Bulletin of the

British Museum (Natural History)

Seaweeds of the western coast of

tropical Africa and adjacent islands:

a critical assessment.

IV. Rhodophyta (Florideae)

2. Genera G

James H. Price, David M. John and
George W. Lawson

Botany series Vol 18 No 3 29 September 1988

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)

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ISBN 0 565 08021 0

ISSN 0068-2292 Botany series

VollSNo 3 pp 195-273
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 29 September 1988

Seaweeds of the western coast of tropical Africa and y
adjacent islands: a critical assessment.
IV. Rhodophyta (Florideae) 2. Genera G

James H. Price

Department of Botany, British Museum (Natural History), Cromwell Road, London
SW7 5BD

David M. John

Department of Botany, British Museum (Natural History), Cromwell Road, London

SW7 5BD

George W. Lawson

23 Sheffield Terrace, London W8 7NQ

Contents

Synopsis 195

Introduction 195

Species list •. •: 197

Acknowledgements 255

References 255

Synopsis

This paper assembles and, so far as is possible without extended field and herbarium studies, examines
critically the validity of records of marine and brackish-water Rhodophyta (Florideae) for the western
coast of tropical Africa. The whole mainland coastline from the northern boundary of Western Sahara
southwards to the southern boundary of Namibia, the oceanic islands from the Salvage Islands southwards
to Ascension and St Helena, and all islands close to the African mainland are included in the area covered.
Each species entry includes all traced records for the species, the names which have previously been
applied to it for the area, and additional comments or evaluation, as necessary. Comments are also
provided at generic or generic group levels in very complex cases. One new combination is effected:
Polycavernosa domingensis (Kiitzing) J. Price & D. John (Sphaerococcus Domingensis Kiitzing).

Introduction

The area dealt with in this part of the work is identical with that covered in parts published
previously (Lawson & Price, 1969; Price, John & Lawson, 1978, 1986; John, Price, Maggs &
Lawson, 1979). Country names employed and their earlier equivalents, and the names of island
groups included, are either listed in the legend or both listed and shown on the map in Fig. 1.
Genera with the initial letter G and constituent species are listed in alphabetical order; space and
organisational requirements within the Bulletin have required subdivision of a text originally
intended to cover as one issue all the remaining genera for the Rhodophyta (Florideae).
Each main entry consists of:

(i) The major bold heading, representing the currently-accepted name and authorities,
(ii) Subsidiary italicised headings at intervals within the entry. These are in square brackets
and essentially subdivide the overall entry. They represent the different ways in which
the species has been referred to throughout the past publication patterns for the area.
Incorrect citations from past literature have been maintained in these subsidiary heads so
that there shall be no doubt as to which record we attribute to which species or lower

Bull. Br. Mus. nat. Hist. (Bot.) 18 (3): 195-273 Issued 29 September 1988

196

J. H. PRICE, D. M. JOHN & G. W. LAWSON

Fig. 1 The coastline of tropical west Africa and the offshore islands.

1, Salvage Islands; 2, Canary Islands; 3, *Western Sahara [- former Spanish Sahara, Spanish West Africa]
(includes the often quoted Rio de Oro, the southern region of the country, but excludes Ifni); 4,
Mauritanie; 5, Senegal; 6, Gambia; 7, Guinea-Bissau [= Portuguese Guinea]; 8, Guinee; 9, Sierra Leone;
10, Liberia; 11, Cote d'lvoire; 12, Ghana; 13, Togo; 14, Benin [= Dahomey]; 15, Nigeria; 16, Cameroun;
17, t Bioko [= Macias Nguema Biyogo, Fernando Poo]; 18, Principe; 19, Sao Tome; 20, t Equatorial
Guinea [= Spanish Guinea]; 21, Gabon; 22, $ Republic of the Congo; 23, Cabinda; 24, Zaire [= Congo
Republic]; 25, Angola; 26, Namibia [= South West Africa]; 27, Ascension Island; 28, Saint Helena; 29,
Pagalu [ = Annobon] . The Cape Verde Islands, which lie immediately to the west of Dakar (Senegal) , have
been omitted from this map but are included in the species list that follows.

* The former colony of Spanish Sahara no longer officially exists, the territory it once covered being divided, by
agreement, between Morocco and Mauritanie. The effective date of the division, Spain concurring, was 28 February
1976, although guerrilla opposition delayed matters until a formal agreement on 14 April 1976. The attempt to
maintain the territory as the Democratic Sahara [Saharan] Arab Republic has apparently entered the 'realm of myth'
(Gretton, 1976).

t Nos 17 (Bioko) and 20 (Spanish Guinea, = Rio Muni) on the original map (part I) are now jointly administered, with
Annobon (29), as Equatorial Guinea. Bioko and Annobon are entered separately, where appropriate, in the species
list.

$ Loango, a name much used by earlier collectors such as Welwitsch, was formerly a coastal region of West Africa. Its
application appears to have included much of the coastline of the Republic of the Congo (22), as well as of Cabinda
(23) and Zaire (24). Because by far the longest and rockiest part of the Loango coast lies now within the Republic of
the Congo we have attributed all marine algal records from Loango to the Congo.

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 197

taxon level; only when clarification was required for comprehension have changes been
made in sub-head citation, in which case explanation is given in intermediary or terminal
notes.

(iii) The distributional data, with countries and island groups arranged in a single alphabetical
order. More generalised but still relevant statements of distribution follow the specific
country list. Complete distribution patterns require a scan of records established under
all names by which a species is known for this or adjacent areas. Hence, generalised
distribution statements are included verbatim since it is not always clear for precisely
which countries within the area they establish records. In all these cases, countries/
islands/generalised statements, numbers within parentheses after the names refer to
corresponding numbers in the references. In the present reference list, for ease of
readjustment for subsequent parts, references have not been renumbered but simply
omitted or added terminally and additionally numbered as appropriate for the present
part. Again, lists of references are therefore only partially interchangeable between
different parts of the overall list. As in 1986, we have considered the alphabetical
sequence of authors' names to be of greater importance than the strict numerical
sequence. References therefore appear in alphabetical order but only cross-referred as
to numerical order, where there is an anomaly for various reasons of rearrangement or
addition. 'References' in the listing cover also manuscript and expeditionary sources, as
well as works currently in press.

(iv) Additional qualifying notes, intermediary or terminal, have proved to be required in
many cases. These notes appear below whole entries or individual parts of entries to
which they specifically refer. Citation of references in the notes depends for its form on
whether those references contain species records (when they consist of authors' names,
followed by the reference number in the terminal list and, where appropriate, the
relevant page numbers after a colon) or do not (when they consist of authors' names, date
of publication, and sometimes page numbers after a colon).

Species nomenclature has been revised as far as possible and the complete author citation is
given for each currently-accepted combination. The subsidiary italicised headings (and any
other discarded combinations that require reference in the context of records of accepted species
for the list area) are included as cross-referencing entries to the currently-accepted names in the
overall list. The necessarily preliminary nature of all parts of the treatments presented has been
emphasised for each previously-published part of the list and applies no less here. As indicated
in previous parts, critical updating of the overall text is kept firmly in mind for the whole work,
although feasibility of that process will remain the final determinant at the appropriate time. We
would appreciate notification of any detected errors and omissions from any of the parts.

Species list

Galaxaura

The complexities of taxonomy/nomenclature in this genus have long been recognised, although
resolution of the problems still leaves much to be desired. Complexities have been hitherto
compounded by a general tendency to apply different epithets to plants later recognised as
tetrasporic and gametophytic generations of the same life-history. The anatomy exhibited by
these different life-history stages is occasionally sufficiently dimorphic for some species to be
entirely misunderstood and the different stages thus to be placed in separate anatomically-
distinguished sections of the genus (cf. Howe, 1917, 1918; Cordeiro-Marino, 108; Papenfuss et
al., 436) -examples are G. obtusata (gametophyte hitherto placed in Spissae*, tetrasporophyte
in Cameratae*) and G. marginata (gametophyte in Vepreculae*, tetrasporophyte in Brachy-
cladia*).
The major earlier work on this genus, that of Kjellman (313), has been subject to major

* Sensu Chou (103), Spissae = Umbellatae; Cameratae = Robustae; Vepreculae is maintained; Brachycladia =
Arboreae.

198 J. H. PRICE, D. M. JOHN & G. W. LAWSON

taxonomic reorganisation of conclusions by Papenfuss & Chiang (435) and Papenfuss et al.
(436) ; this has resulted in the reduction of large numbers of species to synonymy , details of which
will be clear from the species entries. We have generally followed the decisions made by
Papenfuss and colleagues (434; 435; 436), although even then the overall situation remains
complex and confusing. That situation is best summarised by Papenfuss (434: 271) and
Papenfuss et al. (436: 401-402):

The genus Galaxaura has suffered a great deal of misunderstanding. . . . The various
species have been assigned to nine genera if Galaxaura is included.'

Steentoft (535: 121) summarised aspects of the local western Africa situation, outlining the
putative nature of the species distributions deducible from available reports and stating:

're-examination of the S. Tome and Principe material shows that these islands
probably have the richest Galaxaura flora in the Gulf of Guinea.'

On the matter of wider consideration of interrelationships and status of the Galaxauraceae as a
whole, see recent publications by Huisman (especially 1987).

Galaxaura adriatica Zanardini

See Galaxaura oblongata (Ellis & Solander) Lamouroux.

Note. Considered a synonym of G. oblongata by Papenfuss & Chiang (435), Papenfuss et al. (436), and
Magruder(383).

Galaxaura annulata Lamouroux
See Galaxaura rugosa (Ellis & Solander) Lamouroux.
Note. Placed in synonymy of G. rugosa (q.v.) by Papenfuss et al. (436: 421).

Galaxaura coarctata Kjellman

See Galaxaura rugosa (Ellis & Solander) Lamouroux (for records) and Galaxaura decaisnei

J. Agardh (for taxonomic comment in terminal notes).

Note. Supporting information for this synonymy was published in Papenfuss & Chiang (435) and
(especially) in Papenfuss et al. (436: 422), where the full picture is presented.

Galaxaura comans Kjellman

See Galaxaura lapidescens (Ellis & Solander) Lamouroux and Galaxaura oblongata (Ellis &

Solander) Lamouroux.

Note. Relationships between the taxa named above need clarification. According to Papenfuss et al.
(436: 407-8, 427), the type (a tetrasporophyte) of G. comans, from Guadeloupe, West Indies, is actually a
plant of G. lapidescens, not of G. oblongata. G. lapidescens was thought to be the tetrasporophyte of
G. oblongata, but this has more recently been questioned and demonstrated to be untrue for at least one
geographical area (Hawaii); see the notes to G. lapidescens for further detail. Despite the statements,
referred to above, in Papenfuss et al., the same authors elsewhere (p. 410) reiterated Howe's (1918: 197)
suspicion that G. oblongata (gametophyte) and G. comans (tetrasporophyte) represent the same species.

Galaxaura cylindrica (Ellis & Solander) Lamouroux

See Galaxaura oblongata (Ellis & Solander) Lamouroux and Galaxaura lapidescens (Ellis &

Solander) Lamouroux.

Note. According to opinions expressed by Papenfuss et al. (436: 415-418), G. cylindrica is a synonym of
G. oblongata (q.v.); Chou (104) had earlier (1947) indicated that she felt differences between these taxa to
be arbitrary rather than real. Following generally the approach of Taylor (540), who commented that
G. cylindrica and G. oblongata are not too difficult to distinguish despite branch-diameter convergence,
Norris & Bucher (416: 193) were less definite. They maintained the application of the name G. cylindrica
for their Belize material, pending further study, but acknowledged morphological overlap and possible
conspecificity.

Galaxaura decaisnei J. Agardh

See Galaxaura obtusata (Ellis & Solander) Lamouroux [for general concept] and Galaxaura

rugosa (Ellis & Solander) Lamouroux [for records under G. decaisnei for the Canaries].

Note. It has been established that the material representing the type of this taxon, and therefore the basis
for the general concept, is a gametophyte from Barbados, West Indies, and should actually be attributed to

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 199

G. obtusata (Ellis & Solander) Lamouroux (Papenfuss et al. 436: 418-421); Seagrief (570) followed
Papenfuss et al. in accepting this equivalence. By contrast, Vickers's (547) material, from the Canaries and
named as G. decaisnei, was re-examined by B0rgesen (68: 70) and found to be what he then called
Galaxaura squalida Kjellman; this latter was considered to be the sexual generation of Galaxaura
flagelliformis Kjellman, both these names subsequently being placed in the synonymy of G. elongata J.
Agardh, which is now considered a synonym of Galaxaura rugosa (Ellis & Solander) Lamouroux. G.
flagelliformis has also been variously implicated in Galaxaura lapidescens auct. and, through the latter, in
G. cylindrica (q.v.) and G. oblongata (q.v.). Some Despreaux Canaries specimens, originally held in Paris
under the name G. decaisnei}. Agardh (Herb. Thuret), were transferred to Uppsala, where they were then
held as Galaxaura coarctata Kjellman, another name now considered to be a synonym of G. rugosa (q.v.).

Galaxaura elongata J. Agardh

See Galaxaura rugosa (Ellis & Solander) Lamouroux.

Note. Papenfuss et al. (436: 421-424) and Cribb (113: 26-27) both reduced G. elongata J. Agardh to the
synonymy of G. rugosa. Re-attributions of records for the list areas have required the assumption that the
original determination represented a correct application of the previous name. Thus (largely following
Papenfuss et al., 436) records previously attributed in our text to G. elongata have been reallocated
according to the following equivalence:
records as:

G. decaisnei J. Agardh: see G. rugosa (Ellis & Solander) Lamouroux.

G. elongata J. Agardh: see G. rugosa (Ellis & Solander) Lamouroux.

G. flagelliformis Kjellman: see G. lapidescens (Ellis & Solander) Lamouroux.

G. lapidescens auct.: see G. lapidescens (Ellis & Solander) Lamouroux.

G. rugosa auct.: see G. rugosa (Ellis & Solander) Lamouroux.

G. squalida Kjellman: see G. rugosa (Ellis & Solander) Lamouroux.

Galaxaura filamentosa Chou in W. Taylor

[As Galaxaura filamentosa Chou ex W. Taylor]

'tropical West Africa' (561).

[As Galaxaura filamentosa Chou]

Sao Tome (350; 535; 586).

'in tropical parts of the Atlantic and Pacific Oceans' (350; 586).

Tropical Africa (N. Gambia- Congo river)' (598).

[As Galaxaura lapidescens Lamouroux]

Sao Tome (251; 263; 264; 265).

Note. There has been much confusion as to the authorities to be correctly cited for this taxon. The type
derives from a collection by W. R. Taylor in the Galapagos Islands and the first description by R. C. -Y.
Chou was published in Taylor's (1945) paper on the Allan Hancock Expedition collections from there. This
Latin description is verbatim the same as that published later in the same year (September, as opposed to
May) by Chou (103) in her own paper. Thus, the authorities are correctly to be stated as above. In their
most recent revision of Galaxaura, Papenfuss et al. (436: 404, Table I) mentioned a report by Isaac (1971)
of the species from Kenya, but seem not otherwise to have dealt with the matter. The material referred to
above, and published by Hariot (251) and Henriques (263; 264; 265) as representing G. lapidescens
Lamouroux (q.v.), is included here on the basis of the opinion expressed by Steentoft (535) that it was
misdetermined. Steentoft nevertheless observed that Sao Tome plants were more fastigiately branched
and possessed somewhat shorter assimilators. Note that the report by Hariot (251) is merely secondary,
being directly based on the earlier papers of Henriques (263 et seq.).

Galaxaura flagelliformis Kjellman

See Galaxaura lapidescens (Ellis & Solander) Lamouroux [for records] and Galaxaura elongata

J. Agardh [for taxonomic comment].

Galaxaura fragilis (Lamarck) Lamouroux ex Decaisne [and other authority combinations]
See Galaxaura oblongata (Ellis & Solander) Lamouroux.

Note. Authorities hitherto quoted for this combination were usually '(Lamouroux) Decaisne',
'Lamouroux' or 'Decaisne'; critical nomenclatural and taxonomic revision by Papenfuss et al. (436:
415-418) resulted in recognition of the above more accurate citation. Those authors presented also
detailed synonymy, which we have accepted, consequent upon examining the type (a presumed gameto-
phyte from the West Indies, filed under Dichotomaria fragilis, ex Herb. Lamarck, in Herb. Mus. Paris).

200 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Galaxaura frutescens Kjellman

See Galaxaura marginata (Ellis & Solander) Lamouroux.

Note. Papenfuss et al. (436: 413-4) indicated these two taxa (inter alia) to share many attributes, but they
did not have available material in sufficient amounts or state adequate to arrive at a final decision. The type
of G. frutescens Kjellman is ex Herb. Areschoug and derives from Bahia, Brasil; the material, like that of
G. marginata, shows a marginal ridge. Later in the same publication (436: 427), Papenfuss et al. indicated
their opinion that G. frutescens is likely to be shown by future research to be conspecific with G. marginata.

Galaxaura fruticulosa Kjellman

See G. subfruticulosa Chou in W. Taylor and the notes to G. fruticulosa (Ellis & Solander)

Lamouroux.

Galaxaura fruticulosa (Ellis & Solander) Lamouroux

[As ' zoophyta fruticulosa (galaxaura)']
Canaries (219).

Note. See also the entry for Galaxaura subfruticulosa Chou in W. Taylor in the context of the possible
conspecificity of the present taxon with G. fruticulosa Kjellman and with G. subfruticulosa. The above
name is used in referring to the organism that provided the substrate for Conferva villum Agardh (q.v.).
The Galaxaura taxon involved is clearly the same as that recorded earlier in the same work (Gaudichaud,
219: 6-8, 1826) as 'galaxaura lapidescens'. See also the entry for the latter species.

Galaxaura lapidescens (Ellis & Solander) Lamouroux

Canaries (128A; 306B; 436; 598; 604; 605).

Cape Verde Islands (598).

Salvage Islands (598).

[As Galaxaura lapidescens Lamouroux]

Canaries (38; 126; 127; 142; 145; 219; 259; 263; 264; 318; 391; 535).

Note. See also the record from Gaudichaud (219: 156), as Galaxaura fruticulosa (q.v.).
Cape Verde Islands (38; 145; 259).
St Helena (142; 259; 260; 391).
Throughout all tropical seas' (410).
[As Galaxaura lapidescens (Solander) Lamouroux]
Canaries (27; 131).

[As Galaxaura lapidescens (Solander & Ellis)]
Salvage Islands (381; 439).

Note. Piccone (439: 35; 55) noted this taxon as 'G. oblongata (Sol. et Ell.) cf. G. lapidescens (Sol. et
Ell.)', based entirely on the record in Lowe (381).
[As Galaxaura (Microthoe) lapidescens Lamouroux]
Canaries (257).
Tropical Ocean' (257).
[As Galaxaura (Dichotomaria) lapidescens]
Canaries (313).

[As Galaxaura lapidescens (Solander) Lamouroux et var. tomentosa (Kiitzing)]
Canaries (390; 436; 441; 444).

[As Galaxaura lapidescens Lamouroux var. tomentosa Kiitzing]
Canaries (439).

[As Actinotrichia lapidescens Schmitz]
Canaries (547).

Note. See the note at the entry for Actinotrichia lapidescens Schmitz.
[As Galaxaura flagelliformis Kjellman]
Canaries (13; 38C; 68; 191; 226; 227; 379; 490).
Salvage Islands (38C; 556A).
[As Galaxaura flagelliformis (Kjellman) B0rgesen]
Canaries (14; 16; 230; 247; 375; 489; 556).
Cape Verde Islands (556).
Salvage Islands (231; 375; 556).
[As Galaxaura flagelliformis (Kjelin.) B0rgs. [sic!]]

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 201

Canaries (229; 610).

[As Galaxauraflagelliformis (Kjellman) emend. B0rg.]

Canaries (38B; 68; 71; 78; 214; 235).

Salvage Islands (38B; 381).

[As ? Galaxauraflagelliformis (Kjellman) emend. B0rgs. (G. lapidescens Pice.)]

Salvage Islands (452).

[As Galaxauraflagelliformis Foslie]

Canaries (235).

Note. Records published by Henriques (263; 264; 265) for Sao Tome [Ins. Rolas] under the name
Galaxaura lapidescens Lamouroux, based on collections by Quintas, are actually representative of
Galaxaura filamentosa (q.v.) (Steentoft; 535). The latter author (535), nevertheless, found the Sao Tome
plants to be more fastigiately branched and to have somewhat shorter assimilators. Steentoft (535) also,
following Howe (1918: 196), suggested that G. lapidescens represents the asexual (tetrasporophytic) phase
of the sexual Galaxaura cylindrica (Ellis & Solander) Lamouroux, now considered (Papenfuss et al. 436:
415-418) as synonymous with Galaxaura oblongata (Ellis & Solander) Lamouroux (q.v.). The tetra-
sporophyte/gametophyte relationship of G. lapidescens and G. oblongata has been questioned by some
workers and has recently (Magruder; 383) been shown to be untrue for Hawaii, where the G. oblongata
tetrasporophyte is small, inconspicuous, and filamentous. See the notes to G. oblongata and the discussion
in Magruder (383) for additional comment. See also the notes to Galaxaura comans Kjellman, Galaxaura
fruticulosa (Ellis & Solander) Lamouroux, and Galaxaura subfruticulosa Chou in W. Taylor. According to
Audiffred (38C: 175), G. flagelliformis Kjellman is the tetrasporophytic phase of G. squalida Kjellman.

Galaxaura marginata (Ellis & Solander) Lamouroux

Cameroun (288; 350; 586).

Cote d'lvoire (287; 288; 350; 586).

Gabon (288; 294; 350; 586).

Ghana (288; 299; 338; 350; 535; 586).

Liberia (129; 287; 350; 586).

Principe (288; 350; 535; 586).

Sao Tome (251; 288; 350; 535; 539; 586).

Senegambia(296;319).

Warm Atlantic (535).

'in warm temperate and tropical parts of the Atlantic Ocean.' (350; 586).

'Tropical Africa (N. Gambia- Congo river)' (598).

[As Galaxaura marginata (Solander) Lamouroux]

Ghana (153).

[As Galaxaura marginata (Solander in Ellis) Lamouroux emend. Kjellman]

Cameroun (139).

Note. The De Toni (139) record is based directly on data provided in Pilger (454).
[As Galaxaura marginata Lamouroux]
Sao Tome (251; 263; 264; 265).

Note. Reports in 263 and 264 include records from the adjacent islet of Rolas.
[As Galaxaura marginata Kiitzing f. marginata J. Agardh]
Senegambia (390).

[As Galaxaura (Microthoe) marginata Lamouroux]
Tropical Atlantic . . . Ocean' (255).
'Tropical Oceans' (257).
[As Galaxaura tenera Kjellman]
Gabon (350; 435; 570).

'Tropical Africa (N. Gambia - Congo river)' (598).
'widespread in warm temperate and tropical seas' (350).

Note. The Gabon records have often been presented in the form of the phrase 'Gabon River, West
Africa' (435; 570).

[As Galaxaura ventricosa Kjellman]
Gabon (104; 139; 313; 435; 436; 535; 561; 570).
Tropical eastern Atlantic' (277; 561).

202 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Tropical western Africa' (277).

Note. The records from Gabon have often been presented (313; 435; 436; 570) in the form 'Gabon River,
West Africa'. De Toni gave even further detail (139: 198) in his Latin statement: 'in Oceano Atlantico
inferiore in ostio fluminis "Gabon" Africae occidentalis meridionalis (E. Jardin)'. He also repeated
Kjellman's (313) observation that the taxon was most closely allied to G. veprecula.
[As Brachydadia australis Sonder [= Zanardinia marginata (Solander) J. Agardh]]
Gabon (250).

[As Brachydadia marginata (Solander) Schmitz]
Cameroun (454).
Senegambia (131; 390).

[As Brachydadia marginata f. marginata (Kiitzing) J. Agardh]
Senegambia (131).

[As Brachydadia marginata (Solander) Schmitz f. linearis (Kiitzing) J. Agardh]
Cameroun (454).

[As Brachytrichia marginata (Solander) Schmitz]
Sao Tome (251).

[As Zanardinia marginata J. Agardh]
'Warm Atlantic' (410).

[As Zanardinia marginata J. Agardh f. marginata J. Agardh]
Senegambia (27).

Note. Cordeiro-Marino (108) repeated Howe's (1918) comments on dimorphism in Galaxaura; the
overall conclusion then was that G. marginata, for which only the tetrasporophyte was known, could
possibly have G. occidentalis B0rgesen (60: 109) as its sexual phase. A preliminary revision of Galaxaura
by Papenfuss & Chiang (435) concluded that G. tenera Kjellman embraced the 'species' G. stupocaula
Kjellman [stupicaula], G. clavigera Kjellman, G. veprecula Kjellman, and G. ventricosa Kjellman. These
authors further commented that various characteristics of G. tenera agreed very well with those of
G. marginata and that the two may have been conspecific. On the basis of variable, often overlapping, local
Brazilian specimens, Cordeiro-Marino (108) agreed but hesitated to formalize the issue pending study of
types and further field-populations. Subsequently, Papenfuss et al. (436: 411-415) reduced G. tenera
Kjellman to synonymy with G. marginata (Ellis & Solander) Lamouroux, along with G. clavigera
Kjellman, G. occidentalis B0rgesen, G. stupocaula Kjellman, G. ventricosa Kjellman, and G. veprecula
Kjellman.

In work on material from the list area, Steentoft (535: 122) noted three growth forms amongst specimens
from Sao Tome and Principe; one, including sexual material, manifested characteristics intermediate
between G. ventricosa Kjellman (from Gabon) and G. veprecula Kjellman (from the Indian and Pacific
Oceans). Asexual material amongst the specimens had much in common with G. marginata, G. stupicaula,
and G. frutescens (both segregates from G. marginata and created by Kjellman). Steentoft (535) therefore
suggested 'the whole complex should be re-examined together.'

Seagrief (570), dealing with material from nearby South Africa, concluded that Galaxaura marginata
sensu Krauss (1846: 214), Zanardinia marginata sensu Barton (1893: 171, pro parte), Z. marginata b.
diesingiana (Zanardini) J. Agardh, and Brachydadia marginata f. diesingiana (Zanardini) De Toni were
all related to Galaxaura diesingiana Zanardini, not to the true Galaxaura marginata as recorded here.

Galaxaura oblongata (Ellis & Solander) Lamouroux

Canaries (2; 8; 13; 16; 68; 72; 80; 97; 128A; 191; 226; 227; 230; 233; 306B; 375; 435; 436; 489; 490;

535; 584; 598; 604; 610).

Cape Verde Islands (535; 598).

Cote d'lvoire (288; 350; 586).

Gabon (288; 294; 350; 586).

Ghana (153; 288; 299; 338; 350; 535; 586).

Liberia (129; 350; 586).

Principe (288; 350; 436; 535; 586).

Sao Tome (288; 350; 436; 535).

Salvage Islands (38B; 231; 375; 381; 439; 452; 598).

'a tropical Atlantic alga' (2).

'circumtropical' (280; 435).

'cotes occidentals d'Afrique et aux Canaries' (184).

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 203

'in warm temperate and tropical seas' (350; 586).

'Pantropical and subtropical' (535).

Tropical Africa (N. Gambia- Congo river)' (598).

'world wide distribution in tropical and subtropical waters' (383).

[As Galaxaura oblongata (Kjellman) Borgs.]

Canaries (229)

[As Galaxaura oblongata Lamouroux]

Canaries (493).

[As Galaxaura adriatica Zanardini]

Canaries (177).

'cotes occidentales d'Afrique et aux Canaries' (184).

[As Galaxaura cylindrica (Solander [or Ellis & Solander]) Lamouroux]

Canaries (13; 68; 71; 191; 227; 277; 390; 436; 439; 535).

Cape Verde Islands (535).

Principe (251; 265; 436).

Sao Tome (251; 263; 264; 265; 436; 535).

'pantropical' (535).

Note. The Sao Tome records in 263 and 264 include presence on Ins. Rolas, the nearby islet.
[As Galaxaura cylindrica Lamouroux]
Canaries (38).

Cape Verde Islands (38; 145; 260).

[As Galaxaura cylindrica (Solander in Ellis & Solander) Kjellman]
Canaries (108).

Cape Verde Islands (100; 183).
[As Galaxaura fragilis (Lamouroux) Decaisne]
Cape Verde Islands (41).
[As Galaxaura fragilis Lamouroux]
Cape Verde Islands (38; 261; 262; 408; 528; 551).
Gabon (250).

'Mers chaudes en general' (38).
'Tropical seas' (410).
[As Galaxaura fragilis Decaisne]
Cape Verde Islands (41; 42).
[As Galaxaura fragilis (Lamarck) Decaisne]
Gabon (250).

[As Galaxaura fragilis Decaisne var.]
Canaries (439).

Note. Piccone (439) gave no information regarding a varietal name. Levring (375) reported this species
to be well represented in the Salvage Islands, but not to be present elsewhere in the Madeira group.
Piccone (439) noted this taxon as 'G. oblongata (Sol. et Ell.) cf . G. lapidescens (Sol. et Ell.),' based directly
on the record in Lowe (381).

There has been long debate on the possible conspecificity of G. oblongata and G. cylindrica (q.v.).
Steentoft (535: 121) appeared from her comments to be of the opinion that differences between these two
taxa were that G. oblongata is coarser and has more divergent branching. She further indicated that 'G.
cylindrica is perhaps the sexual phase corresponding to the asexual phase known as G. lapidescens.' We
have not followed this - see the details below. Cordeiro-Marino (108: 32) indicated general acceptance of
the close relationships between G. oblongata and G. cylindrica, not only because of the merely narrower
and thinner segments of the latter, but also in a manifest lack of data on tetrasporophytic plants, only the
sexual phases being known. Following data variously presented in Howe (1918) and B0rgesen (68; 80),
Steentoft (535) considered the corresponding tetrasporophyte of the sexual phase G. oblongata to be the
taxon known as G. comans Kjellman (q.v.), not known from the eastern Atlantic. Cordeiro-Marino (108)
also repeated Howe's (1918) comments that G. lapidescens (Ellis & Solander) Lamouroux (q.v.) and G.
comans were possibly the tetasporophytic phases, respectively of G. cylindrica and G. oblongata. Thus,
despite accepting (see above) close relationship between the latter two taxa, Cordeiro-Marino still
maintained them as separate, in common with many other phycologists, including Howe and B0rgesen,
mentioned above. Howe and B0rgesen both considered the demarcation between the entity [G. comans

204 J. H. PRICE, D. M. JOHN & G. W. LAWSON

and G. cylindrical and the entity [G. comans and G. lapidescens] to be arbitrary. None of this accords well
with the more recent opinions of Papenfuss & Chiang (435) and Papenfuss et al. (436), who submerged G.
cylindrica in the synonymy of G. oblongata and believed G. lapidescens to be a good species, with G.
comans as its synonym. We have adopted the views of these more recent workers, especially as outlined in
436, the most recent paper. On other aspects of the differences/distinctness of G. cylindrica and G.
oblongata, see the notes presented at G. cylindrica. It is worth mentioning that B0rgesen also variously
stated (68) that Lamouroux's specimen of G. oblongata, which he (B0rgesen) examined, is a form of G.
obtusata (Ellis & Solander) Lamouroux, and that (80) it is impossible to separate the Red Sea species G.
schimperi Decaisne from G. oblongata. Chou (104) considered that G. oblongata perhaps represented the
sexual phase of more than one species, whilst Itono (277: 14, 16) indicated G. fastigata, G. pilifera, G.
cylindrica, and G. oblongata closely to resemble each other in their external features, leading most
Japanese phycologists to regard the latter three species as G. fastigiata. Despite his arbitrariness in dividing
the four species on the basis of ecologically- or physiologically-affected characteristics, however, Itono felt
they should be maintained pending further work.

Recent work by Magruder (383) has shown that Hawaiian Galaxaura oblongata has a triphasic life
history, with conspicuous gametophytes and small inconspicuous filamentous tetrasporophytes. In this it
accords with data established for two other genera in the Galaxauraceae [= Chaetangiaceae], Scinaia and
Pseudogloiophloea, even having much the same tetrasporophyte form. Most other known species of
Galaxaura and Actinotrichia have conspicuous macroscopic tetrasporophytes, with only a few Galaxaura
spp. (where the tetrasporophyte is not with absolute certainty known) having possibly small filamentous
tetrasporophytes. Magruder (383: 407) commented that this presence of inconspicuous tetrasporophytes
was certainly responsible for difficulties in aligning putative tetrasporophyte 'species' with the correspond-
ing gametophyte 'species' in Galaxaura. Details of both spermatangial conceptacle development and
carpogonial branch development are very similar in G. oblongata, G. obtusata, G. diesingiana, and G.
marginata, according to Magruder (op. cit.).

Galaxaura obtusata (Ellis & Solander) Lamouroux

Canaries (68; 71; 97; 191; 227; 247; 277; 372; 435; 436; 598).

Cape Verde Islands (598).

Gabon (250; 294; 350; 586).

Salvage Islands (598).

Sierra Leone (30; 350; 586).

'eastern Atlantic (Canary Islands) westward to East Africa' (435).

'The tropical Ocean, in all longitudes' (256).

'Tropical Africa (N. Gambia - Congo river)' (598).

Tropical and subtropical oceans' (257).

'widespread in warm temperate and tropical seas' (350; 586).

[As Galaxaura obtusata J. Agardh]

Angola (41; 42).

'Warm Atlantic' (41; 42).

[As Galaxaura umbellata Lamouroux]

Canaries (37; 41; 42; 408).

Cape Verde Islands (408; 528; 551).

'Warm Atlantic' (410).

[As Galaxaura umbellata (Esper) Lamouroux]

Canaries (141A).

Salvage Islands (38B; 215).

[As Galaxaura umbellata Montagne]

Canaries (38).

Cape Verde Islands (38; 551).

Note. Authorities quoted as '(Solander) Lamouroux' (e.g. in Itono (277: 7)) have been treated
throughout the species entry as though the form in the present entry heading had been employed. B0rgesen
(68) examined Lamouroux's specimens labelled as G. umbellata and concluded that they represent G.
obtusata. Papenfuss et al. (436: 418-421) and Seagrief (570: 30) concluded equally that this was correct, at
least pro parte for G. umbellata. Askenasy's (37; 38) records as G. umbellata may really represent G.
rugosa (Ellis & Solander) Lamouroux. In 1897, Askenasy (38: 165), although presenting his entry under
the heading 'G. umbellata Mont, non Lamour.', added a terminal comment in French, which translated as:

RHODOPHYTA (PLORIDEAE) OF TROPICAL AFRICA 205

This species, also found on the Canaries and named by Montagne G. umbellata, seems to me to be
different from the species so named by Espen [sic!], Lamouroux, Kiitzing and J. Ag. (Epicrisis). It is nearer
to G. rugosa (Soland.) Kiitz.' See the entry for the latter species, as well as the note presented at G.
decaisnei J. Agardh, for other comment of relevance to the present taxon.

Itono (278: 8) regarded G. obtusata and G. umbellata as very similar and not separable into two species.
All these forms were suggested as related to the sexual phase of Galaxaura robusta, which is not reported
for coasts of the list area.

Galaxaura rugosa (Ellis & Solander) Lamouroux

Ascension (37; 535).

Cameroun (288; 350; 454; 535; 586).

Canaries (37; 128A; 232B; 306B; 436; 598; 604; 605).

Cape Verde Islands (37; 38B; 535; 598).

Cote d'lvoire (287; 288; 350; 586).

Gabon (586).

Ghana (586; BM (unpublished)).

Liberia (287).

Principe (436; 586).

Salvage Islands (38B; 556A; 598).

Sao Tome (251, pro parte; 263; 265; 288; 350; 436; 535; 586).

'in oceano calidiore atlantico' (27).

'Pantropical' (535).

Tropical Africa (N. Gambia -Congo river)' (598).

'warmeren Atlantischen Ocean (Kanaren, Antillen)' (37).

'widespread in warm temperate and tropical seas' (350; 586).

[As Galaxaura rugosa Lamouroux]

Cape Verde Islands (38; 145; 259; 260).

Sao Tome [including Rolas] (263; 264).

'Ocean Atlantique tropical' (38).

'Warm Atlantic' (410).

[As Galaxaura rugosa (Solander) Lamouroux]

Sao Tome (265 pro parte).

'in warmeren atlantischen Ocean' (503).

[As Galaxaura rugosa Solander]

Canaries (439).

Sao Tome (251 pro parte, 265 pro parte).

[As Galaxaura annulata Lamouroux]

Cape Verde Islands (38; 408; 528).

[As Galaxaura coarctata Kjellman]

Canaries (139; 313; 436; 538).

[As Galaxaura Decaisnei 3. Agardh]

Canaries (547).

'in oceano Atlantico calidiore' (27; 131).

[As Galaxaura elongata J. Agardh]

Canaries (435).

Gabon (294; 350).

Sao Tome (350).

'eastern Atlantic (Canary Islands) westward to East Africa' (435).

'widespread in warm temperate seas and pantropical' (350).

[As Galaxaura squalida Kjellman]

Canaries (13; 38B; 38C; 68; 97; 226; 227; 302; 304; 351; 379; 414; 535).

Cape Verde Islands (38B; 38C; 100; 183).

Salvage Islands (38B; 556 A).

Sao Tome (535).

'pantropical' (535).

206 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Note. The unpublished record from Ghana is based on a specimen (in BM) collected by V. J. Foote in
1946. Records that appear above as pro parte from Sao Tome require reference to the other taxon
involved; in all cases, this appears to be Galaxaura subfruticulosa Chou in W. Taylor (q.v.) - see additional
notes below. The report from the Salvage Islands (38B; 556A) initially (Prud'homme van Reine, 15 Nov.
1982, in litt. to JHP; 556A) indicated the records as new for that island group. The inclusion under this
species entry of the records established as Galaxaura annulata Lamouroux for the Cape Verde Islands is
based on the synonymy deduced and presented by Papenfuss et al. (436); the nature and accuracy of
determination of the actual material involved have not been checked . The correctness of placement here of
the records established as G. Decaisnei J. Agardh may not be firmly authenticated, but has been effected
on the basis of the area concerned (Canaries; warm Atlantic ocean); see the notes to the species entry for
G. decaisnei. Although the type of G. decaisnei (Barbados) proved (cf. 436) to be Galaxaura obtusata,
Vickers's (547) material from the Canaries emerged on examination by B0rgesen (68) as G. squalida [= G.
elongata, now synonymised with G. rugosa]. For a contrasting opinion on G. squalida status, see Audiffred
(38C: 175) and the note to G. squalida here.

For a full recent statement of the synonymy, similarities, and previously employed application of names
to all or parts of this taxon in different geographical areas, see Papenfuss et al. (436). See also Cribb (113:
26-27). Steentoft (535: 123) suggested that G. rugosa and G. subfruticulosa Chou have hitherto been
confused amongst Sao Tome material, hence the 'pro parte' recording of certain previous records,
mentioned above. The significance of such confusion depends on more firm understanding than currently
available of morphological relationships and life-history connections between these taxa, and between
both and G. subverticillata Kjellman. See the detailed notes to G. subfruticulosa Chou and G. obtusata
(Ellis & Solander) Lamouroux; Papenfuss et al. (436: 425) suggested that the tetrasporophyte of G. rugosa
is probably that referred to as G. subverticillata Kjellman, but further work is required to establish the
connection. G. subverticillata is not recorded for the eastern Atlantic.

It is possible (cf. Itono 277: 8-9) that the Pacific Galaxaura pacifica and the Atlantic G. rugosa are
different forms of the same species; both have free assimilatory filaments and similar overall morphologies,
although it seems that the free filaments may persist in older parts of G. rugosa, as opposed to being early
deciduous in G. pacifica.

Galaxaura squalida Kjellman

See Galaxaura rugosa (Ellis & Solander) Lamouroux and G. elongata J. Agardh.

Note. According to Chou (104), the type of G. squalida (from the Virgin Islands) is in Herb. Areschoug
under the name G. rugosa. According to Audiffred (38C: 175), G. squalida is the sexual phase of G.
flagelliformis Kjellman (see G. lapidescens (Ellis & Solander) Lamouroux).

Galaxaura stupicaula ['stupocaula'] Kjellman

See Galaxaura marginata (Ellis & Solander) Lamouroux.

Note. According to Chou (103), the form of the epithet should be stupicaulis, to agree with the generic
substantive form. The sexual phase involved could be G. veprecula Kjellman [= G. marginata, q.v.].
Papenfuss & Chiang (435: 304) rendered the epithet as [G.] stupocaula and considered it to be representa-
tive of Galaxaura tenera Kjellman [? = G. marginata, q.v.]; these data are repeated and elaborated in
Papenfuss et al. (436: 411-415).

Galaxaura subfruticulosa Chou in W. Taylor

Sao Tome (350; 535; 586).

'in tropical parts of the Atlantic and Pacific Oceans' (350; 586).

Tropical Africa (N. Gambia- Congo river)' (598).

[As Galaxaura rugosa (Ellis et Solander) Lamouroux, pro parte]

Sao Tome (251; 263; 264; 265).

Note. See the detailed note to Galaxaura rugosa (Ellis & Solander) Lamouroux. There has been much
debate about the status and relationships of this present taxon. Steentoft (535: 124) observed that G.
subfruticulosa Chou is closely related to G. subverticillata Kjellman, only the latter having been previously
recognised in the Atlantic (and then only the western Atlantic). Itono (278: 5-6) repeated the comment
that G. subfruticulosa is most closely related to G. subverticillata; he concluded, after examining the
supposed differences between the taxa, that 'it is not inconceivable that G. subfruticulosa and G.
subverticillata will be found to be conspecific.'

According to Papenfuss et al. (436: 409), G. fruticulosa Chou in W. Taylor comes close to G. lapidescens
(q.v.). G. fruticulosa was established on the basis of material from Revillagigedo Is., Pacific Mexico, and
said by Chou to be closely similar to G. ramulosa and G. tomentosa. G. lapidescens and G. fruticulosa may
well eventually be thought conspecific. The significance of this is that Chou renamed G. fruticulosa

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 207

Kjellman, calling it G. subfruticulosa and designating a new type (from Clarion Is., Revillagigedo).
Papenfuss et al. (436: 409) went on 'Whether Chou's species and the one of Kjellman actually are
representative of the same species remains to be determined.'

Later in their 1982 publication (436), Papenfuss et al. (pp. 424-5) commented that Galaxaura
subverticillata Kjellman, G. fruticulosa Kjellman, and G. subfruticulosa Chou in W. Taylor all have
swollen basal cells to both long and short assimilatory filaments. The conspecificity of G. fruticulosa
Kjellman, G. fruticulosa (Ellis & Solander) Lamouroux, 1816 (from Bahamas), and G. subfruticulosa
Chou in W. Taylor (Revillagigedo) cannot really be established without detailed examination of the types.
Ellis & Solander's Corallina fruticulosa (1786) illustration shows that G. subverticillata is not of the same
taxon; the branches of the latter do not 'grow smaller towards the ends.' The verticillate arrangement of
assimilatory filaments in East African G. subverticillata was not mentioned by their description of C.
fruticulosa, nor is it clear in Kjellman's (313) treatment of his G. fruticulosa, although it is so in his
description of G. subverticillata. Papenfuss et al. (436: 425) went on to state:

'Extended assimilatory filaments of Galaxaura subfruticulosa Chou are evenly distributed over
the entire surface of the thallus. . . . and slightly verticillate only near the tips. This taxon is
distinct from G. subverticillata Kjellman. On this basis Chou (1945, p. 42) placed
G. subfruticulosa very close to G. ramulosa and G. tomentosa, taxa which we have reduced to
synonymy of G. lapidescens. '

Confusion between G. subverticillata and G. lapidescens may have partially arisen because the
verticillate arrangement in the former is seen clearly only in young branches, the hairs disappearing with
ageing of the branch. G. subverticillata Kjellman is probably the sporophyte of G. rugosa (q.v.); 'Until this
can be established with certainty, however, it seems prudent to retain G. subverticillata as an autonomous
species' (Papenfuss et al. 436: 425).

Galaxaura subverticillata Kjellman

See the terminal notes to the entry for Galaxaura subfruticulosa Chou in W. Taylor.

Galaxaura tenera Kjellman

See the entries for Galaxaura marginata (Ellis & Solander) Lamouroux [records] and for both

G. marginata and G. veprecula Kjellman [notes].

Galaxaura umbellata Lamouroux

See Galaxaura obtusata (Ellis & Solander) Lamouroux [records; notes] and Galaxaura rugosa

(Ellis & Solander) Lamouroux [notes].

Note. The rationale for first attribution of valid publication to Lamouroux (331: 262) is presented in
Papenfuss et al. (436; 419, footnote 11). The combination has sometimes been attributed to (a) Montagne;
(b) (Esper) Lamouroux; (c) (Esper) J. Agardh, as authorities.

Galaxaura ventricosa Kjellman

See Galaxaura marginata (Ellis & Solander) Lamouroux.

Note. Papenfuss et al. (436: 411, 412) referred to the type of G. ventricosa Kjellman, from [near the
mouth of] the Gabon River, West Africa; they placed the taxon in synonymy with G. marginata (Ellis &
Solander) Lamouroux (q.v.). Chou (104) had earlier indicated it to be impossible to distinguish between
G. ventricosa and G. veprecula on the basis of anatomy; both these taxa are in any case now accepted as
lying within the synonymy of G. marginata (Ellis & Solander) Lamouroux.

Galaxaura veprecula Kjellman

See Galaxaura marginata (Ellis & Solander) Lamouroux.

Note. Jaasund (19776: 417) stated that the conspecificity of Galaxaura veprecula (from Madagascar) with
Galaxaura tenera Kjellman (type locality: Mombasa) is still an open question. More recent opinion (e.g.
Papenfuss et al., 436: 411 et seq.) does not, however, confirm this, since many workers accept both these
taxa as lying within the synonymy of G. marginata (Ellis & Solander) Lamouroux; we concur in the latter
view. See also the notes to G. ventricosa Kjellman.

Galaxaura spp.

Canaries (3; 71; 89; 117; 118; 214; 229; 253A; 301; 436; 567).
Cape Verde Islands (191; 500).
Ghana (491).
Senegal (59; 282; 611).

208 J. H. PRICE, D. M. JOHN & G. W. LAWSON

'Gulf of Guinea' (611).
'West Africa' (290).
Note. Bornet (89) stated he had '3 especes' of Galaxaura.

Ganonema farinosa (Lamouroux) Fan & Wang
See Liagora farinosa Lamouroux, and Abbott (1).

Ganonema pinnatiramosa (Yamada) Fan & Wang in Fan et al.
See Liagora farinosa Lamouroux, and Abbott (1).

Gastroclonium clavatum (Roth) Ardissone
Canaries (232B; 253; 583; 598; 604; 605).
[As Gastroclonium ovatum (Hudson) Papenfuss]
Canaries (226: 227).

Note. Haroun Tabraue et al. (253: 273-4) recorded the taxon as 'Nueva cita para Canarias.' The first
citation was of epiphytes on Corallina elongata, later (232B) extended to those on Codium taylorii. Haroun
Tabraue et al. were also (loc. cit.) responsible for transferring the previous records as Gastroclonium
ovatum (Hudson) Papenfuss (q.v.) in 226 and 227 to the present taxon; they stated 'La cita de
Gastroclonium ovatum. . . . para Pta. Pechigueras, Lanzarote. . . . Gil-Rodriguez & Afonso-Carrillo
(1980), tras la revision del pliego, debe ser eliminada del "Catalogo de las Algas marinas bentonicas para el
Archipielago Canario" y ser incluido como Gastroclonium clavatum (Roth) Ardisson.' G. clavatum overall
distribution was cited as "East Atlantic (Cadiz & Canaries) and Mediterranean.' See also the entry for
Gastroclonium ovatum (Hudson) Papenfuss - when originally recording their material as that taxon from
there (Canarias), Gil-Rodriguez & Afonso-Carrillo (226: 64) stated: 'Esta especie, cuya area de distribu-
tion se extiende desde Inglaterra a Mauritania, no habia sido senalada precedentemente para Canarias.'

Gastroclonium kaliforme (Goodenough & Woodward) Ardissone
See Chylocladia verticillata (Lightfoot) Eliding.

Gastroclonium ovatum (Hudson) Papenfuss

Canaries (128A; 226; 598).

Mauritanie (33; 222; 226; 273; 349; 546).

'Atlantique (de 1'Angleterre a la Mauritanie)' (33).

'Atlantique, depuis les cotes anglaises jusqu'en Mauritanie' (222).

'British Isles to Mauritania; Canary Isles' (273).

'desde Inglaterra a Mauritania' (226).

'desde las costas de Inglaterra hasta Mauritania' (546).

[As Chylocladia ovalis (Hudson) Hooker]

'D'Angleterre aux Canaries' (89).

[As Lomentaria ovalis (Hudson) J. Agardh]

'In Oceano atlantico calidiore' (27).

Note. See also the entry for Gastroclonium clavatum (Roth) Ardissone. The Mauritanie record in
Lawson & John (349) is based only on data in 33 and 222. It is probable that the older Canaries references
(in 89) and the more recent secondary statement in Irvine (273) should both be referred (as with the bases
in 226 and 227 for the more recent statement) to Gastroclonium clavatum (Roth) Ardissone. Haroun
Tabraue et al. (253) established that point.

Gastroclonium reflexum (Chauvin) Kiitzing

Canaries (33; 227; 273).

'Atlantique (de 1'Angleterre aux Canaries)' (33).

[As Lomentaria reflexa Chauvin]

Canaries (70; 191).

'English coast southwards to the Canary Islands' (70).

[As Lomentaria reflexa (Chauvin) J. Agardh]

Canaries (441; 444).

[As Lomentaria pygmaea (Lamouroux) Gaillon]

Canaries (401).

[As Lomentaria pygmaea Gaillon]

Canaries (44).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 209

Note. Ardre (33: 141) commented on distinguishing characters for Gastrodonium, indicating that
her material closely approached the Coeloseira of Hollenberg, based on the constant occurrence of
polysporangia in certain species; since this latter distinction lacked supporting characters, she preferred to
retain the material in Gastrodonium. There is slender basis for many of the above records - Piccone (441),
who recorded only a single young plant, formed the direct and only basis for his own repeated records in
(444), as well as for those published in B0rgesen (70) and in Gil-Rodriguez & Afonso-Carrillo (227). This
taxon was frequently referred to as Chylodadia reflexa [auctorum] and the situation as regards the
synonymy is complex (cf. Irvine, 273: 84-85). The record in Montagne (401: 156-7), and therefore also
that in Benitez (44), is attributed here on general nomenclatural grounds. Montagne referred in his
synonymy only to Gigartina pygmaea Lamouroux and Lomentarla pygmaea of Gaillon and of Duby. His
additional comments included: 'Among other algae , to which it adheres, this species, if red, is doubtfully of
the animal kingdom' and he indicated that the species had a great affinity with Lomentaria pusilla De
Notaris [in litt.] and with Chondria nana C. Agardh, all three being very probably only forms of one and the
same species. De Toni (132: 566) recognized Gigartina pygmaea Lamouroux (with doubt) and Lomentaria
pusilla Kiitzing in the synonymy of G. reflexum (Chauvin) Kiitzing; he nowhere in his works referred to
Lomentaria pygmaea Gaillon, to Lomentaria pusilla De Notaris, or to Chondria nana C. Agardh, although
he did place in that same synonymy Lomentaria exigua De Notaris. L. Irvine (273: 85) indicated G.
reflexum to be 'closely related to G. davatum (Roth) Ardiss. . . . but without the reflexed habit.'

(•d id id la acerosa (Forsskal) J. Feldmann & Hamel

Gabon (294; 350; 586).

Sao Tome (350; 535; 586).

Sierra Leone (30; 350; 586).

'in warm temperate and tropical seas' (350; 586).

'most warm seas' (81).

'parait repandue dans toutes les mers tropicales (Atlantique . . .)' (194).

Tropical Africa (N. Gambia - Congo river)' (598).

'tropical and subtropical seas' (614).

[As Gelidium claviferum Kiitzing or its 'forma']

Sao Tome (251; 263; 264; 265).

[As Gelidiopsis rigida (Vahl) Weber-van Bosse]

'Seems to occur in all warm seas' (64).

[As Gelidium rigidum (Vahl) Greville]

'in oceano calidiori atlantico' (27).

'Throughout warm seas' (410).

Note. Problems occasionally attach to accurate determination of material of this taxon from the area (see
Steentoft, 535). B0rgesen (73: 5) earlier (but not acceptedly) recombined this taxon in the genus
Echinocaulon. Of material from the Admiralty Islands (not directly relevant here), Hemsley (260)
commented that This is probably a variety of the almost cosmopolitan Gelidium corneum, Lamour.' [He
was writing under the heading of Gelidium rigidum Greville]. For a general statement of synonymy in
Gelidiella acerosa see Seagrief (570: 30).

Gelidiella pannosa (J. Feldmann) J. Feldmann & Hamel

'sporadically from the French Cote Basque south to Morocco . . . and Senegal' (614).

[As Gelidiella tenuissima J. Feldmann & Hamel]

Canaries (38B;556A).

Cape Verde Islands (38B).

Mauritanie (38B; 122; 556).

Salvage Islands (38B; 556; 556A).

Senegal (38B; 122; 556).

'From Biarritz and Portugal southward to Cape Verde.' (375).

'Portugal hasta Cabo Verde (Ardre 1970)' (518).

[As Gelidiella tenuissima (Thuret) Feldmann & Hamel]

Canaries (38C).

Cape Verde Islands (38C).

Mauritanie (38C; 121).

Salvage Islands (38C).

210 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Senegal (38C).

[As Gelidiella pannosa (Bornet) J. Feldmann & Hamel]

Canaries (598).

Salvage Islands (598).

'Atlantique (Biarritz, Portugal, jusqu'au Cap Vert)' (33).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Note. Fan (1961: 340, note 4) has summarised the nomenclatural and taxonomic history of this taxon in
demonstrating that authority and epithet citation should be as stated in the present entry heading.
Nevertheless, we have considerable sympathy with the change of epithet to tenuissima effected by J.
Feldmann & Hamel (195: 102 et seq. [Revue algol.] or 226 et seq. [collated reprint]) for this taxon in
Gelidiella because of the potential confusion arising from the complexities of both taxonomy and
nomenclature in the difficult Gelidiales; these complexities are well expressed in the recent work by Maggs
& Guiry (614). It is not now possible to reject specific epithets on the grounds of confusion and it is clear
that J. Feldmann's combination in Echinocaulon (?) of pannosum can be treated directly as the basionym
for Gelidiella pannosa, given the homonymy involved in the earlier treatments of Gelidium pannosum
Grunow (now Gelidiopsis pannosa (Grunow) Schmitz) and Gelidium pannosum sensu Bornet, the basis
for much of the J. Feldmann & Hamel (194: 195) reasoning.

Even in the case of the perhaps more desirable Gelidiella tenuissima, there has been frequent
mis-citation of authorities; Thuret's herbarium name was Gelidium tenuissimum and, since Feldmann &
Hamel were not taking up the whole combination in this MS name, there is no requirement to cite Thuret as
authority.

Gelidiella ramellosa (Kiitzing) J. Feldmann & Hamel
Salvage Islands (38B; 231; 375; 598).

Note. Levring (375) considered this species well-separated from Gelidiella tenuissima J. Feldmann &
Hamel (= Gelidiella pannosa (J. Feldmann) J. Feldmann & Hamel, q.v.). G. ramellosa showed good
representation in the Salvage Islands, although Levring (loc. cit.) presented its distribution only as the
Salvage Islands and Tunis. See also details presented under Gelidiella tinerfensis J. Seoane-Camba.

Gelidiella tenuissima J. Feldmann & Hamel

See Gelidiella pannosa (J. Feldmann) J. Feldmann & Hamel.

Gelidiella tinerfensis J. Seoane-Camba
Canaries (227; 518; 574; 598; 614).

Note. Seoane-Camba (518; 574) presented, especially in (518), comments on the distinctions between G.
tinerfensis and its nearest relatives in Gelidiella - G. pannosa, G. ramellosa (q.v.), G. lubrica, and G.
trinitatensis .

Gelidiella sp.

Canaries (38C).
Salvage Islands (38B).

Note. Reported as only sterile material. Audiffred (38C: 175) presented some description of the material
from the Canaries.

Gelidiocolax hemisphaerica Gardner

See Gelidiocolax microsphaerica Gardner.

Gelidiocolax microsphaerica Gardner

Senegal (1A; 575).

[As Gelidiocolax hemisphaerica Gardner]

Senegal (122).

'Subtropical Africa [Senegal (N. Gambia), Mauritania, Former W. Sahara]' (598).

Note. Fan & Papenfuss (575: 33) commented that: 'Dangeard in 1952 reported Gelidiocolax micro-
sphaerica (in error as G. hemisphaerica) from Dakar.' Dangeard (122) did not explain his mis-use of the
specific epithet as 'hemisphaerica'' , but it may possibly relate to his believing it to be a more accurate
reflection of thallus form. Even Abbott & Hollenberg (1A: 342) opened their published description of
Gardner's taxon with the words: Thallus a smooth, more or less hemispherical mound'. Fan & Papenfuss
(575) considered various aspects of the relationships between species currently placed in this genus. In all
probability, the Gelidiocolax sp. reported from Senegal and Mauritanie by Dangeard (121) in 1951 is also
representative of G. microsphaerica; see the following entry.

RHODOPHYTA (PLORIDEAE) OF TROPICAL AFRICA 211

Gelidiocolax sp.

Mauritania (121).
Senegal (121).

Note. Found growing on Gelidiumfoliosum P. Dangeard, particularly on the 'stolons'; the Gelidiocolax
sp. concerned is almost certainly that subsequently (122) referred to by Dangeard as G. hemisphaerica
Gardner [= G. microsphaerica Gardner] (q.v.).

Gelidiopsis gracilis (Kiitzing) Vickers

Senegal (50; 55; 59).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Note. Bodard (50) commented that this species was for long confused with Gelidium crinale (q.v.) in
their collections from Senegal, since the two often occur intermixed. He continued: 'c'est a notre
connaissance la premiere fois que ce Gelidiopsis est signale en Afrique; il etait uniquement connu aux
Antilles, avec Gelidiopsis variabilis [q.v.] dont nous allons parler.' Later, Bodard & Mollion (59: 198)
stated of this species: 'la position systematique est douteuse, il peut etre considere comme une algue
tropicale a faible dispersion geographique.' The present species differs from Gelidiopsis planicaulis (q.v.)
in that the latter lacks the cylindrical axis which is present in at least the distal parts of the former.

Gelidiopsis intricata (C. Agardh) Vickers

Ascension (474; 475).

Canaries (38D; 598).

Cape Verde Islands (38D; 191, 598).

Salvage Islands (38D; 598).

[As Gelidiopsis intricata (Kiitzing) Vickers]

Canaries (38B; 70; 482; 605).

Cape Verde Islands (38B; 100; 183).

Salvage Islands (38B).

'does not extend (southwards) into the Gulf of Guinea' (487).

[As Gelidiopsis intricata (Agardh) Schmitz]

Senegal (542).

[As Gelidium intricatum Kiitzing]

Canaries (68; 118; 486; 489; 495).

Note. It has been questioned (Steentoft, 535: 125-126) that this and G. variabilis (J. Agardh) Schmitz
are really separate species. See the note to the latter for fuller explanation. Material recorded from Sao
Tome by Tandy in Exell (539: 386) and from the Cape Verde Islands by J. Feldmann (183) and Chevalier
(100), as G. intricata, was sterile but, according to Steentoft (loc. cit.), was actually referable to G.
variabilis (q.v.). It is possible that other records here (as, e.g., 38B (Salvage Islands); 70; 100; 183; 482)
should (although sterile/small specimens were again involved) be referred to the Gelidiopsis variabilis
form, whether or not the latter and G. intricata are finally considered to be conspecific. B0rgesen (70)
merely repeated the record from the Canaries given in Reinbold (482: 22), without additional information.
Reinbold's material was also sterile. Audiffred & Weisscher (38B: 29) could not be sure of the
determination of their Salvage Islands material, since their specimen was very small. Trochain's (542: 109)
material from Senegal was merely drift - it, too, was referred by Steentoft (535: 126) to Gelidiopsis
variabilis (q.v.). Sterile specimens may also have caused confusion in routine determination for areas of
Atlantic Africa - G. intricata may have been misidentified with Wurdemannia miniata or Caulacanthus
ustulatus; see the entries for these latter species. Rodriguez (486) misrendered the specific epithet as
intrincatum in recording Gelidium intricatum from the Canaries. Schiffner (495) merely referred to
B0rgesen's earlier (68) statements for Tenerife.

Gelidiopsis planicaulis (W. Taylor) W. Taylor

?Bioko(346;350;586).

Cameroun (346; 350; 586).

Canaries (556A).

Cape Verde Islands (598).

Gabon (294; 350; 586).

Liberia (129; 350; 586).

Sierra Leone (295; 350; 586).

'in tropical parts of the Atlantic Ocean' (350; 586).

212 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Tropical Africa (N. Gambia -Congo river)' (598).

Note. A variable species with regard to flattening of branches; W. Taylor suggested (pers. comm.) that
West African material was very similar to that from the western Atlantic, although flattening in the former
was more localised (294; 350; 586).

Gelidiopsis rigida auctorum [usually (Vahl) Greville or (Vahl) Weber-van Bosse]
See Gelidiella acerosa (Forsskal) J. Feldmann & Hamel.

Gelidiopsis variabilis (J. Agardh) Schmitz

Angola (352).

Bioko (346; 350; 586).

Cameroun (122; 288; 350; 586).

Cape Verde Islands (598).

Cote d'lvoire (288; 350; 586).

Gambia (296; 350; 586).

Ghana (288; 290; 299; 300; 338; 350; 376; 491; 537; 586).

Guinee/Guinea-Bissau (350; ? 529; 586).

Liberia (129; 350; 586).

Mauritanie (349).

Sao Tome (288; 350; 586).

Sierre Leone (295; 350; 586).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Tropical Africa (N. Gambia -Congo river)' (598).

'widespread in warm temperate and tropical seas' (350; 586).

[As Gelidiopsis variabilis (Greville) Schmitz]

Annobon(456;457;535).

Cameroun (500; 535).

Canaries (7535).

Ghana (535).

Guinee (535).

Mauritanie (192).

Sao Tome (93; 535).

Senegal (50; 121; 122; 529).

[As Gelidiopsis variabilis (Greville ex J. Agardh) Schmitz]

Ghana (297).

[As Gelidiopsis variabilis Schmitz]

Cameroun (454).

[As Gelidiopsis intricata (C. Agardh) Vickers]

Sao Tome (539).

Sierra Leone (295).

[As Gelidiopsis intricata (Agardh) Schmitz]

Senegal (542).

[As Gelidiopsis intricata (Kiitzing) Vickers]

Cape Verde Islands (100; 183).

Note. There is doubt as to the distinctions, or indeed the reality of separateness, between G. variabilis
and Gelidiopsis intricata (q.v.). Steentoft (535: 125-126) firmly expressed this in the following terms:
'Larger tuft-forming plants seem to be placed in the former species [G. variabilis], low growing turf-
forming ones in the latter [G. intricata].' In expanding this further, she commented on the Tandy in Exell
(Sao Tome) material (539) and the Cape Verde Islands records (J. Feldmann, 183; Chevalier, 100). Other
records that could possibly concern G. variabilis are discussed in the note to Gelidiopsis intricata (q.v.).
The overall picture seems to be one of difficulty in distinguishing validly between ends of morphological/
anatomical clines. Steentoft's opinions were embodied in the following extract from 535: 125-126:

'Both G. variabilis and G. intricata. . . . are probably in need of critical re-examination. The
latter name is probably invalid, since it is based on Sphaerococcus intricatus C. Ag., the
"cotype" of which [J.] Feldmann has examined and reported as a Gelidium species, since
intercellular rhizines are present. ... it seems to the present writer that there is some doubt as

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 213

to the distinctness of [these species]. . . . However a much greater acquaintance with the
American forms of the so-called G. intricata is required before any conclusions can be
reached.'

Sterile material of G. variabilis may, for Atlantic Africa, have at times also been confused with
Wurdemannia miniata (q.v.) or Caulacanthus ustulatus (q.v.).

The reports from Senegal in Bodard (50) and Sourie (529) have similarly an appreciable degree of
uncertainty about them. Sourie (529: 306) himself expressed doubt about his records of the species from
Guinee/Guinea-Bissau, and his citation from Senegal stated: 'Gelidiopsis sp. (G. variabilis (Greville)
Schmitz?).' Bodard (50: 83-84) indicated that G. variabilis from Senegal was much more robust (but much
rarer) then Gelidlopsis gracilis (Kutzing) Vickers; his single certain specimen of Gelidiopsis variabilis from
Pointe de Sarene, Senegal (February), proved identical with a plant collected by Sourie (529) at Cap
Rosso. At first sight, G. variabilis could easily be confused with Gracilaria augustissima (sic! - for
angustissima) (Bodard 50: 83-84), but the anatomical structure proved to be of Gelidiopsis, rather close to
G. gracilis (Kutzing) Vickers.

Gelidiopsis sp.

Cameroun (337; 344; 393; 394).

Ghana (42 A; 92; 220; 338; 344; 393; 394).

Guinea-Bissau/Guinee (529).

Guinee (269; 344; 384; 393; 394; 529).

Nigeria (344; 393; 394; 415).

Senegal (393; 394; 529).

Note. For a possible attribution of the Senegal record in Sourie (529), see the entry for Gelidiopsis
variabilis (J. Agardh) Schmitz.

Gelidium

Bornet's earlier categorization of Gelidium as 'genre diabolique' is hardly a matter for dispute;
this applies whether the concern is nomenclatural, taxonomic, morphological, or ecological
(Dixon & De Valera, 1961; Dixon & Irvine, 172). This complicated form- variation, often
seasonally different in individual plants and populations, is therefore also reflected in the
excessive names scattered throughout the literature, and probably lies behind the large number
of species reported from South Africa sensu stricto and not from apparently similar areas to the
north and within the check-list area. Cases in point are Gelidium amansii (Lamouroux)
Lamouroux, G. arenarium Kylin, G. caespitosum Kylin, and G. pristoides (Turner) Kutzing. On
the other hand, there are numbers of species names applied within the check-list area that have
never been employed outside it, to north or south. Overall rationalization of the taxonomy and
nomenclature on a pan- Atlantic basis is required, but the difficulties and time-consumption
make the prospect daunting. Furthermore, approaches to species recognition via morphological
characteristics alone are clearly insufficient if a meaningful result is to be obtained; biochemical
and genetic levels will require exploration on a geographically widespread basis to rationalize
the different patterns of understanding of the taxa, and to eliminate contrasting and purely local
traditions of naming.

No such vast and daunting project has been undertaken here and consequently no general
reallocation of records depending on a totally reorganised system of name-allocation is
presented. Transfers of individual records or groups of records have been made on restricted
bases, where there has been good reason to believe that action necessary. In the absence of any
such individual information, the publishing authors' names have been maintained, if for no
other reason than to draw attention to current anomaly requiring further work. Thus, in the
absence of real understanding of limits of taxa considered as species level across virtually the
whole genus, the recording by name (and therefore the reality) within the present list area of
the following 'species' must be considered at best speculative to doubtful, at worst spurious:

Gelidium arbuscula
asperum
attenuatum
canariensis [cartilagineum var. canariensis]

214 J. H. PRICE, D. M. JOHN & G. W. LAWSON

cartilagineum

clavatum

corneum

crinale

elminense

flaccidum

foliosum

latifolium

melanoideum

microdon

micropterum

pristoides

pulvinatum

pusillum

reptans

spathulatum

spinulosum

versicolor

The numerous varieties described in many of the above, where not directly reflected in use by
some authors at species level of names used by other authors at infraspecific levels, also require
rationalisation.

Amongst the above, the names canariensis (q.v.) and versicolor [previously cartilagineum]
(q.v.) are perhaps the least likely to reveal anomalous application, whilst Gelidium sesquipedale
(q.v.) has been omitted entirely from the list. G. sesquipedale is perhaps the least variable
species in the genus and therefore the least likely name to be misapplied. Such confusion as
exists about it has largely resulted from the involvement of Fucus corneus Hudson [Gelidium
corneum auct.] in the situation, since F. corneus of Hudson has been shown (Dixon, 1967) to be
based on Buddie (BM) specimens that proved to be G. sesquipedale. Hence, as a corollary,
derives further confusion about the significance and application of the epithet corneum (q.v.).

Gelidium apiculatum Kiitzing

Note. See Gelidium spinulosum (C. Agardh) J. Agardh, for which this is a possible synonym, and the
note to Gelidium microdon Kiitzing.

Gelidium arbuscula Bory ex B0rgesen

Canaries (306B; 582; 598).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Tropical Africa (N. Gambia- Congo river)' (598).

[As Gelidium arbuscula (Montagne) B0rgesen]

Canaries (2; 229; 269; 270; 351; 374; 393; 394; 486; 490; 529; 567).

Cote d'lvoire (287; 586).

Guinee(350;586).

Mauritanie (59).

Senegal (59; 123; 483; 586).

'remonte du Senegal jusqu'a la Mauritanie.' (59).

'warm temperate and tropical parts of the eastern Atlantic Ocean.' (350; 586).

[As Gelidium arbuscula Bory]

Canaries (4; 5; 68; 191; 227; 379; 489; 493; 547; 581).

Guinea-Bissau (529).

Guinee (529).

Senegal (122; 7529; 7530; 7531).

[As Gelidium arbuscula (Bory) Montagne]

Canaries (253; 583).

Senegal (253).

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 215

[As Gelidium corneum (Hudson) Lamouroux var. nereideuml Lightfoot]
Canaries (401).

Note. See B0rgesen (68: 85-89) for details of the taxon and its background to 1927. Many of the above
records involve considerable doubt as to their accuracy. All Sourie's (529; 530; 531) publications
mentioned that doubt; Feldmann, who reviewed Sourie's material from Senegal, was said by Sourie (529:
116) to have considered specimens of this species 'assez differents de la plante des Canaries'. Essentially
the same message lies behind the Bodard & Mollion statement (59: 198) that their Senegal plants
represented 'une endemique possible que Dangeard avait appelee Gelidium arbuscula et qui semble plutot
une espece nouvelle que J. Feldmann nous a fait remarquer.' Feldmann had earlier (191: 414) expressed
the opinion that (for the Canary Islands) 'Le Gelidium arbuscula, endemique bien caracterise, peut
neanmoins etre rapproche du Gelidium sesquipedale, espece lusitano-africaine.' Acuna Gonzalez (2: 3),
also concerned with Canary Islands plants, commented that there 'tambien existen algunas [especies] que
son endemicas, como. . . . Gelidium arbuscula' . It therefore seems a reasonable possibility that at least two
separate entities are concealed within the application of this name in the list area; it may be possible partly
to utilise ecological characteristics in distinguishing the entities - Santos, Acuna & Wildpret (490) indicated
that what they called Gelidium arbuscula had an 'especial predilection por los acantilados verticales
[vertical or steep cliffs]'.

The record (401) in Montagne is included here on the basis of B0rgesen's comment (68: 86) that G.
arbuscula Bory in herb, 'seems to be a rather common species at the [Canary] Islands. . . . further,
according to a specimen in Herb. Montagne, the Gelidium corneum var. nereideum, too, is this plant'. See
also the note to Gelidium as genus. Haroun Tabraue et al. (583: 111) commented that [on Gomera]:
'Gelidium arbuscula is easily recognisable by its purple coloured thalli, which clearly contrast with the
brown-yellow of Cystoseria abies-marina [amongst which it grows] and the blackish colour of Gelidium
versicolor '.' (Translation from the Spanish.) On p. 112, they added yet another difference - the greater
rigidity of G. versicolor (q.v.).

Gelidium asperum Montagne

Note. See Gelidium spinulosum (C. Agardh) J. Agardh. The present name is probably a manuscript
name appearing only on a specimen in the Herb. Montagne (PC); in any case, it appears much like
Gelidium microdon Kiitzing, itself probably a synonym of G. spinulosum. Data also appear in B0rgesen
(68: 90).

Gelidium attenuatum (Turner) C. Agardh

See Gelidium latifolium (Greville) Bornet in Bornet & Thuret.

Gelidium caespitosum Kylin

Note. This name was originally applied by Fox to BM specimens from Ghana (April 1952) and from
Nigeria (March and November 1954); the records were subsequently published by her (1957) under the
name Gelidium pusillum (Stackhouse) Le Jolis. Specimens were afterwards reassigned by Lawson & John
(350) to Gelidium corneum sensu B0rgesen.

Gelidium canariensis (Grunow) Seoane-Camba

Canaries (518; 598).

[As Gelidium versicolor (S. Gmelin) Lamouroux]

Canaries (172; 180; 227; 253; 352).

[As Fucus versicolor Gmelin]

Canaries (90).

Note. The taxon was recorded as both Fucus cartilagineus L. and Fucus versicolor Gmelin by Bory (90),
who also (90: 304) stated the former to be a synonym of the latter. The alga was said to be: Tres comun sur
toutes les pierres et les roches de la rade de Sainte-Croix.'. De Toni (131: 152 et seq.) also placed G.
cartilagineum in the synonymy of G. versicolor Lamouroux.
[As Fucus cartilagineus L.]
Canaries (90).

Note. See the note above, to Fucus versicolor Gmelin.
[As Gelidium cartilagineum (L.) Gaillon]

Canaries (25; 68; 81; 89; 131; 170; 229; 254; 269; 305; 318; 351; 386; 393; 394; 401; 482; 486; 489;
490; 567).

Note. The records in Montagne (401: 158), J. Agardh (25), and De Toni (131) are based on Bory's (90)
data.

216 J. H. PRICE, D. M. JOHN & G. W. LAWSON

[As Gelidium cartilagineum var. canariense Grunow in Piccone or var. canariensis Grunow in

Piccone]

Canaries (68; 131; 191; 439; 482; 489; 567; 581).

[As Sphaerococcus cartilagineus (L.) C. Agardh]

Canaries (19).

Note. Recorded by C. Agardh (19: 286-288) as 'Ad insulas Canarias; sec. Bory Voy.'
[As Gelidium cartilagineum Gaillon]
Canaries (44; 385; 493).

Note. The record in Benftez (44) is directly based on Montagne (401). That in von Martens (385) is based
(385: 38) on a record of Mertens, from Tenerife.
[As Gelidium cartilagineum (L.) Greville]
Canaries (37; 242).
[As Corallina rubens Tournefort]
Canaries (548).

Note. For explanation of attribution of this record, see the notes to Jania rubens (L.) Lamouroux.
[As Gelidium cartilagineum Greville]
Canaries (493).

Note. See the entry for Gelidium versicolor (S. Gmelin) Lamouroux for records from other geographical
areas. The rationale for recognition of Gelidium canariensis is documented in the notes to G. cartilagineum
(L.) Gaillon.

Gelidium capillaceum Kiitzing [or (S. Gmelin) Kiitzing]
See Pterocladia capillacea (S. Gmelin) Bornet & Thuret.

Gelidium cartilagineum (L.) Gaillon

See Gelidium canariensis (Grunow) Seoane-Camba and Gelidium versicolor (S. Gmelin)

Lamouroux.

Note. The name Gelidium cartilagineum (L.) Gaillon has to be rejected for that genus since it was based
on material that has proved to be representative of the genus Plocamium (Dixon, 1967). Thus, the
southern African species of Gelidium to which the epithet had largely by custom been applied required
another name, the earliest available being shown by Dixon (1967: 58) to be Gelidium versicolor
(S. Gmelin) Lamouroux, based on Fucus versicolor of Gmelin (1968).

The most northerly attached records of the taxon to which the name G. versicolor (S. Gmelin)
Lamouroux is thus to be applied are from the Canary Islands (Dixon, 170: 47, and subsequent publications,
especially 172: 134). For that same area, Seoane-Camba (518) recently concluded that materials from
'mainland' Spain and from the Canary Islands were sufficiently different to represent distinct taxa, despite
hitherto having been collectively called Gelidium cartilagineum (i.e. correctly G. versicolor). He raised the
var. canariensis Grunow of G. cartilagineum to specific status as Gelidium canariensis (Grunow) Seoane-
Camba; all previous Canaries records are here transferred to that latter species entry, although material
behind all the earlier records has not been checked to confirm their validity. Previously, J. Feldmann (191:
414) had considered (without effecting it) that the Canaries material of then so-called Gelidium cartila-
gineum was 'represente par une variete endemique (var. canariensis} assez differente du type pour meriter
peut-etre d'etre consideree comme une espece distincte. Le G. cartilagineum type est une espece
indo-pacifique, tres abondante en particulier en Afrique du Sud et en Californie.' See also the general note
to Gelidium.

Gelidium claviferum Kiitzing

See Gelidiella acerosa (Forsskal) J. Feldmann & Hamel.

Gelidium corneum sensu B0rgesen

Canaries (486).

Gabon (350; 586).

Gambia (350; 586).

Ghana (350; 377; 586).

Liberia (350; 586).

Nigeria (350; 586).

Togo (350; 586).

'widespread in boreal-antiboreal to tropical parts of the Atlantic Ocean.' (350; 586).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 217

[As Gelidium corneum (Hudson) Lamouroux]

Angola (352).

Canaries (66; 401).

Gabon (294).

Gambia (296).

Ghana (288; 491).

Liberia (129; 288).

Nigeria (288).

Togo (288).

'in [warmeren] atlantischen [Ocean]' (504).

'warmer parts of the Atlantic Ocean' (60).

[As Gelidium corneum (Linn.) Lamour.]

Canaries (331; 332).

[As Gelidium corneum (Turner) Lamouroux]

Canaries (68; 71; 499).

Cape Verde Islands (499).

'Nordwestafrika' (499).

'Westafrika'(499).

'[Madeira, but] . . . probably widely distributed.' (375).

'Seems to be widespread.' (68).

[As Gelidium corneum (Lamouroux)]

Canaries (196).

[As Gelidium corneum Lamouroux]

Angola (41; 42).

Canaries (44; 547).

Cape Verde Islands (38; 223; 578; 596).

'Throughout all oceans.' (410).

[As Gelidium corneum Hudson]

Cape Verde Islands (145).

[As Gelidium corneum [sensu - as vgl.] Feldmann et Hamel]

Angola (500).

[As Gelidium arbuscula Bory]

Liberia (286).

Togo (293).

[As Gelidium pusillum}

Nigeria (213).

[As Sphaerococcus corneus (Hudson) C. Agardh]

Canaries (19).

Note. C. Agardh (19: 279-285) recorded the 'parent' species as 'Ad Teneriffam; Bory'. He also
recognised 17 varieties/subspecies, mostly with names still employed somewhere in Gelidium. Amongst
those is ft. sesquipedalis, recorded from the Mediterranean and Cadiz. The Bory record and the
significance of this S. corneus entry will need careful rationalization.
[As Fucus corneus L.]
Canaries (90).

Notes. See the note at Sphaerococcus corneus (Hudson) C. Agardh subhead, above.

General notes. For records under names often attributed as varieties of Gelidium corneum (Hudson)
Lamouroux, refer as follows in this work:

var. capillaceum (S. Gmelin) C. Agardh

Pterocladia capillacea (S. Gmelin) Bornet et Thuret
var. clavatum (Lamouroux) C. Agardh

Gelidium pusillum (Stackhouse) Le Jolis
var. crinale (Turner) C. Agardh

Gelidium crinale (Turner) Desmazieres
var. nereideuml Lightfoot

Gelidium arbuscula Bory ex B0rgesen

218 J. H. PRICE, D. M. JOHN & G. W. LAWSON

var. plnnatum (Hudson) Montagne

Pterodadia capillacea (S. Gmelin) Bornet & Thuret
var. sesquipedale Clemente

Gelidium sesquipedale (Clemente) Thuret in Bornet & Thuret
var. spinulosum C. Agardh

Gelidium spinulosum (C. Agardh) J. Agardh.

Amongst the attributive 'species' so involved, some (e.g. G. pusillum, G. crinale) are often considered as
conspecific (e.g. by Wynne, 1986).

Records established under the name Gelidium corneum in most cases present considerable interpretive
problems, whatever authorities have been quoted for the combination. G. corneum represents perhaps the
classic extreme of general confusion. Overlying all that is the fundamental difficulty of typification of the
basionym Fucus corneus of Hudson. For general background to nomenclatural and morphological
problems with this taxon, see Dixon (1967), together with the notes to the entries here for the genus
Gelidium and for Gelidium sesquipedale (Clemente) Thuret in Bornet & Thuret. The name as utilized at
present for the list area is only in the sense applied by B0rgesen (68: 85), and even that brings with it the
need for a complete revision of whatever backing material is available for all the records involved.
Therefore, what we offer here is merely a statement of rationalization through what others have said, not of
what we conceive as taxonomic/nomenclatural truth, even assuming the latter to be approachable. If
B0rgesen's taxon does finally turn out to be a good one, it will require whatever is the earliest available
name, or a new one, as the case may be. At presents corneum as an epithet is completely confused in all
respects.

It is clear that even the reduction of the general chaos of names/taxa in Gelidium to the two aggregates
'latifolium' and ' pusillum' (leaving aside for now the more extreme morphologies of G. versicolor, q.v.,
and G. sesquipedale, q.v.), as has been suggested by Dixon & Irvine (172: 126 et seq.) leaves a plethora of
problems in that these two aggregates are themselves not satisfactorily morphologically distinguishable
apart at all times and from all locations; the artificial key presented in 172: 127 admits as much from the
statement that it 'should be used with caution', and from the nature of the distinguishing characteristics
provided. Since the epithet corneum has variously been applied, and conceptually recognized as applying,
for our area to forms that could be placed in either or both of the aggregates, the point made by Lawson &
John (350: 177) regarding Fox's (213) Nigerian plants reported as G. pusillum being more closely related to
G. corneum sensu B0rgesen may have relatively little overall significance, although morphologically
accurate for their material at that time.

As observed by Seagrief (570), some applications of the name corneum (those sensu Harvey) for
material from Sea Point, South Africa, do not represent Gelidium corneum (Hudson) Lamouroux (nor,
apparently, Gelidium corneum sensu B0rgesen), but are attributed to Gelidium micropterum Kutzing
(q.v.). On the basis of BM holdings of material collected during the her Benguellense of F. Welwitsch
[those specimens circulated under the numbers 73 (Benguela); 74 (Mo9amedes); 75 (drift, from near
Benguela)], it seems highly likely by comparison with the illustration presented by Kutzing (326: 21, Tab.
59 (4137), 1868) that this attribution to Gelidium micropterum Kutzing is also true for at least some
material from Angola. It probably equally applies for the coast of Namibia, whence Gelidium micropterum
Kutzing has been firmly reported. There has, however, been doubt about the way in which the name
micropterum Kutzing has been applied in Namibia; it has been suggested that Gelidium pristoides (Turner)
Kutzing was perhaps involved. For some of the supposed distinctions between G. corneum and G.
micropterum, see P. S. Rao (1970: 71-73).

Material recorded by Lawson, John & Price (352: 311), from Angola, as patches of short red algal 'turf
of Gelidium corneum (Hudson) Lamouroux, included a mixture of forms that, in isolation, would have
been considered as representing variously G. corneum sensu B0rgesen, G. pusillum (Stackhouse) Le Jolis,
and Gelidium micropterum Kutzing.

All the above complications are, of course, compounded by varying local traditions of epithet
application in Gelidium such that, other than generic validity, the use of an epithet in one country or region
may present no biogeographic information in any way of use for another area. This is so for most of the
epithets of other than very local application and markedly so for 'corneum'.

Gelidium coronopifolium (Goodenough & Woodward) Lamouroux

See Sphaerococcus coronopifolius (Goodenough et Woodward) Stackhouse.

Gelidium crinale (Turner) Desmazieres
Gabon (350; 586).
Gambia (296; 350; 586).

RHODOPHYTA (FLORIDEAfi) OF TROPICAL AFRICA 219

Ghana (350; 586).

Liberia (350; 586).

Mauritania (349).

Sao Tome (350; 586).

Sierre Leone (350; 586).

Western Sahara (349; 476).

[As Gelidium crinale (Turner) Lamouroux]

Angola (352).

Canaries (2; 68; 108; 118; 191; 214; 252; 486; 517; 546; 547; 584).

Gabon (294).

Ghana (152; 338; 491; 535).

Guinee-Frangaise (529).

Liberia (129).

Mauritanie(252;500;535).

Salvage Islands (38B; 231; 375).

Sao Tome (251; 265; 535).

Senegal (33; 121; 122; 529).

Sierra Leone (30; 295).

'Atlantic coast of Europe, N. Africa and N. America' (375).

'Atlantico de Inglaterra a Canarias' (517).

'Atlantique (de 1'Angleterre aux Canaries, Cap Vert . . .)' (33).

'English coast to the Canary Is.' (68).

'Gulf of Guinea' (269).

'in [warmeren] atlantischen Ocean' (504).

'repandu dans toutes les mers chaudes' (188).

[As Gelidium crinale Lamouroux]

Canaries (547).

[As Gelidium crinale Lamouroux in Bory]

Canaries (89).

'D'Angleterre aux Canaries' (89).

[As Gelidium corneum (Huds.) Lamour. var. crinale (Turner) C. Agardh]

Canaries (401).

[As Gelidium crinale (Thuret) Lamouroux]

Senegal (530).

Note. The epithet rendering in Prime (476: 23) is as 'orinale\ a purely typographic error that has been
ignored. It is now generally accepted that Lamouroux, despite employing the combination G. crinale (in
Bory, Diet, class. d'Hist. nat. 7: 191 (1825)), presented neither basionym data and source, nor descriptions
of the taxon, thus not effecting a proper transfer; the first to achieve correct recombination of Turner's
epithet crinale in Gelidium was Desmazieres.

Opinions on this taxon and the often-confused Gelidium pusillum (Stackhouse) Le Jolis have varied
widely, from absolute synonymy to at least usage in very different and identifiable senses. Except where
both the concepts used and the mode of application of the names have both been entirely clear and
consistent, the possible equivalence of G. crinale with G. pusillum agg. has not been here taken as
accepted. Names are therefore otherwise maintained as used by the publishing author, but it should be
borne in mind that, given reasonable consistency of view and application, it is strongly likely that many
records under the two names represent the same taxon. See also the entry for G. pusillum (Stackhouse)
Le Jolis.

Gelidium elminense Dickinson

Cote dTvoire (288; 350; 586).

Ghana (152; 288; 350; 586).

'in tropical parts of the eastern Atlantic Ocean.' (350; 586).

'Tropical Africa (N. Gambia-Congo river)' (598).

Note. This species was originally described on the basis of plants from Elmina and Iture, both Ghana,
collected in April 1945 (25th and 26th, respectively). According to Lawson & John (350; 586), the species
falls within the Gelidium crinale complex sensu Dixon. From her own comments in the original description

220 J. H. PRICE, D. M. JOHN & G. W. LAWSON

(152: 43), Dickinson was markedly reluctant to indulge in describing yet another new species in an already
ill-understood and cluttered genus, but could see no other way out of such a tediously complex situation.

Gelidium flaccid urn P. Dangeard

Guinea-Bissau (529).

Mauritanie (121).

Senegal (121; 122; 529).

'Subtropical Africa [ Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Note. In publishing the original description of G. flaccidum, Dangeard (121: 23) noted that 'La coupe
transversale de la fronde montre la grande rarete des rhizines. Par ce caractere ce Gelidium se rapproche
du G. melanoideum.'

Gelidium foliosum P. Dangeard

Mauritanie (121).

Senegal (50; 121; 122; 123; 529).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Note. In the process of describing new taxa from Senegal and Mauritanie, Dangeard (121 : 22) stated that
'Deux especes enfin de Gelidium [G. flaccidum; G. foliosum] de ses regions sont nouvelles et bien
caracterisees'. Despite that, he went on to state [in both cases of G. foliosum]: 'rappelle un peux le
Gelidium pusillum var. pulvinatum' and, later: 'ressemble un peu par son port au G. galapagense decrit par
R. TAYLOR [sic!] (1945)'. He added that G. foliosum 'il en differe par ses ramules a tetrasporanges
retrecis a la base et comme pedicelles.'

The resemblance to Gelidium pusillum, especially var. pulvinatum, seems to have been too close for
many workers to take the present taxon seriously. Soon after G. foliosum was published, Sourie (529: 289
and footnote 1) commented: 'Selon J. Feldmann (opinion inedite), ce Gelidium [foliosum] pourrait n'etre
qu'une forme de G. pusillum.'' Even Bodard (50) doubted the validity of it, stating: 'Je pense que le
Gelidium foliosum ne peut etre considere que comme une variete de Gelidium pusillum'. He did not give
reasons for this opinion. More recently, one of us (DMJ) annotated a collection (John & Seku; no. 6949)
from Harper, Liberia, 2 Jan. 1972, determined as Gelidium pusillum var. pulvinatum, 'Similar to
description of G. foliosum by Dangeard (121: 22) from Senegal'.

Gelidium galapagense W. Taylor

See the entry to Gelidium foliosum P. Dangeard.

Gelidium intricatum Kiitzing

See Gelidiopsis intricata (C. Agardh) Vickers.

Gelidium latifolium (Greville) Bornet & Thuret

Canaries (253; 306B; 598; 604).

Cape Verde Islands (598).

Senegal (59).

Western Sahara (253; 349; 393; 394).

'Atlantico (Noruega - Rio de Oro)' (604).

'Atlantico Oriental (Noruega - Rio de Oro)' (253).

'Atlantique: depuis les cotes anglaises jusqu'au Rio de Oro' (222).

'Nordwestafrika' (499).

'Southern Norway to Rio de Oro' (172).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Tropical Africa (N. Gambia- Congo river)' (598).

[As Gelidium latifolium (Greville) Thuret & Bornet]

Senegal (59; 7529).

'Atlantic Ocean (European and African coasts, Canary Islands)' (177).

'Atlantique (de 1'Angleterre au Rio de Oro)' (33).

[As Gelidium latifolium Bornet]

'Atlantic Ocean: . . . African coast.' (566).

[As Gelidium attenuatum (Turner) C. Agardh]

Western Sahara (393; 394; 476).

Note. Despite the convenient recognition, in some parts of the world, of the contrasting morphologies of
the 'species aggregates' known as G. latifolium and G. pusillum, it is by no means certain that the two

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 221

groups do not represent merely extremes of a morphological cline representing environmental or racial
characteristics within the range of expression of a single genome. Haroun Tabraue et al. (253: 259) referred
to their Canary Islands report as: 'nueva para el Archipielago Canario.' Dangeard (117; 118: 184-5) had
previously stated that Gelidium attenuation was missing from the Canaries. It remains to be shown whether
this and the other records above are significant or not. The expression of difficulty in determination and
doubt occurs throughout many of the comments associated with records: Gayral (222: 373) indicated the
wide range in morphology of the 'species', from 'var. luxurians" (wide branches; distichous) to 'var. hystrix'
(cylindrical; minor laterals spiralled on axes), with all kinds of intermediates. Sourie (529) recorded his
determination as doubtful, though the reason for doubt is not clear. Perhaps the extreme in confusion over
this taxon was shown clearly by Dixon (170: 47), who revealed previous misdetermination as Gelidium
sesquipedale (Clemente) Thuret in Bornet & Thuret (q.v.) of 'the entity known as G. attenuatuni '; G.
sesquipedale is one of the few species in the genus with a very characteristic appearance. Two of us (Lawson
& John, 349) were unable to find material in Western Sahara; our record is based on previous comments, of
which the only original one is probably that in Primo (476).

Gelidium melanoideum Schousboe ex Bornet

Senegal (52; 529).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania Former W. Sahara]' (598).

[As Gelidium melanoideum var. pseudopulvinatum J. Feldmann]

Senegal (529).

Note. For an analysis of characteristics that distinguish between G. melanoideum and the Gelidium
crinalelG. pusillum complex, see Dixon & de Valera (1961). The species, originally recognised from the
Tangier region, is principally of Mediterranean origin. See also the note to Gelidium flaccidum P.
Dangeard.

Gelidium microdon Kiitzing

Western Sahara (349; 393; 394).

'Atlantique (. . . Canaries . . .)'. (33).

[As Gelidium spinulosum (C. Agardh) J. Agardh]

Western Sahara (476).

Note. It is possible that the taxa G. microdon and G. spinulosum may be wholly conspecific, in which case
the above records should all appear under the latter, being the older valid name. Ardre (33: 68), Bornet
(89: 112 [272]) and B0rgesen (68: 90) all comment on conspecificity of these taxa; Bornet is the most
unequivocal in his opinion, whilst Ardre suggests conspecificity and B0rgesen merely quotes Bornet's
comments, under his heading of G. spinulosum (C. Agardh) J. Agardh. See also Gelidium asperum
Montagne. Bornet's comment was: 'Les Gelidium apiculatum Kiitz. . . . et microdon Kiitz. . . . qui sont
figures d'apres des echantillons de Tanger et de Cadix, ne sont que des formes de cette espece.' 'Cette
espece' refers to the entry under which the statement appears, Gelidium spinulosum J. Agardh.

The record above in 349 is based only on Michanek (393; 394) and Primo (476), Lawson & John not
themselves having detected specimens.

Gelidium micropterum Kiitzing

Namibia (348; 522; 522 A).

Senegal (121; 122).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gelidium corneum (Hudson) Lamouroux]

Angola (352).

Note. The Lawson & Isaac (348) record from Namibia was based directly on the data from 522 (Simons),
since they did not locate material. Applications of the epithet micropterum, the concept behind the taxon,
and any synonymy involved are all highly complex and opinions on the matter have varied greatly. For a
detailed consideration of some aspects of this, see the notes to Gelidium corneum sensu B0rgesen as
regards South Africa, Angola, and Namibia. Gelidium pristoides (Turner) Kiitzing (q.v.) is often
considered to be involved, whilst De Toni (131: 152 et seq.) placed G. micropterum (albeit with '?') in the
synonymy of Gelidium cartilagineum (L.) Gaillon [now Gelidium versicolor (S. Gmelin) Lamouroux or
G. canariensis (Grunow) Seoane-Camba].

Gelidium pannosum Bornet

See Gelidiella tenuissima J. Feldmann & Hamel.

222 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Gelidium pannosum Grunow

See Gelidiella tenuissima J. Feldmann & Hamel.

Gelidium pectinatum Montagne

Canaries (68: 191; 392; 547; 584).

'Ad . . . litora Africae borealis' (318).

[As Gelidium pectinatum (Schousboe) Montagne]

Canaries (177; 227).

Note. B0rgesen (68) knew of a single specimen found in the Canaries; he had not seen it. Citations of
authorities that include the name of Schousboe are in error (as, e.g., in Boudouresque et al., 1984). The
form used by the latter, 'Schousboe ex Montagne', is inaccurate since Montagne (403: 108) did not use the
whole of the name by which Schousboe had referred to the taxon (Teloedema pectinatum Schousboe) on a
specimen in Webb's herbarium.

Gelidium pristoides (Turner) Kiitzing
Namibia (348).

Note. See the notes to Gelidium micropterum Kiitzing and to Gelidium corneum sensu B0rgesen. There
has been past suggestion that Gelidium pristoides and Gelidium corneum auct. are directly related taxa;
Montagne (404: 77), for example, referred to pristoides as 'variete pristoides du G. corneum', having more
or less contemporaneously (403: 108) included 'Sphaerococcus corneus var. pristoides C. Ag.' in the
synonymy of his Gelidium pectinatum. Gelidium pristoides is widely and abundantly reported for many
localities in South Africa (Simons, 523: 259; Carter & Anderson, 579: 117; and others), so that whatever
the real status of the morphology to which the name is applied, it occurs often in the general area here
considered. A recent treatment of all non-parasitic genera of Gelidiaceae founded primarily on surface cell
morphology (Akatsuka, 1986) resulted in the establishment of a new genus, Onikusa, to include both
Gelidium pristoides (Turner) Kiitzing (the type species of Onikusa) and Gelidium japonicum (Harvey)
Okamura. Onikusa is derived from the Japanese name for G. japonicum. Since the study affects also the
entries for other genera of Gelidiaceae (Pterocladia; Suhria) found in the list area, it will be dealt with fully
later.

Gelidium pulchellum (Turner) Kiitzing

Gambia (296).

Senegal (121; 122).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gelidium pulchellum var. claviferum]

Canaries (38C).

Salvage Islands (38C).

Note. It is commonly the case with most authors that Gelidium pulchellum is taken to be a synonym of
Gelidium pusillum (Stackhouse) Le Jolis (q.v.), or at least to fall morphologically within the range taken to
represent the 'Gelidium pusillum-G. crinale-G. pulchellum series' (see John & Lawson, 296: 289-290).
Boudouresque et al. (1984: 44) and Wynne (1986) both believed at least pulchellum and pusillum to be
synonymous, Boudouresque et al. further not distinguishing for the latter the var. pulvinatum (C. Agardh)
J. Feldmann. Audiffred (38C: 173) cited the records from Canaries and Salvage Islands only in the context
of the entry for Ceramium echionotum J. Agardh, the 'host'. The species list provided in the same
publication included only Gelidium pusillum Stackhouse) Le Jolis, perhaps tending to imply that a
nomenclatural and taxonomic change had been overlooked.

Gelidium pulvinatum (Kiitzing) Thuret ex Bornet

Canaries (68; 490).

[As Gelidium pulvinatum Thuret]

Canaries (493).

Note. B0rgesen (68) found no specimens of this form amongst Canaries material, but it was listed by
Sauvageau (493). It seems probable (B0rgesen, 68) that G. pulvinatum is closely related to G. pusillum
(q.v.) and G. spathulatum (q.v.), all being forms of a single species. Schmidt (497: 110-111) relegated
Gelidium pulvinatum Kiitzing to Gelidium pusillum f. pulvinata (Kiitz.), a process initially carried out by
J. Feldmann (in Hamel, 1927) on the basis of C. Agardh's (19: 284) Sphaerococcus corneus o. pulvinatus.

Gelidium pusillum (Stackhouse) Le Jolis

Angola (352).

Ascension (7474; 475; 567).

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 223

Benin (288; 293; 350; 586).

Bioko (346; 350; 586).

Cameroun (269; 288; 337; 350; 374; 454; 460; 484; 500; 537; 586).

Canaries (13; 16; 38B; 38C; 38D; 68; 70; 71; 108; 128A; 191; 226; 227; 229; 236; 237; 253; 306B;

336; 351; 375; 379; 392; 486; 497; 499; 517; 556; 584; 585; 598; 604; 605; 608).

Cape Verde Islands (213).

Cote d'lvoire (287).

Gabon (213; 350; 586).

Gambia (350; 586).

Ghana (213; 288; 297; 350; 377; 491; 586).

Liberia (129; 287; 288; 350; 586).

Mauritanie (38B; 38C; 38D; 349; 7529; 556).

Salvage Islands (38B; 38C; 38D; 231; 375; 556; 556A; 598).

Sao Tome (288; 350; 586).

Senegal (38B; 38C; 38D; 59; 121; 213; 253; 336; 529; 556).

Sierra Leone (30; 213; 295; 336; 339; 350; 374; 378; 586).

Togo (288; 293; 350; 586).

Western Sahara (38B; 38C; 38D).

'Atlantico (Noruega - Cabo Verde. . . .)' (253; 604).

'Atlantique (de 1'Angleterre aux Canaries)' (33).

'Atlantischer Ozean, von den englischen Kiisten an siidwarts bis zur afrikanischen Kiiste und

den Kanaren' (499).

'Central Norway to Cape Verde' (172).

'extensive in warm & tropical seas' (81).

'Nordwestafrika' (499).

'Semble etre repandu dans toutes les mers chaudes' (188).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Tropical Africa (N. Gambia- Congo river)' (598).

Tropical West Africa' (611).

'Von den englischen Kiisten an sudwarts bis zur afrikanischen Kiiste' (497).

'Westafrika' (499).

[As Gelidium pusillum var. pulvinatum (C. Agardh) J. Feldmann]

Angola (352).

Canaries (518; 519; 584).

Gabon (294).

Gambia (296).

Liberia (129; 287; 288).

Mauritanie (349).

Senegal (121; 122; 529).

Sierra Leone (295).

[As Gelidium pusillum (Stackhouse)]

Canaries (5).

[As Gelidium pusillum Le Jolis]

Canaries (547).

[As Gelidium pusillum (Stark) le jolis var. pulvinatum (Ag.) Feldm.]

Senegal (50).

[As Gelidium pusillum var. conchicola Pice. & Grun.]

Senegal (121; 122; 529).

[As Gelidium pusillum var. minusculum Weber van Bosse]

Senegal (121; 122; 529).

[As Gelidium crinale (Turner) Lamouroux]

Mauritanie (252; 500; 535).

Western Sahara (476).

224 J. H. PRICE, D. M. JOHN & G. W. LAWSON

[As Gelidium reptans (Suhr) Kylin]

Mauritania (121; 122).

Namibia (348).

Sao Tome (93; 535).

Senegal (121; 122; 296; 529).

[As Gelidium corneum (Hudson) Lamouroux]

Angola (352).

[As Gelidium corneum (Hudson) Lamouroux var. davatum (Lamouroux) C. Agardh]

Canaries (401).

Note. According to P. S. Dixon (pers. comm., 1974), the taxon Gelidium pusillum (Stackhouse) Le Jolis
var. pulvinatum (C. Agardh) J. Feldmann should correctly be known as Gelidium crinale (Turner)
Lamouroux. Although Gelidium reptans (Suhr) Kylin has been recorded from Mauritanie, Namibia, Sao
Tome, and Senegal, and tentatively from Gambia, we here follow B0rgesen (81) and others in considering it
as no more than a variety of Gelidium pusillum. Sourie (529: 116), in commenting on material collected and
recorded as G. reptans, stated 'les exemplaries recoltes ne seraient peut-etre qu'une forme de G. pusillum'.
There is a similarly close relationship between G. foliosum of P. Dangeard and G. pusillum var.
pulvinatum, as indicated by John (288); see also the notes in the entry for G. foliosum. Material recorded
by Lawson, John & Price (352) from Angola as patches of short red algal 'turf of G. corneum (Hudson)
Lamouroux included a mixture of forms that, in isolation, would have been considered as representing
G. corneum sensu B0rgesen, G. pusillum (Stackhouse) Le Jolis and G. micropterum Kiitzing (q.v.).

Gelidium reptans (Suhr) Kylin
Note. See Gelidium pusillum Stackhouse) Le Jolis, especially the note to that species entry.

Gelidium rigidum (Vahl) Greville [or simply Greville]
See Gelidiella acerosa (Forsskal) J. Feldmann & Hamel.

Gelidium senegalensis Bodard (nomen) [or J. Feldmann mscr.]

Senegal (59).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gelidium senegalensis J. Feldm. mscr.]

Senegal (59).

Note. Included for completeness only, since no description or further data have been traced, and the
relationships of material to which the name has been applied are not clear. Bodard & Mollion (59) included
the form of the name used as heading here in their Table I (Baie de Goree), without description or further
mention in the text. Later, in their Table IIIB, they referred to what must be taken as the same taxon as 'G.
senegalensis J. Feld. mscr.' The name was presumably annotated on to a specimen or specimens by
Feldmann and taken up temporarily by Bodard.

Gelidium serrulatum J. Agardh
[As Gelidium serrulatum Ag. var.]
Angola (261; 262).

Note. Originally described from Venezuela and known from Trinidad. Accepted by Taylor (540: 357)
without comment, but not frequently employed as a name in the literature. Relationships of the material
here involved are not clear.

Gelidium sesquipedale (Clemente) Thuret in Bornet & Thuret

Canaries (68; 117; 139; 191; 227; 598).

Cape Verde Islands (122; 252; 598).

Guinea-Bissau (529).

Mauritanie (121; 122; 500; 529).

Senegal (122; 252).

Western Sahara (393; 394; 476; 529).

'de Inglaterra a Canarias' (517).

'does not extend [southwards] into Gulf of Guinea' (487).

'English coast southwards to Canary Islands' (68).

'French Channel coast to Mauretania' (172).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Siidengland bis Mauritanien.' (567).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 225

[As Gelidium sesquipedale (Clemente) Bornet & Thuret]

Mauritanie (349).

Western Sahara (349).

[As Gelidium sesquipedale (Turner) Thuret in Bornet & Thuret]

'D'Angleterre aux Canaries' (89).

[As Gelidium sesquipedale (Turner) Thuret]

Cape Verde Islands (191; 423).

Mauritanie (252).

'Atlantique: . . . sur les cotes anglaises et descend jusqu'a la pointe du Cap Blanc, en

Mauritanie.' (222).

'Atlantique (de I'Angleterre a la Mauritanie' (33).

'du sud de I'Angleterre a la Mauritanie' (195).

[As Gelidium sesquipedale Thuret in Bornet & Thuret]

'dall' Inghilterra alle Canarie' (390).

[As Gelidium sesquipedale Thuret]

Canaries (117; 118).

Cape Verde Islands (38).

'De I'Angleterre aux Canaries.' (38).

[As Gelidium resquipedalex]

Canaries (237).

[As Gelidium corneum Lamouroux var. sesquipedale J. Agardh]

Cape Verde Islands (38).

'De I'Angleterre aux Canaries' (38).

[As Gelidium corneum (Hudson) Lamour. var. sesquipedale Clemente]

Canaries (401).

[As Gelidium corneum (Hudson) Lamouroux var. sesquipedale Agardh]

Cape Verde Islands (408).

[As Gelidium corneum (Hudson) Lamouroux var. sesquipedale Greville]

Canaries (141A).

Note. B0rgesen did not find this species on the Canaries; he had only two previous records, from
Montagne (401: 158) and from the herbarium of H. C. Lyngbye. He therefore referred to the species there
as 'probably rare'. Although the Western Sahara record in Lawson & John (349) is supported by an original
discovery from the Cap Blanc area, the statement for Mauritanie is secondary, being based on various of
the data included here. See also the notes to Gelidium arbmcula Bory ex B0rgesen.

Gelidium spathulatum (Kiitzing) Bornet

Canaries (16; 68; 191; 375; 517; 584).

'Atlantique (de I'Angleterre aux Canaries . . .)' (188).

'Atlantique (du golfe de Gascogne aux Canaries)' (33).

[As Gelidium spathulatum Kiitzing]

Canaries (493).

Note. Bornet (89: 108 [268]) suggested that this taxon covered a form of G. crinale (q.v.), not a good and
separate species. B0rgesen (68) expressed general agreement with that view. For extended comments on
the status of G. spathulatum, see Feldmann & Hamel (195: 115) and Ardre (33: 71-72 [203-204]).

Gelidium spinulosum (C. Agardh) J. Agardh

Canaries (2; 68; 191; 221; 227; 401; 431; 499).

Western Sahara (393; 394; 476).

'Nordwestafrika' (499).

'outre le Maroc aux Canaries' (221).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gelidium spinulosum J. Agardh]

Canaries (242).

'De Cadix aux Canaries' (89).

226 J. H. PRICE, D. M. JOHN & G. W. LAWSON

[As Gelidium spinulosum J. Agardh ex J. Agardh]

'In Oceano atlantico calidiore' (27).

[As Gelidium corneum (Hudson) Lamouroux var. spinulosum C. Agardh]

Canaries (318; 401).

'Ad oras Africae' (318).

Note. See the notes to Gelidium microdon Kiitzing for comments on conspecificity of these two and other
species. Although Acuna Gonzalez (2) stated 'perteneciente al continente Africano', Papenfuss (431: 174)
excluded the species from the South African marine flora.

Gelidium versicolor (S. Gmelin) Lamouroux

Angola (298; 352; 611).

Canaries (253; 306B; 582; 583).

'West African coast to the north of the Gulf of Guinea' (352) .

[As Gelidium cartilagineum (L.) Gaillon]

Angola (298; 393; 394; 424; 425).

Note. For records of what was previously deemed the same taxon from the Canary Islands, see Gelidium
canariensis (Grunow) Seoane-Camba. Although Simons in Day (523: 528) recorded the South African
distribution as only 'Port Nolloth to Port Elizabeth', as usual in conditions of heavy surf on rock, the
presence of strong populations in Angola and of the acknowledged records from Port Nolloth, near the
Namibia/South Africa border, would certainly suggest that any rocky areas with surf along the Namibian
shore-line could also bear populations. See also the general note to Gelidium and that to Gelidium
arbuscula Bory ex B0rgesen.

Gelidium spp.

Angola (298; 352).

Ascension (474).

Cameroun (337; 344; 393; 394; 533).

Canaries (2; 5; 8; 66; 71; 214; 237; 351; 393; 394; 486; 493; 583).

Cape Verde Islands (423).

Cote d'lvoire (287; 290; 394).

Ghana (42A; 287; 290; 297; 299; 338; 340; 344; 376; 393; 394; 487; 567).

Guinee (344; 384; 393; 394; 529).

Liberia (287).

Mauritanie(349;529).

Namibia (348; 522).

Nigeria (344; 393; 394).

Senegal (121; 123; 344; 393; 394; 411; 529).

Sierra Leone (339; 344; 393; 394).

Western Sahara (349; 393; 394; 395; 476).

'Tropical west Africa' (611).

West Africa (290; 344).

Gigartina acicularis (Roth) Lamouroux

Bioko (346; 350; 586).

Cameroun (172; 243; 269; 337; 344; 350; 447; 537; 586).

Canaries (2; 38B; 38C; 38D; 128A; 172; 227; 253; 306B; 392; 7490; 584; 598; 604).

Gambia (350; 586).

Ghana (42A; 153; 269; 291; 335; 338; 340; 344; 350; 374; 537; 586).

Guinee (344; 350; 537; 586).

Liberia (350; 586).

Mauritanie (38B; 38C; 38D; 349; 500).

Salvage Islands (38B; 38C; 38D; 598).

Sao Tome (93; 350; 586).

Senegal (38B; 38C; 38D; 53; 55; 59; 344; 537).

Sierra Leone (350; 586).

Western Sahara (36B; 38C; 38D).

'[Associations on] , , , African west coast' (374).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 227

' Atlantico (Inglaterra - Sudafrica . . .)'(253).

'Atlantik Kiiste von Afrika (. . . Golf von Guinea)' (270).

'Atlantique (de 1'Angleterre a la Mauritanie)' (33).

'British Isles to the Cameroons' (172).

'British Isles south to Cameroun' (243).

'from the British Isles . . . south to Cameroun' (447).

'Gran Bretana a Camerun' (604).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara].' (598).

Tropical Africa (N. Gambia- Congo river)' (598).

Tropical West Africa' (611).

'widespread in warm temperate and tropical seas' (350; 586).

[As Gigartina acicularis (Wulfen) Lamouroux]

Angola (352).

Cameroun (173).

Canaries (13; 16; 70; 108; 189; 191; 375; 379; 439; 441; 444; 489; 499; 517).

Gambia (296).

Ghana (173).

Guinea-Bissau (7529).

Guinee(?529).

Liberia (129).

Mauritanie (349; 529).

Salvage Islands (215).

Sao Tome (252).

Senegal (529).

Sierra Leone (30; 295).

Western Sahara (349).

'Atlantico (de Inglaterra a Canarias)' (517).

'Atlantique: depuis les cotes anglaises jusqu'en Mauritanie' (222).

'Atlantique (de 1'Angleterre aux Canaries . . . Sans doute repandu dans toutes les mers

chaudes.'(189).

'Cosmopolite/subcosmopolite' (529) .

'D'Angleterre aux Canaries' (89).

'de 1'Irlande et de la Grande-Bretagne au Cameroun' (178).

'From the English coast southwards to the Canary Islands' (70).

'Golf von Guinea' (270).

'Nordwestafrika' (499).

'Seems to occur in all warmer seas.' (63).

[As Fucus acicularis Turner]

Ghana (271).

[As Sphaerococcus acicularis (Wulfen) C. Agardh]

'In mari Atlantico, ab oris Galliae septentrionalis ad Africam.' (588).

Note. B0rgesen's (70) specimens were all small and sterile. The Hornemann (271) record from 'Danish
Guinea' [present day Ghana] is attributed here on nomenclatural equivalence only, but there is no reason
to doubt its presence in Ghana, since modern records are plentiful. The P. E. Isert collection examined by
Hornemann needs examination for full confirmation, but the relevant material may have been destroyed at
C [Copenhagen] during the 1807 fire that resulted in the loss of some Isert specimens. There may be a very
early (1803) Canaries record concealed amongst the heterogeneity of material described under Fucus
perforatus Bory (90: 305, pi. V, figs 1 A, B, C); see the entries for Carpococcus perforatus (Bory) J. Agardh
and Laurencia perforata Montagne.

Gigartina bracteata (S. Gmelin) Setchell & Gardner
Namibia (36B;522A).

Note. See 36B: 314 for discussion by Wynne of various morphological, taxonomic, and nomenclatural
aspects.

228 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Gigartina burmannii (C. Agardh) J. Agardh
See Gigartina stiriata (Turner) J. Agardh.

Gigartina confervoides (L.) Lamouroux [or simply Lamouroux]
See Gradlaria verrucosa (Hudson) Papenfuss.

Gigartina cylindrica Despreaux

Note. See Gradlaria dura (C. Agardh) J. Agardh. The entry in Montagne (401: 160) for Gigartina dura
Desmazieres states under 'Hab.': 'Gigartina cylindrica Despr. in schedula.'

Gigartina dura Desmazieres

See Gradlaria dura (C. Agardh) J. Agardh.

Gigartina elegans Greville in St-Hilaire
See Gigartina teedii (Roth) Lamouroux.

Gigartina fastigiata J. Agardh

See Gigartina scutellata (Hering) Simons in Seagrief .

Gigartina flagelliformis (Sonder) Sonder

'Ad oras Africae occidentales: Lenormand!' (326).

Note. Neither the alga being referred to nor the location from which it came are clear, 'West Africa'
being referred to. Sonder (1845: 55) originally described his taxon in Polyides and certainly the plant
illustrated in Kiitzing's (326: pi. 5, no. 4030, figs c, d) 1868 presentation has much in common with
Poly ides/ Furcellaria, rather than with Gigartina, where Sonder (1846) recombined it. The 1868 reference
to the species is the first time that West Africa is referred to, all previous details concerning material being
from 'Novam-Hollandiam'. Presumably Kiitzing received or saw Lenormand Herbarium material citing
the African data. If Kiitzing's drawing is an accurate representation of the Lenormand material, then it has
to be taken that the entry is generically mis-placed in Gigartina. In 1869, when Kiitzing again (pi. 18, figs c,
d, no. 4248) employed the name Gigartina flagelliformis of Sonder, the plant illustrated appears closer to
Gradlaria then to either Gigartina or Poly ides/ Furcellaria. Kutzing then stated (p. 7) lnec Kg. Tab. ph.
XVIII. tab. 5. ' and cited only 'Novam Hollandiam', remarking that the structure was still nearer to Polyides
or Furcellaria. J. Agardh (24: 283, no. 38) maintained the taxon in Gigartina, but commented that the
internal structure was like Poly ides/ Furcellaria and cited only New Holland as location. De Toni (131: 227)
commented 'Videtur forsan Rhabdoniae species'.

Gigartina griffithsiae (Turner) Lamouroux [or simply Lamouroux]
See Gymnogongrus griffithsiae (Turner) Martius.

Gigartina perforata (Bory) De Toni [or (Bory) J. Agardh]
See Carpococcus perforatus (Bory) J. Agardh.

Gigartina pistillata (S. Gmelin) Stackhouse
Canaries (70; 191; 227; 229; 306B; 375; 489; 490; 499; 517; 598).
Mauritanie (245B; 252; 349; 500; 529).
Senegal (59; 394).
Western Sahara (349).
'Atlantico (de Inglaterra a Canarias)' (517).
'Atlantique (de 1'Angleterre au Rio de Oro)' (33).
'Atlantique: depuis les cotes britanniques jusqu'au Rio de Oro' (222).

'Atlantischer Ozean, von der englischen Kiiste an siidwarts bis zur nordwestafrikanischen.'
(497).

'British Isles to South Africa' (172).

'depuis le sud de la Grande-Bretagne jusqu-au Senegal (Bodard, com. verb.)' (173).
'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).
'From the English coast southwards to the Canary Islands.' (70).
[As Gigartina pistillata Stackhouse]
'de 1'Angleterre aux Canaries' (89).
Note. The record by two of us (Lawson & John, 349) of this taxon from Western Sahara is based on Ardre

RHODOPHYTA (PLORIDEAE) OF TROPICAL AFRICA 229

(33); we did not collect the alga there. B0rgesen (70) based his comments solely on the report by Bornet
(89: 273); there were no specimens in Herb. Thuret, nor did he find it.

Gigartina pygmaea Lamouroux

See Chylocladia reflexa Lenormand in Desmazieres [Gastrodonium reflexum (Chauvin) Kiitz-

Gigartina radula (Esper) J. Agardh
Namibia (161; 348; 437; 500; 522; 522A; 523).

Note. Unusually, with the exception of the purely secondary reports in Schmidt & Gerloff (500), all the
above statements of record are at least partly based on original observation and collection. This taxon is
also well known from South Africa (570) and from many parts of the Southern Ocean system.

Gigartina scabiosa (Kiitzing) Papenfuss

See Gigartina scutellata (Hering) Simons in Seagrief .

Gigartina scutellata (Hering) Simons in Seagrief
Namibia (522 A).

[As Gigartina scabiosa (Kiitzing) Papenfuss]
Namibia (348; 522).

Note. Both the above recordings are based, at least in part, on original observations and collections. The
species is also widely known in South Africa, whence it was first described; there are nomenclatural
problems, the nature of which is clear from the entry on G. scutellata in Seagrief (570: 31). Clearly, Simons
had decided that, despite contrasting opinions held elsewhere in regard to synonymy in at least parts of the
application (and perhaps the concept) of the name Chondrus scutellatus Hering, it is in fact the same plant
that is being referred to. If this is so, then the plant hitherto mostly known as Gigartina scabiosa (Kiitzing)
Papenfuss has an earlier valid name that must be invoked. Hence, the recombination proposed by Simons
of Sphaerococcus (Chondrus) scutellatus Hering (1841: 91) in Gigartina, resulting in the binomial used as
heading to the present entry. The inclusion of these data in Seagrief s list (570: 31) was clearly based on
information provided by Simons in a letter to Seagrief, doubtless with Simons's permission. He (the latter)
presumably expected his paper on Trematocarpus (Sarcodiaceae) in southern Africa and the exclusion of
Chondrus scutellatus Hering to have appeared before that of Seagrief. Despite the delay in publication of
the Seagrief check-list (the correspondence with Simons was in 1982; Seagriefs paper did not appear until
1984) no further publication details of the Simons work were given, implying that it had not appeared by
then. In view of both previous correspondence and proximity, it is likely that Seagrief would have had his
attention drawn by Simons to its appearance. We have searched carefully for any sign of the publication but
have been unable to trace it. Hence, the recombination is cited as above, details provided by Seagrief
having been adequate to validate Simons's proposed changes. Seagriefs suggested synonymy of G.
scutellata included Gigartina fastigiata J. Agardh [non Postels et Ruprecht].

Gigartina stellata (Stackhouse in Withering) Batters

See Mastocarpus stellatus (Stackhouse in Withering) Guiry in Guiry, West, Kim & Masuda.

Gigartina stiriata (Turner) J. Agardh

Namibia (348; 437; 522; 522A; 523).

Note. A further, previously separate, taxon in Gigartina. G. burmannii (C. Agardh) J. Agardh, has been
shown to be the tetrasporophyte of G. stiriata.

Gigartina teedii (Roth) Lamouroux

Angola (500).

Cape Verde Islands (589; 598).

Guinea-Bissau (529).

Mauritanie (7344; 349; 7529).

Namibia (36B; 161 ;522A).

Senegal (47; 50; 52; 55; 59; 399; 529; 531).

Western Sahara (349).

'British Isles to Angola . . . Cape Verde Islands' (172).

'Nordwestafrika' (499).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Westafrika' (499).

230 J. H. PRICE, D. M. JOHN & G. W. LAWSON

[As Gigartina teedii Lamouroux]
Cape Verde Islands (38).
Senegal (38; 122; 408).
[As Gigartina teedii Roth]
Cape Verde Islands (259).
[As Gigartina teedii J. Agardh]
Angola (41; 42).

Note. There seems no real evidence for the presence of this species in the Canary Islands; Dangeard
(117; 118) specifically stated that it was missing from there. A certain number of expressed uncertainties
have been ignored here - Bodard (47), for instance, recorded what may well have been the taxon as
'Gigartina aff. teedii', and Dangeard (122), more verbosely, commented for Senegal 'un Gigartina a
ramification assez regulierement pennee, a branches et rameaux aplatis, qui se presente, soit a 1'etat sterile,
soit porteur de nombreux cystocarpes . . . rapelle beaucoup le G. teedii . Askenasy's record was
rationalised from probabilities, not fully recorded; he (38) stated, 'n'ont pas ete rapportees des iles du Cap
Vert; il est tres probable qu'elles y croissent aussi et qu'on les trouvera plus tard'. Ardre (33: 130-131)
stated the southern limit of the G. teedii distribution range to be Morocco. On the subject of the
relationships between G. teedii and Gigartina elegans Greville in St.-Hilaire, see Cordeiro-Marino (108:
77-78). For general background, see Guiry (589).

Gigartina spp.

Cameroun(393;394).
Ghana (344; 393; 394).
Guinee(393;394).
Mauritanie(393;394).
Namibia (348; 438; 522; 611).
Nigeria (393; 394).
Senegal (393; 394).
Sierra Leone (393; 394).

Note. The records from Namibia could well include any (or a mixture) of Gigartina radula, G.
scutellata, or G. stiriata (q.v.) in addition to other unidentified forms.

Ginnania furcellata Montagne

Note. See Scinaia forcellata Bivona-Bernardi. The genus Ginnania is based on material from the
Canaries, published by Montagne (401: 162), repeated elsewhere by the same author (402) and subse-
quently by others (e.g. Kiitzing, 318). Montagne later (408) cited the species with the addition '(Turn, sub
Viva).' According to Papenfuss (434: 281), the report of Ginnania furcellata from the Cape of Good Hope
(Harvey, 354: pi. 69) related to Pseudogloiophloea (now Scinaia} capensis. This agrees with the absence of
records more southerly than the Congo for any species of Scinaia in this area, 5. capensis never having been
reported. It is possible that the latter occurs in Namibia, and perhaps Angola.

Gloiophloea verae Dickinson

See Scinaia verae (Dickinson) Huisman.

Goniolithon

Afonso-Carrillo et al. (582: 25) accepted the characterization of this genus in the original
(Foslie, 1898a) sense, as proposed by Cabioch (1972). All statements emanating from Afonso-
Carrillo and/or colleagues must therefore be viewed in that light. The status of the genus
Goniolithon Foslie has been explained by Johansen (1981: 218); Foslie's original concept of 1898
was later rejected by its author (1900<f). The later changed concept was very different from the
first version, and Setchell & Mason (1942) suggested that the second concept should be
segregated as the new genus Neogoniolithon (type N. fosliei). Goniolithon, the first concept,
remains (type G. papillosum) . For a fuller explanation, see Woelkerling (1988). There is little or
no authentic recording remaining for the list area under Goniolithon.

Goniolithon accretum Foslie & Howe

See Neogoniolithon accretum (Foslie & Howe) Setchell & Mason.

Goniolithon boergesenii Foslie

See Hydrolithon boergesenii (Foslie) Foslie.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 231

Goniolithon brassica-florida (Harvey) Foslie

Note. Often referred to Neogoniolithon, as N. brassica-florida (Harvey) Setchell & Mason. See the note
to Neogoniolithon mamlllare (Harvey) Setchell & Mason.

Goniolithon byssoides (Lamarck) Foslie [or simply Foslie]

See Titanoderma byssoides (Lamarck) Chamberlain & Woelkerling.

Goniolithon mamillare (Harvey) Foslie

See Neogoniolithon mamillare (Hauck) Setchell & Mason.

Goniolithon mamillosum (Hauck) Foslie [and formae]

See Neogoniolithon mamillosum (Hauck) Setchell & Mason and notes to Neogoniolithon

mamillare (Hauck) Setchell & Mason.

Goniolithon orotavicum Foslie

See Neogoniolithon orotavicum (Foslie) Lemoine.

Goniolithon papillosum (Zanardini ex Hauck) Foslie

See Titanoderma papillosum (Zanardini ex Hauck) J. Price, D. John & G. Lawson.

Goniolithon polycephalum (Foslie) Afonso-Carrillo

See Titanoderma polycephalum (Foslie) Woelkerling, Chamberlain & Silva.

Goniolithon subtenellum Foslie

See Lithophyllum subtenellum (Foslie) Foslie.

Goniolithon sp.

Canaries (212).

Note. Recorded by Foslie & Printz (212: pi. XLIII) as almost entirely covering Archaeolithothamnion
africanum Foslie [Sporolithon africanum (Foslie) Afonso-Carrillo] 'by Goniolithon or Lithophyllum sp.'.
In practical terms, the alga referred to could well have been of Neogoniolithon spp. (q.v.). We are currently
unable to assign the plants to one or other genus/species.

Gottoniella fusiformis B0rgesen
See Cottoniella fusiformis B0rgesen.

Gracilaria and Polycavernosa

Gracilaria Greville is a very large, widely distributed, and complex genus, with more than 100
species of recent citation. Taxonomically difficult, the genus has been subject over the years to
the occasional vogue of new species recognition and description, largely on the grounds of
occurrence in different geographical areas. The literature therefore abounds with specific
epithets, many of which will probably in due course prove to be quite superfluous. This general
situation has been worsened by partial revisions of which many have appeared more obscuratory
than elucidatory. Nevertheless, many valuable and detailed studies have been undertaken in
recent years, and are still in progress, on various aspects of generic or specific taxonomic
practice. See, for example, Bird & McLachlan, 19820, 45, 46, 1984; Edelstein et al., 1978;
Yamamoto, 1978; McLachlan & Bird, 318A; Oliveira, 1984; Oliveira filho & Plastino, 1982,
1984; Oliveira, McLachlan & Bird, 1982; Oliveira, 1983; Oliveira, Bird & McLachlan, 1983;
Abbott & Norris, 1985; Bird, van der Meer & McLachlan, 1982; Gargiulo, De Masi & Tripodi,
1985; Bird & Oliveira, 592; and many others. A useful bibliography is presented in McLachlan &
Bird, 593. The extent of current work reflects both widespread and abundant occurrence of the
genus and its importance as a source of phycocolloid. Because of the existence of such detailed
studies, rationalization here of records is only undertaken to the extent dictated by elimination
of clear nomenclatural anomaly or by application of synonymy unequivocally established by the
recent studies. Further development of work currently in progress may result in considerable
change in both the basic criteria employed and the names to be applied as a consequence. Bird &
McLachlan (1984: 41) have cogently summarised problems in this genus in stating: 'Although
Gracilaria Greville is a reasonably well-defined genus of the Gigartinales, the taxonomy of its
species is notoriously chaotic. It is becoming increasingly apparent that certain minor but
fundamental differences among morphologically similar taxa merit consideration if species are

232 J. H. PRICE, D. M. JOHN & G. W. LAWSON

to be correctly defined. ... it is equally important to sample widely for such features, to
determine their uniformity throughout populations and under a variety of environmental
conditions'. More recently, McLachlan & Bird (593: 27-28) have suggested that: 'We can expect
little meaningful progress, either in understanding the genus or being able to exploit its species,
until there has been a proper resolution of many taxonomic problems . . . for most of the species
of commerce it is difficult to assign a specific name with any assurance'. This applies equally to
non-commercial species.

Sources of much useful information on the generic group including Gracilaria and the
segregate Polycavernosa are the Sections IV and V (pp. 67-162) of Abbott & Norris (1985). We
have generally followed the ideas expressed there and have made use of appropriate comments
from the text. On the current opinions concerning acceptability of Polycavernosa as distinct
from Gracilaria, see the generic notes to the entry for Polycavernosa. Because of the close,
complex and not unanimously-accepted nature of the relationships between Gracilaria and
Polycavernosa, the latter has been included here, with all its relevant potential species, although
out of its alphabetical order. Cross-references will be given as appropriate later, in alphabetical
position.

Gracilaria angustissima (Harvey) Bodard

Senegal (51; 52).

[As Gracilaria angustissima]

Senegal (50).

Note. The relationships of the material on which these records are based and that often referred to as
Gracilaria foliifera var. angustissima (Harvey) W. Taylor are not clear. Bodard (52) presented the
combination used as species entry heading here as the main head for his species entry, qualifying it as 'nom.
prov.'. He included in his synonymy both Gracilaria foliifera var. angustissima (Harvey) W. Taylor [1937]
and Gracilaria multipartita var. angustissima J. Agardh [1876]. Since Bodard had no fertile specimens
available, he took for reference F. S. Collins no. 610, a cystocarpic Gracilaria close in form to Bodard's
other material and clearly neither of the taxa with which the latter was contrasting 'G. angustissima''; these
taxa were known by him as Gracilariopsis tridactylites and G. disputabilis . Although he deduced that 'G.
angustissima' was much closer to G. dentata than to the other species treated, Bodard's last sentence stated:
'Cette espece est encore mal connue, c'est pourquoi nous maintenons provisoirement ce nom.'

Other complications exist, in that Bodard both previously (51) and subsequently (Bodard & Mollion, 59)
had in places referred to what must (presumably) be the same entity as that here concerned as G. dentata
var. angustissima (51) and 'Gracilaria dentata P. Dang. var. angustissima Bodard comb, nov.' (59: Table I -
Goree). Since the latter paper is the latest (1974), it could be considered that this statement represents
Bodard's final view of the matter, in which case acceptance as a valid taxon would require transfer to
Polycavernosa dentata (q.v.). Since re-examination of original specimens is required to establish both firm
relationship to Gracilaria dentata sensu stricto and validity of any transfer to Polycavernosa, we have
retained the present arrangement pending further work.

A little earlier than any of the works cited above, Bodard (50: 83-84, 1966) had passingly referred to a
record for Pointe de Sarene (Senegal) under Gelidiopsis variabilis: 'A premiere vue, on pourrait confondre
cette espece avec des Gracilaria augustissima'. We have taken the view that this is orthographic or
typographic error for Gracilaria angustissima; the record therefore appears here.

Gracilaria urinata (C. Agardh) J. Agardh

Canaries (70; 128A; 184; 191; 227; 252; 375; 392; 547; 598; 610).

Mauritanie(184;252;349).

Western Sahara (349).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gracilaria armata J. Agardh]

Canaries (547).

Note. See also the notes to Gracilaria dura (C. Agardh) J. Agardh. For recent comments on the taxon G.
armata in the NE. Atlantic/Mediterranean area, see Gargiulo, De Masi & Tripodi (1985).

Gracilaria augustissima sine auct.

See Gracilaria angustissima (Harvey) Bodard.

Gracilaria bursa-pastoris (S. Gmelin) Silva

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 233

Cape Verde Islands (598).

Mauritanie (349).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'British Isles to Senegal . . . Cape Verde Islands' (172).

[As Gracilaria compressa (C. Agardh) Greville]

Cape Verde Islands (252 ; 448 ; 449) .

Mauritanie (252).

Sao Tome (25).

Senegal (529).

'warmeren atlantischen Ocean' (506).

[As Gracillaria compressa Greville]

Cape Verde Islands (38).

[As Gracilaria compressa C. Agardh]

'Warmer Atlantic' (410).

Note. J. Agardh (25: 594) recorded G. compressa as 'ad insul. S:t Thomas (Hb. Binder!)'. Piccone (449)
indicated that the systematic position of his material from Cape Verde Islands was uncertain. The mention
of this species from Mauritanie by two of us (see Lawson & John , 349) was secondary and based on that for
G. compressa in Hariot (252). For recent comments on aspects of this taxon in the NE. Atlantic/
Mediterranean area, see Gargiulo, De Masi & Tripodi (1985).

Gracilaria camerunensis Pilger

Cameroun (139; 350; 454; 500; 586).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Senegal (59; 394; 399; 611).

Tropical Africa (N. Gambia- Congo river)' (598).

'in warm temperate and tropical parts of the eastern Atlantic Ocean' (350; 586).

Taire se limite au golfe du Benin et a la Mauritanie' (59).

'plus au sud dans le golfe du Benin' (59).

[As Gracilariopsis camerunensis (Pilger) Bodard]

Cameroun (51).

Senegal (51; 52).

'surement presente dans tout le Golfe du Benin' (52).

Note. De Toni (139: 250-251) based his entry directly on Pilger's (454) treatment of the Ledermann
collection from 'Bodje', Cameroun. Bodard (52) indicated the taxon to be very near to Gracilariopsis
sjoestedtii [= Gracilaria lemaneiformis (Bory) Weber-van Bosse], although he did not see the type of
Pilger's Gracilaria camerunensis.

Gracilaria cervicornis (Turner) J. Agardh

?Bioko(346;586).

Cameroun (586).

Ghana (586).

'in warm temperate and tropical seas' (586).

Tropical Africa (N. of Gambia - Congo river)' (598).

[As Gracilaria ferox J. Agardh]

?Bioko (350).

Cameroun (269; 337; 350; 537).

Ghana (108; 350; 418; 419).

'in warm temperate and tropical seas.' (350).

[As Gracilaria sp.]

Cameroun (337).

Note. Oliveira filho, McLachlan & Bird (1982) and Oliveira filho, Bird & McLachlan (1983) have
indicated that G. cervicornis (Turner) J. Agardh and G. ferox J. Agardh represent a morphological
complex with entities distinguishable at extremes but with overlapping gradations even within populations.
Apart from the vegetative appearance, similarities include structure of the tetrasporangia, spermatangia,
cystocarps, and internal anatomy, and poor gelling of agar extracts. Hence, Oliveira f. et al. have reduced
the complex to the single species G. cervicornis, G. ferox being reduced to synonymy.

234 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Morphologically, Gracilaria domingensis Sender ex Kiitzing [in syn.] shares certain similarities with G.
cervicornis, but the two taxa are said to be separable as distinct species since G. domingensis has its
spermatangia in deep crypts [cf. shallow conceptacles], with its cortex being thin and of subquadrate cells
[cf. multilayered cortex of radially elongated cells]. This has led us to decide that G. domingensis should be
recombined in Polycavernosa (q.v., also P. dentata J. Agardh).

There has not been total agreement on the conspecificity of G. cervicornis and G. ferox. Cordeiro-
Marino (108: 61) had expressed an opinion that these taxa were both different from each other and from G.
domingensis, by virtue of the ferox subcylindrical principal axis and its branching in more than one plane.
G. ferox was described from Pernambuco (Brasil). Cordeiro-Marino's (108) Ghana record was derived
directly from Ohmi (419). Similarly, Hoppe's (269) Cameroun report was based on Lawson (337).

Recent comments on G. cervicornis in the north-east Atlantic and Mediterranean are presented in
Gargiulo, De Masi & Tripodi (1985).

See also the notes at Gracilaria multipartita (Clemente) Harvey.

Gracilaria compressa (C. Agardh) Greville [or Greville]
See Gracilaria bursa-pastoris (S. Gmelin) Silva.

Gracilaria confervoides (L.) Greville

See Gracilaria verrucosa (Hudson) Papenfuss.

Gracilaria corallicola Zanardini

See Gracilaria multipartita (Clemente) Harvey and Gracilaria lacinulata (Vahl) Howe.

Note. This taxon is not the Rhodymenia corallicola of Ardissone (q.v.). For recent comments on the
taxon in the north-east Atlantic and Mediterranean areas, see Gargiulo, De Masi & Tripodi (1985).

Gracilaria corticata J. Agardh
[As Fucus aeruginosus Turner]
Ghana (271; pro parte)

Note. For the background to the inclusion here of this taxon, see the notes to Gracilaria multipartita
(Clemente) Harvey [= Gracilaria foliifera (Forsskal) B0rgesen]; these notes also present explanation of
the use above of 'pro parte'. The situation may require revision in the light of any established or deduced
identity of the P. E. Isert Ghana collections, on which the above published information was based.
Unfortunately, the Isert material was probably destroyed in a fire in K0benhavn (1807). What is really
represented by the taxon G. corticata J. Agardh is not clear; from available information, it seems highly
likely to be little more than a form of G. multipartita, although that equally difficult concept Gracilaria
[Polycavernosa] dentata could also have been involved. Many authors have expressed reservations as to
significance and/or name application in some or all of these taxa. Jaasund (19776: 420) outlined the
confusion in application of the names G. corticata and G. multipartita/ G. foliifera, at least in Indian Ocean
material. Askenasy (38: 168), commenting under his G. dentata J. Agardh heading, indicated: 'Je possede
un exemplaire de cette algue provenante de 1'herbier Lenormand et etiquete: Gracilaria corticata J. Ag.
Ins. Sal. Cap de Verde, Forbes I.' See also the notes to Polycavernosa (Gracilaria) henriquesiana (Hariot)
Chang & Xia.

Gracilaria damaecornis J. Agardh
Senegal (51; 52).
[As Gracilaria type D]
Senegal (529).

Note. Bodard's (52: 36) entry is wholly based on Sourie's plants collected from Cap de Naze. Specimens
are apparently quite rare. Bodard's (loc. cit.) final sentence reads 'C'est le Gracilaria type D de Sourie'.
The latter has also been entered at Gracilaria spp. for completeness.

Gracilaria debilis (Forsskal) B0rgesen

See the notes to Polycavernosa debilis (Forsskal) Fredericq & Norris and Polycavernosa

(Gracilaria) henriquesiana Hariot.

Gracilaria dendroides Gargiulo, De Masi & Tripodi

See the note to Gracilaria verrucosa (Hudson) Papenfuss.

Gracilaria dentata J. Agardh

Note. See also Polycavernosa dentata (J. Agardh) G. Lawson & D. John and P. henriquesiana (Hariot)
Chang & Xia. Xia & Abbott (571: 161, footnote 1) have indicated that: 'If Gracilaria dentata J. Agardh as

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 235

interpreted by Lawson and John (350) proves to be the same species as P. henriquesiana , its transfer to
Polycavernosa should be made and that binomial used as it will be the earlier name'. This course of action
has been carried out by Lawson & John (586), but according to Abbott later (in litt. to DMJ, 11/6/1986) the
isotypes of (and customs of application of names of) G. dentata and G. henriquesiana do not represent the
same entity.

It seems likely that workers generally on West African material will not entirely agree with Abbott as to
the customs of name application there, more especially as a rather small proportion of the material
determined usually emerges as male and as vegetative characteristics of specimens so overlap that intuitive
evaluation is often the principal basis for naming. Ghanaian populations, especially, show clinal morpho-
logical variation resulting from collecting season; the correlation of low tidal periods in the day with season;
growth on abrupt scarp or gentle dip slopes relative to aspect and wave-impact; microniche (e.g., pools;
crevices) within major habitat; extent and form of perennation.

Similarly, there may be little typificatory basis for the so-definite statement of difference by Abbott (in
litt.). Steentoft (pers. comm., March 1987), examining the type material of Gracilaria henriquesiana
Hariot (Ribeiro 14; Coimbra [COI]), found both male and female plants present; the males possessed
spermatangia developed within deep, variably- but often multi-locate crypts in the thallus, so that the
species has to be referred to Polycavernosa rather than Gracilaria if the generic division is accepted. If
Ohmi's (419) study of the Ghanaian material available to him is representative of the whole West African
range, attribution by name application confirms placement in Polycavernosa.

Since G. dentata has already been recombined in Polycavernosa (586), it therefore appears there as
records in this listing; earlier establishable misdeterminations as G. henriquesiana (Polycavernosa henri-
quesiana) also appear under Polycavernosa dentata, with the exception of Sao Tome material, for which
evidence is thus far lacking as regards spermatangial form. Sao Tome elements therefore remain as
Polycavernosa (Gracilaria) henriquesiana, pending clarification.

Gracilaria dichotomo-flabellata P. Crouan & H. Crouan in Schramm & Maze

Note. See the entries for Gracilaria feldmannii Bodard and Gracilaria mammillaris (Montagne) Howe.
Schneider (1975: 643, 645) has shown that the above authority citation is a nom. nud., the correct
publishing authorities form being 'P. & H. Crouan ex Collins & Hervey'.

Gracilaria disputabilis (Bodard) Bodard

Cote d'lvoire (288; 350; 586).

Ghana (288; 350; 586).

Senegal (51; 350; 586).

Senegal (Casamance) (51; 350; 586).

'in warm temperate and tropical parts of the eastern Atlantic Ocean' (350; 586).

Tropical Africa (N. Gambia - Congo river)' (598).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gracilariopsis (?) disputabilis Bodard]

Senegal (Casamance) (51; 52)

[As Gracilariopsis (?) tridactylites Cr. (= J. Ag.?) M. Bodard, comb, nov.]

Senegal (Casamance) (52).

's'etend audela de la Casamance [Senegal] jusqu'en Cote d'lvoire (recolte Sourie).' (52).

[As Gracilaria tridactylites Bodard]

Senegal (51).

Note. The type of confusion that applies in at least certain taxa in this and adjacent genera is clear from
the comment made by two of us (Lawson & John, 350; 586) that Polycavernosa [Gracilaria] dentata occurs
in much the same situations as the present species 'but the two species are impossible to separate on the
shore.' In such circumstances, it is not surprising that misapplication of names has been a consistent aspect
of all but the most critical taxonomic work conducted in the most well-endowed institutions. In speaking of
his Gracilariopsis (?) [Gracilaria] tridactylites, Bodard (52: 39) stated it to be 'une espece qui est souvent
confondue avec Gracilaria foliif era (sensu lato) et particulierement avec 1'ancien Gracilaria lacinulata." He
later (51 : 869) commented that 'La modification de la precedente etude [= 52] tient essentiellement dans la
suppression d'une espece. Les noms provisoires de Gracilariopsis tridactylites et Gracilariopsis (?)
disputabilis donnes pour deux especes ne correspondent qu'a une seule espece dont le developpement du
cystocarpe merite une attention toute particuliere.' Subsequently (in litt. to DMJ, 18/2/1971), Bodard had
carried simplification in use of at least the quantity of names further still, in stating 'II faut rassembler

236 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Gracilariopsis tridactylites et Gracilariopsis disputabilis dans la meme espece Gracilariopsis henriques-
iana.' This latter view has never been generally accepted, even taking account of the deep confusions
attaching to taxon limits, and has not been further employed here.

Gracilaria domingensis Sender ex Kutzing [in synonymy]

Note. We have concluded that this taxon should be recombined in the genus Polycavernosa, if the latter
is accepted. See details in the note to Gracilaria cervicornis (Turner) J. Agardh and the recombination
entry for Polycavernosa domingensis (Kutzing) J. Price & D. John.

Gracilaria dura (C. Agardh) J. Agardh

Canaries (70; 89; 191; 226; 390; 598).

'Atlantic Ocean (warmer regions)' (177).

[As Gigartina dura Desmazieres]

Canaries (44; 401).

'warmer parts of the Atlantic Ocean . . . seems to be widespread.' (70).

[As Gracilaria dura (C. Agardh) J. Agardh var. j3 Lyra J. Agardh]

Canaries (439).

Note. According to De Toni (132: 433-434), the Gigartina dura Desmazieres record presented in
Montagne (401: 160) is actually referable to Gracilaria armata (C. Agardh) J. Agardh (q.v.). Another
name involved seems to have been Gigartina cylindrica Despreaux; see the note at the latter. Piccone's
(439) record of var. j3 Lyra was named because of the secondary branching present. B0rgesen (70) did not
find the species, but mentioned it because Bornet (89: 283) indicated it was a Canarian species 'referring
most probably to a specimen in Montagne's Herb.' For recent comments on the taxon in the north-east
Atlantic and Mediterranean see Gargiulo, De Masi & Tripodi (1985). For an additional record probably to
be referred here, see the entry for Gracilaria rubra (C. Agardh) J. Agardh.

Gracilaria feldmannii Bodard ['(nom. prov.)']

Note. Although Bodard (52: 44) named this taxon as 'nom. prov.', he did present a Latin diagnosis,
stating also that on first collection it was taken as a Rhodymenia. Subsequently (51: 881 and 885), Bodard
decided that the species was really one already known as ' Gracilaria dichotomoflabellata (Crouan MS)
Maze et Schramm', a mis-statement of authority form (see the latter entry). G. dichotomoflabellata was
itself included in Gracilaria mammillaris (Montagne) Howe (q.v.), a fact confirmed by Bodard (in litt.,
18/2/1971): 'Quant au Gracilaria Feldmannii, c'est tout simplement Gracilaria mammillaris d'Amerique.'

Gracilaria ferox J. Agardh

See Gracilaria cervicornis (Turner) J. Agardh.

Gracilaria foliifera (Forsskal) B0rgesen

See Gracilaria multipartita (Clemente) Harvey.

Gracilaria henriquesiana Hariot

See Gracilaria dentata J. Agardh, Polycavernosa dentata (J. Agardh) G. Lawson & D. John, and

Polycavernosa henriquesiana (Hariot) Chang & Xia.

Gracilaria lacinulata (Vahl) Howe

Note. See the entry for Gracilaria multipartita (Clemente) Harvey. Hauck (1885) indicated that the
present species was possibly a form of Gracilaria foliifera [= G. multipartita for the eastern Atlantic], a
likelihood borne out by B0rgesen's (70) and Howe's (1920) examination of Canary Islands material,
previously identified by Piccone (439) as Gracilaria corallicola Zanardini, and of Vahl's Fucus lacinulatus
specimens.

See also the notes to Gracilaria disputabilis (Bodard) Bodard.

Gracilaria lemaneiformis (Bory) Weber-van Bosse

Senegal (592).

[As Gracilariopsis sjostedtii (Kylin) Dawson]

Senegal (51; 52; 59; 182).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'surement presente dans tout le Golfe du Benin' (52).

[As Gracilariopsis sjoestedtii Dawson]

Senegal (55).

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 237

[As Gracilariopsis sjoestedtii Kylin]
Senegal (399).

Note. Renderings of the specific epithet as 'sjostedtW and 'sjoestedtii1 are treated as equivalents here.
Bodard (52: 32-34) stated that this was by far the most frequent Gracilarial Gracilariopsis species in
Senegal, and was often referred to as Gracilaria confervoides , in error. Such reference is not surprising;
Hoyle (1984: 48, 50), basing his comments on Abbott (1983), indicated that currently there are available no
reliable vegetative characteristics 'by which what is called G. fracilaria] verrucosa (Hudson) Papenfuss by
Abbott & Hollenberg (1976) can be distinguished from G. sjostedtii Kylin'. Abbott, the editor of the work
in which Hoyle (1984) published, added in parenthesis that although both Hoyle and she realised that
verrucosa as an epithet was incorrectly applied to the Calif ornian entity, they had not at that stage enough
evidence to apply another name.

Since that time, the name Gracilaria lemaneiformis ['lemanaeiformis'] has often been utilised for the
western Atlantic specimens and, indeed, Dawson, Acleto & Foldvik (1964: 59-60) had earlier already
recombined that epithet in Dawson's Gracilariopsis. Recently (1986), Bird & Oliveira (592: 319-320), in
describing their new species Gracilaria tenuifrons and providing details of the distinctions between it and
the seemingly close G. lemaneiformis, commented:

'We have previously questioned the wide distribution of some species of Gracilaria. . . . and
now suggest that reports of G. sjoestedtii/ G. lemaneiformis outside the Pacific Ocean may be
erroneous. It would, however, be equally incorrect to equate all western Atlantic records of
G. sjoestedtii with G. tenuifrons. . . . It is appropriate here to urge strongly that identification of
G. lemaneiformis and related species be authenticated by the presence of reproductive structures
and especially [Gracilariella] chorda-type spermatangia.'

See also the notes to the entry for Gracilaria verrucosa (Hudson) Papenfuss.

Gracilaria nianiniillaris (Montagne) Howe

Senegal (51; 55; 59).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gracilaria mamillaria]

Senegal (59).

[As Gracilaria feldmannii Bodard '(nom. prov.)']

Senegal (51; 52).

Note. See Schneider (1975: 643, 645) on the rendering of the epithet with the doubled radical consonant;
we, too, have followed Montagne's basionym usage, despite the existence of the alternative (single 'm')
form. For further data on attribution of records here, see the note to the entry for Gracilaria feldmannii
Bodard.

Gracilaria multipartita (Clemente) Harvey

Canaries (70).

Cape Verde Islands (145; 252).

Ghana (586).

Mauritanie (252).

Senegal (529, pro parte).

'British Isles south to Senegal' (243 A).

'probably widespread in warm temperate and tropical seas.' (586).

[As Gracilaria multipartita Harvey]

Cape Verde Islands (38).

[As Gracilaria multipartita Clemente]

Cape Verde Islands (145).

[As Gracilaria multipartita Agardh]

Canaries (117; 118).

[As Rhodymenia multipartita (Clemente) Montagne, or simply Montagne]

Cape Verde Islands (38; 408).

[As Gracilaria foliif era (Forsskal) B0rgesen]

Canaries (191; 226; 227; 267 A; 517; 598).

Cape Verde Islands (408; 598).

Ghana (290; 299; 350; 376; 377; 611).

238 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Mauritanie (52; 267 A; 344; 349; 393; 394; 529; 537).

Senegal (48; 51; 52; 55; 56; 59; 267 A; 399; 529, pro parte; 542).

Western Sahara (349).

'Atlantique (de 1'Angleterre au Maroc, Canaries . . .)' (33).

'British Isles to Senegal' (172).

'connue de tout 1' Atlantique tropical' (52).

'Liberian-Cameroon Coast' (381A).

'Pantropical' (529).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'probably widespread in warm temperate and tropical seas.' (350).

'Tropical Africa (N. Gambia- Congo river)' (598).

'warmer parts of Atlantic' (81).

'warmere Teile des Atlantik' (224).

[As Gracilaria corallicola Zanardini]

Canaries (439).

[As Gracilaria lacinulata (Vahl) Howe]

Canaries (70).

[As Fucus aeruginosus Turner]

Ghana (271, pro parte)

Note. Guiry & Freamhainn (243A) have presented reasoning as to why Gracilaria foliif era should now be
recognized under the name G. multipartha (Clemente) Harvey for material from the eastern Atlantic. This
is a resuscitation of a name much used previously, as a result of the separation of materials from the western
Atlantic (G. tikvahiae), eastern Atlantic (G. multipartita) and Red Sea/Arabian Sea/Indian Ocean (G.
foliif era). For other recent comments on the taxon in north-east Atlantic/Mediterranean areas, see
Gargiulo, De Masi & Tripodi (1985). See Lawson & John (350; 586) and Bodard (52) for comments on the
reallocation here, or to a new species, of African (e.g., Senegalese) and Canaries plants hitherto called
Gracilaria [Polycavernosa] dentata. See also the latter entry.

The G. foliiferal multipartita records in Sourie (529) are stated by the author to be equivocal - 'Sa
presence a Daker est tres douteuse' - hence the use of 'pro parte'. Sourie (529: 116) noted that Feldmann,
who examined the plants, considered them possibly to represent Gracilaria [Polycavernosa] dentata,
possibly Gracilaria foliif era.

For the possible relationships between the taxa and name applications in G. multipartita and G. corticata
J. Agardh, see the notes at the entry for the latter.

The occasional statement in the literature that Gracilaria domingensis Sender ex Kiitzing is a morpho-
logical variant of Gracilaria multipartita var. polycarpa (Greville) J. Agardh and the name G. domingensis
therefore a synonym of G. foliif era [G. multipartita] is a misconception. See Oliveira filho, Bird &
McLachlan (1983). G. domingensis is similarly not synonymous with G. cervicornis; see the notes to the
latter.

According to De Toni (132: 447-448), the Fucus aeruginosus of Dawson Turner (Hist. Fuc., t.147),
referred to in Hornemann (271) for Ghana, is to be attributed in part to Gracilaria multipartita (Clemente)
Harvey and in part to Gracilaria corticata J. Agardh (q.v.). With this kind of nomenclatural equivalence
and in the absence of examination of the P. E. Isert Ghana specimens, which may or may not have been
destroyed in the 1807 fire in K0benhavn (see 350; 586), it is possible that the original material represented
any or a mixture of G. multipartita, G. corticata, and G. [Polycavernosa] dentata. See the fuller note at
G. corticata.

Gracilaria occidentalis (B0rgesen) Bodard

Senegal (48; 51; 52; 59; 182).

'doit se trouver ca et la dans 1' Atlantique tropical' (59).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Note. Recombined in Gracilaria by Bodard (48) because of the nature and structure of the cystocarps.
Described in Rhodymenia by B0rgesen.

Gracilaria poitei (Lamouroux) C. Agardh

See Laurencia poitei (Lamouroux) Howe and notes to the entry for Polycavernosa henriques-

iana (Hariot) Chang & Xia.

RHODOPHYTA (PLORIDEAE) OF TROPICAL AFRICA 239

Gracilaria rubra (C. Agardh) J. Agardh
Canaries (227).

Note. Gracilaria rubra is a recently-described Chinese species for which the correct authorities are
Chang & Xia. This is the only traced citation of the binomial with the present authorities for either the list
area or any other geographical region. Even Gil-Rodriguez and Afonso-Carrillo do not refer to it
elsewhere. The alphabetical order of entries around it in (227) shows that G. rubra is out of sequence and
probably represents an orthographic error for "dura\ the authorities for which are '(C. Agardh)
J. Agardh'.

Gracilaria sjostedtii Kylin

See Gracilaria lemaneiformis (Bory) Weber-van Bosse.

Gracilaria tenuifrons Bird & Oliveira f .

See Gracilaria lemaneiformis (Bory) Weber-van Bosse.

Gracilaria tridactylites Bodard

See Gracilaria disputabilis (Bodard) Bodard.

Gracilaria verrucosa (Hudson) Papenfuss

Angola (352; 535).

Cameroun (288; 350; 535; 586).

Canaries (38D; 128A; 227; 392; 584; 598).

Congo (535).

Cote d'lvoire (287; 288; 350; 586).

Ghana (288; 299; 350; 376; 418; 419; 535; 586).

Liberia (129; 350; 586).

Mauritanie (38D; 349; 535).

Namibia (36B; 274; 348; 393; 394; 522; 522A; 612).

Sao Tome (288; 350; 535; 586).

Senegal (38D; 535).

Senegal (Casamance) (350; 586).

Sierra Leone (295; 350; 586).

Togo (287; 288; 350; 586).

Western Sahara (38D; 349).

'Atlantique (de la Scandinavie au Rio de Oro)' (33).

'Atlantique: depuis la Scandinavie jusqu'au Rio de Oro' (222).

'Luderitz [S.W.A.] to Port Elizabeth [S.A.]' (523).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Tropical Africa (N. Gambia- Congo river)' (598).

'West Africa' (345).

'widely distributed ... in the Gulf of Guinea' (535).

'widespread from boreal-antiboreal to tropical seas' (350; 586).

'widespread in all except the coldest waters . . . widely distributed species in the Gulf of Guinea'

(535).

[As Gracilaria confervoides (L.) Greville]

Angola (239).

Cameroun (239; 454; 500).

Canaries (70; 191; 252).

Congo (239; 249; 250).

Guinea-Bissau (529).

Mauritanie (252).

Namibia (162; 274; 500).

St Helena (269).

Sao Tome (93).

Senegal (99; 252; 529; 531).

Senegal (Casamance) (99).

Sierra Leone (30).

240 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Western Sahara (476).

'Atlantic Ocean (. . . African . . . coasts, Canary Islands . . .)' (177).

'Cosmopolite dans les mers temperees et chaudes' (189).

'der ganzen westafricanischen Kiiste' (239).

[As Gracilaria confervoides Lamouroux]

Cameroun (239).

Ghana (153; 338).

Senegal (Casamance) (122).

[As Gracilaria confervoides Greville]

Angola (41; 42).

[As Gigartina confervoides (L.) Lamouroux]

Canaries (401).

Note. Drift material only.
[As Gigartina confervoides Lamouroux]
Canaries (44).

[As Sphaerococcus confervoides (L.) C. Agardh]
'In mari Atlantico, ab Anglia usque ad Africam, vulgaris.' (19).
'In mari . . . atlantico . . . Africano' (318).

[As Sphaerococcus confervoides (L.) C. Ag. /3 Procerrimus (Esper) Turner]
'Cum priore' [= entry above] (19).
[As Sphaerococcus confervoides Ag.]
Canaries (385).

Note. The Senegal record first established in Dangeard (122) may be doubtful; Bodard & Mollion (59),
for the same coast, stated: 'nous n'avous pas trouve le G. verrucosa contrairement a ce qui a ete ecrit
precedemment, c'est vraisemblablement Gradlariopsis Sjostedtii [= Gracilaria lemaneiformis (Bory)
Weber-van Bosse] qui a ete donne sous ce nom, sa repartition geographique est surement beaucoup plus
vaste que ne 1'indiquent les references actuelles.'

Recent studies by Bird, van der Meer & McLachlan (1982) and by Gargiulo, De Masi & Tripodi (1985)
have established that records of Gracilaria verrucosa (Hudson) Papenfuss throughout the world need
reservation in their acceptance and, often, critical re-evaluation. The recent description by Gargiulo, De
Masi & Tripodi (loc. cit.) of the segregate species Gracilaria dendroides from the Bay of Naples is a case in
point. Certain of the records noted above are clearly doubtful and require re-analysis; many of the
remainder would benefit from re-affirmation. For the present, we merely repeat all detected sources of
data.

Gracilaria wrightii (Turner) J. Agardh [or simply J. Agardh]
See Polycavernosa henriquesiana (Hariot) Chang & Xia.

Gracilaria spp.

Canaries (66).

Cape Verde Islands (445; 446).

Cote d'lvoire (288).

Ghana (92; 297; 299; 376; 377; 394; 487; 491; 537; 567).

Mauritanie (349; 395; 476; 533).

Sao Tome (93; 535).

Senegal (48; 529; 531; 542).

Sierra Leone (374).

Western Sahara (349; 393; 394; 395; 476).

'West Africa' (290; 345; 479).

'African west coast' (374).

Note. Bodard (52: 36) considered the Sourie (529) Senegal records of 'Gracilaria type D' to relate
to Gracilaria damaecornis J. Agardh (q.v.) and of 'Gracilaria sp. 2' to relate (Bodard, 52: 38) to
Polycavernosa [Gracilaria] dentata (J. Agardh) G. Lawson et D. John (q.v.). The records are also entered
here for completeness of recording.

All the records of Lawson & John (349) from Western Sahara are secondarily based on extant
information in the literature. Piccone (445: 72) gave an entry headed 'GratilariaT , in which he attributed
with doubt two young sterile individuals found on larger algae. Material collected by John (288) from Cote

RHODOPHYTA (FLORIDEAfi) OF TROPICAL AFRICA 241

d'lvoire was suggestive, from its flattened nature and its branching, of Gracilaria foliifera (now G.
multipartita, q.v.).

Gracilariopsis

Differences in the Gracilaria cystocarp ontogeny were recognized and presented by Sjostedt
(1926), when he dealt with the structure of the gonimoblast. The differences were emphasised by
Dawson (1949) in erecting the genus Gracilariopsis, although this latter was subsequently
synonymized with Gracilaria by Papenfuss (433). Authors have not universally accepted that
synonymy (e.g. Umamaheswara Rao, 1972).

Gracilariopsis camerunensis (Pilger) Bodard
See Gracilaria camerunensis Pilger.

Gracilariopsis ? disputabilis Bodard

See Gracilaria disputabilis (Bodard) Bodard.

Gracilariopsis lemanaeformis (Bory) Dawson, Acleto & Foldvik
See Gracilaria lemaneiformis (Bory) Weber-van Bosse.

Gracilariopsis sjoestedtii (Kylin) Dawson

See Gracilaria lemaneiformis (Bory) Weber-van Bosse.

Gracilariopsis (?) tridactylites Cr. fr. (= J. Ag.?) M. Bodard, comb. nov.
See Gracilaria disputabilis (Bodard) Bodard.

Gracilariopsis sp.

Senegal (48).
Note. Bodard (48: 877) indicated this to be a dorsiventral growth-form.

Polycavernosa Chang & Xia

For detailed elucidation of the background to the distinguishing features of this genus, and
analysis of its relationship to Gracilaria Greville (from which it is a potential segregate), see Xia
& Abbott (571; 600) and Chang & Xia (572). The distinctions between the genera are not
regarded by all workers as sufficient to justify separation, although Xia & Abbott (600: 415-417)
have recently strongly reiterated their belief in that separation, at least at subgeneric level.

The major distinctions involved are two-fold and clearly laid out in tabular form by Xia &
Abbott (600: Table 1). They concern the spermatangial ontogeny and the origin of the
gonimoblast. In this 1987 paper (600), Xia & Abbott also introduced a previously unremarked
spermatangial difference between Polycavernosa and Gracilaria - the production of clusters of
spermatangia isolatedly spread over the thallus surface in species attributable to Polycavernosa
(a distribution pattern easily missed in sections, leading to a conclusion of sterility), as opposed
to the numerous spread-out (not clustered) spermatangial conceptacles occurring densely over
the plant surface in Gracilaria (verrucosa - type spermatangia). Xia & Abbott (600: 416-417)
acknowledged that the spermatangial differences are the more important for placement of
plants in Polycavernosa or Gracilaria, and that gominoblastic characteristics then best separate
the species in Polycavernosa. Remarking that most of the nomenclatural and taxonomic
problems that exist are within the limits of Gracilaria (sensu stricto), they added that these more
easily demonstrable spermatangial characteristics 'are an important companion ... to the
equally or more important, but more difficult to demonstrate, origin of the gominoblast.'

Difficulties of allocation of species to one or the other of these two groups arise principally
from a lack of critical definitive data on aspects of the developmental morphology, and it will be
interesting to see if future extension of available information continues to support the postulate
of two well-founded genera (or sub-genera) within the overall grouping of gracilarioid plants.

Polycavernosa debilis (Forsskal) Fredericq & Norris

Note. See the entry for Polycavernosa henriquesiana (Hariot) Chang & Xia. The present taxon, hitherto
combined in Gracilaria, has been transferred to the segregate genus Polycavernosa Chang & Xia. For
details, see Fredericq & Norris (1985), Xia & Abbott (571), and Chang & Xia (572). For data on
life-history phases of P. debilis, see Littler et al. (1987). For possible complications and variable
applications of the epithet debilis in these genera, see Xia & Abbott (600: Table 2 and footnote, p. 416).

242 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Polycavernosa dentata (J. Agardh) G. Lawson & D. John

Cameroun (586).

Cote d'lvoire (586).

Gabon (586).

Gambia (586).

Ghana (586).

Liberia (586).

Nigeria (586).

Sao Tome (586).

Senegal (Casamance) (586).

'mainland coast of tropical West Africa' (586).

[As Gracilaria dentata J. Agardh]

Angola (298; 352).

Cameroun (52; 213; 288; 350; 454; 535).

Cape Verde Islands (38; 74; 139; 140; 213; 598).

Congo (249; 250; 535).

Cote d'lvoire (52; 287; 288; 350).

French Equatorial Africa [probably Gabon] (213).

Gabon (294; 350).

Gambia (296; 350; 535).

Ghana (213; 288; 291; 292; 297; 350; 418; 491; 535; 579).

Liberia (129; 287; 288; 350).

Nigeria (213; 288; 350; 535).

Sao Tome (93; 213; 288; 350; 535).

Senegal (38; 48; 50; 51; 52; 59; 74; 122; 213; 249; 529; 535).

Senegal (Casmanace) (52; 350).

Senegambia (25; 27; 132; 139; 296; 410; 454; 483).

'connue de tout 1'Atlantique tropical' (52).

'Gulf of Guinea' (535).

Taire se limite au golfe du Benin et a la Mauritanie' (59).

'Northwestern Africa' (540).

'probably in most warm temperate and tropical seas' (350).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Tropical Africa (N. Gambia- Congo river)' (598).

[As Gracilaria dentata Greville]

Cape Verde Islands (483).

[As Gracilaria dentata P. Dangeard]

Senegal (55; 59; 399).

[As Gracilaria corticata J. Agardh]

Cape Verde Islands (38).

[As Gracilaria henriquesiana Hariot]

Cote d'lvoire (59).

Ghana (45; 153; 335; 338; 340; 344; 350; 374; 418; 419; 535; 537; 593; 594; 609).

Sao Tome (419; 535).

Senegal (55; 59; 394; 398; 399).

'West Africa' (45).

'coast of the Gulf of Guinea' (269) .

Taire se limite au golfe du Benin et a la Mauritanie' (59).

[As Polycavernosa henriquesiana (Hariot) Chang & Xia]

Ghana (571; 572).

[As Gracilaria foliif era (Forsskal) B0rg.]

?Guinea-Bissau (529).

?Senegal(529).

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 243

[As Sphaerococcus rangiferinus Kiitzing]

Congo (249).

[As Sphaerococcus oligacanthm Kiitzing]

Congo (249).

Senegambia (326).

[As Gracilaria sp. 2]

Senegal (529).

Note. Bodard (52: 38) considered this 'sp. 2' of Sourie (529) to be 'Gracilaria dentata J. Ag.' [see
Polycavernosa dentata (J. Agardh) G. Lawson & D. John]. Even the attribution here of the records
established in Sourie (529) as Gracilaria foliifera (Forsskal) B0rgesen [- Gracilaria multipartita
(Clemente) Harvey] must remain in some doubt, since Sourie stated (p. 116): 'les echantillons rapportees a
cette espece ne le sont qu'avec incertitude, selon Feldmann ils pourraient etre aussi des formes de
G. foliifera (Forsk.) Boergs., tres polymorphic.'

Askenasy (38: 168) noted under his entry for Gracilaria dentata J. Ag.: 'Je possede un exemplaire de
cette algue provenant de 1'herbier Lenormand et etiquete: Gracilaria corticata J. Ag. Ins. Sal. Cap de
Verde, Forbes I.'

Of the Kiitzing (326) record for Senegambia, Pilger (454: 302) commented 'von Senegambien beschrieb
Kiitzing Sphaerococcus oligacanthus , der zu unserer Art gezogen wird'.

The Hornemann record (271), under Fucus aeruginosus Turner, from Danish Guinea [= Ghana] and
published in 1819, may have involved the present species, or Gracilaria multipartita (Clemente) Harvey, or
both plus Gracilaria corticata J. Agardh. See the fuller note to G. multipartita (Clemente) Harvey for
details.

See also the entries for Gracilaria angustissima (Harvey) Bodard and Gracilaria spp.
General note.

Abbott (see 571; 572) has accepted Polycavernosa as a good genus; she (in litt. to DMJ, 1986) believes,
on the basis of examination of types, that Sphaerococcus rangiferinus Kiitzing and Gracilaria dentata J.
Agardh are synonymous, but not so with the type of Gracilaria henriquesiana Hariot [Polycavernosa
henriquesiana (Hariot) Chang & Xia]. The first publication of the name Sphaerococcus rangiferinus
(Kiitzing, 318: 779), the basionym for Abbott's proposed (in litt.) recombination in Gracilaria, cites
Kiitzing's own Sphaerococcus cervicornis (Kiitzing, 316: Tab, 62, Fig. II) in synonymy. This, in turn, refers
back to C. Agardh (19: 292-293) and thence to Dawson Turner (Fuci . . ., tab. 121). Kiitzing's own 5.
cervicornis is illustrated as a tetrasporangial plant in apical longitudinal section (Fig. II, 2) and in gross
morphology (fig. II, 1); no indication is given of spermatangial material, even if such were present. Abbott,
in recently commenting (litt., 11 Dec. 1986) on her examination and acceptance of the equivalence of
application in the epithets rangiferinus Kiitzing and dentata J. Agardh (see above), made no mention of
spermatangial material being present. There is thus insufficient evidence from the available rangiferinus
material of the latter being the earliest epithet to apply in Polycavernosa and further work is required if
such is ever to be established. By contrast, evidence for the application of dentata in the genus
Polycavernosa is provided by material referred to by Ohmi (1968), in error as Gracilaria henriquesiana,
following Lawson's and Dickinson & Foote's earlier publications. For further detailed data, see Lawson &
John (586). It is perhaps for this reason that Abbott (in litt.) has proposed the transfer of rangiferinus not
(as might be expected) to Polycavernosa, but to Gracilaria. If G. dentata is correctly transferred to
Polycavernosa, on acceptance of the distinctness of the latter genus, and if Abbott is correct as to the
rangiferinus I dentata synonymy, then rangiferina will require transfer to Polycavernosa and, being the
earliest epithet, will replace the name P. dentata (J. Agardh) G. Lawson & D. John by the combination P.
rangiferina. This should perhaps be carried out for rationalisation purposes until (if ever) the type material
of S. rangiferinus Kiitzing can be definitively shown to be correctly associated with one or the other genus.
We hesitate currently to effect the recombination here, even provisionally, in view of the obvious intention
toward further work on the matter expressed in letters from Abbott (supr. cit.). As indicated above, this
presupposes acceptance of a real generic distinction between Gracilaria and Polycavernosa, a matter on
which there is no unanimity of opinion. We remain willing to be convinced fully, but are currently
sufficiently so to effect support at least the recombination of dentata in Polycavernosa (586).

In the matter of the identity of material referred to by Hornemann (271) as Fucus aeruginosus Turner,
which could have involved Gracilaria / Polycavernosa] dentata (inter alia), see the notes to Gracilaria
multipartita (Clemente) Harvey and Gracilaria corticata J. Agardh. See, finally, the entry for Gracilaria
angustissima (Harvey) Bodard.

Polycavernosa domingensis (Kiitzing) J. Price & D. John, comb. nov.

[Basiomyn: Sphaerococcus Domingensis Kiitzing, Tab. Phyc. 19: 8, no. 4254, Tab. 22 (1869).]

244 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Note. See the note to Gradlaria cervicornis (Turner) J. Agardh. This present recombination in
Polycavernosa is carried out for rationalisation purposes, the basionym having been applied to plants
which share certain morphological similarities with G. cervicornis but separable because of the occurrence
of spermatangia 'in deep crypts' and the possession of a thin cortex of subquadrate cells.

Polycavernosa henriquesiana (Harlot) Chang & Xia

'Atlantic Africa' (600).

[As Gradlaria henriquesiana Hariot]

Sao Tome (59; 137; 139; 213; 251; 350; 390; 485; 535; 586).

'in the tropical eastern Atlantic Ocean.' (586).

[As Gradlaria henriquesii P. Hariot]

Sao Tome (265).

[As Gradlaria wrightii (Turner) J. Agardh]

Sao Tome (251; 264; 265; 535).

[As Gradlaria wrightii J. Agardh]

Sao Tome (263; 264).

Note. Steentoft (535: 133) stated this taxon to be endemic to Sao Tome. Records under this name from
elsewhere in the list area have been taken, from experience, to have been inaccurately determined and to
relate correctly to finds of Polycavernosa dentata (q.v.). In the absence of either spermatangial material
required for confirmation, or sufficiently definite and clear statements for dependable reallocation without
examination, the Sao Tome records are retained until further study, but they could also prove to be
P. dentata.

Reallocation here of records for Sao Tome reported under the name Gradlaria wrightii (Turner) J.
Agardh is a matter of redetermination, not of nomenclatural equivalence. G. wrightii is currently
considered to be a synonym of Gradlaria debilis (Forsskal) B0rgesen (q.v.) [= Polycavernosa debilis
(Forsskal) Fredericq & J. Norris].

Hariot (251), Hariot in De Toni (137) and De Toni (139) all include the comment 'affine a G. potei e G.
corticata\ either as succinctly as that (137), or in greater detail but with the same background (251; 139).

More recent work by Steentoft (pers. comm., 3 March 1987) has shown that the type material of
Gradlaria henriquesiana (Ribeiro 14) in Coimbra (COI) includes both male and female plants. The males
possess spermatangia in deep crypts in the thallus surface - hence the acceptance here of the combination in
Polycavernosa.

Grallatoria tingitana (Schousboe ex Bornet) Abbott

See Callithamniella tingitana (Schousboe ex Bornet) Feldmann-Mazoyer.

Grateloupia

For cogent comment on generic and some specific criteria within the family Cryptonemiaceae,
see Kraft (1977).

Grateloupia dichotoma J. Agardh

Canaries (33; 70; 89; 108; 184; 188; 191; 214; 227; 273; 350; 379; 403; 499; 547; 584; 586; 598).

'Atlantique (de 1'Angleterre aux Canaries . . .)' (33).

'Atlantique tempere et chaud (de la Manche aux Canaries . . .) (188).

'British Isles southwards to Canary Isles; . . . probably more widespread than present records

suggest.' (273).

'cotes occidentales d'Afrique et aux Canaries' (89).

'Im Atlantischen Ozean von der englischen und normannischen Kiiste siidwarts bis nach

Kamerun'(498;499).

'Nordwestafrika.' (499).

'Tropical Africa (N. Gambia - Congo river)' (598).

'warmer parts of the Atlantic Ocean' (60; 70).

[As Chondrus crispus Lyngbye]

Canaries (44; 401; 403).

Note. Montagne (403: 103) reattributed the material he previously (401) called Chondrus crispus
Lyngbye to the present taxon in stating (under his entry for Grateloupia fimbriata Montagne):

'cette Algue ne pourrait pas etre une forme contractee du G. dichotoma J. Ag. mais je n'ai vu
aucune transition entre les deux especes . . . ce que j'ai donne autrefois avec doute

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 245

(Cryptogamie des Canaries, p. 157) comme le Chondrus crispus appartient au G. dichotoma. Le
G. fimbriata a encore des caracteres communs avec le G. Proteus Kiitz. trop incompletement
decrit.'

B0rgesen (70), who examined a small specimen from Montagne's Herbarium, concluded that Mon-
tagne's later assessment was correct and his opinion was requoted by Ardre (33: 125) in her entry for
Chondrus crispus (L.) Lyngbye, although she was in error in implying that B0rgesen had been the first to
show this:

'rappelons que Borgesen (1929) a montre que le Chondrus crispus des lies Canaries (Montagne,
1840 [sic!]) etait un Grateloupia dichotoma.'

De Toni (134: 1559, no 6) had the situation assessed correctly when, under G. dichotoma synonymy, he
commented: 'Chondrus crispus Mont. Canar. p. 157 (fide auctoris).' Since the list in his work is based
directly on Montagne's 1841 data (401), Bem'tez's (1928) record (in 44) is also attributed here.

Lawson & John (350; 586) have correctly observed that doubt attaches to Schmidt's (498) comment
(repeated in 499) on distribution southwards to Cameroun. Although there is no very good theoretical
reason for its not being present throughout the mainland Gulf of Guinea area, intensive and widespread
collecting throughout both mainland and islands coastlines has failed to establish authentic records. No
supporting specimens in any collections with connection to Schmidt have been located.

Grateloupia doryphora (Montagne) Howe

Angola (273; 577).

Canaries (18; 598).

Cote d'lvoire (288; 350; 586).

Gambia (18; 296; 350; 586).

Ghana (18; 273; 288; 299; 350; 376; 577; 586).

Liberia (129; 288; 350; 586).

Mauritanie (349).

Senegal (577).

Senegambia(24;296).

'Portugal to Ghana; Angola' (273).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

'Tropical Africa (N. of Gambia - Congo river)' (598).

'widespread in warm temperate and tropical seas' (350; 586).

'entre las costas de Gambia y Ghana' (18).

[As Grateloupia lanceola J. Agardh]

Angola (41; 42; 273).

Guinea-Bissau (529).

Mauritanie (529).

Senegal (47; 50; 59; 122; 221; 529; 530; 542).

Senegambia (24; 134).

'Atlantic (North Africa)' (410).

'Atlantique tropicale' (529).

'Du sud de 1'Espagne au Senegal' (89; 542).

'In atlantico ad littus Africae superioris' (27).

'pantropicale' (59).

[As Grateloupia lanceolata]

Senegal (411).

[As Grateloupia gibbesii Harvey]

Ghana (153).

'de 1' Atlantique oriental chaud' (588).

Note. Specimens attributed under this name are often very variable, from narrow lanceolate plants with
marginal proliferations to wider, almost undivided fronds.

The conspecificity of Grateloupia lanceola]. Agardh emend. Ardre & Gayral was finally established and
properly published by Dawson, Acleto & Foldvik (1964), after Ardre & Gayral (588) had carried out
preliminary rationalisation of many previous combinations into the synonymy of the there-accepted
Grateloupia lanceola J. Agardh. See also the notes for Grateloupia senegalensis Bodard.

246 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Grateloupia filicina Lamouroux) C. Agardh

Angola (298; 352).

Benin (288; 293; 350; 586).

Cameroun (288; 350).

Canaries (50; 142; 145; 227; 584).

Cape Verde Islands (50).

Cote d'lvoire (50; 59; 287; 288; 290; 350; 394; 586).

Gambia (296; 350; 586).

Ghana (153; 287; 288; 290; 291; 344; 350; 586).

Liberia (129; 287; 350; 586).

Mauritanie (349).

Namibia (348; 522).

Nigeria (288; 350; 586).

St Helena (142).

Senegal (50; 59).

Senegambia (24).

Togo (288; 293; 350; 586).

'Atlantique (de 1'Angleterre a la Mauritanie)' (33).

'de 1'Angleterre jusqu'en Cap de Bonne Esperance' (553).

'pantropicale' (59).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara] (598).

Tropical Africa (N. Gambia - Congo river)' (598).

'widespread in warm temperate and tropical seas' (350; 586).

[As Grateloupia filicina (Wulfen) C. Agardh]

Cameroun (213; 500).

Canaries (24; 50; 70; 134; 191; 213; 499; 500).

Gambia (296).

Ghana (154; 213).

Namibia (522A).

Nigeria (213).

Senegal (50; 55; 122; 213).

Senegambia (296).

'Atlantique: depuis la cote meridionale des iles britanniques jusqu'en Mauritanie' (222).

'Atlantique, des cotes d'Angleterre jusqu' au Cap de Bonne Esperance' (553).

'Nordwestafrika' (499).

'Sans doute cosmopolite dans toutes les mers chaudes' (188).

'Seems to occur in all warmer seas' (60; 70).

'von Marokko bis zum Kap der Guten Hoffnung' (239).

'warmeren atlantischen Ocean bis zum Cap der guten Hoffnung' (509).

'Westafrika' (499).

[As Grateloupia filicina Wulfen]

Canaries (142; 391).

St Helena (142; 391).

[As Grateloupia filicina (Wulfen in Jacquin) C. Agardh]

Canaries (24; 108).

[As Grateloupia filicina C. Agardh]

St Helena (260).

'Atlantic (as far as the Cape of Good Hope)' (410).

'Du sud de 1'Angleterre aux Canaries' (89).

'Probablement cosmopolite dans les mers chaudes' (221).

[As Grateloupia filicina (Wulfen) J. Agardh]

Cameroun (454).

Togo (454).

[As Grateloupia filicina (Lamouroux) C. Agardh var. luxurians A. & E. S. Gepp]

RHODOPHYTA (PLORIDEAE) OF TROPICAL AFRICA 247

'Gulf of Guinea to South Africa' (273).

'Some of the large and well-developed plants found in the Gulf of Guinea are close to variety

luxuriatis* (350; 586).

[As Grateloupia filicina (Lamouroux) C. Agardh var. filicina]

'British Isles to S. Africa' (273).

[As Grateloupia filicina [vars. authority combinations] var. ramentacea]

St Helena (142; 260; 391).

[As Grateloupia filicina forma filiformis (Kiitzing) Pilger]

Cameroun (139; 454).

Grateloupia fimbriata Montagne

See the notes at Grateloupia dichotoma J. Agardh.

Grateloupia gibbesii Harvey

See Grateloupia doryphora (Montagne) Howe.

Grateloupia lanceola J. Agardh

See Grateloupia doryphora (Montagne) Howe.

Grateloupia proteus Kiitzing

See the notes at Grateloupia dichotoma J. Agardh.

Grateloupia scutellata Kiitzing

Cape Verde Islands (38; 134; 325; 597; 598).

Note. Kiitzing's (1867) original description (325: 8) stated the location simply as '"Cap. vert." Bolle.'
This was reported in De Toni (134: 1572) as ' "Caput Viride" (Bolle)', the species being recorded as one of
the 'Species minus cognitae', with the terminal comment 'An reverea Grateloupia?' Askenasy (38: 173)
indicated that the specimens collected by Cardoso were determined by Bornet, apparently with little doubt
as to generic attribution because some of the material was cystocarpic. Prud'homme van Reine & Lobin
(597) similarly omitted any indications of doubt in indicating the species as endemic to the Cape Verde
Islands.

Grateloupia senegalensis Bodard
Senegal (47; 50).

Note. Bodard (47) presented in key and descriptive forms the distinctions between his new species and
the very similar G. lanceola J. Agardh [= G. doryphora (Montagne) Howe]. According to Bodard (I.e.),
this species and the rest of the interesting associated tropical flora do not go further north along the African
coast than Pointe de Sarene, therefore not reaching the Cape Vert peninsula.

Grateloupia spp.
Senegal (50).

Note. The listings in Bodard (50) refer to Grateloupia filicina, Grateloupia lanceola, and Grateloupia
senegalensis. The collective Grateloupia spp. is also used elsewhere, implying that there may have been
other taxa suspected as present. We refer here to the record simply to draw attention to that possibility.

Griffithsia

The limits of subgeneric taxa, in all cases where only vegetative material is available for
determination amongst routine field or laboratory determinations, appear difficult at best and
often impossible to establish with certainty. Thus, many of the records listed in the entries which
follow are necessarily subject to considerable reservation, aside from any definite differences of
taxonomic opinion. Comments are made where appropriate, but it may be taken that confirma-
tion would be desirable in most cases.

Griffithsia arachnoidea C. Agardh

Canaries (44; 71; 177; 318; 320; 401).

[As Griffithsia furcellata J. Agardh]

Canaries (89; 128A; 133; 227; 547; 584; 598).

Cape Verde Islands (598).

[As Corynospora furcellata (J. Agardh) Levring]

Canaries (38D; 375).

248 J. H. PRICE, D. M. JOHN & G. W. LAWSON

[As Neomonospora furcellata (J. Agardh) G. Feldmann & Meslin]
Canaries (33; 177; 190; 191; 196; 221; 222).
'Atlantique (cote de France et Canaries . . .)' (33; 190).
'Atlantique: cotes de France jusqu'aux Canaries et Maroc' (222).

Note. The taxon (as Cory nospora furcellata (J. Agardh) Levring) has also been reported from various
locations in Madeira. It is necessary to avoid confusion with Corynospora arachnoidea Harvey, now known
as Mazoyerella arachnoidea (Harvey) Gordon-Mills & Womersley.

The conspecificity of G. arachnoidea C. Agardh and G. furcellata J. Agardh has often been stated in the
literature, but it was not until B0rgesen's (71: 29) treatment that use of the earlier name G. arachnoidea
was resumed for the combined taxon. Even then (as will be clear from the above), there has been little
consistency in the approach. It has more recently been suggested that Griffithsia furcellata J. Agardh
should be transferred to Anotrichium furcellatum (J. Agardh) Baldock (see Baldock, 1976: 560), perhaps
indicating that reconsideration as to the conspecificity of G. arachnoidea and G. furcellata may here be
required on an individual record basis. The entry for Monosporus pedicellatus (I.E. Smith) Solier will
present reasoning on the segregate genus Anotrichium. Accuracy of determination on an individual record
basis may be further complicated by the possibility that A. furcellatum (J. Agardh) Baldock may be
conspecific with Anotrichium multiramosum (Setchell & Gardner) Baldock.

Griffithsia argus Montagne

See Wrangelia argus (Montagne) Montagne.

Griffithsia barbata (J. E. Smith) C. Agardh [and other authority combinations]
See Anotrichium barbatum (C. Agardh) Nageli.

Griffithsia capitata B0rgesen

Canaries (4; 71; 191; 227; 375).

Salvage Islands (598).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Note. According to B0rgesen (71: 37-38), some high degree of overlap with Griffithsia globifera
(Harvey) J. Agardh, Griffithsia ? phyllamphora Kiitzing, Griffithsia schousboei Montagne, Griffithsia
opuntioidesL Agardh, and Griffithsia corallinoides (L.) Batters occurs. However, all these taxa show some
differences and are therefore not synonymous (B0rgesen, 71).

Griffithsia confervoides Suhr
Namibia (522 A).

Note. See the notes to the entry for Griffithsia cymosa Simons and those to Griffithsia corallinoides (L.)
Batters. The present taxon in all probability includes Griffithsia cymosa Simons, as indicated by Simons et
al. (522 A) and Stegenga (599: 114).

Griffithsia corallina (Lightfoot) C. Agardh [or C. Agardh]
See Griffithsia corallinoides (L.) Batters.

Griffithsia corallinoides (L.) Batters

Canaries (227).

'Atlantique [de la Scandinavie au Maroc (?)]' (33).

[As Griffithsia corallina (Lightfoot) C. Agardh]

Canaries (318; 401).

[As Griffithsia corallina C. Agardh]

Canaries (44; 401).

[As Griffithsia corallina (J. E. Smith) C. Agardh]

Canaries (268).

[As Griffithsia corallina (Lightfoot) J. Agardh]

Canaries (24).

[As Griffithsia corallina var. ? tenuior]

'in mari Atlantico et Mediterraneo' (21).

Note. Considerable doubt attaches to the validity of Canaries reports of this taxon. All those trace back
directly or indirectly to that in Montagne (401), which record was based on sterile (i.e. non-reproductive)
material. B0rgesen (71) was unable to trace specimens in Herb. Montagne and therefore doubted the
accuracy of the determination. Many other expressions of that same doubt have been published, both
before and subsequent to B0rgesen's statement - for example, in J. Agardh (1851), based directly on

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 249

Montagne's (1841) record, 'sed sterilis et hinc forsan specie dubia'; in Ardre (33: 170), who similarly
repeated the doubt already expressed by Dangeard (118) in establishing his record from Morocco; and in
others. The real attribution of material to which the name G. corallina [corallinoides] has been applied
probably varied; both Griffithsia schousboei Montagne and Neomonospora furcellata (J. Agardh) G.
Feldmann & Meslin [= Griffithsia arachnoidea C. Agardh] have been invoked at times (e.g. in Ardre; 33).
Seagrief (570) considered G. corallina of Barton, from the Cape, to be a synonym of Griffithsia
confervoides Suhr. See also the notes to G. capitata B0rgesen.

Griffithsia cymosa Simons

Namibia (348; 521; 522; 599).

Note. Simons (521: 1; 4-5) indicated his newly-described taxon as clearly a Griffithsia but with some
atypical features in regard to the nature of branchlets on the cystocarp-bearing dwarf-shoot and the
arrangement of the cystocarps. Procarps and cystocarps were all the reproductive stages seen. Stegenga
(599: 114), referring to Simons' G. cymosa, commented: 'the morphology of the Namibian Griffithsia
cymosa Simons (Simons 1970) is such as to make conspecificity with G. confervoides [Suhr] likely.'
Stegenga had already indicated G. confervoides as the most common species of Griffithsia around the Cape
Peninsula and vicinity . The Simons data for Namibia (522) were repeated by Lawson & Isaac (348) . See the
entry for Griffithsia confervoides Suhr.

Griffithsia flosculosa (Ellis) Batters

Canaries (33; 128A; 191; 227; 306B; 517; 584; 598).

'Atlantico de Inglaterra a Canarias' (517).

'Atlantique (de 1'Angleterre aux Canaries)' (33).

[As Griffithsia setacea (Ellis) C. Agardh]

Canaries (71; 133; 318; 401).

St Helena (401).

'English coast down to the Canary Islands' (71).

[As Griffithsia setacea C. Agardh]

Canaries (44; 89; 401).

St Helena (260).

'D'Angleterre aux Canaries' (89).

[As Griffithsia setacea Ellis]

Canaries (24; 142; 391).

St Helena (142; 391).

[As Griffithsia setacea J. Agardh var. sphaerica (Schousboe ex C. Ag.) G. Feldm.-Mazoyer]

Canaries (89).

[As Griffithsia setacea (Ellis) C. Agardh var. sphaerica}

Canaries (71).

Note. Many of the records cited for the Canaries are based solely and directly on the early report by
Montagne (401). Since Montagne stated 'in rupibus maritimis canariae sed sterilis lecta', there inevitably
exists some doubt (sometimes openly expressed; e.g. in 71; 227) as to the determinations. Similar doubt
applies for St Helena (e.g. 391). The 'var. sphaerica' seems likely to be of doubtful validity. Baldock (1976:
510; 563-566) considered that this species (cited as Griffithsia setacea (Ellis) C. Agardh) 'should probably
be referred to Halurus'.

Griffithsia furcellata J. Agardh

See Griffithsia arachnoidea C. Agardh.

Griffithsia globifera (Harvey) J. Agardh

See the notes to Griffithsia capitata B0rgesen and Griffithsia schousboei Montagne.

Griffithsia opuntioides J. Agardh

Angola (352).

Canaries (3; 4; 5; 16; 33; 71; 89; 128A; 191; 225; 227; 229; 375; 379; 547; 584; 598; 604; 605).

Guinea-Bissau (529).

Senegal (529).

Western Sahara? (349).

'[Africa] north of the Gulf of Guinea' (352).

'Angola . . . occurs to the north of the Gulf of Guinea . . . but not in the Gulf itself (487).

250 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Note. Also known from Madeira (375). See the notes for the entries on Griffithsia capitata B0rgesen,
Griffithsia schousboei Montagne and Griffithsia sp.

Griffithsia phyllamphora J . Agardh

Canaries (5; 13; 71; 89; 139; 191; 196; 227; 392; 489; 490; 499; 547; 584; 598; 604; 605).

'nordwestafrikanische Kiiste' (499).

Note. According to B0rgesen (71), this taxon is not the same as G. phyllamphora of Kiitzing, which may
be the same as G. capitata of B0rgesen (q.v. as to the notes).

Griffithsia radicans Kiitzing
Canaries (13; 227; 598).

Note. This taxon was based on material from Brasil and clearly left at least De Toni (133; 1287) in some
doubt as to placement, since he included it in his 'Species . . . quod sectionem dubiae' with the comment
'An SpermothamnionT . Taylor (540: 515-516) merely described the taxon, without further comment, and
indicated it as from Brasil.

Griffithsia schousboei Montagne

Canaries (24; 33; 38D; 44; 71; 89; 128A; 133; 177; 191; 196; 227; 306B; 375; 401; 405; 406; 407;

493; 547; 584; 598).

Cote d'lvoire (287; 288; 350; 586).

Ghana (287; 288; 350; 586; 590).

Guinee (287).

Western Sahara? (349).

'Atlantic Ocean (from the Bay of Biscay to Canary Islands)' (177).

'Atlantic (S. Europe & N. Africa)' (410).

'Atlantique, de la cote Basque aux Canaries' (191; 196).

'Du golfe de Gascogne aux Canaries' (89).

'Golfe de Gascogne southwards to the Canary Islands' (71).

'in vicino Atlantico ad Tingin et ins. canarias' (133).

'in warm temperate and tropical parts of the Atlantic Ocean' (350; 586; 590).

Tropical Africa (N. Gambia - Congo river)' (598).

[As Gr. [iffithsia] Schousboei Mont, in Park. Webb.]

'In mari adriatico, mediterraneo et atlantico ad oras Europae meridionalis et Africae' (318).

[As Griffithsia schousboei var. minor}. Feldmann]

'Atlantique tempere chaud, de la cote basque aux Canaries' (190).

'Du Golfe de Gascogne aux Canaries' (89).

Note. Apparently widely recorded, and also known from Madeira (375). Some doubt attaches to the
original Montagne record for the Canaries (401); B0rgesen (71), without having seen the plants, stated that
Montagne's material was sterile and the determination therefore doubtful. Sauvageau (493) believed it to
be either G. schousboei or Griffithsia opuntioides J. Agardh (q.v.). See also the notes to Griffithsia capitata
B0rgesen and Griffithsia sp. Norris & Bucher (416: 203) also indicated a relationship between G. globifera
(Harvey) J. Agardh and G. schousboei; these, morphologically similar, are most easily separated by the
form of the spermatangia (the former lacks an involucre; the latter has one).

Griffithsia secunda Harvey ex J. Agardh

See Anotrichium tenue (C. Agardh) Nageli (notes to the entry).

Griffithsia setacea (Ellis) C. Agardh [or C. Agardh]
See Griffithsia flosculosa (Ellis) Batters.

Griffithsia tenuis C. Agardh

See Anotrichium tenue (C. Agardh) Nageli.

Griffithsia sp.

Ghana (299; 300; 350; 376; 377; 491; 586).
Senegal (59; 529).
Sierra Leone (31).
Western Sahara (349).
Note. Lawson & John (349: 114) originally indicated close vegetative correspondence to Griffithsia

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 251

schousboei and G. opuntioides (branches of both with large obovoid to barrel-shaped cells) ; they were
unable to decide further since all the material lacked reproductive organs (the position of spermatangial
sori is the determinant).

Ghanaian material of the genus also presents difficulties (350: 330; 586: 275) since narrow elongate
vegetative filament cells on some specimens indicate the existence of a further species to G. schousboei.
Again, absence of reproductive structures does not permit certainty, even (in this case) of generic
attribution (cf. also 300; 376).

Gulsonia annulata Harvey

See the notes to Gulsonia ecorticata G. Lawson & D. John.

Gulsonia ecorticata G. Lawson & D. John

Ghana (350; 377; 586).

'in the tropical eastern Atlantic Ocean' (350; 586).

Tropical Africa (N. Gambia - Congo river)' (598).

[As Gulsonia sp.]

Ghana (299; 376; 491).

Note. Distinctions between Gulsonia ecorticata, G. annulata Harvey, and G. medlterranea Kylin are
outlined in Lawson & John (350; 586).

Gulsonia mediterranea Kylin

See the notes to Gulsonia ecorticata G. Lawson & D. John.

Gulsonia sp.

See Gulsonia ecorticata G. Lawson & D. John.

Gymnogongrus

For a recent consideration of the generic background, see Schotter et al. (514). For more recent
details on the connections between some known Gymnogongrus spp. and crustose life-history
stages often hitherto called Erythrodermis see Decew & West (1977), Candia & Kim (1977), and
Ardre(1978).

Gymnogongrus capensis (C. Agardh) J. Agardh

See the notes to Gymnogongrus complicatus (Kiitzing) Papenfuss.

Gymnogongrus complicatus (Kiitzing) Papenfuss
Namibia (36B; 348; 522; 522A).

Note. Also well known from South Africa (Papenfuss, 1943; Seagrief, 570); see the latter reference for
suggested synonymy, which includes Gymnogongrus capensis (C. Agardh) J. Agardh.

Gymnogongrus corymbosus J. Agardh
Namibia (36B).

Note. Wynne (36B: 314) has recorded cystocarpic material under this name from the drift near
Swakopmund. Primarily a South African seaweed, the species has been placed in the synonymy of
Gymnogongrus glomeratus J. Agardh by Seagrief (570: 33); the latter species was also recorded attached
by Wynne (36B: 315). See the entry for G. glomeratus.

Gymnogongrus crenulatus (Turner) J. Agardh

Canaries (214; 227).

Cape Verde Islands (598).

Mauritanie (349).

'British Isles to Mauretania' (172).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. Sahara]' (598).

[As Gymnogongrus norvegicus (Gunnerus) J. Agardh]

Cape Verde Islands (191; 252; 408; 423).

Mauritanie (252; 500).

'Atlantique (de la Norvege a la Mauritanie)' (33).

'Atlantique nord: des cotes anglaises jusqu'au cap Blanc (Mauritanie)' (222).

[As Gymnogongrus norvegicus J. Agardh]

Cape Verde Islands (38).

252 J. H. PRICE, D. M. JOHN & G. W. LAWSON

[As Sphaerococcus norvegicus (Gunnerus) C. Agardh]

'In mari Atlantico, ab oris Angliae usque ad caput bonae spei' (19).

Note. Feldmann (191: 426) noted that the determination of this species from the Cape Verde Islands
'meriterait-elle confirmation'; subsequent records seem to have effected that confirmation. Comments on
the biology, including life-histories, of this entity are presented in many recent studies, including Gayral
(222: 455) and Ardre (1978).

Gymnogongrus devoniensis (Greville) Schotter
Cape Verde Islands (598).

Note. Schotter et al. (514) considered plants otherwise attributable to Gymnogongrus crenulatus
(Turner) J. Agardh, but possessing internal cystocarps, to be a separate species - G. devoniensis. Dixon &
Irvine (172: 219) indicated such plants to be rare 'found only in Cornwall, Devon, Galway, Mayo, Down
and France.' The present record is a considerable extension and probably indicates a widespread range for
such occasional forms, whether or not one accepts their species status.

Gymnogongrus dilatatus (Turner) J. Agardh
Namibia (36B; 162; 348; 453; 522; 522A; 523).

Note. This taxon is better known from South Africa, its currently accepted distribution having been
described (523) as 'Luderitz to False Bay'. Previous Liideritzbucht and Swakopmund records are repeated
in more recent draft texts (e.g. 348) since additional records have not been established, save from the drift
(36B).

Gymnogongrus glomeratus J. Agardh

Namibia (36B; 162; 298; 312A; 348; 437; 438; 522; 522A).

Note. Also well-known from the western coastline of South Africa, the concept includes Gymnogongrus
corymbosus J. Agardh (q.v.). The earliest record in the sequence above is that of Dinter (162) in 1922, in
turn supposedly based partly on an earlier one in Pilger (453), in 1908. No such record is actually presented
by Pilger, who reported only Gymnogongrus dilatatus (Turner) J. Agardh (q.v.).

Gymnogongrus griffithsiae (Turner) Martius

Canaries (38B; 38D; 70; 108; 188; 191; 227; 229; 252; 254; 306; 375; 379; 392; 517; 546; 598).

Mauritanie (38B; 38D; 252; 350; 500; 586).

Salvage Islands (38B; 38D; 231; 375; 598).

Sierra Leone (30).

'Atlantico (de Inglaterra a Canarias . . .)' (517).

'Atlantique (de 1'Angleterre aux Canaries . . .)' (33).

'Atlantique, de 1'Angleterre aux Canaries . . .)' (140).

'Atlantique nord (de 1'Angleterre aux Canaries . . .)' (189).

'British Isles to Mauretania and Canary Isles' (172).

'de 1'Angleterre aux Canaries' (514).

'English coast southwards to the Canary Islands' (70).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gymnogongrus Griffithsiae Martius]

'D'Angleterre aux Canaries' (89).

[As Gigartina griffithsiae (Turner) Lamouroux]

Canaries (401).

[As Gigartina griffithsiae Lamouroux]

Canaries (44).

[As F[ucus] Griffithsiae Turner]

Ghana (271; 7350; 7586).

Note. The locational data for the Ghana record were presented as 'Danish Guinea'. Both the more
recent treatments of that record were quite rightly expressing doubt as to the modern translation of the
record; confirmation of both location and determination may be impossible since at least part of the Isert
collection was burnt in 1807. A degree of doubt seems to apply to many records named as this species- that
for Sierra Leone in Aleem (30), for example, is questioned by Lawson & John (350; 586), whilst the
material available to Audiffred & Weisscher (38B) for the Salvage Islands was only determinable with
reservation, since basal fragments of the thallus were all that was discovered. The species is clearly not
well-represented nor frequent in the southern part of its range around the Canaries, Salvage Islands, and
Madeira; many of the records from there are of small, sterile specimens.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 253

Gymnogongrus interim-dins Kylin
Cote d'lvoire (7288; 350; 586).
Nigeria (213; 288; 350; 586).

'in warm temperate and tropical parts of the eastern Atlantic Ocean.' (350; 586).
Tropical Africa (N. Gambia- Congo river)'. (598).
Note. Also well-known from South Africa (Seagrief, 570).

Gymnogongrus nigricans P. Dangeard

Angola (352).

Benin (288; 293; 350; 586).

Bioko (346; 350; 586).

Cameroun (269; 288; 292; 337; 350; 537; 586).

Cote d'lvoire (288; 350; 586).

Ghana (288; 292; 350; 586).

Liberia (7129; 288; 350; 586).

Sso Tome (93; 586).

Senegal (122; 292; 529).

'in warm temperate and tropical parts of the eastern Atlantic Ocean' (350; 586).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Tropical Africa (N. Gambia- Congo river)' (598).

[As Gymnogongrus sp.]

Cameroun (484).

Note. In common with other species in this morphologically 'plastic' genus, there is considerable form
variation according to the environment; many of the references above (e.g. 350; 586) emphasise that. Some
of the doubts so raised (e.g., those in 288 for Cote d'lvoire material) have now been eliminated (cf. 350;
586), but others (e.g. those from Liberia in 129) remain, resulting in tentative determinations.

Gymnogongrus norvegicus (Gunnerus) J. Agardh
See Gymnogongrus crenulatus (Turner) J. Agardh.

Gymnogongrus patens J. Agardh

Canaries (229).

'Atlantique nord: des cotes anglaises jusqu'au cap Blanc (Mauritanie)' (222).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

[As Gymnogongrus patens (Gooden. et Woodw.) J. Agardh]

Mauritanie (349).

Western Sahara (349).

Note. Dangeard on two occasions (117; 118) stated this taxon to be absent from the Canary Islands.
However, the recent (229) record in Gil-Rodriguez & Wildpret de la Torre (1980) has refuted that. See
Dixon & Irvine (172: 217) on the status of species and its assignment to Gymnogongrus.

Gymnogongrus tenuis (J. Agardh) J. Agardh

Angola (352).

Cote d'lvoire (288; 350; 586).

Gambia (288; 296; 350; 586).

Guinea-Bissau (529).

Liberia (129; 288; 350; 586).

Nigeria (213; 288; 350; 586).

Senegal (55; 59; 122; 213; 529).

'Atlantique tropicale' (529).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).

Tropical Africa (N. Gambia - Congo river)' (598).

'widespread in warm temperate and tropical seas' (350; 586).

Note. The specific epithet in (55) is rendered as 'tennis' in error. Elsewhere (e.g. 529), the authorities are
often inaccurately presented as simply 'J. Agardh'. See also the notes to the entry for Gymnogongrus spp.

Gymnogongrus vermicularis (C. Agardh) J. Agardh

Namibia (348; 522; 522A).

254 J. H. PRICE, D. M. JOHN & G. W. LAWSON

Note. This species is well-known from South Africa (Seagrief, 570) and appears at many accessible
appropriate locations along the Namibia coastline, cited in the above references. See also comments in the
entry for Chondrus elongatus Montagne.

Gymnogongrus spp.

Benin (293).

Cape Verde Islands (7445; 7446).

Cote d'lvoire (287; 288).

Gabon (250).

Ghana (344).

Guinea-Bissau (529).

Liberia (129; 287; 288).

Namibia (348; 522).

Note. Reservations that commonly attach to such limited records are again apparent here. The doubt
expressed for Cape Verde Islands records in Piccone (445; 446) relates directly to generic attribution.
Plants noted by John (288) in Cote d'lvoire were of at least two contrasting morphologies, the larger being
perhaps no more than a form of Gymnogongrus tenuis (J. Agardh) J. Agardh (q.v.), widely known in the
Gulf of Guinea region. Materials of most collections (e.g. that from Gabon) were of difficult sterile
specimens, usually detected in very small amounts (e.g. those from Liberia in 129). Material from
Namibian localities reported by Simons (522) was repeated without comment in 348.

Gymnophlaea canariensis Kiitzing
Canaries (24; 318; 324).
[As Halymenia capensis Montagne]
Canaries (44; 401; 403).

Note. The attribution of these records is not clear. In the original description, Kiitzing (318: 712) gave as
synonymy:

'Halymenia capensis Mont. Canar. p. 164 (excl. Syn. Ag.) Gigartina dichotoma Despr. (sec.
Mont.) ... In mari Canariensi . . . Spec, dedit cl. Montagne.'

The reference to Montagne (401: 164) concerns the original description, including as synonymy both
Gigartina dichotoma of Despreaux and Halymenia furcellata var. capensisl Ag. Sp. alg. I. p. 214, of
Halymenia capensis (q.v.).

Records of the latter taxon from the Cape, as opposed to those from the Canaries dealt with here, were
synonymised later (p. 738) in Kutzing (318) with Chondrus capensis (Montagne) Kiitzing; this is a change
from the treatment earlier (Kutzing, 317: 25), where Halymenia capensis of Montagne was directly the
source of Gymnophlaea capensis Kutzing.

A Canaries specimen of apparently the same species, whatever name should be correctly applied, was
collected by Broussonnet and given by Bouchet of Montpellier to P. B.-Webb; it was given the name
Ceramium fucoides by Broussonnet.

Gymnothamnion elegans (Schousboe ex C. Agardh) J. Agardh
[As Gymnothamnion elegans (Schousboe) J. Agardh]
Ascension (475).

Canaries (13; 33; 191; 227; 375; 584; 598).
Senegal (529).

'Atlantique (de la cote Basque aux Canaries . . .)' (33; 190; 196).
Tantropical' (529).

'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598).
[As Gymnothamnion elegans (Schousboe in C. Agardh) J. Agardh]
Canaries (108).
Senegal (529).

[As Gymnothamnion elegans J. Agardh]
Canaries (191).

[As Plumaria schousboei (Bornet) Schmitz]
Canaries (39; 71; 489; 490).
'Nordwestafrikanische Kiiste' (499).
Note. This taxon, despite the difficulties of being certain as to identification when only vegetative

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 255

material is available, is in all probability more widespread in the area than the current recording pattern
would suggest. Close scrutiny of all potential hosts collected is normally required to locate specimens.
Available data from outside the list area suggest that generalised statements indicating a pantropical
Atlantic distribution are probably correct - Levring (375), for example, indicated records for Madeira,
Bermuda, and the West Indies.

Acknowledgements

We acknowledge with many thanks the assistance received from many sources. The principal benefactors
are (i) British Museum (Natural History), Department of Botany, for provision of the necessary research
facilities during construction of the series of papers that includes the present part ; (ii) Mr J . R. Laundon for
editorial expertise; (iii) Ms Sandra Davie, who willingly and promptly typed the original version of a
somewhat concentrated manuscript; (iv) Dr W. F. Prud'homme-van Reine and Mr R. H. Simons, who
provided hitherto unpublished data from expeditions, respectively to CANCAP (Canaries and Cape
Verde Islands) and Namibia.

References

This single list results from the application of three major criteria:

1. The basis is the list previously published in 1986 (Price et al., Florideae 1 (Genera A-F), this
series). Numbers used there for references remain in general unchanged if the same reference also
appears here. The only case of change is the introduction of a number for a reference that (a) was
previously relevant only through appropriate taxonomic comment and lacked records for any
Florideae 1 taxa, but (b) does contain records that are relevant to taxa in the present part. The reverse
is also true - a number is dropped (although it will be reapplied as required in subsequent parts) if a
reference is here only included for taxonomic reasons.

2. References previously included are omitted here if they are not relevant for either taxonomic
purposes or through provision of records.

3. Publications of relevance that have been either first detected or actually published in the time since
Florideae 1 (i.e., since 30/10/86) are included here for the first time. In many cases, they require a
number because of the presence of appropriate records. Numbers allocated lie in the sequence from
567 onwards, the point at which bulk allocation in the 1986 list ceased.

Since the list is arranged overall alphabetically on authors' surnames, the application of the above criteria
produces numerical irregularity, which has been overcome by insertion at the appropriate point of a
numerical cross-reference. The reference system is therefore approachable with facility from various
directions - with startpoints in biogeography, in taxonomy, or simply through appropriate authors.

585. Abbott, I. A. 1962. Some L/agoro-inhabiting species oiAcrochaetium. Occ. Pap. Bernice P. Bishop
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1 . 1984a. Two new species of Liagora (Nemaliales, Rhodophyta) and notes on Liagora farinosa

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2. Acuna Gonzalez, A. 1970. Algunos aspectos de la vegetacion submarina de las Islas Canarias.

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3. - 1972. Observaciones ecologicas sobre las algas de la zona literal de Las Galletas, Tenerife.
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4. - 1972? ['1968-1970']. Cinco nuevas citas de algas Rhodophyceae en la Isla de Tenerife. An.
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5. Santos G[uerra], A. & Wildpret [de la Torre], W. 1970. Algunos aspectos de la vegetacion

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8. Afonso-Carrillo, J. 1980o. Algunas observaciones sobre la distribution vertical de las algas en la
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576. 1985. Conexiones intercelulares entre diferentes talos de Neogoniolithon absimile (Foslie et

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256 J. H. PRICE, D. M. JOHN & G. W. LAWSON

13. — & Gil-Rodriguez, M. C. 19806. Datos para la flora marina de la Isla de Fuerteventura. Vieraea
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14. - 1981. Sobre el limite meridional de Sauvageaugloia chordariaeformis (Crouan) Kylin

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16. — & Wildpret de la Torre, W. 1979 ['1978']. Estudio de la vegetation algal de la costa del

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582. Afonso-Carrillo, J., Gil-Rodriguez, M. C. & Wildpret de la Torre, W. 1985 ['1984']. Algunas
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18. - Haroun Tabraue, R., Villena Balsa, M. & Wildpret de la Torre, W. 1984 ['1983'].
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19. Agardh, C. A. 1822. Species algarum rite cognitae ... 1 (2). Lund.

Note. There is another version of this first volume beside that issued at Lund. The parts issued at
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issues.
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24. Agardh, J. G. 1851. Species genera et ordines algarum, . . . floridearum, . . . 2(1). Lund.

Note. Facsimile reprint, J. Cramer, 1977.

25. -1852. Species genera et ordines algarum, . . .floridearum, . . . 2(2). Lundae.

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Pars II: 2, 1852. 3. There are also internal differences of numbering of pages between copies. BMNHcopyis
numbered straight through from 337-720, including therein the Addenda [701-706] and Index [707-720]. In
the copy from which the Cramer (1977) reprint was prepared, the Index [unnumbered, of 14 sides] is placed
immediately after p. 700 and is followed by six sides [also unnumbered] of Addenda. Content of these
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787 to 1291 are in both cases in Vol. 2, Part 3, 1863. Despite this, the Index in the end of the Cramer (1977)
facsimile of Vol. 2, Part 3 (pp. 1279-1291) indicates the same page numbers as does the BMNH version.
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RHODOPHYTA (PLORIDEAE) OF TROPICAL AFRICA 257

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132. - 1900. Sylloge algarum omnium hucusque cognitarum ... 4. Sylloge Floridearum . . . Sectio

II - Familiae I-IV. Patavii.

133. - 1903. Sylloge algarum omnium hucusque cognitarum ... 4. Sylloge Floridearum . . . Sectio

III - Familiae V-VI. Patavii.

134. - 1905. Sylloge algarum omnium hucusque cognitarum ... 4. Sylloge Floridearum . . . Sectio
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137. _ 19096. Hariot, P. - Les algues de San Thome (cote occidentale d'Afrique. - Journal de
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139. - 1924. Sylloge algarum omnium hucusque cognitarum ... 6. Sylloge Floridearum . . . Sectio

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140. - & Forti, A. 1913. Contribution a la flore algologique de la Tripolitaine et de la Cyrenaique.

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260 J. H. PRICE, D. M. JOHN & G. W. LAWSON

141 A. - — & Levi, D. 1888. L'algarium Zanardini. Venezia.

142. Dickie, G. 1872. On the marine algae of the island of St. Helena. /. Linn. Soc. (Bot.) 13: 178-182.
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152. Dickinson, C. 1. 1952. Marine algae from the Gold Coast: IV. Kew Bull. 7: 41-43.

153. - & Foote, V. J. 1950. Marine algae from the Gold Coast I. Kew Bull. 5: 267-272.

154. - 1951. Marine algae from the Gold Coast: II. Kew Bull. 6: 133-138.

161. Dinter, K. 1921. Index, der aus Deutsch-Siidwestafrika bis zum Jahre 1917 bekannt gewordenen

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162. - 1922a. Index, der aus Deutsch-Siidwestafrika bis zum Jahre 1917 bekannt gewordenen
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168. - 1926ft. Index der aus Deutsch-Siidwestafrika bis zum Jahre 1917 bekannt gewordenen

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170. Dixon, P. S. 1958. The occurrence of Gelidium sesquipedale (Clem.) Thur. in the British Isles. Br.
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172. — & Irvine, L. M. 1977. Seaweeds of the British Isles Volume 1 Rhodophyta Part 1 Introduction,
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173. Dizerbo, A.-H. 1974. La repartition des Gigartina (Gigartinales, Gigartinacees) du Massif Armori-

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180. Farnham, W. F. 1980. Studies on aliens in the marine flora of southern England. In J. H. Price, D.
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182. Feldmann, G. & Bodard, M. 1965. Une nouvelle espece de Botryocladia des cotes du Senegal.

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183. Feldmann, J. 1935. Algues marines des Isles du Cap Vert recoltees par M. le Professeur Aug.

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and a new sequence (pp. 1-358) at the bottom of the page. See also no. 100.

184. — 1937. Recherches sur la vegetation marine de la Mediterranee. La Cote des Alberes.

Note. Originally published as Revue algol. 10: 1-339. Printed 28 October 1937, but published with '1938'
on title-page of part. The separate form was published with '1937' on title-page and attributed inside as
extracted from the Revue algol. Tome X, Nov. 1937. The BMNH copy of the journal was received 22 June
1938. No textural differences exist between the two versions.

188. 1939. Les algues marines de la Cote des Alberes. IV - Rhodophycees. Revue algol. 11:

247-330.

Note. Includes Bangiales, Nemalionales, Gelidiales and Cryptonemiales.

189. —1941. Les algues marines de la Cote des Alberes. IV -Rhodophycees (suite). Revue algol. 12:
77-100.

Note. Covers the Gigartinales and Rhodymeniales.

190. 1942. Les algues marines de la Cotes des Alberes. IV. Rhodophycees (fin). Trav. algol. 1:

29-113.

Note. Covers Ceramiales. Travaux algologiques 1 replaced volume 13 of the original series of Revue
algologique.

191. - 1946. La flore marine des lies Atlantides. Mem. Soc. Biogeogr. 8: 395-435.

192. - 1951. La flore marine de 1'Afrique du Nord. C. r. somm. Seanc. Soc. Biogeogr. 28: 103-108.

194. — & Hamel, G. 1934. Observations sur quelques Gelidiacees. Revue gen. Bot. 46: 528-549.

195. - 1936. Floridees de France VII. Gelidiales. Revue algol. 9: 85-140.

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 261

196. Feldmann-Mazoyer, G. 1941. Recherches sur les Ceramiacees de la Mediterranee Occidental.

Alger.
Foslie, M. 18980. Systematical survey of the Lithothamnia. K. nor. Vidensk. Selsk. Skr. 1898 (2):

1-7.
198. - 19006. New or critical calcareous algae. K. nor. Vidensk. Selsk. Skr. 1899 (5): 1-34, 1900.

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1-22.

212. — & Printz, H. 1929. Contributions to a monograph of the Lithothamnia . . . after the author's
death collected and edited by Henrik Printz. Kgl. norske Vidensk. Selsk. Museet. Trondheim.

213. Fox, M. 1957. A first list of marine algae from Nigeria. J. Linn. Soc. (Bot.) 55: 615-631.
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214. Fremy, P. 1936. Marine algae from the Canary Islands especially from Teneriffe and Gran Canada

IV. Cyanophyceae. K. dansk. Vidensk. Selsk. Skr. 12(5): 1-43.

215. Gain, L. 1914. Algues provenant des campagnes de VHirondelle II (1911-1912). Bull. Inst.

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Gargiulo, G. M., De Masi, F. & Tripodi, G. 1985. A study on Gracilaria dendroides sp. nov.
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219. Gaudichaud [-Beaupre], C. 1826; 1827. Botanique [pp. viii + [l] + 522] . . . [Part 4, Livre II.

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Note. This represents one volume of an eight volume work. It is stated in the preface (p. vii) that the
'thalassiophytes' were determined by 'M. Agardh' (doubtless C. Agardh). Part 4, pp. 129-168, was
published in June 1827. There are earlier geographically arranged chapters, pp. 3-146, that make occasional
mention throughout of seaweeds for relevant areas. The Atlas is not relevant to the present work; there are
no algal plates.

220. Gauld, D. T. & Buchanan, J. B. 1959. The principal features of the rock shore fauna in Ghana.

Oikos 10: 121-132.

221. Gayral, P. 1958. La nature au Maroc II Algues de la cote Atlantique marocaine. Rabat.

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578. - 1912. Marine algae of the Scottish National Antarctic Expedition [Part VI, pp. 73-83].

In W. S. Bruce & R. N. R. Brown (Eds), Report on the scientific results of the voyage of S. Y.
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224. Gerloff, J. 1957. Einige Algen aus der Bucht von Daressalaam. Willdenowia 1(5): 757-770.

225. Gil-Rodriguez, M. C. 1980 ['1979']. Revision taxonomica-ecologia del genero Cystoseira C. Ag. en

el archipielago Canario. Vieraea 9(1-2): 115-148.

226. — & Afonso-Carrillo, J. 1980. Adiciones a la flora marina y catalogo ficologico para la Isla de
Lanzarote. Vieraea 10 (1-2): 59-70.

227. - 1981 ['1980']. Catalogo de las algas marinas bentonicas (Cyanophyta, Chlorophyta,
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229. — & Wildpret de la Torre, W. 1980a. Contribucion al estudio de la vegetacionficologica marina
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230. - 19806. Contribucion a la ficologia de la Isla del Hierro. Vieraea 8(2): 245-260.

231 . — Acebes Ginoves, J. R. & Perez de Paz, P. L. 1978. Nuevas aportaciones a la flora ficologica de
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232B. - Haroun Tabraue, R. [J.], Afonso-Carrillo, J. & Wildpret de la Torre, W. 1985. Adiciones al
catalogo de algas marinas bentonicas para el Archipielago Canario. II. Vieraea 15(1-2):
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233. Gomez Garreta, A., Ribera Siguan, A. & Seoane-Camba, J. 1979. Nuevas citas para la flora
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262 J. H. PRICE, D. M. JOHN & G. W. LAWSON

235. Gonzalez Henriquez, M. N. 1976. Contribution al estudio del epifitismo en Zostera marina L.

(Zosteraceae) en la playa de Las Canteras (Gran Canada). Botanica Macaronesica 2: 59-67.

236. Gonzalez, N. 1977a. Estudio de la vegetation literal de la zona de Maspalomas. Botanica

Macaronesica 4: 23-30.

237. - 1977b. Estudio de la vegetation bentonica literal del noroeste de la Isla de Gran Canada
(Banaderos, San Felipe, Sardina, Las Nieves). Botanica Macaronesica 4: 85-104.

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Botanica Macronesica 5: 47-60.

239. Goor, A. C. J. van 1923. Die Hollandischen Meeresalgen (Rhodophyceae, Phaeophyceae und
Chlorophyceae) insbesondere der Umgebung von Helder, des Wattenmeeres und der
Zuidersee. Verh. K. Akad. Wet., Amst., Tweedesectie, 23(2): I-IX + [1] + 1-232.
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242. Grunow, A. 1868. Algae. In E. Fenzl (Ed.), Reise der Osterreichischen Fregatte Novara urn die

Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wullerstorf-
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589. Guiry, M. D. 1984. Structure, life history and hybridization of Atlantic Gigartina teedii (Rho-

dophyta) in culture. Br. phycol. J. 19: 37-55.

447. —1985? ['1984']. Photoperiodic and temperature responses in the growth and tetrasporogenesis

of Gigartina acicularis (Rhodophyta) from Ireland. Helgoldnder wiss. Meeresunters . 38: 335-
347.
Note. Reverse of front cover states printed 31. XII. 1984- hence publication date suggested as 1985.

243. — & Cunningham, E. M. 1984. Photoperiodic and temperature responses in the reproduction of
north-eastern Atlantic Gigartina acicularis (Rhodophyta: Gigartinales). Phycologia 23: 357-
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243A. - - & T.-Freamhainn, M. 1986 ['1985']. Biosystematics of Gracilaria foliifera (Forsskal)

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246. Hamel, G. 1927. Recherches sur les genres Acrochaetium Naeg. et Rhodochorton Naeg. Saint Lo.

247. - 1924-1930. Floridees de France. I-II. Revue algol. 1: 278-292, 427-457, 1924; III-IV. Rev.
algol. 2: 39-67, 280-309, 1925; V. Revue algol. 3: 99-158, 1928; VI. Revue algol. 5: 61-109,
1930.

Note. Reprint of I-II, repaged continuously l-46 + [l]; III, repaged [2] + 50-80; IV, repaged [in error],
69-98; V, repaged 99-158 [repeats original]; VI, repaged 1-49.

249. Hariot, P. 1895. Liste des algues recueillies au Congo par M. H. Lecomte. J. Bot. Paris 9:

242-244.

250. 1896 ['1895']. Contribution a la flore algologique du Gabon et du Congo franc,ais. C. r. Ass. fr.

Avanc. Sci. 24(2); 641-643.

251. - 1908. Les algues de San Thome (cote occidentale d'Afrique). J. Bot. Paris II, 1: 161-164.

252. 1911. Algues de Mauritanie recueillies par M. Chudeau. Bull. Soc. hot. Fr. 58 [= IV, 11]:

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253. Haroun Tabraue, R. J., Gil-Rodriguez, M. C., Afonso-Carrillo, J. & Wildpret de la Torre, W. 1984

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13: 259-276.
253A. - 1985? ['1984']. Estudio ecologico y fenologico de algunas especies del genero

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254. Harvey, W. H. 1846-1851. Phycologia britannica: . . ., vols II, III, Rhodospermeae , . . . vol. IV.

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255. 1860. Phycologia australica; or, a history of Australian seaweeds; ... 3 London.

256. — 1862. Phycologia australica; or, a history of Australian seaweeds; ... 4. London.

257. 1863. Phycologia australica; or, a history of Australian seaweeds ... [5] ... and a synopsis of

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RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 263

command of Captain George S. Nares, R.N., F.R.S. and the late Captain Frank Tourle
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260. - 18856. Ill - Report on the botany of the Bermudas and various other islands of the Atlantic
and Southern Oceans. In C. W. Thompson & J. Murray, Report on the scientific results of the
voyage of H. M.S. Challenger during the years 1873-76 under the command of Captain George S.
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261. Henriques, J. [A] 1885 ['1884']. Contribuicao para o estudo da flora d'algumas possessoes

portuguezas I Plantas colhidas por F. Newton na Africa occidental. Bolm Soc. broteriana 3:
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Note. See also no. 262.

262. Henriques, I. [= J.] [A.], [De Toni, G. B., & Levi, D.] 1886. Contribugao para o estudo da flora

d'algunas possessoes portuguezas. Plantas colhidas por F. Newton na Africa occidental, (dal
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Note. This work was published in April 1886. An extract from Henriques (1885) (261); the reference has
been cited as here since there is definite evidence that the text was affected by editing before reproduction in
Notarisia. Mistakes present in the original have been corrected and new ones introduced.

263. Henriques, J. [A] 1886. Algae [pp. 217-221]. In J. [A.] Henriques, Contributes para o estudo da

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264. - 1887. Flora de S. Thome. - [130] [pp. 381-383]. In G. B. De Toni & D. Levi, Contributiones
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Note. A complete extract from Henriques (1886) (263); the present text has been attributed to Henriques
solely, as there appear to be no alterations in the algal text.

265. — 1917. Catalogo das especies de animais e plantas ate hoje encontradas na ilha de S. Tome.
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267A. Hoek, C. van den 1982. The distribution of benthic marine algae in relation to the temperature

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268. Hooker, J. D. [& Harvey, W. H.] 1847. LV. Algae, L. In J. D. Hooker, The botany of the antarctic

voyage of H.M. Discovery ships Erebus and Terror, in the years 1839-1843 ... 1. Flora
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269. Hoppe, H. A. 1969. Marine algae as raw materials. In T. Levring, H. A. Hoppe & O. J. Schmid,

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270. — & Schmid, O. J. 1962. Meeresalgen als moderne Industrieprodukte. In H. A. Hoppe (Ed.),
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271 . Hornemann, J. W. 1819. Anniversaria in memoriam reipublicae sacrae et litterariae cum universae,

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Note. According to Hepper & Neate (1971) and Hepper (1976), Isert (the source of the material studied by
Hornemann) collected both along the Ghanaian coastal part of what was Danish Guinea and at Whydah ( =
Ouidah), in present day Benin. Since the whole coast of Benin and Togo is, with the exception of more recent
artificial installations and rocks exposed due to their interference with the natural west-east longshore drift of
sand, sandy and/or lagoon in configuration and structure, we attribute the algal records (clearly from rocky
substrata, if involving attached material) solely to the Ghanaian stretches. This ignores the possibility that
Isert could have collected drift algal material in Togo and Benin.

Howe, M. A. 1917. A note on the structural dimorphism of sexual and tetrasporic plants of
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1920. Algae [pp. 553-618]. In N. L. Britton & C. F. Millspaugh, The Bahama Flora. New

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Phycologia 25: 271-296.

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273. Irvine, L. M. 1983. Seaweeds of the British Isles 1 Rhodophyta Part 2 A Cryptonemiales (sensu

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277. Itono, H. 19776. Studies on the southern Japanese species of Galaxaura (Rhodophyta). Micro-

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278. — 1980. The genus Galaxaura (Rhodophyta) in Micronesia. Micronesica 16: 1-19.
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Note. A pair of extracts from the July 1850 and May 1851 numbers of Nouvelles Annales de la Marine et des
Colonies. Texts clearly repaged, but division of present text probably represents at least the break between
the parts as originally published, pp. 1-8 and pp. 9-19.
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286. John, D. M. 1972a. A new species of Botryocladia (Rhodophyceae, Rhodymeniales) from the Gulf

of Guinea. Phycologia 12: 33-36.

287. - 19726. The littoral ecology of rocky parts of the north-western shore of the Guinea Coast.
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288. - 1977 ['1976']. The marine algae of Ivory Coast and Cape Palmas in Liberia (Gulf of Guinea).
Revue algol. II, 11: 303-324.

290. - 1986. Littoral and sub-littoral marine vegetation. In G. W. Lawson (Ed.), Plant ecology in
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291. - & Asare, S. O. 1975. A preliminary study of the variations in yield and properties of
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292. - & Lawson, G. W. 1972 ['1971']. Additions to the marine algal flora of Ghana I. Nova
Hedwigia2l:8ll-841.

293 . — 1972. The establishment of a marine algal flora in Togo and Dahomey (Gulf of Guinea) .
Botanica mar. 15: 64-73.

294. — 1974. Observations on the marine algal ecology of Gabon. Botanica mar. 17: 249-254.

295. — I911a. The marine algal flora of the Sierra Leone Peninsula. Botanica mar. 20: 127-135.

296. - 1977 'b. The distribution and phytogeographical status of the marine algal flora of
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487. Round, F. E. 1981. The ecology of algae. Cambridge.

Santelices, B. & Stewart, J. G. 1985. Pacific species of Gelidium Lamouroux and other Gelidiales
(Rhodophyta), with keys and descriptions to the common or economically important species
[pp. 17-31]. In I. A. Abbott & J. N. Norris (Eds), Taxonomy of economic seaweeds With
reference to some Pacific and Caribbean species. La Jolla, California.

489. Santos Guerra, A. 1972. Contribucion al estudio de la flora marina de la Isla de La Gomera. Vieraea

2(1): 86-102.

490. — Acuna G.[onzales], A. & Wildpret [De La Torre], W. 1970. Contribucion al estudio de la flora
marina de la Isla de La Palma. Cuad. Bot. canaria 9: 20-29.

491 . Sanusi, S. S. 1980. A study of grazing as a factor influencing the distribution of benthic littoral algae.

M.Sc. thesis, University of Ghana.

270 J. H. PRICE, D. M. JOHN & G. W. LAWSON

493. Sauvageau, C. 1912. A propos des Cystoseira de Banyuls et Guethary. Bull. Stn biol. Arcachon 14:
133-556.

Note. There also exists a separate, repaged 1-424.
495. Schiffner, V. 1931. Neue und bemerkenswerte Meersalgen. Hedwigia 71: 139-160; 161-205.

497. Schmidt, O. C. 1929a. Beitrage zur Kenntnis der Meeresalgen der Azoren I. Hedwigia 69: 95-

113.

498. — I929b. Beitrage zur Kenntnis der Meeresalgen der Azoren II. Hedwigia 69: 165-172.

499. — 1931. Die marine Vegetation der Azoren in ihren Grundziigen dargestellt. Biblthca hot. 102:
1-116.

500. - & Gerloff, J. 1957. Die marine Vegetation Afrikas in ihren Grundziigen dargestellt.
Willdenowia 1:709-756.

503. Schmitz, F. & Hauptfreisch, P. 1896. Chaetangiaceae. In A. Engler & K. Prantl, Die naturlichen

Pflanzenfamilien 1(2): 335-339. Leipzig.

504. - 1896. Gelidiaceae. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien 1(2):
340-349. Leipzig.

506. — 1896-97. Sphaerococcaceae. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien

1(2): 382-396. Leipzig.
509. - 1897. Gloiosiphoniaceae. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien

1(2): 505-508. Leipzig.
Schneider, C. W. 1975. Taxonomic notes on Gracilaria mammillaris (Mont.) Howe and Gracilaria

veleroae Dawson (Rhodophyta, Gigartinales). Taxon 24: 643-646.
514. Schotter, G., Feldmann, J. & Magne, M.-F. 1968. Recherches sur les Phyllophoracees. Bull. Inst.

oceanogr. Monaco 67(1383): 1-99.
570. Seagrief, S. C. 1984. A catalogue of South African green, brown and red marine algae. Mem. hot.

Surv.S.Afr.47:i-vi + l-12.
Note. For the purposes of this part of the list, this reference has been allocated a number since it contains

records for the area.

517. Seoane-Camba, J. 1965. Estudios sobre las algas bentonicas en la costa sur de la Peninsula Iberica

(literal de Cadiz). Investigation pesq. 29: 3-216.

574. — 1977. Sur une nouvelle espece de Gelidiella trouvee aux lies Canaries: Gelidiella tinerfensis

nov. sp. Bull. Soc. phycol. France 22: 127-134.

518. — 1979. Sobre algunas Gelidiaceas nuevas o poco conocidas de las costas Espanolas. Acta hot.
malacitana 5: 99-112.

Soli-hell, W. A. & Mason, L. 1943. New or little known crustaceous corallines of Pacific North
America. Proc. NatlAcad. Sci. U.S.A. 29: 87-92.

519. Simons, R. H. 1964. Species of Plocamium on the South African coast. Bothalia 8: 183-193.

521. - 1970. Marine algae from southern Africa 1. Six new species from the inter- and infra-tidal
zones. InvestlRep. Div. Sea Fish. Rep. S. Afr. 88: [4] + 1-13.

522. — 1973. Unpublished list (in litt.) of species from South West Africa, principally collected during
a Graves/Isaac/Lawson/Simons field trip in 1957. Known to be incomplete.

522A. - — [Lawson, G. W. & Isaac, W. E.] 1982. Namibian seaweeds and their biogeographic affinities.
Unpublished manuscript based on the same trip and data as in 348 and 522 (q.v.).

523. - 1974. Algae, (including diatoms and seaweeds) [pp. 239-261]. In J. H. Day, N. A. H. Millard,
& M.-L. Penrith, A guide to marine life on South African shores. 2nd ed. Cape Town &
Rotterdam.

Note. The first edition (not seen) was probably 1969, since that is the copyright date and the Preface is
dated April 1968. Overall tripartite editorship is obvious from both Copyright allocation and statements in
Preface. Latter also clearly indicates responsibility of R. H. Simons for algal section. Presumably algal
content of first edition was as in second, since preface statement unchanged from past. Relevant records are
for Namibia.

Sjdstedt, L. G. 1926. Floridean studies. Lunds Univ. Arsskr. N.F. Avd. 2, 22(4): 1-95.
Sonder, O. W. 1845. Nova algarum genera et species quas in itinere ad oras occidentales Novae
Hollandiae, collegit L. Preiss, Ph. Dr. Bot. Ztg 3: 49-57.
— 1846. Algae. In C. Lehmann, Plantae Preissianae 2: 148-160. Hamburg.

528. Sonder, [O. W.] 1852. Algae. In J. A. Schmidt, Beitrage zur Flora der Cap Verdischen Inseln. Mit
Beriicksichtigung alter bis jetzt daselbst bekannten wildwachsenden und kultivirten Pflanzen.
Nach eigenen Untersuchungen und mit Benutzung der gewonnenen Resultate anderer Reisenden:
125/127. Heidelberg.

Note. The algal material was sent to Sonder in Hamburg; he worked on the plants and provided both the
determinative data and text as printed.

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 271

529. Sourie, R. 1954a. Contribution a 1'etude ecologique des cotes rocheuses du Senegal. Mem. Inst. fr.

Afr. noire 38: 1-342 + [!].

Note. From the note on p. 117, it is clear that the algae were worked on mainly by J. Feldmann, but that
Sourie took account of some of the views of Dangeard as expressed in the latter's memoir on the Cap Vert
(Dakar) peninsula algae. Since the exact contribution of the various people involved is in doubt, we have left
the reference in the name of Sourie, who seems to have exercised overall authorship.

530. - 19546. Principaux types de zonations verticales des algues sur le littoral rocheux de la
presqu'ile du Cap Vert (Zone intercotidale). Rapp. Commun. int. hot. Congr. 8(17):
151-153.

531. — 1954c. Etude ecologique sommaire des fonds sableux en Baie de Dakar. Annls EC. sup. Sci.
Dakar 1:141 -155.

Note. Sourie stated (p. 141) that many of the specific determinations of algae were by J. Feldmann.
533. Southward, A. J. 1958. The zonation of plants and animals on rocky sea shores. Biol. Rev. 33:

137-177.
Stafleu, F. A. & Cowan, R. S. 1976. Taxonomic literature. A selective guide to botanical publications

and collections with dates, commentaries and types 1: A-G. 2nd ed. [Regnum veg. 94].

Utrecht.
535. Steentoft, M. 1967. A revision of the marine algae of Sao Tome and Principe (Gulf of Guinea). /.

Linn. Soc. (Bot.) 60: 99-146.
599. Stegenga, H. 1986. The Ceramiaceae (excl. Ceramium) (Rhodophyta) of the South West Cape

Province, South Africa. Biblthca phycol. 74: 1-149.

537. Stephenson, T. A. & Stephenson, A. 1972. Life between tidemarks on rocky shores. San Francisco.
Sunding, P. 1972. A botanical bibliography of the Cape Verde Islands. Oslo [Botanical Garden,

University of Oslo]. Stencilled.

- 1973. A botanical bibliography of the Canary Islands. 2nd ed. Oslo [Botanical Garden,
University of Oslo]. Stencilled.

538. Svedelius, N. 1911. Florideae. In A. Engler & K. Prantl, Die natiirlichen Pflanzenfamilien 1(2):

200-276. Leipzig.

539. Tandy, G. 1944. Algae. In A. W. Exell, Catalogue of the vascular plants ofS. Tome (with Principe

and Annobon): 386, Appendix I. London.
Taylor, W. R. 1939. Algae collected on the presidential cruise of 1938. Smithson. misc. Collns 98:

1-18.
1945. Pacific marine algae of the Allan Hancock expeditions to the Galapagos Islands. Allan

Hancock Pacific Expedns 12: [8] + i-iv + 528.

540. 1960. Marine algae of the eastern tropical and subtropical coasts of the Americas. Ann Arbor.

542. Trochain, J. 1940. Contribution a 1'etude de la vegetation du Senegal. Mem. Inst. fr. Afr. noire 2:

[1-6] + 1-433 + [63].
Note. J. Feldmann clearly had a great deal to do with the main determinations on which the algal list (pp.

108-110) was based; since the extent to which the data were accepted or amended by Trochain is not clear,

and since there are other parts to the text which seem definitely to have been attributable to Trochain, we

have accepted the latter as overall author. For individual comments on species, the more correct authorship

citation would undoubtedly be 'Feldmann, J., in Trochain, J.,' etc.
Turner, D. 1809. Fuci sive plantarum fucorum generis a botanicis ascriptarum ... pis 121 (Vol. II;

May 1809) and 147 (Vol. Ill; October 1809). London.
Umamaheswara Rao, M. 1972. On the Gracilariaceae of the seas around India. J. mar. biol. Ass.

India 14: 671-696.

546. Varo, J., Ramirez, J. & Renteria, J. 1979. Estudio de la vegetacion bentonica del litoral granadino.

Acta hot. malacitana 5: 79-98.

547. Vickers, A. 1897? ['1896']. Contribution a la flore algologique des Canaries. Annls Sci. nat. (Bot.)

VIII, 4: 293-306.

Note. The date is somewhat difficult to cite as there is some confusion regarding the dates of various issues.
It does seem possible that pre-prints were issued in 1896; this is the date usually cited (see Lawson & Price,
1969: 345-346).

604. Viera-Rodriguez, M. A. 1987. Contribucion al estudio de la florula bentonica de la isla de La

Graciosa. Canarias. Vieraea 17: 237-259.

608. — & Wildpret de la Torre, W. 1986. Contribucion al estudio de la vegetacion bentonica de la isla

de La Graciosa. Canarias. Vieraea 16: 211-231.

605. — Gil-Rodriguez, M. C., Audiffred, P. A. J., Prud'homme van Reine, W. F., Haroun-Tabraue,
R. & Wildpret de la Torre, W. 1987. Contribucion al estudio de la florula bentonica del islote de
Montana Clara. Canarias. Vieraea 17: 271-279.

272 J. H. PRICE, D. M. JOHN & G. W. LAWSON

548. Viera y Clavijo, J. de 1866; 1869. Dlccionario de historia natural de las islas Canarias, ... 1 (A-G).
Gran Canada.

Note. The background to this work is explained in detail by Martin Aguado (1957) , who outlined (p. 8) the
career of Viera y Clavijo and the progress of the work. The MS was completed in 1799, with the title
Diccionario de Historia Natural de las Canarias, but was not published until the indicated dates. See also
references 549 and 386. A further version of the work appeared in 1942 under the Tublicaciones de la
Biblioteca Canada' series (Tenerife). A more recent (1982) new edition was edited by M. Alvar and included
an introduction and appendix, with purely historical/literary data.

551. Webb, P. B. 1849. Spicilegia Gorgonea; or a catalogue of all the plants as yet discovered in the Cape
de Verd Islands . . . In W. J. Hooker, Niger Flora . . .: 89-197. London, Paris & Madrid.

553. Weber-van Bosse, A. 1921. Liste des algues du Siboga II Rhodophyceae Premiere partie Proto-
florideae, Nemalionales, Cryptonemiales. In M. Weber, Siboga-Expeditie . . . Monographic
LIXb. Uitkomsten op Zoologisch, Botanisch, Oceanographisch en Geologisch Gebied. . . . Livr.
LXXX1X: [6] + 187-392+ [4].

556. Weisscher, F. C. M. 1983. Marine algae from Selvagem Pequena (Salvage Islands). Bolm Mus.

munic. Funchal35: 41-80.

556A. - - Audiffred, P. A. J. & Duineveld, G. C. A. 1982 [15 November 1982]. MS list (in litt.) from
Prud'homme van Reine on Netherlands CANCAP Expeditions to the Canaries and Salvage
Islands. (See also entry 557.)

557. — Prud'homme van Reine, W. F. & Duineveld, G. C. A. 1985. Marine algal vegetation of Bahia
del Confital near Las Palmas de Gran Canaria. Unpublished manuscript [from Prud'homme van
Reine] on the findings of the Netherlands CANCAP Expeditions (see entry 556 A).

Woelkerling, W. J. 1988. The coralline red algae: an analysis of the genera and subfamilies of

nongeniculate Corallinaceae (Rhodophyta). London.
561. Womersley, H. B. S. & Bailey, A. 1970. Marine algae of the Solomon Islands. Phil. Trans. R. Soc.

B, 259: 257-352.

36B. Wynne, M. J. 1986. Report on a collection of benthic marine algae from the Namibian coast
(southwestern Africa). Nova Hedwigia 43: 311-355.

- 1986. A checklist of benthic marine algae of the tropical and subtropical western Atlantic.
Can.J. Bot. 64:2239-2281.

566. See Zaneveld, J. S. 1956.

567. See Liming, K. 1985.

570. See Seagrief, S. C. 1984.

571. Xia, B. & Abbott, I. A. 1985. The genus Polycavernosa Chang et Xia (Gracilariaceae, Rhodo-

phyta): a comparison with Gracilaria Grev., and a key to the species [pp. 157-162]. In I. A.
Abbott & J. N. Norris (Eds), Taxonomy of economic seaweeds With reference to some Pacific
and Caribbean species. La Jolla, California.

572. See Chang, C. F. & Xia, B. M. 1963.

574 . See Seoane-Camba, J . 1 977 .

575. See Fan, K.-C. & Papenfuss, G. F. 1959.

576. See Afonso-Carrillo, J. 1985.

577. See Riouall, R. et al. 1985.

578. See Gepp, A. & Gepp, E. S. 1912.

579. See Carter, A. R. & Anderson, R. J. 1986.

581 . See Cardell Cristellys, E. et al. 1977.

582. See Afonso-Carrillo, J. et al. 1985.

583. See Haroun Tabraue, R. J. et al. 1985.

584. See Ribera Siguan, M. A. et al. 1985.

585. See Abbott, I. A. 1962.

586. See Lawson, G. W. & John, D. M. 1987.

588. See Ardre. F. & Gayral, P. 1961 .

589. See Guiry, M. D. 1984.

590. See John, D. M. & Lawson, G. W. (unpublished).

592. See Bird, C. J. & Oliveira filho, E. C. de 1986.

593. See McLachlan, J. & Bird, C. J. 1986.

594. See Lawson, G. W. 1954.

596. See Bailey, J. W. & Harvey, W. H. 1862.

597. See Prud'homme van Reine, W. F. & Lobin, W. 1986.

598. See Prud'homme van Reine, W. F. (in litt. 10/4/87).

599. See Stegenga, H. 1986.

RHODOPHYTA (pLORIDEAE) OF TROPICAL AFRICA 273

600. Xia, B. & Abbott, I. A. 1987. New species of Polycavernosa Chang & Xia (Gracilariaceae,
Rhodophyta) from the western Pacific. Phycologia 26: 405-418.

604 . See Viera-Rodriguez , M . A . 1 987 .

605. See Viera-Rodriguez, M. A. et al. 1987.

608. See Viera-Rodriguez, M. A. & Wildpret de la Torre, W. 1986.

609. Yamamoto, H. 1978. Systematic and anatomical study of the genus Gracilaria in Japan. Mem. fac.

Fish., Hokkaido Univ. 25: 97-152.

566. Zaneveld, J. S. 1956. Economic marine algae of tropical south and east Asia and their utilisation.
Indo-Pacific Fisheries Council, Serial publications No. 3: [2] + 1-55.

610. See Gonzalez, N. 1979.

61 1 . See John, D. M. & Lawson, G. W. 1988.

612. See Rotmann, K. W. G. 1987.

614. See Maggs, C. A. & Guiry, M. D. 1988.

British Museum (Natural History)

MACROLICHENS OF EAST AFRICA

T. D. V. Swinscow and H. Krog

Dr Swinscow was formerly Deputy Editor of the British Medical Journal.
Dr Krog is Professor of Taxonomic Botany at the University of Oslo.

This book is based mainly on collections made in the field by the authors. It covers
77 genera and 629 species. It is the first substantial study of a tropical lichen flora to
be undertaken by modern research methods. Thin-layer chromatography has been
used throughout, and the great majority of species have been studied by
microscopic examination of microtome sections. The nomenclature has been
thoroughly revised, and in all cases the basionym is given. The book will be
indispensable to students of the lichens of the African continent and valuable to
readers interested in lichens throughout the tropics.

Summer 1988, viii -I- 384pp, 185 figs., 16pp colour illustrations.
Hardback. 0 565 01039 5. £20.00

Titles to be published in Volume 18

An illustrated catalogue of the type specimens in the Greville diatom herbarium

By David M. Williams

Erik Acharius and his influence on English lichenology

By David J. Galloway

Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV.
Rhodophyta (Florideae) 2. Genera G

By James H. Price, David M. John and George W. Lawson

A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent

By Christopher R. Fraser- Jenkins

Some Cretaceous and Paleogene Trinacria (diatom) species

By Patricia A. Sims and Robert Ross

Corydalis (Papaveraceae: Fumarioideae) in Nepal

By Magnus Liden.

Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester