection). Collector not known (?Epffenhausen) s.n., St. Thorn.,
1838 (in Hb. Binder, HBG).
Distribution Africa : ?South Africa, Angola, Rdpublique du Congo, S. Tom& Cameroun,
Nigeria, Ghana, Sierra Leone, Cab0 Verde Is., Mauritanie, Canary Is., Maroc, Tanger,
Agores. Widespread in warmer waters.
Hypnea spinella (c. Ag.) Kiitz., Spec. alg., 759 (1849); Bnrgesen, Dansk Bot. Ark., 3 :
384 (1915-20); Henriques (1886: 220; 1917: 166); Hariot (1908: 163).
The distinction between H. spinella and H . cervicornis has long been in doubt. Quintas’s
plants are of the mat-forming type mentioned by Taylor (1960 : 465), but they are almost
completely sterile and the few tetrasporangial branches present are insufficient to give a
reasonable idea of their range of form. The plants resemble the basal parts of what the
present writer calls H. cerviwrnis except that there is less tendency for the main and side
branches to differ in width, and the form of the tetrasporangial branches is nearer to
that illustrated by Bsrgesen (1915-20, fig. 369).
Dangeard (1952: 282) mentions H. ctenomycas J. Ag. and it seems possible that this is
the same form as the above. For Maroc, Dangeard (1949: 162) mentions H. spinella,
though Gayral (1958) does not. The record for South Africa rests on a single Krauss
collection, listed by Barton (1893 : 173).
If H. spinella is maintained as distinct from H. cerviwrnis, then it would still appear to
be widely distributed on the Atlantic coast of Africa.
Exsiccata s . T O : Quintas
~
s a . , Equador, Canaldas Rolas. Alga da superficie (caldreo)
banhada pela enchenb-ccr violhcea. 1885. Quintas 11, Ilh6u das Rolas,
1885 [with Laurencia perforuta].
Distribution Africa : ?South Africa, Annobon, S. Tom6, Sierra Leone, ?S8n6gal, ?Cab0
Verde Is., Canary Is., ?Maroc.
Champiaceae
Champia Demaux 1808
Champia parvula (C. Ag.) Harv., Ner. Bw.-Am., 2: 76 (1853).
Carpine (1959:78 and 87) reports this species for both S. Tom6 and Principe. It appears
to be widely spread in warmer waters.
Champia tripinnata Zanard., Regemb. Flora, 1851, 34 (1851); PI. Mar. rub.: 69 and
pl. 9, figs 2 and 2a (1858).
There is a single plant of this species, 5 om high, translucent and almost colourless. It
possesses sessile, urn-shaped ostiolate pericarps within which, around the cystocarp, is a
mesh of very slender cells.
Pilger (1920: 5) identified an Annobon specimen (Nildebrued 6779) as belonging in C.
tripinwtu, a species otherwise reported only from the Red Sea.
Newton’s plant is oppositely or alternately branched, the branches of somewhat
variable width (between 1 and 2.5 mm) with the segmentsat least, as long as, and generally
twice as long as, wide. The diaphragms are very difficult to see.
Marine algae of S h Tomd and Principe
135
Zanardini (1858: 69) gives the height of his plant as ‘4.5 decim.’ but from the illustration
and measurements given this should obviously be ‘ 4 to 5 cm’. The record for Principe is
new.
Exsiccata
P R ~ N C I P E Nezoto,~
:
s.n., Praia
de Sundy, 1887.
Distribution Africa : Annobon, Principe. Red Sea.
Ceramiaceae
Centroceras Kiitz. 1841
Centroceras clavulatum (C. Ag.) Mont., Explor. scient. Algerie: Algues 1 : 140 (1846).
As C. hyalacanthum in Henriques (1886 : 219) ; as Ceramium clavulosum in Hariot (1908 :
164) and Henriques (1917: 166).
This is probably the most widespread of the Ceramiaceae in the Gulf of Guinea, though
not necessarily occurring in quantity. Part of Quintas’s material came from the sublittoral
association he investigated.
Carpine (1959 : 82) reports this species from the littoral region on S. Tom&
Exsiccata s. TOMB: Fox Nielsen 226, sin. loc., August 1956. Quintas s.n. (Norstedt 12),
Equador, Canal das Rolas. Alga do fundo (calcheo) c6r carminada, 1885
[with Bryothamnion seaforthii]. Quintas 9, Ilh6u das Rolas, 1885.
Distribution Africa: Portuguese East Africa; see also Fox (1957: 626). Congo, Gabon,
Togo and Dahomey are the only countries not represented so far.
Ceramium Roth 1779
Ceramium gracillimum var. byssoideum (Harv.) Mazoyer, Bull. Soc. Hist. nut.
Afr. N . 29: 323 (1938).
This species is recorded for S. Tom6 for the first time by Carpine (1959: 77), who
collected it in 11 m of water.
Spyridia Harvey 1833
Spyridia aculeata (Schrimper) Kutz., Phyc. gen. : 377 (1843); Bsrgesen, Dansk Bot.
Ark., 3 : 237 (1915-20); as S. filamentosa in Henriques (1886: 219; 1917: 166).
Newton is recorded as having collected S.filamentosa on S. Tom6 in 1881 (Henriques,
1885: 133) and although this plant has not been seen, i t is likely that i t is the same as
Quintas’s, i.e. S. aculeata.
Quintas’s plant lacks the slender ramuli, corticated only a t the nodes, usually found
in S. aculeata. Instead i t has numerous short, stiff spine-like branchlets, heavily corticated
a t the nodes and bearing terminally the two or three recurved spines distinctive of the
species. There are no hook-like ends to the branches as reported by Berrgesen (1915-20:
237).
The form of the branchlets suggests S. complanata J. Ag. but there is no trace of flattening of the thalli nor of distichous arrangement of the branchlets. Taylor (1960: 540) says
that S. complanata is ‘doubtfully distinct from S . aculeata var. distich’.
It is a pity there is so little material, for Quintas’s plant is unusual and closely resembles
S . a r m t a Kutz., which was sunk in 8.aculeata by De Toni (1903: 1433).
The S. Tom6 record is new for the island and i t is interesting that Carpine (1959: 77)
also reports f. typica, i.e. f. aculeata, collected a t the northern end of SHo Tom6 in 11 m of
water. He also reports f. inermis from Principe for the first time.
Var. hypeodes is reported washed up on the coast of SBnBgal by Trochain (1940 : log),
136
MARQARETSTEENTOFT,
F.L.S.
and var. distich is reported growing attached*by Sourie (1954: 108). From Mauritanie
var. h&neoides is reported (presumably growing attached) (Feldman, 1938 : 111).
Ezsiccata s. TOXI%:Quintas s.n., Equador, Canal das Rolas. Alga da superficie (basalto)
banhada pela enchente-chr violhcea. 1885.
Distribution Africa: S. Tom6, Principe, S6n6ga1, Cab0 Verde Is., Canary Is., Maroc,
?Tanger. Most warmer waters except, apparently, the Pacific.
Spyridia clavata Kiitz., Linnea 15: 744 (1841); Brargesen, Damk Bot. Ark., 3: 235
(1915-20); Hariot (1908: 164); Henriques (1917: 166).
There are very few clavate branches on this specimen, though the branching is
distichous.
Bergesen (1915-20 : 237) gives a record for Senegambia probably taken from De Toni
(1903: 435). There are specimens from Gambia in Hb. Binder, and Chevalier (1920: 789)
has collected this species in Sdnkgal. Barton (1897 : 373) reports it from Angola, though
not growing attached.
Emiccata s. TOME: Ribeiro 16, Roga Praia da Nazareth, 1894,
Distribution Africa : ?Angola, S. Tom6, Gambia, SBn6gal. Widely distributed on the
tropical and subtropical coasts of Bmerica.
Rhodomelaceae
Acanthophora Lamour. 1813
Acanthophora muscofdes (L.) Bory, Voy. Coquille, Bot.: 156 (1828). Non Henriques
(1886: 221; 1917: 106), Hariot (1908: 163).
Recorded by Carpine (1959 : 77) in 11 m of water off S. Tom6.
Acanthophora ramulosa (Lindenb. ms.) Kiitz., Steentoft emend. ; Kutzing, Phyc.
g e m :437 (1843);as A. muswides in Hariot (1908 : 163) and Henriques (1917 : 166) both p.p.
Ribeiro’s plant is a large one, about 16 om high, much branched and with spines on
both the short branchlets and on the long branches, on the latter both young and mature.
The short branches are terminally developed in many cases into rather wrinkled spineless
portions in which tetrasporangia formation is incipient. Except for their position they
resemble the stichidia illustrated by Falkenberg (1901, pl. 22, fig. 3) for A. delilei. From
the form of the stichidia, and from the distribution of spines independent of the origin of
branches and branchlets, Ribeiro’s plant cannot be placed in A. muscoides.
The difference between the 8 . Tom6 plant and A. apicijera is also clear. The long
branches are much more highly developed than in A. spicifera, they bear spines, and the
stichidia and some of the branchlets are of a different form. Finally Ribeiro’s plant swells
up much more in water, is fairly soft and has a deep pink or ruby colour.
These observations led to comparison with the plant known as A. ramulosa (Lindenb.)
Kutz., from Angola, one of the species sunk in A. muscoides by De Toni (1903: 818) and
illustrated in Kiitzing (1865: 27 and pl. 76a,b). I n Kutzing’s description (1843: 437) A .
ramulosa is much branched, with elongate, filiform and slender branchlcts, and with
minute spines. Kutzing adds that i t differs from A. delilei only in being more slender.
Specimens of A. ramubsa found in Hb. Binder and in C P show however that
A. ramulosa differs considerably from A. delilei, both in distribution of spines and in
position of stichidia. Nor is A. ramulosa particularly slender when soaked out.
Finally a comparison with A. dendroides was made. A. ramulosa has, on some of the
older main axes, side branches strongly reminiscent of those seen in Chondria. They are
Marine algae of Sao I’omC and Principe
137
ovoid or cylindrical, with a t most one or two spines, and their insertion on the branch is
pinched. I n A . dendroides the branchlets are normally of this form, and the stichidia also
resemble ordinary vegetative branchlets. Thus although A . dendroides also has branches
and branchlets not subtended by spines, i t seems to be a far less spiny plant and the
stichidia are o f a different form from those of A . ramulosa.
A . ramulosa is thus better considered as an independent species in the opinion of the
present writer. In vegetative structure Ribeiro’s plant agrees very well with the specimens
of A . ramulosa examined. In habit both are bushy, much branched plants, with welldeveloped branches (of unlimited growth) bearing numerous spines. The branchlets are
of two kinds. On the younger parts of the plant they resemble simple short branches and
obviously in due course elongate. On the older main axes both spines and short Chondriatype branchlets are present (see above). Neither branches nor branchlets are in the axils
of spines. The branches of unlimited growth predominate in determining the habit and
appearance of the plant.
While Ribeiro’s plant is tetrasporangial, the Angola plants are female, and apparently
somewhat older. They are larger, about 20 cm high, brown when soaked out rather than
red, and with rather less well-developedbranchlets. The similarity of texture when soaked
out is noticeable, none of the plants being so hard as seems usual among many other
species of the genus.
Chamberlain (Isaac & Chamberlain, 1958: 151) refers to a plant from Portuguese East
Africa which has subglobose cystocarps, but from the description it seems very unlikely
that this plant could be placed in A . ramulosa. As shown in Text-fig. 3, A . ramulosa does
not, in any case, have conspicuously and constantly subglobose cystocarps.
A . ramulosa is a new record for S. TomB, and the species is apparently confined to the
Gulf of Guinea.
Text-fig. 3. Acnnthopitorn rcimzdosa (Lindenb.) Kutz. emend. Steentoft. Range of form of
cystocarps.
Type Acunthophora ramulosa (Lindenb. ms. ex Hb. Binder) Kiitz., Kiiste von Angola
(Kiitzing, 1843: 437) (L).There are isotypes in HBG and C, the latter sub. nom. A . delihi
and A . muscoides.
The specimens in Hb. Binder (HBG) as well as the type, are labelled ‘Angola, 1839’ in
Binder’s handwriting, and appear to be part of the material collected by Epffenhausen
in that year.
In P1.2, fig. 2 the isotype from Hb. Binder is shown.
~ O A R E STEENTOFT,
T
F.L.S.
138
Exsiccata s.
TOM&:
Ribeiro 12, Ropa Praia da Nazareth, 1894.
Distribution Africa: Angola, S. Tom&.Apparently endemic to the Gulf of Guinea.
Acanthophora spicifera (Vahl) B s g . , Bot. Tidsskr., 30: 201 (1910); Damk Bot. Ark.,
3 : 259 (1915-20).? A .orientalis J. Ag., Spec. alg., 2 (3):816 (1863).As A . muscoides Grev.
in Henriques (1886:221) and p.p., in Hariot (1908:163)and Henriques (1917:166);as
A . thierrii Lamour. in Barton (1897:373)and Pilger (1911: 303).
The plants here considered are very different from A . muscoides. Not only do they
have very few spines on the main axes (and then only on the oldest parts), but the short
spiny lateral branches are not subtended by spines. The main axes entirely dominate the
habit of the plant as stated by Falkenberg (1901: 231).
Moller’s plants have tetrasporangia, and these are borne in the various ways described
by Bsrgesen (1915-20: 259). One of the commonest ways is in small spineless dome-shaped
lateral branches on the stichidia, while the terminal portion of the stichidium can also be
so formed. This is said to be a distinguishing feature of A . orientalis.
Both Weber v. Bosse (1913-28: 347)and Bsrgesen (1945:61)have pointed out that the
distinction between A . spicifera and A . orientalis is a doubtful one. Papenfuss (1943: 92)
reports A . orientalis for South Africa and this record is included in the distribution given
below.
A. spicifera is a new record for both S. Tom6 and Principe. The SBn&galrecord is based
on Jardin’s specimen (Jardin, 1850-51 : 10).
Exsiccata
9. TOMI%:Fox Nielsen 234, sin. loc., August 1956 (duplicate in BM spirit
collection, also =Thorold 612, sub. nom. A . muscoides,inBM spirit collection).
Moller 8, sin. loc., 1885.Moller 12, Bahia de Anna Chaves, 8-1885.
PRfNCrPE: Newton s.n., Praia de Sundy, Agosto de 1887.
Distribution Africa : ?South Africa, Anogla, S. Tom6, Principe, Cameroun, Sierra
Leone, SBn6gal. More or less pantropical.
Bostrychia Montagne 1838
Bostrychia binderi Harv., Ner. Austral. : 68 and pl. 28 (1847-49); Post, Int. rev. ges.
Hydrobiol., 48 (1): 76 (1963);as B . tenella in Dickinson & Foote (1950:272).
This is a new record for S. Tom& De Toni (1903:1167)suggests that B . pilifera Kiitz.
from Senegambia, is a form of B . binderi, but earlier (p. 1163)he has already suggested
that it was a form of B . tenella, as does Dr Post (in litt.).
Exsiccata s. TO&: Fox Nielsen 249, sin. loc., August 1956.
Distribution Africa : South Africa, Angola, S. Tom&,Cameroun, Ghana, GuinBe, Sierra
Leone. Apparently tropical and subtropical Atlantic and Pacific Oceans.
Bostrychia radicans Mont. in Sagra, Hist. Cuba: 419 (1842);Post, Int. rev. ges. Hydrobiol.,48(1):58(1963)and
Rev.algol.,4:270(1963);asB.simpliciusculuinFox(1957:626).
The plants are very much mixed with Murrayellu periclados, Symplocu atlantica and
Hydrocoleum lyngbyaceum.
Welwitsch 59 from Angola is labelled as a mixture of B. radicans and B . mixta, but
published (Barton, 1897:373)as B. tenella.
Exsiccata s. TO&: Fox Nieken 231,232,233, sin. loc. August 1956 ( =T?mrold 609,611
in BM spirit collection, sub. nom. B . moritziana).
Distribution Africa : Portuguese East Africa, South Africa, RQpublique du Congo,
S. Tom&,Gabon, Cameroun, Nigeria, Ghana, Guide, Sierra Leone. Probably pantropical.
Marine algae of S h Tom6 and Principe
139
Bryothamnion Kutz. 1843
Bryothamnion seaforthii (Turn.) Kiitz., Phyc. gen.: 433 and pl. 52, fig. 11 (1843);
Bergesen, DanskBot. Ark., 3: 284 (1915-20); Henriques (1886: 221; 1917: 166); Hariot
(1908: 163).
According to De Toni (1903 : 975) this species was first collected on Ilhdu das Rolas by
Griife (Greeffe). Taylor (1960: 587) considers that the species is probably a deep-water
one, and i t has been found among an unnumbered sublittoral collection of Quintas’s.
Moller’s and Ribeiro’s plants were probably also sublittoral.
The apparent rarity of this species in the Gulf of Guinea may arise from its intolerance
of low salinities, but i t must be remembered that very few collections have so far been
made in the sublittoral zone.
There are plants of this species, in Hb. Binder, from Angola.
TOMB : Moller 14, sin. loc., 1885. Ribeiro 13, Ropa Praia da Nazareth, 1894.
Distribution Africa : Angola, S. Tom& ?Pantropical.
Exsiccuta
S.
Bryothamnion triquetrum (Gmel.) Howe, Jl N . Y . Bot. gdn., 16: 222 (1915) ;Brargesen,
Dansk Bot. Ark. : 3 : 282 (1915-20) ; Taylor, Mar. alg. . . . Americas: 587 and pl. 72, fig. 6,
pl. 73, fig. 4 (1960), as B. triangulare in Henriques (1886: 221; 1917: 166); Hariot (1908:
163), as Alsidium triangulara in Barker Webb (1849: 197).
This species was first collected on Ilhdu das Rolas by Grafe (see above). I n addition to
Quintas’s plants, Carpine (1959 : 77) has also found this species in 11 m of water.
Material from Angola has been found in Hb. Binder, though the species is not mentioned
in Barton (1897).
Emiccuta s. TOMB : Quintas [27], I l h h das Rolas, 1885. [ ?Quintas]s.n., Ilhdu da Rolas
[1885].
Distribution Africa : Angola, S. Tomd, Cabo Verde Is. ?Pantropical.
Chondria C . Ag. 1817
Chondria sp.
Reported by Carpine (1959: 78) in 11 m of water off S. Tomd.
Digenea C . Ag. 1823
Digenea simplex (Wulf.) C. Ag., Spec. ah.: 389 (1821); Bergesen, Dansk Bot. Ark., 3:
281 and 469 (1915-20); Taylor, Mar. alg. Florida: 174 and pl. 24, fig. 20, pl. 33, fig. 7
(1928);Hariot (1908: 163); Henriques (1886: 221; 1917: 166).
The plants are all much branched, the sublittoral ones up t o 7 cm high, the others
shorter. These latter are probably from an intertidal habitat if the remarks of Bsrgesen
(1915-20: 281) and Taylor (1960: 689) apply. Among the epiphytes present is Centroceras
clavulatum.
D. simplex seems t o be rather uncommon in the Gulf of Guinea, and certainly of sporadic
occurrence.
Exsiccuta s. TO^ : Quintas 18, Ilhdu das Rolas, 1885. Quintas 19, I l h h das Rolas, 1885.
Quintas s.n., Equador, Canal das Rolas. Alga do fundo. 1885. Quintas s.n.,
Equador, Canal das Rolas. Alga do fundo (calchreo) c6r carminada. 1885.
Distribution Africa: Portuguese East Africa, S. Tom&,Ghana, Cabo Verde Is. Pantropical.
140
MARGARETSTEENTOFT,
F.L.S.
Laurencia Lamour. 1813
.
Laurencia intricata Lamour., Essai . . Thalass.: 43 (1813); Taylor, Mar. alg. . . .
Americas: 626 (1960); Mar. alg. Florida: 179 and pl. 34, fig. 8 (1928).
This species was found mixed with Quintas’s material of Bryothumnion tripuetrum.
According to Yamada (1931: 225 and pl. 17b) L.intricata is a variety of L . obtusa.
Exsiccata s. TOMB: Quintas 27, Ilh6u das Rolas, 1885. Quintas s.n., Ilh6u das Rolas,
1885.
Distribution Africa: 8 . Tom& Warm western Atlantic from Bermuda and Florida to
Panama, Pacific Mexico.
Laurencia obtusa (Huds.) Lamour., Essai . . . Thulass.: 42 (1813); Okamura, Icon.
Jap. Alg.,4:175andpl. 193(1923);Henriques(1886:221;1917:166);Hariot(1908:163).
Two varieties are present among the S. Tome material available. Quintas’s plants most
resemble Mildbraed 6616 (in Hb. Bsrgesen) from Annobon, and these seem t o agree most
closely with the description of var. rigidula Grunow (18734: 45). Cardoso’s plant,
together with two found in Hb. Hamburg, more resembles L . obtusa as described and
figured by Taylor (1928: 180 and pl. 33, fig. 3), though in parts L. natalensis Kylin (i.e.
L . obtusa var. natalensis (Kylin) Bsrgesen, 1945: 59) is called to mind.
The specimen of L . brachyclados Pilger (Mildbraed 6719 from Annobon) in H b Bsrgesen
would seem to represent the basal parts of a young plant of var. rigidula, but it is so
minute, and a t such variance with the published description (Pilger, 1920: 6) that i t would
seem better.to leave i t alone for the time being.
Carpine (1959: 77, 80, 87, 88) also reports L. obtusa in both S. Tom6 and Principe.
Exsiccata s. TOM^: Cardoso s.n., sin. loc. [?Epffenhausen] St Thorn., 1839 (in Hb.
Binder, HBG). Quintas 7, Ilh6u das Rolas, 1885. Quintas s.n., Equador,
Canal das Rolas. Alga da superficie banhada pela enchente colhida sobre
zoophytos. 1885.
Distribution (of the species as a whole) Africa: ?South Africa, Annobon, S. Tome,
Principe, Cameroun, Sendgal, Cab0 Verde Is., Mauritanie, Canary Is., Maroc, Madeira,
Tanger, Agores. I n all but the coldest waters.
Laurencia perforata (Bory) Mont. in Barker Webb & Berthelot, Hist. nat. Iles Canaries,
3 (2): 155 (1840); Yamada, Univ. Calif. Pub18 Bot., 16 (7): 193 and pl. 3 b (1931); Hariot
(1908: 163); Henriques (1917: 166); Carpine (1959: 83).
This species was first reported by Hariot from Gravier’s collection, but has since been
found by the present writer among several other collections. All these plants show considerable resemblance to Bsrgesen’s from the Canary Islands.
I n 1957, when the Coimbra material was first examined, a n unidentified Quintas-specimen of this species was found, but i t has not been seen again in the present examination.
From Pilger’s description of his species L. brachyclados from Annobon, one would be
inclined to assume that the latter was but a dwarf, or a t least immature form, of L.
perforata. An isotype of L . perjorata in Hb. Agardh (no. 36616) has been examined. These
plants are 2 to 3 cm high, blackened and totally non-adherent to paper, just as Pilger
describes L. brachyclados. Yamada (1931: 216) has examined the holotype of the latter
species however, and maintains that it is quite different from L. perforata.
However, an isotype of L. brachyclados (Mildbraed 6719) in Hb. Bsrgesen shows a
totally different type of plant, more slender, less branched, of different habit, bleached
out and quite adherent to the mica on which i t is mounted. Even discounting the effects
of bleaching and pressure, i t is hard to reconcile this specimen with Pilger’s description
and there is a t least the possibility that some confusion has arisen.
Marine algae of S6o TontC and Principe
141
Two new records for L . perforata outside S. Tomi. have been found. One is for Gabon,
leg. Chollon (PC), the other is for Gambia, sin. leg. 1837 (in Hb. Binder, HBG). This is
probably one of Epffenhausen’s plants.
Exsiccuta s. TOMP:Gravier s.n., Ribeira Peixe, 12 avril 1906. MarBe basse. (PC). Fox
Nielsen 250, sin. loc., August 1956. Moller 8, sin. loc., 1885 [with Acanthophora spicifera]. Quintas 11, 1lhi.u das Rolas, 1885 [with Hypnea spinella].
Quintas 12, Equador, Canal das Rolas. Alga da superficie (calchrio) banhada
pela enchente, c6r verde. 1885.
Distribution Africa : Gabon, ?Annobon, S. Tom& Gambia, Canary Is., Madeira. Indian
and Pacific Oceans.
Laurencia pinnatifida (Gmel.) Lamour., Essai . . . Thalass.: 42 (1813); Henriques
(1885: 134); Carpine (1959: 77).
Newton’s plant from 1881 is untraced, but Carpine has collected this species in 11 m of
water. The South African record rests on Barton (1893 : 174).
Exsiccata 5. Tom6 : non. vid.
Distribution Africa : South Africa, Annobon, S. Tomi., SQnBgal,Mauritanie, Canary Is.,
Maroc, Madeira, Tanger, Aqores. Mediterranean and most eastern Atlantic coasts south
to SBndgal, an ‘uncertain’ record for the western Atlantic (Taylor, 1960: 628))Malaysia,
western Pacific.
Laurencia poitei (Lamour.) Howe, Bull. Torrey Bot. Club, 32: 583 (1905); Bsrgesen,
Dansk Bot. Ark., 3: 245 (1915-20) ; Taylor, Mar. alg. Florida: 181 and pl. 34, figs 4 and 10.
(1928); Yamada, Univ. Calif. Publ. Bot., 16 (7): 219 andpl. 15a (1931). As L . tuberculosa
in Hariot (1908: 163) and Henriques (1917: 166). ? as Gracilaria poitei in Henriques
(1886: 221; 1917: 166).
Laurencia tuberculosa was reduced to the synonymy of L . poitei by Yamada (1931 :
219). Cfracilariapoitei Lamour. is the basionym, but it is no longer clear to which of the
S. Tom6 plants Henriques (1886 : 221) is referring. It must have been a Moller or Quintas
plant, though it could not be considered a correct identification for any of their Laurencia
specieR seen in the present revision.
Exsiccata s. TOMP: Ribeiro 15 [sub. norn. Laurencia tuberculosa J. -4gh.1, Roqa Praia da
Nazareth, 1894.
Distribution Africa: S. Tomi.. Otherwise on the warmer Atlantic coasts of America,
Malaysia.
Murrayclla Sclimitz, 1893
Murrayella periclados (C. Ag.) Schmitz, Ber. dt. bot. Ges., 11: 227 (1893); Bsrgesen,
Dansk Bot. Ark., 3: 314 (1915-20); Post, Int. rev. Ges. Hydrobiol, 48 (1): 142 (1963).
The material was determined by Dr Post. The record is new for S. Tom&
Exsiccuta s. TO&: Fox Nielsen 231,232,233, sin. loc., August 1956 (=Thorold 609 and
611 in BM spirit collection). Plants sterile, mixed with Bostrychia radicans
and Bachelotia antillarum, Hydrocoleum lyngbyaceum etc.
Distribution Africa : Portuguese East Africa, South Africa, RBpublique du Congo, S.
TomB, Cameroun, Ghana, GuinBe, Sierra Leone. Pantropical and subtropical.
142
MARGARET
STEENTOFT,
F.L.S.
Ricardia Derb. & Sol. 1856
Ricardia montagnei Derb. et Sol., Annls Sci. nat., Bot., s6r. 4, 5 : 209 and pl. 14, figa 1
to 7 (1856); Bsrgesen, K. damke Vidensk. Selsk., Biol.,9 (1):74 (1930).
Reported by Carpine (1959: 78) upon Laurencia obtusa, dredged in 11 m of water.
Cyanophyceae
Oscillatoriaceae
Hydrocoleum Kiitz. 1843
Hydrocoleum lyngbyaceum Kiitz., Spec. alg. : 259 (1849); Sampaio, Est. Ens. Doc.,
Ministr. Ultramar, 47: 30 (1958);Publ. Inst. Bot. Gonculo Sarnpaio, 2 s&.,48: 24 (1962);
Est. cient. . . . Carington da Costa: 246 (1962).
Both var, typim, with firm sheaths to the trichomes, and var. rupestre, with diffluent
sheaths, were first reported for S. Tom6 by Sampaio (1958:30). Thorold’s specimen is var.
typim.
Exsiccuta
Fox Nielsen 231, sin. loc., August 1956 (=Thorold 609 in BM spirit
collection) (mixed with Syrnploca atlantica, Murrayella periclados etc.).
9. TOM$:
Distribution Africa : Gabon, S. Tomb, Ghana, Canary Is., Maroc, Tanger. Cosmopolitan.
Lyngbya C . Ag. 1824
Lyngbya majuscula (Dillw.) Harv., Phyc. brit., pl. 62 (1846-51).
Found growing with Centroceras clavulatum in ‘turf’. This is a new record for S. Tom&
Exs:siccata s. TOM$: Fox Nielsen 226, sin. loc., August 1956.
Distribution Africa : Gabon, S. Tomb, ?Cameroun,Canary Is., Maroc, Tanger, Acores.
Cosmopolitan.
Symploca Kiitz. 1843
Symploca atlantlca Gom., Monog. Oscill.: 129 (1892).
Exsiccuta
S. TOMI%:
Fox Nielsen 231, sin. loo., August 1956 ( =Titorold 609 in BM spirit
collection) (mixed with Hydrocoleum lyngbyaceum, Murrayelb pericladoa
etc.)
Distribution Africa : S . Tom6, Nigeria, ?Ghana.
ACKNOWLEDGEMENTS
The author wishes to thank the following, who have most kindly made identifications
in the general indicated: Dr C. Bliding (Enteromorpha),Dr J. Soderstrom (Chdophora),
Dr E. Post (Bostychiuand Murrayelb),Dr F. Drouet (three genera of Cyanophycerte).
Grateful thanks are also due to the Directors of the Staatsinstitut fiir allegemeine
Botanik, Hamburg, the British Museum (Natural History), the Botanical Museum and
Herbarium, Lund, the Laboratoire de Cryptogamie of the Museum National d’Histoire
naturelle, Paris, and the Mdaeum of the Royal Norwegian Society for Science and Letters,
Trondheim.
In particular the author wishes to thank Prof. D.phi1. Morten Lange and the Danish
State Science Fund Foundation for the financial support which has made this revision
possible.
Marine algue of S6b Tmn6 and Priiicipe
143
REFERENCES
The main sources of information on the occurrence of species on the Atlantic coast of Africa (South
African to Tanger) are to be found in Fox (1957). I n addition Portuguese East Africa is included below,
together with some additional sources. Otherwise only references from the foregoing text are mentioned.
ANON, 1958 (reprinted 1964). Tables of temperature, relative humidity and precipitation for the world,
4, pp. xii+220, 2 maps. M.O. 617 d. H.M.S.O., London.
ADMIRALTY,
1964. Tide tables, 2, 1965. Atlantic and Indian Ocean. Pp. x x x + 4 3 1 . London.
AYKENASY,
E., 1888. Forschungareise 'Gazelle', 4, Bot., pp. 58, pls. 12.
AYICENASY,
E., 1896. Enumeration des algues des fles du Cap Vert. Bol. SOC.Broteriu, 13: 150-175.
BARKER
WEBB,P., 1849. Spicilegia gorgonea in W. J. Hooker (ed.), Niger Flora, 196-197.
BARTON,
E. S., 1893, 1896. A proviional list of the marine algae of the Cape of Good Hope. J. Bot..
Lond., 31: 53-56, 81-84, 110-114, 138-144, 171-177, 202-210 (1893); 34: 193-198, 458-461 (1896),
BARTON,E. S., 1897. Welwitsch's African marine algae. J. Bot., Lond., 35: 368-374.
BERRIT,G. R., 1961, 1962. Contribution la connaissance des variations saisonnibres dans le golfe de
GuinBe. Cahiers ocianogr., 13: 715-727 (1961); 14: 718-729 (1962).
RCIHGESEN,
F., 1913, 1915-20. The marine algae of the Danish West Indies. 1. Chlorophyceae, Dansk
Bot. Ark., 1 ( 4 ) : pp. 158+2, map (1913); 3. Rhodophyceae, Dansk Bot. Ark., 3 ( 6 annual parts):
pp. 504 (1915-20).
BORCESEN,
F., 1927. Marine algae from the Canary Islands. 3. Rhodophyceae 1, K . danske Vidensk.
Sebk., Biol., 6 ( 6 ) :pp. 97. (See also FrBmy, P., 1936, and Lemoine, Mme. P., 1929.)
BORGESEN,
F., 1942, 1943, 1945. Some marine algae from Mauritius. 3. Rhodophyceae 1, K . danake
Vidensk. Sebk., Biol., 17 ( 5 ) : pp. 64, pls. 2 (1942); 3. Rhodophyceae 2, K . danske Vidensk. Selak.,
Biol., 19 ( 1 ) :pp. 85, pl. 1 (1943); 3. Rhodophyceae 4, K. danske Vidensk. Selak., Biol., 19 (10):pp. 68
(1945).
CARPINE,
C., 1959. Aperpu sur les peuplements littoraux. RBsultats scientifiques des Campagnes de la
Calypso 10. Annls Inst. ocianoyr., Monaco, 37: 75-89.
CHEVALIER,
A., 1920. Exploration botanique de l'Afrique Occidentale Franpzise, 1, algues (det. P. Hariot),
785-789.
CHEVALIER,
A., 1935. Les lles du Cap Vert. Flore de 1'Archipel. Revue Bot. appl. A r k . Trop. 15, algues
(det. J. Feldmann): 1069-1071.
CHOW,R., 1945, 1947. Pacific species of Galaxaura. 1. Asexual types. Pap. Mich. Acad. Arts. 30 (1):
35-56, pls. 11 (1945); 2. Sexual types. Pap. Mich. Acad. Arts. 31 (1): 3-24, pls. 13 (1947).
COTELONEIVA,J. M., 1958a, b. Contribuipb o estudo geolbgico e geomorfol6gico da Ilha de SBo Tomb
e dos Ilhbus das Rolas e das Cabras. Be Conf. Int. Africaniet. occident., S6o T m L , 1 9 5 6 , 2 : 147-153 +
map (1958a); ContribuiqIo para a geologia e geomorfologia da Ilha do Principe. 6e Conf. Znt.
Africankt. occident., Sdo Tomb, 1956, 2 : 167-162+map (1958b).
DANCEARD,
P., 1949. Les algues marines de la cBte occidentale du Maroc. Botaniate, 34: 89-189.
DANGEARD,
P., 1952. Algues de la presqui'ile du Cap Vert (Dakar)e t 88s environs. Botaniste, 36:
195-329, pls. 14-21.
DE TONI,G. B., 1889, 1895, 1903. Sylloge algarum..
1.
Chlorophycearum 1, 2 (1889); 3.
Fucoidearum (1895); 4. . Floridearum 3 (1903).
DICKIE,G., 1873. On the marine algae of St. Helena. J . Linn. Soc. (Bot.), 13: 178-182.
DICKIE,G., 1873-5. Enumeration of the algae collected at the Cape Verde Is.,
.,J . Linn. Soc. (Bot.),
14: 344-349.
DICKINSON,
C. I., 1952, 1953. Marine algae from the Gold Coast. 3. Kew Bull., 1951: 293-297 (1952);
4. Kew Bull., 1952: 41-43 (1953).
DICKINSON,
C. I. & FOOTE,
V. J., 1951, 1952. Marine algae from the Gold Coast. 1. Kew Bull., 1950:
267-272 (1951); 2. Kew Bull., 1951: 133-137 (1952).
EGEROD,L. E., 1952. An analysis of the siphonous Chlorophycophyta. Univ. Calif. Publ. Bot., 25 ( 5 ) :
325-454, PIS. 29-42.
ENCLER,
A., 1895. Die PfEanzenwelt Oat Afrikas ., 5, theil C.
EXELL,A. W., 1944. Catalogue of the vascular plants of S . Tomb (with Principe and Annobon). Algae
(det. G. Tandy), 386.
P.. 1901. Die Rhodomelaceen
.,Fauna und Flora dea Bolfea won Neapel Monogr., 25:
FALKENBERG,
pp. 754, pls. 24.
FELDMANN,
J., 1931. Rbmarques sur les genres Gelidium etc. Recueil des trav. cryptog.
Mangin
pp. 151-166.
FELDMA",J., 1938. Sur la &partition de Diplanthem uwightii Aschers. sur la cBte occidentale d'Africa.
Bull. SOC.Hist. nat. Afr. Nord., 29 (1-2): 107-112.
FOSLIE,
M., 1897. On some Lithothamnk. K. norske Vidensk. Selak. Skr.. 1897 (1): 1-20.
FOSLIE,M., 1900a. New or critical calcareous algae. K. norake Videmk. SeLk. Skr., 1899 ( 5 ) : 1-34.
FOSLIE,M., 1900b. Five new calcareous algae. K. nor8ke Vidensk. Selek. Skr.,1900 (3): 1-6.
...
.. .. .
..
..
..
..
...
144
MARQARETSTEENTOFT,
F.L.S.
FOSLIE,
M., 19OOc. Revised systematical survey of the Melobesieae. K . norske Vidensk. Selsk. Skr.,
1900 (5): 1-22.
FOSLIE,M., 1907. Alg. not. 3. K. nwake Videnak. Selak. Skr., 1906 (8): 1-34.
FOSLIE,M., 1909. Alg. not. 6. K. nwake Videnak. Selak. Skr., 1909 (2): 1-63.
FOSLIE,
M., 1929. Contributions to a monograph ofthe Lithothamnia. ed. H. Printz, pp. 60, pls. 75.
Fox, M., 1957. A first list of marine algae from Nigeria. J . Linn. SOC.
(Bot.),55 (no, 362): 615-631.
pl. 54.
P. A., 1934.Pugillus algarumMaroccanum. Bull. SOC.Sci. nut. Maroc., 14 : 4-8.
F R ~ M P.
Y ,& BUROLLET,
FR~MY
P.,, 1936. Cyanophyceaein Bsrgesen, F., Marine algae from the Canary Is., K . danske Videnak,
Selak., Biol., 12 (5): 43.
GAYRAL,
P.. 1958. Algues de In cBte athntiqw marocaine, pp. 523, pls. 152.
GERLOFF,J., 1959. Bachebtia (Bornet) Kuckuck ex Hamel oder Backlotia (Bornet) Fox? Nov.
Hedwigia, 1 (1): 37-39.
GRAVIER,C., 1906. Sur les formations coralliennee de 1% San-Thorn6 (golfe de Guinbe). C.r. hebd.
Seanc. Acad. Sci., Paris, 143 (23): 929-931.
GRAVIER,
C., 1907. Rapport sur me mission scientiflque a l’ile de San ThomB. Nouv. A r c h Missiona
scient. litt. etc., 15 (2): 5-70.
GRUNOW,A., 1873-4. Algen der Fidschi- Tonga- und Samoa-Inseln. J . Mus. Oodefroy,3: 23-50.
HAMEL,G., 1931. Chorophycbe deaet)tesfrawaiaes, 1: XV+168+pls. 1; 2: pp. 34+pls. 1.
HAMEL,Q., 1931-9.Phdophyc6ea de France. Pp. 80+XVII+81-32., pls. 10.
HARIOT,P., 1895. Lista des algues receuillies au Congo par M. H. Lecomte. J . Bot., Pa&, 9: 242-244.
HARIOT,
P., 1908. Les algues de St. Thorn&J . Bot.,Parie, 2 dr., 1: 161-164.
HAUCE,F., 1886. Die Meereealgen Deutachlaida und Oeateweich. Pp. VIII+575, pls. 5.
HAUCE,F., 1888. Meeresalgen von Puerto-Rico. Bot. Jb., 9: 467-470.
HENRIQUES,
J., 1885. Contribui@opara o estudo d8 flora d’algumaa poeaessoes portuguezaa 1. Plantas
colhidas por F. Newton, na Africa occidental. Bol. SOC.Broteria, 3, algae: 131-134.
HENRIQUES,
J., 1886. Contribuiqoea para o estudo da Flora d’Africa. Flora de 8. Thorn& Bol. SOC.
Broteria, 4, algae: 217-222.
HENRIQUEB,
J., 1917. Catalogo das especies de anhais e plantas ate hoje encontrads no Ilha de 5.
Thom6. BoZ. SOC.Broteria, 27, algae: 165-166.
HOECE,C. Van den, 1963. Rezrision of the European apeciea of Cladophora. Pp. vii + 248, pls. 55.
HOW, M. A., 1917. A note on the structural dimorphism of sexual and tetrasporic plants of Qakzxaura
obtmata. BUZZ.TOW~Ybot. Club, 43: 621-624.
HOWE,M. A., 1918. Further notes on the structural dimorphism of sexual and tetrasporic plants in the
genus Qalmxura. Mem. Brooklyn bot. adn, 1: 191-196, pls. 3-4.
ISAAC,
W. E., 1956. Marine algae of Inhaca Island and of the Inhace Peninsula, 1. S. Afr. J . Bot., 22
(4): 181-193, pls. 38-43.
ISAAC,
W. E., 1957. Some marine algae from Xai-Xai. 8.Afr. J . Bot., 23 (3): 75-102, pls. 28-33.
ISAAC,
W. E. & CHAMBERWIN. Y.M., 1968. Marine algae of Inhaca Island and of the Inhaca Peninsula,
2. S. Afr.J. Bot., 24 (3): 123-158, pis. 3-6.
JARDIN,
E., 1860-1. Herborisations
Cdte occidentale d’Afrique 1845-8. Nouv. annla Mar. et Colon.,
July, 1850: 3-4; May, 1851: 9-10.
KJELLMAN,
F. R., 1900. Om Florid6-slagtet Q a k a u r a , dew organografi och systematik. K. Suenaka
VelenakaAcad.hundl., 33 (1): 1-99, pls. 20.
KYLIN,H., 1938. Verzeichnis einiger Rhodophyceen von Sud Afrika. Acta Univ. Lund. N.F. Avd. 2,
34 (8): pp. 26, pls. 8.
F. T., 1843. Phycologia generalis. Pp. xxxii+459, pls. 79.
KUTZINQ,
KUTZINQ,
F. T., 1849. Species abarum. Pp. vi+922.
KUTZINO,
F. T., 1861, 1865, 1869. Tabulaephycobgicicae,11: pp. 32, pls. 100 (1861); 15: pp. 36, pls. 100
(1865); 19: pp. 36, pls. 100 (1869).
H., 1958. Esboco da carta de aptideo agricola de SEo Tom6 e Principe. Garcia de Orto,
LAINSE SILVA,
6 (1): 61-86, pls. 1-4, maps.
LANJOUW,
J. & STAFLEU,
F. A., 1964. Indez Herbarioruna, Part 1. Utrecht.
LAWSON,
G. W., 1954. Seaweeds from Sierra Leone. JZ W. Afr. Sci. Ass., 1: 63-67.
LAWSON,
G. W., 1955. Rocky shore zonation in the British Cameroons. J1 W. Afr. Sci. Ass., 1: 78-88,
pl. 7.
LAWSON,
G. W., 1957. Some features of intertidal ecology of Sierra Leone. JZ W. Afr. Sci. Ass., 3,
166-1 74.
LAWSON,
G. W., 1960. The Caulerpw of West Africa. Nigerian Field, 25 (1): 23-31.
LEMOINE,
Mme. P., 1911. Structure anatomique des MBlobbsiBes. Annk Inat. ocdanogr., Monaco, 2
(2): pp. 213 inc. 3 tables, pls. 6.
Mme. P., 1929, Corallinweae: Melobesieae in Bargesen, F., Marine algae from the Canary
LEMOINE,
Islands. K. dunake Vidensk.Selsk., Biol., 8 (1): 19-62, pls. 4.
LINDAUER,
V. W. & CHAPMAX,V. J., 1961. The marine algae of New Zealand 11.Nov. Hedwigia, 3
(2 + 3) : 129-350, PlS. 57-97.
.. .
J . Linn. SOC. (Bot.) Vol. 60, N o . 382
MARGARET STEENTOFT, F.L.S.
Plate 1
(Fncing p . 144)
J . Linn. SOC.(Bot.) Vol. 80, No. 382
MARGARET STEENTOFT, F.L.S.
Plate 2
Marine algae of SCo Tom&and P h c i p e
145
MARTENS,G. yon, 1866. Die Tango in Die Preussische Expedition nach Ost Asien, Bot. theil, pp. 152,
pls. 8.
MONOD,T., 1960. Note botanique sur les iles de Siio Tomb et de Principe. Bull. In&. j r . Ajr. noire, s6r.
A, 22: 19-83, PIS. 11.
MURRAY, G., 1888-9. Catalogue of the marine algae of the West Indian region. J. Bot., Land., 26:
193-196 (1888); 27:237-242,257-262,289-305 (1889).
NYLANDER,
W.,1888. ContribuipGes para o estudo da flora da costa occidental d’Africa. Lichenes
nonnulli. Hol. Soc. Broteria, 5: 221-224.
G. F., 1940. Notes on South African marine algae 1. Bat. not. 1940: 200-226.
PAPENFUSS,
PAPENFUSS,
G. F., 1943, 1951. Notes on South African marine algae 2. S. Afr. J . Bot., 9 ( 3 ) : 79-92
(1943); 3 . S . A j r . J . Bat., 17 ( 4 ) : 167-188 (1951).
PIc(.oNE, A., 1884. Alghe in Crociera del Corsclro ulle Isole Marlera e C‘anarie del Capitano Enrico
d’Albertis, pp. 60, pl. 1.
PILGF:R,
R., 1911. Die Meeresalgen von Kamerun. Bat. Jb., 46:294-323.
PILGER,
R., 1919. Ueber Corallinaceae von Annobon. Bot. Jb., 5 5 : 401-435.
l ’ i i i ; ~ R.,
~ , 1920. Algae Mildbraedianae Annobonensis. Bot. Jb., 57:1-14.
POST,E.,1963a. Bostrychia radicans im Siisswasser Westafrikas. Rev. algol., 4: 270-281.
r o w , E.,1963 b. Zur Verbreitung und C)kologie der Bostrychia-Caloglossa Assoziation. Znt. rev. Ges.
Hydrobiol., 48 ( 1 ) : 47-152.
RODRIOL~ES,
J. E. de M., 1960. HevisLo das algas de S. Tom6 e Principe de herbhrio do Instituto
Botinico de Coimbra 1. Phaeophyta. Garcia de Orto, 8 ( 3 ) : 583-595, pls. 6.
SAMPAIO,
J., 1958. Cian6fitas de S. Tom6 e Principe. Estudos Ensaios Docum. .7ta Invest. Ultramar,
47:pp. 80, pls. 10.
SAMPAIO,
J., 1962a. Cian6fitas de S. Tom6 e Prlncipe ( 2 : s6r.). Estudos Gent. . C‘arington du
Costa: 243-259.
SAMPAIO,
J., 19626. Cian6fit,asde S. Tom6 e Principe, 1 and 2. Publea Inst. Bat. ‘Or. Qoncalo Sampaio’
(2a Bplr.), 48 : pp. 1-74, pls. 10; 49 : 243-259, pls. 3.
SAMPAIO,
J., 1963. Cian6fitas de S. Tom6 e Principh ( 3 : ser.). Estudos Ensaios Docum. Jta Invest.
~iltramar,108:pp. 52, pls. 6.
SCHMIDT,
0 . C . , 1929. Die marine Vegetation der Azoren. Nov. Hedwigia, 68:327-346.
SCHMIDT,
0. C., 1930. Beitriige zur Kenntnis der Meeresalgen der Azoren 1 and 2. Nov. Hedwigia, 69:
95-113, 165-172.
SCHMIDT,
0. C., 1931. Die marine Vegetation der Azoren. Bib. bot., 102:pp. viii+ 116, pls. 10.
SCHRAMM,
P. E., 1950. Deutschland um! Ubersee.
SODERSTROM,
J., 1963. Studies in C’kidophora.Bat. Qothoburg., 1 : pp. 47 +map.
SOURIE,
R., 1954. Contribution a 1’6tude Bcologique des cdtes rocheuses de SBnBgal. Mdm. Inst. f r . Afr.
noire, 38:pp. 342, pls. 23.
STEENTOFT
NIELSEN,M., 1959. Common seaweeds at Lagos 1. Nigerian Field, 23 (1): 34-44.
STEPHENSON,
T. A., 1948. The constitution of the intertidal fauna and flora of South Africa 3. Ann.
Natal Mus., 11 ( 2 ) : 207-324, pls. 15, 16.
SVEDELIUS,
N., 1945. Critical studies on some species of Galaxaura from Ceylon. Ark. Bot., 32A ( 6 ) :
pp. 74, PIS. 9.
SVEDELIUS,
N.,1953. Critical studies on some species of Gakuxaura from Hawaii. Nova Acta R . Sac.
Scient. upsal., ser. 4, 15 ( 9 ) :pp. 91.
TAYLOR,
W. R., 1928. The marine algae of Florida.. ., C’arnegie Inst. Wash. Publ., 379:pp. v+219
PIS. 37.
TAYLOR,
W. R., 1937. Marine algae of the northeastern coast of North America. UnSv. Michigan Stwl.
wient. Ser., 13 : pp. vii + 427, pls. 60.
TAYLOR,
W. R., 1960. Marine ukae of the eastern tropical and subtropical coasts of the Americas, pp.
xii + 870, pls. 80, photographs 14.
THORNTHWAITE,
C. W., 1948. An approach toward a rational classification of climate. Geog. Rev., 38
( 1 ) : 55-94, figs 13, map.
TRocHArN, J., 1940. Contribution ii 1’6tude de la vegetation du S6n6gal. Mdm. Inst. f r . Afr. noire, 2:
pp. 433, pls. 30.
WATTS,J. C. D., 1960. A summary of the meteorological and hydrological observations made in tho
Sierra Leone River Estuary area. Bull. Inst. j r . Afr. noire, ser. A. 22: 1159-1164.
VICKERS,
A., 1905. Liste des algues marines de la Barbade. Annla Sci. nut., g8 s6r., Bot., 1: 45-66.
WEBERvan BOSSE,A., 1913-28. LiHte des algues du Siboga 1-4. Siboga-Expeditie Monogr., 59a-d:
pp. 533, pls. 16.
YAMADA,
Y . , 1931. Notes on Laurencia, with special reference to the Japanese species. Univ. C?alif.
E’ztbls Bat., 16 ( 7 ) : 185-310, PIS. 20.
ZANARDINI,
G . . 1858. Plantae maris rtcbri, pp. 101, pls. 12.
. .
146
MARGARETSTBENTOFT,
F.L.S.
EXPLALNATLTION OB PLATES
PLATE 1
Ormilaria henriquesiam Hariot. Syn- and isotypes.
Fig. 1. Syntype, Ribeiro 14 (PC).
Fig. 2. Isotype, leg. Ribeiro (14P)(COI).
Fig. 3. Isotype, Ribeiro 14 (COI).
Fig. 4. As Fig. 3.
Fig. 5. Syn- (or iso-P)type, Moller 15 (COI).
Fig. 6 As Fig. 5.
Figs 2 and 3 are on the same scale as Fig. 4.
PLATE
2
Fig. 1. Gracilariahenriquesiana Hariot. Syntype (?partof Ribeiro 14, (PC).
Fig. 2. Acanthophma ramulosa (Lindenb.)Kiitz. emend. Steatoft. Isotype. x 2 approx. (Hb. Binder,
HBG).