ISSN 0968-0446

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THE NATURAL
HISTORY MUSEUM

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Botany Series

THE

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HISTORY
MUSEUM

VOLUME 24 NUMBER 1 23 JUNE 1994

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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.)
The Natural History Museum, 1994

Botany Series
ISSN 0968-0446 Vol. 24, No. 1, pp. 1-100

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Bull. not. Hist. Mus. Land. (Bot.) 24(1): 1-34

Issued 23 June 1994

Pre-Linnaean references for the Macaronesian
flora found in Leonard Plukenet's works and
collections

THE NATURAL

tiCTOR

12 JUL tta-r

PRESENTED
GENERAL LIBRARY

JAVIER FRANCISCO-ORTEGA*

Department of Botany, The University of Texas, Austin,
Texas 78713-7640, USA

ARNOLDO SANTOS-GUERRA

Jardin de Aclimatacion de La Orotava, Calle Retama Num. 2, E-38400 Puerto de la Cruz, Tenerife,
Canary Islands, Spain

CHARLES E. JAR VIS

Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD

CONTENTS

Introduction 1

Taxa found in Plukenet's works 3

Taxa found in Plukenet's herbarium but not in his works 29

Discussion 30

References 31

Systematic index 33

Bibliography abbreviations found in Plukenet's works 33

SYNOPSIS. A review of early references to the flora of the Macaronesian region has been carried out through the
study of Plukenet's publications and his herbarium collection, now part of the Sloane Herbarium in The Natural
History Museum in London. A total of 97 descriptions and 54 drawings of Macaronesian plants has been located in
the four published works of this English herbalist that appeared between 1691 and 1705. 131 specimens from
Macaronesia representing 87 taxa have been found in his collection; 33 of them do not have obvious descriptions in
his published works and five descriptions, supposedly of Canarian plants, seem to have been incorrectly assigned to
this region. Phrase-names described by earlier authors that were cited as synonyms by Plukenet have also been
studied but few of them proved to be clearly related to Macaronesian taxa. This study reveals that Plukenet's work
provides the single most important pre-Linnaean account of the Macaronesian flora, and his herbarium contains one
of the oldest known collections of herbarium specimens from this region. The name Campanula canariensis L. (=
Canarina canariensis (L.) Vatke) is lectotypified.

INTRODUCTION

The Macaronesian region comprises the archipelagos of the
Canary Islands, Salvages, Azores, Madeira and Cape Verde
and has strong links with the northwestern parts of Africa
from southern Morocco to Cape Verde. Its flora contains a
high number of endemics and has been traditionally regarded
as a relic of the flora which existed in the Mediterranean
basin during the Tertiary age (Sunding, 1979).

The earliest known reference to Macaronesian natural
history was given in the first century AD by Pliny 'the elder'
who mentioned the abundance of palm and pine trees in
'Canada' and how, in what seems to be the island of El
Hierro, there were 'Ferulae' trees which precipitated water

* Corresponding author

The Natural History Museum, 1994

(Pliny, 1826a; for reviews of Pliny's work and the eastern
Atlantic islands see Steffen (1944), Alvarez-Delgado (1945)
and Martinez-Hernandez (1992)).

The first European visitors who described the region
noticed the peculiarities of its flora in accounts that dated
from the fourteenth to sixteenth centuries (e.g. Niccoloso da
Recco in 1341 (Bonnet, 1943); Bontier & Le Vernier in the
early fifteenth century (Cioranescu, 1980); Gomez de Cintra
in the mid fifteenth century (Bonnet, 1940); P. Gomez
Escudero also in the mid fifteenth century (Morales-Padron,
1978); A. de Palencia in the late fifteenth century (Lopez de
Toro, 1970), Fernandes in 1507 (Santiago, 1947); Diaz-Tanco
in 1520 (Rodrfguez-Monino, 1934; Del Rio-Ayala, 1935);
Nicols (1583); Frutuoso (Serra-Rafols et al., 1964) and Torri-
ani in 1590 (Torriani, 1978) and Espinosa (1594)). Some of
these descriptions were based on the original names given by

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

the pre-Hispanic inhabitants of these islands. As early as 1341
Niccoloso da Recco found that the bark of some Canarian
trees could be used to produce red dyes. Similar accounts
were also reported in the early fifteenth century as extensive
areas of scrub of 'higuieres' (Euphorbia balsamifera Aiton) in
Lanzarote and of 'tarajal' (Tamarix canariensis Willd.) in
Fuerteventura were found by Bontier and Le Vernier respec-
tively (Cioranescu, 1980).

Nevertheless, the earliest published and most complete list of
endemics of the Macaronesian region, appears to have been
provided by Diaz Tanco in 1520. This Spanish traveller visited the
Canary Islands between 1505 and 1520 (Rodriguez-Monino,
1934) and published a list of at least 12 Canarian endemics, using
the original pre-Hispanic names such as 'bales' (Plocamapendula
Aiton), 'thabbaybas', actually 'tabaibas' (Euphorbia spp.) and
'tabinaste', actually 'tajinaste' (Echium spp.) (Del Rio-Ayala,
1935). Analogous descriptions were given by the English trader
T. Nicols who mentioned the local Spanish names of 'taybayba',
'barbusano' (Apollonias barbujana (Cav.) Bornm. and 'vinatico'
(Persea indica (L.) Spreng.), for the latter noting that it was a tree
'exceeding heavie, and will not rot in anie water' (Nicols, 1583).
However, the most comprehensive account of the Macaronesian
flora from the sixteenth century was given by G. Frutuoso in 1590
(Serra-Rafols et al. , 1964). This Portuguese naturalist apparently
visited the Canary Islands late in the sixteenth century and not
only listed many of their endemics but also briefly described the
vegetation of some of the islands.

Most of these works provided descriptions of the Canary
pine (Pinus canariensis C. Sm.) and the Lauraceae forests
and put special emphasis upon the orchil lichen (Rocella
spp.), the Canary palm (Phoenix canariensis Chab.), the
dragon-tree (Dracaena draco L.) and the rain-tree or 'garoe'
(Ocotea foetens (Aiton) Baill.) of El Hierro. In addition,
these references gave unique records concerning the use of
plants by the pre-Hispanic inhabitants of the islands (Garcia-
Morales, 1989) which have been confirmed through the study
of archaeological remains. There is clear archaeological evi-
dence of the pre-Hispanic use of Scirpus holoschoenus L.
(syn. Holoschoenus vulgaris Link) and Phoenix canariensis as
textile species (Galvan-Santos, 1980; Rodriguez-Santana,
1989), cropping of barley, wheat, lentil, broad bean and grass
pea (Del Arco-Aguilar et al., 1991; Martin-Rodriguez, 1992)
and the gathering of Pinus canariensis seeds, Pteridium
aquilinum (L.) Kuhn rhizomes (Mathiensen, 1960) and Vis-
nea mocanera L. f. fruits (Del Arco-Aguilar et al., 1991). The
latter was quoted by early chroniclers as having its fruits
eaten raw or used to make a kind of 'honey' or 'wine' known
as 'cuche' or 'chacerquen' (P. Gomez Escudero in Morales-
Padron, 1978; Espinosa, 1594).

Furthermore these early Canarians were able to make
weapons and tools using wood of Apollonias barbujana
(Cav.) Bornm., Juniperus phoenicea L., Neochamaelea pul-
verulenta (Vent.) Erdtman, Olea europaea L. ssp. cerasifor-
mis (Webb & Berth.) G. Kunkel & Sunding and Pinus
canariensis (Diego-Cuscoy, 1961).

Although a definitive study of the European knowledge of
the Macaronesian flora between the fourteenth and eigh-
teenth centuries has not yet been undertaken, there are
reports which suggest that some of the endemic taxa were
well-known by naturalists during this pre-Linnaean period.
An example can be found in the Historia general y natural de
las Indias by Fernandez de Oviedo (1548). This renowned
naturalist gave an extensive description of the 'rain-tree' on
the island of El Hierro. Later, in 1590, Torriani reproduced

an original drawing of one of its branches, which facilitated
the botanical identification of this species as Ocotea foetens
(Maynar, 1943). Similarly, in 1576 Clusius gave a detailed
description and drawing of Dracaena draco based on a tree
which he found growing in Lisbon in the garden of the
monastery of 'S. Maria a Gratia' (Arber, 1938).

Besides these references, it is also known that soon after
the conquest of the islands some species were used as wood
for building and firewood for sugar mills (Parsons, 1981).
Plants were also exploited for products such as dyes (from
Parmelia perlata (Huds.) Ach., Roccella spp., Laurus azorica
(Seub.) Franco), soda (from Zygophyllum fontanesii Webb &
Berth., Mesembryanthemum crystallinum L.), perfumes
(from Convolvulus scoparius L. f.), medicines (from Dra-
caena draco} and pitch (from Pinus canariensis) (Schenck,
1907; Viera y Clavijo, 1808; 1866-1869; Lobo-Cabrera,
1988). A strong trade based on these products was estab-
lished between the Canaries and the most important Euro-
pean ports during the fifteenth and sixteenth centuries
(Gonzalez- Yanes, 1953; Fernandez- Armesto, 1982; Lobo-
Cabrera, 1988) and many references to the granting of
permission for their exploitation are given in official resolu-
tions from the island council or 'cabildo' (Serra-Rafols, 1949;
Serra-Rafols & De la Rosa, 1952, 1965, 1970). Mention of
Canarian endemics can also be found in those documents that
were issued to establish the division of the land after the
conquest (Serra-Rafols, 1978; Moreno-Fuentes, 1988). These
documents are known as 'datas' and in them names of
Canarian plants are usually given, as the plants were often
used to indicate boundaries between different areas.

These few examples are drawn from only a small number of
pre-Linnaean references on Macaronesian endemic plants.
Further research is needed in order to provide a clearer
picture of the state of knowledge of early explorers, herbalists
and naturalists of the flora of this region.

Leonard Plukenet was one of the most outstanding bota-
nists of the seventeenth century. With its approximately 2000
plant drawings, his Phytographia (Phyt.) was one of the most
important pre-Linnaean works containing illustrations of
plants. This work was issued in four volumes between 1691
and 1694 (Plukenet, 1691o, 16916, 1693, 1694) and he also
had an extensive herbarium with approximately 8000 speci-
mens (Pulteney, 1790) which was the basis for many of his
drawings (Dandy, 1958). This large collection was the result
of his enthusiastic acquisition of specimens, particularly of
those exotic plants then grown in the most important gardens
of Britain. He was also in close contact with most of the
British herbalists of the time such as Doody, Cuningham,
Ray, Sloane and Petiver and was in correspondence with
other botanists from abroad (Pulteney, 1790). Most of his
collection is now at The Natural History Museum in London
(BM) where it comprises Volumes 84-105 of the Sloane
Herbarium. There are also two unnumbered volumes con-
taining his specimens which are titled 'Herbarium Vivum
Plukenetianum' (HVP) and Thesaurus Botanicus'. Between
1696 and 1705, Plukenet produced three other works
(Plukenet, 1696, 1700, 1705) in which he gave descriptions of
the plates illustrated in his Phytographia and also listed many
more plant species for which drawings were not published.
These works are Almagestum botanicum (Aim.), Almagesti
botanici mantissa (Mant.) and Amaltheum botanicum
(Amalth.) in which the taxa are enumerated alphabetically
under their polynomial names. Additional illustrations were
included in the last two of these works, and so about 2740

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

figures can be found in Plukenet's works as a whole. It seems
that only those plants regarded as exotic were included in the
Phytographia, and the other three publications contain the
rest of his collection (Pulteney, 1790).

The work of Plukenet was greatly admired by Linnaeus
who considered his Phytographia to be one of the most
extraordinary pieces of botanical illustration then published
(Linnaeus, 1736, 1751). Linnaeus himself frequently referred
to Plukenet's polynomials in describing and naming many of
his own species.

In this paper we will review the publications of this English
herbalist in relation to the Macaronesian flora. The aim of
this study has been to assess what knowledge of the flora of
the region was available to the European herbalists in the
seventeenth century.

Manuscripts from Plukenet's collection which are held in
the Sloane Collection at the Department of Manuscripts in
the British Library (BL) have also been studied (see Scott
(1904) for the location of Plukenet's manuscripts in the
Sloane Collection).

For most of the species listed in his works, Plukenet
referred to plant names given by previous herbalists. How-
ever references to these authors were given in an abbreviated
form and a key for them was presented in the final part of the
Almagestum. In order to locate possible previous accounts of
the Macaronesian taxa, earlier names cited by Plukenet have
also been studied. Nevertheless some of the abbreviated
references were not listed in Plukenet's key and we have
therefore attempted to trace them through the reviews of
botanical literature carried out by Linnaeus (1736), Pritzel
(1872), Jackson (1881) and Heller (1959) and through the
resources of the British Library and the Libraries of the
Royal Botanic Gardens, Kew and The Natural History
Museum. A list clarifying the abbreviations encountered in
those of Plukenet's descriptions that are covered in our study
appears at the end of this paper. It is hoped that this will help
other researchers to trace some of the early references
mentioned in Plukenet's works.

The possible influence of Plukenet's works on subsequent
studies of the Macaronesian flora was analysed by reviewing
the utilization of Plukenet's names by Linnaeus (Richter,
1835-1840) and Webb & Berthelot (1836-1850). Further
research in this area was undertaken through the study of
those identifications of Plukenet's drawings provided by
Giseke (1779) and Tenzel (1820) and of Linnaeus' annotated
copies of Plukenet's works held at the Linnean Society of
London.

Taxonomic determination of phrase-names or polynomials
given by Plukenet was undertaken after study of his her-
barium collection at the Department of Botany (BM). This
was facilitated by the fact that many of the specimens were
labelled by Plukenet with references to the plate and figure
numbers of his works. Plukenet's handwriting was compared
with that found in Dandy (1958) and in his correspondence
from the Sloane Collection at the Department of Manuscripts
in the BL. For those labels which apparently were not written
by Plukenet, specimens were checked with the original illus-
trations found in his works. Labelled specimens which do not
seem to be similar to their respective illustrations are also
reported in this study. There are also some collections that do
not have an obvious description or illustration in Plukenet's
published work and there are also Macaronesian specimens
which have no label.

It is hoped that the results presented here will help to

encourage further research concerning early references to the
flora of the Macaronesian region and will contribute to the
understanding of how the natural history of this area was
perceived by European naturalists of the seventeenth cen-
tury.

List of abbreviations. BL = British Library, BM = The
Natural History Museum, London (formerly the British
Museum (Natural History)), HS = Sloane Herbarium, HVP
= Herbarium Vivum Plukenetianum. The following abbre-
viations refer to the most frequently used references in this
study: Aim.: = Plukenet (1696), Amalth.: = Plukenet
(1705), Linn. = Linnaeus (1753), Gis. - Giseke (1779),
Mant. - Plukenet (1700), Phyt. - Plukenet (1691a, 1691ft,
1693, 1694), Ten. = Tenzel (1820), W.B. - Webb & Berthe-
lot (1836-1850).

TAXA FOUND IN PLUKENET'S WORKS

A total of 97 descriptions of Macaronesian plants has been
found in Plukenet's works, and each is listed below. In this
list each description is given an entry number followed by the
polynomial name used by Plukenet in bold. Text not relevant
to the actual description has been omitted, and is indicated by
'[. . .]'. Reference to Plukenet's published accounts and
illustrations is given after the phrase-name including the
appropriate page or plate number (abbreviated as 't.'), the
latter being followed by the figure number (abbreviated as
'f.'). Plates from Almagesti botanld mantissa and Amaltheum
botanicum are abbreviated using the code 't.'

Information compiled for each entry is treated under five
headings:

(1) Earlier names and references mentioned by Plukenet
(coded as Syn.:). Names are given in italics followed by the
reference author in parenthesis. Untraced names and refer-
ences are cited as they are found in Plukenet's text followed
by '[?]'.

(2) Citation of Plukenet's descriptions in works by subse-
quent authors dealing with the Macaronesian flora (coded as
His.:). The binomial name with which Plukenet's polynomial
has been identified by Linnaeus (1753), Giseke (1779),
Tenzel (1820) and Webb & Berthelot (1836-1850) (coded as
Linn., Gis., Ten. and W.B. respectively) is given in square
brackets.

(3) Taxonomic determinations of Plukenet's polynomials
(coded as Del.:). Labelled specimens used in establishing
these determinations (made by us) and their location in HS
are given in square brackets. Additional information is given
in square brackets. Nomenclature and taxonomy follows
Hansen & Sunding (1993); endemic taxa are marked with an
asterisk. Illegible handwriting is indicated as [illeg.]. Deter-
minations which are only based on Plukenet's description and
not on drawings or specimens from his collection are indi-
cated with '[?]'.

(4) Other herbarium specimens (coded as Oth.:). Under this
heading we list those specimens of Macaronesian taxa found
in Plukenet's collection which are not labelled with a corre-
sponding polynomial or a reference to Plukenet's works.

(5) Comments (coded as Com.:). Under this heading we
provide supplementary information including Spanish com-
mon names reported by Plukenet, origin of material and
other relevant details found in the original description or in

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

Plukenet's herbarium. This study of Spanish common names
given by Plukenet has drawn on the studies of Viera y Clavijo
(1866-1869), Ceballos & Ortuno (1951), Kunkel (1971) and
Santos-Guerra (1983).

1. Acetosa arborescens, subrotundo folio; ex Insulis
Fortunatis Aim.: 8 (1696), Mant.: 3 (1700), Phyt.: t.
252, f. 3 (1694).

Syn.: Lunaria magorum Arabum Ital. Lunaria di Magi Arabi
(Lobelius, 1576), Lunaria magorum Arabum (Lobelius,
1581), Lunaria Oxalidis rotundae folio (Bauhin, 1596), Oxalis
rotundi folia frequentissima: a Rovillio picta, sed non
descripta: sorte Lunaria Magorum Arabum (Pona, 1601),
Lunaria magorum Arabum quid? (Bauhin, 1623), Lunaria
magorum Arabum (Bauhin & Cherler, 1651).
His.: Rumex lunaria L. [Linn.; Gis.; Ten.; W.B.].
Det.: * Rumex lunaria L. [HS 93: 9, 95: 12].
Oth.:HS99: 14, HVP: 4.

8. Apocynum arboreum ad Elaeagni faciem accedens
Canariense, siliquis binis Nerii aemulis, (Cornicar
I it su km is vulgo) a p id bus recurvis Aim.: 35 (1696),
Phyt. :t. 260, 13(1694).

Syn.: Nelem-pala (Rheede tot Draakenstein, 1689).

Det.: Periploca laevigata Aiton [HS 99: 83].

Oth.:HS95:55.

Com.: Plukenet reported the common name 'Cornicar';

plants of this species are known as 'comical' in the Canaries.

This species was described twice by Plukenet (see entry 10),

this description being based on adult plants.

9. Apocynum scandens, angustis Rosmarinae foliis, e
Maderaspatan. Nannary-chedde Malabarorum. Huic
multum convenit Illucl, quod ex Insulis Canarinis, olim
nobis transmittebatur, & in Almagesto nostro
memoratur Amalth.: 19, t. 361, f. 1 (1705).

2. Adianthum album Canariense, ramosius Aim.: 11
(1696).

3. Alsine spuria repens ex Insulis Fortunatis folio
Hederae terrestris, molli, & incano Aim.: 24 (1696),
Phyt.: 1.256(1694).

Com.: None of the drawings published by Plukenet in Phyt.:
t. 256 appears to correspond with the description in the
Almagestum and we believe that the citing of this plate was an
error.

4. Amaranthus Siculus spicatus, radice perenni [. . .]
ex Insula Maderensi Aim.: 26 (1696), Phyt.: t. 260, f. 2
(1694).

Syn.: Amaranthus Siculus spicatus, radice perenni (Boccone,

1674).

His.: Achyranthes aspera L. [Linn.; Gis.; W.B.], A. argentea

Lam. [Ten.].

Det.: Achyranthes aspera L. [HS 95: 41; 99: 61].

Oth.:HS97:78.

5. A n a gal I id is facie Frutex Canariensis Aim.: 29
(1696).

6. Anonis viscosa lutea, non spinosa, minor. Ex Insula

Pico Amalth.: 15(1705).

Syn.: Anonis viscosa lutea non spinosa minor Lusitanica
(Hermann, 1698).

7. Anonis viscosa lutea mil is capreolata erecta foliis
splendentibus glabris [. . .] ex Insulis Fortunatis ad nos
allataestMant.: 15(1700).

Syn.: Plukenet gave the following synonym: Anonis (sorte)
annua lutea, siliqua glabra breviori (Morison, 1680). How-
ever, the name found in Morison's work is 'Anonis lutea
annua recta hirsuta viscosa siliquis hirsutis brevioribus, nobis\

10. Apocynum scandens angusto Rorismarini folio, ex
Insulis Fortunatis [. . .] Henio Hispanis vulgo Aim.: 37
(1696), Mant.: 17 (1700), Phyt.: t. 261, f. 2 (1694).

Syn.: Naru-nindi (Rheede tot Draakenstein, 1690), Apo-
cynum Hispanicum frutescens Linariae folio (Tournefort,
1694), Apocynum fruticosum scandens, Genistae, Hispanicae
facie, floribus luteis odoratis (Sloane, 1696). According to the
description (Mant.: 17) the name Apocynum (forte) caule
tenui alte scandens capsulis echinatis was used by Banister
(1693). However, we have been unable to find this name in
this work.

Det.: Periploca laevigata Aiton [HS 99: 85].
Com.: The Spanish common name 'Henio' is mentioned by
Plukenet. However, plants of this species are known as
'comical' in the Canary Islands. The species was described
twice by Plukenet (see entry no. 8), this second entry being
based on young plants which are morphologically rather
different from adult individuals.

11. Aquifolium laeve non spinosum, angustiore folio
Lauri; ex Insula Palma Amalth.: 19 (1705).

Det.: * Ilex canariensis Poir. [?].

Com.: Determination based on the description and on a

specimen found in HS 189: 12 in J. Cuningham's collection

from La Palma, made on the island in the late seventeenth

century.

12. Aquifolium amplissimis foliis Minus corrugatum ex
Insulis Fortunatis Aim.: 38 (1696), Mant.: 18 (1700),
Phyt.: t. 262, f. 1(1694).

His.: Ilex platyphylla Webb & Berth. [W.B.].

Det.: * Ilex perado Aiton ssp. platyphylla (Webb & Berth.)

Tutin [HS 95: 60].

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

13. Arbor Americana amplioribus subrotundis duris &
nervosis foliis, fructu pugni majoris magnitudine.
Insulis Fortunatis & Jamaicensibus nostratibus Mamee
nuncupatur [. . .] Mommina, s. Mamee arbore [. . .]
Hujus autem fructus ut Mala-granata sunt putamine
corticoso. Mammee Canarinae folijs, & facie, Arbor ex
Insula Johanna Aim.: 39 (1696), Mant.: 125 (1700),
Phyt.: t. 268, f. 1 (1694), Phyt.: t. 204, f. 2 (1694).

Syn.: Mamey (Nieremberg, 1635), Anda Brasiliensibus
(Marggraf, 1648), Arbor vinifera Couton Juglandi similis
(Bauhin & Cherler, 1650).

Det.: Mammea americana L. [a herbarium specimen for this
species is found in HS 96: 170. It has a label which states
'Mammee vera Arboris loium ex Insula Canarina' and does
not have any reference to Plukenet's works. However it
resembles the illustration depicted in Phyt.: t. 268, f. 1. There
is another specimen in HS 99: 91 which refers to this plate but
without any mention of a collection site. The plant given in
Phyt.: t. 268, f. 1 (HS 99: 110) is not of M. americana and it is
not a species from the Macaronesian flora. We can only
assume that Plukenet incorrectly assigned the illustration
from Phyt.: t. 268, f. 1 to his description of M. americana.
Plukenet also indicated that the drawing found in Phyt.: t.
204, f. 2 was of this species. However this plate and its
corresponding specimen (HS 96: 171) are of Bosea yervamora
L. which is a Canarian endemic. A description of this species
was also provided by Plukenet (for further discussion see
entry 15)].

14. An Arbor mirabilis ex Insula Ferro aquam still a us
Mant.: 171(1700).

Syn.: Sagapeni est liquor fruticis ferulacei oleandro montano
similis, bonum quede color e ex albo (Manadi & Sylvij, 1598),
Arbor aquam stillans (Bauhin & Cherler, 1650), Ombrion
nullis aedificiorum vestigiis, habere, in montibus stagnum
arbores similis ferulae ex quibus aqua exprimatur, e nigris
amara ex candidioribus potui iucunda (Pliny, 1826o), In
prima earum, cui nomem est Embrion, aedificia nee sunt nee
fuerunt, iuga montium stagnis madescunt ferulae surgunt ad
arboris magnitudinem: earum quae nigrae sunt, expressae
liquorem reddunt amarissimum, quae candidae, aquas revo-
munt etiam potui accomodatas (Solinus, 1958), Ferulae sunt
arborescentes, afferente Vossio, Not, in Pompom, Melam [?],
Sagapenum enim Ferulaceae Plantae gummi Dioscorides tra-
dit. lib. 3. cap 95 [?]. Plukenet also quoted the following
description from Galenus (1587) 'Sagapenum enim Ferulaceae
Plantae gummi esse. Quam. Panaci similem esse aif . However
we have been unable to trace it in this publication.
Det.: * Ocotea foetens (Aiton) Benth. & Hook.f. [there is no
herbarium specimen for this species in Plukenet's collection,
determination has been based on the sixteenth century illus-
tration given by Torriani (1978)].

15. Arbuscula baccifera Canariensis, Syringae
caeruleae foliis, purpurantibus venis, fructu
monopyreno. Yerva-mora Hispanorum [. . .]
Yerva-Mora ab Hispanis quoque dicitur Solanum
baccis rubris & aureis; Ipsum tamen Solani ethymon
ob quandam in quibusd partibus similitudinem longe
pluribus iisque diversissimis stirpium generibus ab
eodem Populo imponi consuevit: Affinitas ergo formae
& coloris baccarum eo sorsan Hispanos Canariense
induxit, ut hujus Arbusculam Yerva Mora (i.e.) Solani
titulo insigniverint Aim.: 42 (1696), Mant.: 21 (1700).

Syn.: Tilia sorte arbor racemosa, folio longiori subtus albi-
cante nervis purpureis insignito, flore pentapetalo purpureo
(Sloane, 1696).

His.: Bosea yervamora L. [Although Linnaeus published this
name originally in 1753, he cited Plukenet's polynomial as a
synonym only in the twelfth edition of his Systema naturae
(Linnaeus, 1767); W.B.].

Det.: * Bosea yervamora L. [a herbarium specimen found in
HS 96: 171 belongs to this species. Plukenet associated this
specimen with the figure in Phyt.: t. 204, f. 2 and the
description of entry 13 (Aim.: 39). However his description
of B. yervamora does not refer to this illustration. In fact in
Aim.: 39 he identified it as a species of Mammea; this is also
confirmed by one of the labels of the specimen which states:
'A Mamee Mamaya s. Momin'. Plukenet's accounts for B.
yervamora (Aim.: 42, Mant.: 21) are extremely precise in, for
example, the citation of the common name used in the
Canaries. It is therefore surprising that he did not assign the
illustration from Phyt.: t. 204, f. 2 (HS 96: 171) to the
description].

Com.: Plukenet reported the use of the name 'Yerba-Mora'
in the Canary Islands, where it is known as 'hierbamora',
'yervamora' and 'hediondo'. This name is also mentioned in
his description from Aim.: 181, Mant.: 99, namely Hedera
arbor ea ex argenteo & viridi foliis eleganter variegatis [. . .]
Neque prorsus abhorres a Yerva Mora Canariensibus Hispa-
nis. Hujus [. . .] non autem scandit haec sed erigitur (=
Hedera helix L.).

16. Arbuscula Canariensis Salicis, aut potius Oleae
Sylv. Barbadensium foliis & facie, seminibus ad tactum
(quando maturis) e capsulis cum strepitu profilientibus,
Snap Tree nostratibus vulgo. in Hort. Reg. Sancti
Jacobi apud Westmonasterium praeterito anno cura D.
Georg. London ex Seminibus natae sunt quamplurimae
hujus adolescentes arbusculae [. . .] Quoad Vascula
seminalia magnam habet convenientiam cum Curini
Speciebus Malabarorum Aim.: 44 (1696), Phyt.: t.
313, f. 1 (1694).

Det.: * Justicia hyssopifolia L. [HS 93: 140].
Com.: From the account of this species it seems that Plukenet
based his description on material grown by George London,
who was Master Gardener and Deputy Superintendent of the
Royal Gardens under William III. The description found in
entry 34 also seems referable to /. hyssopifolia.

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

17. Arbutus angustiori folio non serrato ex Insulis
Fortunatis. Aut potius Cerasus Canariensis Adrachne
foliis magis acuminatis, fructu parvo, Fragiformi,
monopyreno & Cerasis nostratibus plumirum divers!

Aim.: 49 (1696).

Syn.: Morocoks (Ogilvy, 1671).

Det.: Myrica faya Alton [a specimen of this species is found
in HS 95: 104; it does not bear any reference to Aim.: 49
except a label which states 'Arbutus non [illeg.] ex Canarinis';
a tentative determination has been made based on the
resemblance of this specimen to Plukenet's original descrip-
tion. It seems that this species was described twice as the
description is virtually identical to that found in entry 23].

18. Arbutus serratis foliis Canarina, cortice
circumrupto, sive duodecies Anni spat io, corticem
abjiciens, ex Insula Tenerifa nobis est allata Aim.: 49
(1696).

His.: Arbutus canariensis Veill. [W.B.].
Det.: * Arbutus canariensis Veill. [there is a specimen of A.
canariensis in HS 95: 104 which, although not explicitly linked
to any page number of the Almagestum, bears three labels
which agree with the original description from Aim.: 49].
Com.: One of the main features of this species is that
individuals change their bark regularly. This was noticed by
Plukenet and was mentioned both in the description (Aim.:
49) and on the specimen labels.

19. Atriplex angustifolia Canariensis, maritima,
dentata, repens. Flos de A In mo Hispanis dicta Aim.:
61, 399 (1696), Phyt.: t. 326, f. 3 (1694).

Syn.: Atriplex angustifolia maritima dentata (Ray, 1686),
Atriplex argentea dentata curassavica (Hermann, 1689).
Det.: Atriplex glauca L. [there is now no specimen upon
which Plukenet's account was based; a tentative determina-
tion has been made based on the drawing from the Phy-
tographia] .

Com.: The use of the common name 'Flos de Alumo' is
reported by Plukenet, though plants of A. glauca are known
in the Canary Islands by the names 'saladillo', 'salado' and
'marisma'.

20. It uph thai in urn Canariense Leucanthemum, Cotulae
foetidae crassioribus foliis, radice, acrisapore, &
fervido [. . .] Magala ab Insulanis nuncupatur Aim.:
73 (1696), Phyt.: t. 272, f. 6 (1694).

Syn.: Pyrethrum flore Bellidis (Bauhin, 1623).
His.: Chrysanthemum frutescens L. [Linn.; Gis.], Pyrethrum
frutescens (L.) Gaertn. [Ten.], Argyranthemum frutescens
(L.) Sch. Bip. [W.B.].

Det.: * Argyranthemum frutescens (L.) Sch. Bip. ssp. frute-
scens [there is a specimen in HS 95: 200 of A. frutescens which
bears a label which states 'Cotula e Canarinis Ins.' but the
material resembles the figure found in Phyt.: t. 272, f. 6.
Plukenet incorrectly cited Phyt.: t. 272, f. 5 for this species,
and it is Phyt.: t. 272, f. 6 which agrees with the description.
Subsequent taxonomic works from Linn., Gis., Ten. and
W.B: noted this error and established that the correct draw-
ing for this name is that in Phyt.: t. 272, f. 6].

Oth.: HVP: 28.

Com.: The use of the common name 'Magala' is indicated in

the description. This species is known locally as 'magarza'.

21. Campanula Canariensis Regia. s. Medium radice
tuberosa, foliis sinuatis, coefiis, Atriplicis aemulis,
ternis circa caulem ambientibus, flore amplo pendulo,
colore flammeo rutilante [. . .] inter Convolvulorum
species per incuriam posita, in Hort. Regio Hampton,
nunc viget, ubi mende Januario, flores editit Aim.:
76-77 (1696), Phyt.: t. 276, f. 1 (1694).

Syn.: Cachruiaqua (Hernandez, 1651), Totoncaxoxo coyollin

(Hernandez, 1651).

His.: Campanula canariensis L. [Linn.; Gis.; Ten.; W.B.].

Det.: * Canarina canariensis (L.) Vatke [HS 95: 133; 99: 161;

102: 160].

Oth.: HS 87: 79; 92: 38; 104: 59; HVP: 33.

Com.: Plukenet reported that plants of this species were

cultivated in the Royal Garden at Hampton Court Palace, in

south-west London.

Lectotypification of Campanula canariensis
Linnaeus

Linnaeus' protologue for this name (Fig. 1) in Species plan-
tarum (Linnaeus, 1753) comprises a new diagnostic phrase
name (C. foliis hastatis dentatis oppositis petiolatis, capsulis
quinquelocularibus) , the citation of two synonyms from Lin-
naeus (1738, also cited via van Royen, 1740) and Plukenet
(1694, 1696), and the statement 'Habitat in insulis Canariis'.
The diagnosis is a modification of that used in his earlier
Hortus cliffortianus where Linnaeus published an illustration
(see Fig. 2), but unfortunately no material exists in either the
Clifford or Linnaean (LINN, S, UPS, H, MW) herbaria.
Apart from Linnaeus' figure, the only other visual element in
the protologue is the Plukenet illustration (see Fig. 3).
Although it is perhaps a little more stylized than that
executed by Ehret for Clifford (for Hortus cliffortianus), it
shows the form of the corolla much more clearly, and has the
considerable advantage that there is a voucher specimen in
the Plukenet Herbarium upon which the illustration was
evidently based. Both illustrations undoubtedly belong to the
plant known as Canarina canariensis (L.) Vatke, so there are
no taxonomic complications associated with this choice.
Although the Clifford illustration would have been more
familiar to Linnaeus, the existence of the voucher material
makes us favour instead Plukenet's illustration and we for-
mally choose Campanula Canariensis regia s. Medium radice
tuberosa, foliis sinuatis,. . . Plukenet, Almagestum Bot. 76
(1696); Phytographia t. 276, f. 1 (1694) as the lectotype

* Cattle fubJivifo.

ctnarinft. if. CAA/IPANULA foliis haftatis dcntatis oppofitls pe-
tiolatis, capfulis quioquelocularibus.

Campanula foliis hnftatis dentatis, cr.ule determinate fo-
lioib. Hort. clijf. 6$. t. 8. Roy. lugdb. 147.

Campanula canartcnfis regia f. Medium radice tubcrofa,
foliis finuatis czfiis atriplicis xmulis ternis circum
ciulem arabientibus, flore amplo pendulo: colore
flammeo rutilante. Pink. aim. 76. 1. 176 /. i..

Hnbttat in ixfulis Canariis. Tf

Fig. 1 The original protologue of Campanula canariensis Linnaeus
(1753).

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

Fig. 2 Ehret's illustration of Campanula foliis hastatis dentatis, caule determinate foliose from Linnaeus' Hortus Cliffortianus (1738).

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

(typotype in Herb. Sloane 99: 161, BM) of Campanula
canariensis L.

22. Carduus Acanthoides, s. incanus minor, elegans, ex
Insula Maderensi Aim.: 85 (1696), Phyt.: t. 274, f. 1
(1694).

Det.: Galactites tomentosa Moench [HS 99: 170].

23. Cerasus Canariensis, mucronato Lauri angustiore
folio, fructu parvo Fragiformi, Ossiculo tuberculoso,
monopyreno Aim.: 95 (1696), Mant.: 43 (1700).

Syn.: Morococks (Ogilvy, 1671).

Det.: Myricafaya Alton [?].

Com.: Description virtually identical to that found in entry

17.

24. Cicutae fatuae nostrati similis, Planta Maderensis

Aim.: 104(1696).

25. Cistus latifolia major trinervis incano folio, floribus
purpureis ex Insula Pico Mant.: 49 (1700).

Syn.: Ledon (Belon, 1553), Ledon tertium Cyprium (Clusius,
1601a), Cistus Ledon cretense (Bauhin, 1623), Cistus Ledon
latifolium Creticum (Bauhin & Cherler, 1651), Cistus Ledon
latifolium Creticum Triumfet (Cupani, 1696).

26. Convolvulus Canariensis, longioribus foliis
mollibus, & incanis. Ahilo-porro Salvages Hispanis
nuncupatur Aim.: 114 (1696), Mant.: 54 (1700), Phyt.:
t. 325, f. 1 (1694).

Syn.: Convolvulus marinus Catharticus foliis Acetosae flore
niveo (Plumier, 1693).

His.: Convolvulus canariensis L. [Linn.; Gis.; Ten.; W.B.].
Det.: * Convolvulus canariensis L. [HS 93: 110].
Com.: The common name 'Ahilo-porro Salvages' is men-
tioned in the description. However Convolvulus species bear
the common names 'corregiiela', 'chaparro', 'guaydil', 'lena
noel' whereas the name 'ajo porro' is used for plants of
Allium spp.

27. Convolvulus Canariensis minimus, flore
ochroleuco, semine nigro. Hartelauena Indigenis dictus

Aim.: 400 (1696), Phyt.: t. 324, f. 4 (1694).

His.: Sarothra gentianoides L. [Gis.].

Det.: * Convolvulus fruticulosus Desr. [there is a specimen
without a label of this Canarian endemic in HS 93: 110. It
resembles the drawing published by Plukenet and a tentative
determination has been made based on it].
Com.: Plukenet mentioned the use of the common name
'Hartelauena' (perhaps a derivation of 'corregiiela'). For
common names of Convolvulus species see entry no. 26. It is
worth mentioning that according to the description the spe-
cies bears yellow flowers, however C. fruticulosus does not
bear flowers with this colour.

28. Cupressus nana, Canariensis fructu minore. In
Palma quoque Insula una ex Fortunatis oritur haec
arbos Aim.: 125 (1696), Mant.: 61 (1700).

Syn.: Arbor cujus fructus Abhel (Clusius, 1576), Habhel
(Clusius, 1601a, Cupresso similis Arbor in Syria (Bauhin,
1623), Uyt Persien kont daar de Com Taxa (Rauwolf, 1707).
Det.: Juniperus cf. phoenicea L. [?]

Com.: Determination based on the description. It is notewor-
thy that a specimen of this species is also found in HS 189: 32
in J. Cuningham's collection from La Palma. We were unable
to find an earlier edition of Rauwolf s work.

29. Cytisus arboreus, Canariensis, oblongo folio,
argentea & holosericea lanugine subtus villoso, flore
pallidiori [. . .] Texo Insulanis nuncupatur Aim.: 128
(1696), Phyt.: t. 277, f. 4 (1694).

Syn.: Cytisus albus sylvestris (Cordus, 1561), Cytisus Alpinus

(Dalechamps, 1586), Cytisus albicans, folio Trifolii vulgaris

(Bauhin, 1623).

His.: Cytisus proliferus L. f. [W.B.].

Det.: * Chamaecytisus proliferus (L. f.) Link ssp. proliferus

var. proliferus [HS 96: 2].

Com.: Plants of this species are known as 'escobon',

Plukenet, however, reporting the common name Texo'. The

name 'tejo' is used for plants of Erica scoparia L. ssp.

platycodon (Webb & Berth.) Hansen & G. Kunkel.

30. Cytisus Canariensis, microphyllos, angustifolius,
prorsus incanis [. . .] Esta Insulanis nuncupatur Aim.:
128 (1696), Phyt.: t. 277, f. 5 (1694).

Syn.: Cytisus minor ibus foliis, ramulis tenellis villosis (Bau-
hin, 1623), Cytisus Hispanicus, primus Clusii folio virescente
(Bauhin & Cherler, 1650).

His.: Teline canariensis (L.) Webb & Berth. [W.B.].
Det.: * Teline canariensis (L.) Webb & Berth. [HS 96: 2.
Plukenet did not give a figure number for this species in his
Almagestum, but there is a herbarium specimen of T. canar-
iensis which bears Plukenet's handwriting and indicating that
it relates to Phyt.: t. 277, f. 5].
Oth.: HS 87: 117, 93: 132, 100: 14.

Com.: T. canariensis is known in the Canary Islands as
'retamon', 'retama de cumbre' or 'gildana'. However
Plukenet reported the common name of 'Esta'.

31. Cytisus Canariensis, microphyllos, cauliculis
villosis angustis viridibus foliis [. . .] Mysalva
Insulanis, dicta Aim.: 128 (1696), Mant.: 63 (1700),
Phyt.: t. 277, f. 6(1694).

Syn.: Cytisus Hispanicus siliquis Ornithopodij (Bauhin,
1620), Cytisus Montis Calcaris (Bauhin & Cherler, 1650).
His.: Genista canariensis L. [Linn.; Gis.; Ten.], Adenocarpus
foliolosus (Aiton) DC. [W.B.].

Det.: * Adenocarpus foliolosus (Aiton) DC. [HS 96: 2.
Plukenet did not give a figure number for this species in his
Almagestum, but there is a herbarium specimen of A. foliolo-
sus which bears Plukenet's handwriting indicating that it
corresponds to fig. 6, and therefore the identifications pro-
vided by Linnaeus, Tenzel and Giseke were incorrect. Webb
& Berthelot (1842) realised this error in their account of
Genista canariensis L. and assigned Phyt.: t. 277, f. 5 for G.

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

Fig. 3 Plukenet's illustration of Campanula Canariensis regia s. Medium radice tuberosa, foliis sinuatis, from his Phytographia, t. 276, f. 1
(1694), the lectotype of Campanula canariensis L., Sp. PI. 1: 168 (1753), designated here.

10

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

Fig. 4 Plukenet's illustration of Digitall affinis Canariensis Solidaginis acutis foliis leviter pilosis,
from his Phytographia, t. 325, f. 2 (1694).

. . (= Isoplexis canariensis (L.) Loud.)

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

11

Fig. 5 Voucher material of Digitali affinis Canariensis Solidaginis acutis foliis leviter pilosis,
Plukenet's herbarium (now part of Herb. Sloane, vol. 100: 18 (BM)).

. . (= Isoplexis canariensis (L.) Loud.) in

12

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

canariensis (see entry 30) and Phyt.: t. 277, f. 6 for A.
foliolosus. This has been confirmed in our research after the
study of the labels found with Plukenet's specimens].
Oth.: HS 100: 14, HVP: 53.

Com.: The common name 'Mysalva' is used by Plukenet, but
Adenocarpus species are known as 'codeso'.

32. Digital! at finis Canariensis Solidaginis acutis foliis
leviter pilosis, flore aureo cucullato, staminibus croceis
cristae cavo accumbentibus ornato [. . .] Matera
Insulanis vulgo, Hujus folia sunt impense amara Aim.:
400 (1696), Phyt.: t. 325, f. 2 (1694); see Fig. 4.

Syn.: Adel-odagam (Rheede tot Draakenstein, 1689), Bahel-

tsjulli (Rheede tot Draakenstein, 1689).

His.: Digitalis canariensis L. [Linn.; Gis.; Ten.], Callianassa

canariensis (L.) Webb & Berth. [W.B.].

Det.: * Isoplexis canariensis (L.) Loudon [HS 100: 18]; see

Fig. 5.

Oth.:HS87: 118.

Com.: Plukenet reported the use of the common name

'Matera', however plants of this species are commonly called

'cresta de gallo' or 'dedalera'. The species described in entry

33 also seems to refer to this name.

33. Digitalis lutea, flore magno, Canariensis Aim.: 131
(1696).

Syn.: Digitalis lutea magno flore (Bauhin, 1623), Digitalis
lutea flore maiore folio latiore (Bauhin & Cherler, 1651).
Det.: * Isoplexis canariensis (L.) Loudon [?]
Com.: It seems that Plukenet described this species twice (see
entry 32).

34. Ecbolii Indici, s. Adhatodae cucullatis floribus
aemula, Hyssopifolia, Planta ex Insulis Fortunatis

Aim.: 132 (1696), Phyt.: t. 280, f. 1 (1694).

Syn.: Tsjanga-puspam (Rheede tot Draakenstein, 1689).

His.: Justicia hyssopifolia L. [Linn.; Gis.], Gendarussa hys-

sopifolia (L.) Webb & Berth. [W.B.].

Det.: * Justicia hyssopifolia L. [HS 96: 13, 100: 20].

Oth.:HS87: 122.

Com.: It seems that Plukenet described this species twice (see

entry 16).

35. Echium album Maderense, & Echium Tingitanum
procerius, floribus immaculatis Aim.: 133 (1696),
Phyt.:t. 278, f. 5(1694).

Det.: * Echium cf. leucophaeum Sprague & Hutch. [HS 100:

22].

Com.: Although the description is given for a plant from

Madeira, the specimen found in HS 100: 22 is a Canarian

endemic.

36. Ficoides Africanum Mesembrianthemum, s. Ficus
Aizoides teretifoliis, fuccesis, micis argenteis
interspersis, flore carneo ex Insulis Fortunatis. Cosca
Insulanis vulgo Aim.: 149 (1696).

Det.: Mesembryanthemum nodiflorum L. [determination
based on the Spanish common name reported by Plukenet. It
is noteworthy that in HS 96: 123 there is a herbarium

specimen without labels of this species].
Com.: Plants of this species are known as 'cosco' in the
Canary Islands, a name extremely similar to that reported by
Plukenet - 'Cosca'.

37. Ficus sylvestris [. . .] Hujus ramuli ex Insulis
Fortunatis allati Aim.: 144 (1696), Phyt.: t. 281, f. 1
(1694).

Syn.: De Caprifichi (Anguillara, 1561), Caprificus (Cordus,
1561), Ficus sylvestris (Bauhin, 1623), Caprificus voratur e
sylvestri genere ficus nunquam maturescens (Pliny, 1826b),
Ficus sylvestris Dioscoridis [?].
Det.: Ficus carica L. [HS 100: 43].

38. Filicula crispa lanugine hepataci coloris vestita, ex
Insulis Fortunatis Aim.: 150 (1696), Mant.: 77 (1700),
Phyt.: t. 281, f. 4(1694).

Syn.: Filix minor russa lanugine tola obducta inpinnas tantum

divisa raras non crenatas subrotundas (Sloane, 1696).

His.: Acrostichum marantae L. [Linn.; Gis.], A. velleum

Willd. [Ten.].

Det.: Cheilanthes catanensis (Cos.) H.P. Fuchs [HS 100: 52].

39. Filix Hemionitis dicta, Maderensis Hederae
arboreae aliquatenus aemula, s. foliorum basi auriculis
binis, utrinque donato Aim.: 155 (1696), Mant.: 82
(1700), Phyt.: t. 287, f. 4 (1694).

Syn.: Hemionitis peregrina (Clusius, 16016), Hemionitis per-
egrina Clusii & aliorum (Ray, 1686), Hemionitis peregrina
foliorum segmentis sinuatis longioribus & magis acuminatis
seu hederae folio anguloso (Sloane, 1696).
His.: Asplenium palmatum Lam. [Ten.].
Det.: Asplenium hemionitis L. [HS 100: 51. In HS 96: 44
there is a label without a specimen which states: 'Hemionitis
Maderensis hederaefolio base auriculis binis [illeg.] donate';
it seems that it refers to this description].

40. Filix Hemionitis dicta Maderensis pediculis
splendentibus nigris, crenatis foliis Asari rotundioribus
cr en aru in segmentis oblongo quadratis, ob semina
adnascentia per ambitum circumcirca reflexis Aim.:
155, 400 (1696), Mant.: 82 (1700), Phyt.: t. 287, f. 5
(1694); see Fig. 6.

His.: Adiantum reniforme L. [Linn.; Gis.; Ten.; W.B.].
Det.: Adiantum reniforme L. [HS 100: 51; see Fig. 7. In HS
96: 44 there is a label without a specimen which states:
'Hemionitis Maderensis pediculis nigris folys Asari rotun-
dioribus' which seems to refer to this species].

41. Filix ramosa Canariensis Rutae murariae pinnulis
angustis, altius incisis, mediae costae alternatim
alligatis [. . .] Hujus folium totale, (circumscriptione)
est sere triangulatum Aim.: 156 (1696), Phyt.: t. 291,
f. 2 (1694).

His.: Trichomanes canariense L. [Linn., Gis., Ten.].

Det.: Davallia canariensis (L.) J. E. Sm., [HS 93: 169, 100:

61].

Oth.:HS87: 143,95: 19.

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

13

Fig. 6 Plukenet's illustration of Filix Hemionitis dicta Maderensis pediculis splendentibus nigris, . . . (= Adiantum reniforme L.) from his
Phytographia, t. 287, f. 5 (1694).

14

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

. / ^4-

Fig. 7 Voucher material of Filix Hemionltis dicta Maderensis pedlculis splendentibus nigris, . . . (= Adiantum reniforme L.) in Plukenet's
herbarium (now part of Herb. Sloane, vol. 100: 51 (BM)).

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

15

Fig. 8 Plukenet's illustration of Horminum hastatis amplioribus foliis, s. Ari modo alatis, . .' . (= Salvia canariensis L.) from his
Phytographia, t. 301, f. 2 (1694).

16

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

jyjyjjjy^yyyiyilllllUlI^

Fig. 9 Voucher material of Hormlnum hastatis amplioribus foliis, s. Ari modo alatis, . . . (= Salvia zanariensis L.) in Plukenet's herbarium
(now part of Herb. Sloane, vol. 100: 138 (BM).

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

17

42. Filix Saxatilis pervenusta, foliis punctatis, s.
Adianthum Maderense folio Filicis, caule tenui
candicante Aim.: 150 (1696), Phyt.: t. 284, f. 4 (1694).

Syn.: Filix pulverulenta pinnulis obtuse dentatis (Plumier,

1693).

Det.: Cystopteris cf. fragilis (L.) Bernh. [HS 96: 40].

43. Foeniculum dulce Azoricum; a com muni dulci
differ! radiorum umbellae magnitudine, & longitudine,
umbella concava, seminibus grandioribus Aim.: 157
(1696).

44. Geranium Cretico simile minus incanum acu
pusilla, ex Insulis Fortunatis Aim.: 169 (1696).

45. Geranium Hispanicum magna radice, Cicutae folio
crassiori, [. . .] Insulis etiam Fortunatis reperitur, in
quibus collect urn suit, & nobis allatum Aim.: 169-170
(1696).

Syn.: Geranium Cicutae folio aculongissima (Bauhin, 1620,
1623), Geranium Hispanicum magna radice Cicutae folio
crassiori (Sherard, 1689).

46. Geranium saxatile procumbens, foliis subtus
canescentibus, flore majore candicante, ex Insula Pico.
Cura industrij Hortulani Johannis Adams in Hort.
Comptoniano, ex seminibus enutritam conspeximus

Mant.: 89-90 (1700).

Syn.: Tlatlauh capatli Oxygeranio Mex. (Hernandez, 1651).
Com.: It seems that Plukenet based his description on plants
from John Adams, Gardener to the Duke of Beaufort,
presumably obtained from Henry Compton's Garden at
Fulham Palace in London (Rohde, 1932).

47. Gramen Canarium Ischaemi paniculis Aim.: 175
(1696).

Syn.: Gramen dactylon folio arundinaceo majus: aculeatum
sorte Plinij (Bauhin, 1623; Ray, 1688a), Gramen Canarium
Ischaemi paniculis (Parkinson, 1640), Gramen repens cum
panicula Graminis Mannae (Bauhin & Cherler, 1651), Gra-
men legitimum (Magnol, 1676).

48. Helleborine similis Canariensis. Flos Spiritus
Sancti, vulgo Aim.: 183 (1696).

Syn.: Flos Spiritus Sancti (Hermann, 1689).
Com.: The common name 'Flos Spiritus Sancti' is given.
Plukenet referred to a Helleborine similar to that from the
Canary Islands, though as there is no mention of a Hellebo-
rine Canariensis in any of his works (but only to a 'Hellebo-
rine Canadensis'), it is likely that this citation was a
typographic error.

49. Horminum hastatis amplioribus foliis, s. Ari modo
alatis, caulibus & pediculis araneosa lanugine villosis,
ex Insula Gomera quae una est ex Fortunatis [. . .]
Mustazi Insulanis & Salvia arborea vulgo Aim.: 185
(1696), Mant.: 103 (1700), Phyt.: t. 301, f. 2 (1694);
see Fig. 8.

Syn.: Sclarea Lusitanica glutinosa amplissimo folio (Tourne-

fort, 1694).

His.: Salvia africana L. [Linnaeus (1762); Gis.], 5. canariensis

L. [Ten.; W.B.].

Det.: * Salvia canariensis L. [HS 100: 138; see Fig. 9].

Oth.: HS 100: 99, 102: 47.

Com.: Linnaeus (1762) cited Plukenet's reference in the

synonymy of 5. africana L. He also identified the drawing

from Phyt.: t. 301, f. 2 as S. canariensis in his annotated copy

of Plukenet's Phytographia. The common names 'Mustazi'

and 'Salvia arborea' are mentioned by Plukenet for the

Canary Islands. However this species is known as 'garitope',

'salvia morisca' or 'salvia'.

50. Horminum latifolium Canariense pilosum, foliis
alterno situ pofita non repugnarent, eandem hane esse
plantam suspicarer Aim.: 185 (1696), Mant.: 103
(1700), Phyt. :t. 301, f. 3(1694).

Syn.: Marum Aegyptiorum (Alpino, 1627).
Det.: * Convolvulus canariensis L. [HS 100: 138].

51. Hypericum, s. Androsaemum magnum Canariense
ramosum, copiosis floribus fruticosum Aim.: 189
(1696), Phyt.: t. 302, f. 1 (1694).

His.: Hypericum canariense L. [Linn.; Gis.], H. floribun dum

Aiton [Ten.], Webbia floribunda (Aiton) Webb & Berth.

[W.B.].

Det.: * Hypericum canariense L. [HS 96: 104].

52. Jasminum album trifoliatum flore magno ex Insula
Maderensi Aim.: 195 (1696), Phyt.: t. 303, f. 1 (1694).

Syn.: Jasminum sorte Azoricum (Grisley, 1661).

His.: Jasminum azoricum L. [Linnaeus (1762); Gis.; Ten.].

Det.: * Jasminum azoricum L. [HS 100: 149].

Com.: A description for this species was also provided by

Plukenet in entry 54.

53. Jasminum Canariense foliis amplioribus laeto
virentibus venosis, hirsutis & asperis, umbellatis
Schetti floribus purpureis, ex caliculis inflatis,
prorumpentibus. Ex Insulis Fortunatis nobis advecta.
At ex India Orientali; ad D. Petiver allata est Aim.:
196 (1696).

Det.: * Viburnum tinus L. ssp. rigidum (Vent.) P. Silva. [our
determination is based on a specimen found in HS 100: 150
whose label states 'From Ins. Canaries, Jasminum'. The
description provided by Plukenet agrees with most of the
morphological features of this specimen] .

18

54. Jasmin u in Azoricum trifoliatum lucidum,
acuminatis foliis, flore albo odoratissimo, flexilibus &
obsequiosis Virgulis. A Jasmino flavo odoratissimo
maxime diversum Amalth.: 123, t. 423, f. 6 (1705).

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

refers to Phyt.: t. 199, f. 3, but it cannot be identified with any
Macaronesian species.

Det.: * Jasminum azoricum L. [HS 93: 210].

Com.: The description found in entry 52 also refers to this

species.

55. Kali aizoides Canariense procumbens, Portulacae
pallescentibus succulentis foliis, aspergine rorida
perpetuo in ad id is [. . .] Poita ( amilo Insulanis dictum;
Haec species est ad caulium nodos prolifera cum
floribus sere inconspicuis Aim.: 202 (1696), Phyt.: t.
303, f. 4 (1694).

His.: Aizoon canariense L. [Linn.; Gis.; Ten.; W.B.].
Det.: Aizoon canariense L. [there is a herbarium specimen in
HS 100: 42 which is similar to the figure given by Plukenet
(Phyt.: t. 303, f. 4), it has a label which follows the descrip-
tion (Aim.: 202) but states that the specimen is from 'Ind.
Or.'. Furthermore in HS 100: 92 there is a label without a
specimen that states 'Kali aizoides procumbens lichenides
folio ex Insulis Fortunatis'; it is likely that this label belongs
to the specimen found in HS 100: 42].

Com.: The use of the common name 'Poita Camilo' (it may
be a derivation of 'pata (de) camello') is indicated by
Plukenet. Actual common names for this species are 'pata
perro' or 'patilla'.

56. Laurifolia Canarina diphyllos [. . .] Sorte IN i mho
Javanensium prima Aim.: 211 (1696), Phyt.: t. 305, f.
1 (1694).

Syn.: Lauro Indica (Bontius, 1658).

Com.: There is a herbarium specimen in HS 96: 143 associ-
ated with Plukenet's figure, but the specimen does not belong
to any species of the Macaronesian flora.

57. Laurotaxa epiphyllocarpos, crenatis foliis, maxima,
e singulis foliorum crenis, baccifera. Ex Insula Palma;
nuper allata est Mant.: 114 (1700).

His.: Ruscus androgynus L. [Linnaeus published this name in
1753. However, he referred to Plukenet's polynomial only in
his earlier Hortus diffortianus (Linnaeus, 1738)], Danae
androgyna (L.) Webb & Berth. [W.B.].
Det.: * Semele androgyna (L.) Kunth [?].
Com.: Specimens of this species have not been found in
Plukenet's herbarium; this tentative determination is based
on the original description and on information provided by
Linnaeus (1738) and Webb & Berthelot (1847). Also, J.
Cuningham collected plants of this species during his stay in
La Palma in the late seventeenth century, one of which is
found in HS 189: 25.

58. Laurus Azorica pallidioribus & latioribus foliis,
inodora Aim.: 210 (1696), Phyt.: t. 199, f. 3 (1693).

Syn.: Laurus [. . .] Pompeius Lenaeus adjecit quam mustacem
appellavit, quoniam mustaceis subjiceretur. Hanc esse folio
maxima, flaccido que et albicante (Pliny, 1826c).
Com.: The specimen found in HS 96: 139 has a label which

59. Laurus Indica Aim.: 210 (1696), Mant.: 115
(1700), Phyt. :t. 304, f. 1(1694).

Syn.: Laurus Indica (Aldino, 1625), Laurus Indica. Indica

sive Americana Laurus (Ferrarius, 1633), Laurus Americana

cujus cortex Cassia ligneae multum assimilatur (Parkinson,

1640), Quauh eloxochitl (Hernandez, 1651), Laurus Regia.

Laurier Royal (Vallot, 1665), Laurus Americana sive Persea

Clusij (Schuyl, 1672), Laurus [. . .] Accessit et regia, quae

coepit Augusta appellari, amplissima et arbore et folio (Pliny,

1826c).

His.: Laurus indica L. [Linn.; Gis.; Ten.], Persea indica (L.)

Spreng. [W.B.].

Det.: * Persea indica (L.) Spreng. [HS 100: 167].

60. Laurus Maderensis angustifolia pa 1 1 id a
odoratissima, venis foliorum aversa parte magis
extantibus Aim.: 210 (1696), Phyt.: t. 199, f. 2 (1693).

Com.: The herbarium specimen which corresponds with this
figure is found in HS 96: 138 but it does not belong to any
Macaronesian species.

61. Laurus Regia odoratissima Maderensis Aim.: 210
(1696).

Syn.: Laurus latifolia Azorica, Cinamomi odore (Commelin,
1689), Laurus Regia odoratissima Maderensis (Hermann,
1689).

62. Lavandula maritima Canariensis spica multiplici
caerulea Aim.: 209 (1696), Phyt.: t. 303, f. 5 (1694).

His.: Lavandula multifida L. [Gis.], L. abrotanoides Lam.

[Ten.].

Det.: * Lavandula buchii Webb [HS 96: 134, 96: 135].

63. Limonium parvum Hell id is minoris folio Aim.: 221
(1696).

Syn.: Limonium pumilum (Clusius, 1601ft), Limonium par-
vum bellidis minoris folio (Bauhin, 1623), Limonium minus
bellidis minoris folio. Petit Limonium a feuilles de Marguerite
(Dodart, 1676).

His.: Statice limonium L. [Linn.].

Det.: * Limonium pectinatum (Aiton) Kuntze [HS 96: 149.
The original description does not mention any of the Macaro-
nesian Islands. However the specimen has a label (i.e.
'Limonium parvum Bellidis minoris folio CBP Limon. pumi-
lum Clus. ex Insula Canarina') which follows the phrase-
name from Aim.: 221 and also refers to the Canary Islands].

64. Linariae similis, Arbuscula Canariensis, latiore
folio viridi Amalth.: 133 (1705).

Det.: * Kleinia neriifolia Haw. [?].

Com.: This description is similar to that of entry 65, and we

believe that it refers to vigorous plants with broader leaves.

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

65. Linariae si mi I is Arbuscula Canariensis, folio
longiore carnoso fragili, fubtus purpurascente
Crithimmi recipiens Aim.: 223 (1696), Mant.: 118
(1700), Phyt.: t. 304, f. 3 (1694).

Syn.: Plantae Lav endulae folio (Clusius, 1605), Frutex Indiae
Orientalis Lavendulae folio (Bauhin, 1623), Arbor Laven-
dulae folio (Bauhin & Cherler, 1650), Texioquahoitl (Her-
nandez, 1651).

His.: Cacalia kleinia L. [Linnaeus published this name in
1753. However he quoted Plukenet's polynomial only in his
earlier Hortus cliffortianus (Linnaeus, 1738)].
Det.: * Kleinia neriifolia Haw. [this determination is based on
Plukenet's drawing and description. However there is also a
herbarium specimen in HS 102: 86 without a label which
belongs to this species].

66. Lupinus major villosis foliis Maderensis flore albo
& purpureo [. . .] nostra tamen folia, baud proprie
angusta did merentur Aim.: 229 (1696), Mant.: 120
(1700).

Syn.: Lupini albi in quorum genere reperiuntur maximi, flore
antequam aperiatur, subcaeruleo intus vero albo (Camerarius,
1588), Lupinus sativus major, & quartus Clusii, flore e
coeruleo purpurascente. Lupinus Aethiopicus quorundam
(Besler, 1613), Lupinus Indicus medius caerulens (Parkinson,
1640), Lupinus caeruleus minor per ennis Virginianus repens,
nobis (Morison, 1680), Lupinus medius caeruleus (Ray,
1686), Lupinus incarnatus (Hermann, 1687), Lupinus angus-
tifolius, flore e candido purpureo (Cupani, 1696).
Det.: Lupinus albus L. [?].

67. Melissa Canarina multifido folio spicata, odorem
(amphorae spirans, penetrantissimum Mant.: 128, t.
430, f. 2. (1705); see Fig. 10.

Syn.: Katu-kurka (Rheede tot Draakenstein, 1690), Moldav-

icae Species trifolia, ex America, nuper ad nos missa Turne-

fort. de opt. Meth. instituend in re Herber epist. ad Sherard.

pag-18[?].

His.: Chrysanthemum indicum L. [Ten.].

Det.: * Cedronella canariensis (L.) Webb & Berth. [HS 101:

13, upper specimen].

Com.: It appears that this species was described twice by

Plukenet (see entry 69).

68. Melissa (forte) an Mentha viridis. Haec autem
Melissa non est, sed re vera ad Mentharum genera
accenseri meretur, & nominari licet Mentha pi In I if era
Betonicae forma, & odore aromatico, ex Insulis
Fortunatis. Polihomons Insulanis dicta Aim.: 247
(1696), Mant.: 127 (1700), Phyt.: t. 307, f. 1 (1694).

Syn.: Yxiayaboal Chapaltepecensis (Ray, 16886), Melissa
elatior foliis magnis dentatis glabris ad genicula binis, /lores
odoratos luteos patulos stamina bina quasi cornua protru-
dentes, in summitate caulium ramatim serens (Banister, 1693).
Det.: * Bystropogon canariensis (L.) L'Her. [there is an
unlabelled specimen corresponding with this species in HS
101: 15 which resembles the drawing depicted by Plukenet in
Phyt.: t. 307, f. 1; a tentative determination is based on this
specimen].

19

Oth.:HS88:73;96: 187.

Com.: Bystropogon species are known in the Canaries as
'poleo de monte' o 'poleo monte', this agrees with Plukenet's
description as he reported on the use of the word 'Poli-
homons'. Two different descriptions which seem referable to
this species have been found in Plukenet's works (see also
entry 70).

69. Melissa forte Canarina triphyllos odorem
Camphorae spirans penetrantissimum Aim.: 401
(1696), Phyt.: t. 325, f. 5 (1694).

Syn.: Aztaxochitl (Hernandez, 1651).

His.: Dracocephalum canariense L. [Linn.; Gis.], Cedronella

canariensis (L.) Webb & Berth. [W.B.].

Det.: * Cedronella canariensis (L.) Webb & Berth. [HS 101:

13, lower specimen].

Oth.: HS 88: 71; 94: 14; HVP: 120.

Com.: The description found in entry 67 also refers to this

species.

70. Melissophyllum citratum ex Insulis Fortunatis,
Lingo-veha ab Insulanis dictum Aim.: 247 (1696),
Phyt. : t. 306, f. 4 (1694).

Det.: * Bystropogon cf. canariensis (L.) L'Her. [determina-
tion based on the drawing provided by Plukenet; however,
there is also an unlabelled specimen in HS 101: 14 which
resembles this figure].

Com. : The use of the common name 'Lingo-veha' is indicated
for this entry. However plants of this species are known as
'poleo de monte' or 'poleo monte'. Another description
which seems to refer to this species can be found in entry 68.

71. Mentha Maderensis I us u lac. minus odorata Aim.:
248 (1696).

72. Mentha sylvestris Azorica longioribus foliis incanis,
spica longiore & crassiore Aim.: 248 (1696).

73. Mentha Canariensis frutescens, foliis subtus
lanugine candidissima, villosis floribus glomeratis e
sinu foliorum longioribus pediculis insidentibus Aim.:
248 (1696), Phyt.: t. 307, f. 2 (1694).

His.: Mentha canariensis L. [Linn.; Gis.], Bystropogon canar-
iensis (L.) L'Her. [Ten.; W.B.].

Det.: * Bystropogon plumosus (L. f.) L'Her. [HS 96: 186].
Oth.: HS 101: 18 [although this specimen is badly preserved,
it resembles the drawing given by Plukenet in Phyt.: t. 307, f.

2].

Com.: The phrase-name found in entry 74 also seems to refer
to this species.

74. Mentha Canariensis, minore folio subtus incano,
ramosissimus Aim.: 248 (1696).

Det.: * Bystropogon plumosus (L. f.) L'Her. [HS 96: 186.
The specimen is not explicitly linked with Aim.: 248 but it
does have a label which states 'Mentastrum incanum canar-
iensis minore folio ramosiss' which follows this entry].
Com.: It seems that Plukenet provided another description
for this species in entry 73.

20

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

JP/

u

Fig. 10 Plukenet's illustration of Melissa Canarina multifido folio spicata. odorem Camphor ae spirans, penetratissimum (= Cedronella
canarlensis (L.) Webb & Berth.) from his Phytographia, t. 430, f. 2 (1705).

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

75. Muscus arboreus aurantiacus, staminibus
tenuissimis, ex Insulis Fortunatis [. . .] Totus citrino
colore nitet peramaeno Aim.: 254 (1696), Mant.: 132
(1700), Phyt. :t. 309, f. 1(1694).

Syn.: Laricus Muscus (Bauhin & Cherler, 1650). Plukenet
claimed that the name Muscus arboreus rutilans was given by
Bauhin (1623); there are five polynomials commencing 'Mus-
cus arboreus' in Bauhin's work but none of them appears in
exactly this form.

Del.: Lethariella canarlensis (Ach.) Krog [HS 97: 4, 101: 31].
Oth.:HS92:94.

76. Muscus cinereus, e i amis Arbor um dependens,
Canariensis, ex staminibus crassioribus geniculatis, in
tenuissima & longissima Ilia ramulosus. [. . .] Tanta
copia inveniuntur hi musci apud Virginienses iis
praesertim locis, ubi in Maris accessu aquis abundant,
ut Hiberno tempore Totas arbores quibus adnascuntur
dejiciant, eo quod pabulum Vaccis & ovibus suis
praebent gratissimum Aim.: 254 (1696).

Syn.: Muscus capillaceus longissimus (Bauhin, 1623).
Plukenet claimed that Clusius (1601) used the name Muscus
capillaceus cineri colons, e ramis Ilicis dependens. However,
we have been unable to trace this polynomial in either part of
this publication.

Del.: Usnea articulata (L.) Hoffm. [determination based on
Plukenet's description and on an unlabelled specimen found
in HS 101: 31 which follows this description].

77. Origanum Maderense nostrati simile odoratius
capitulis albicantibus. Gratissimo odore nares
perstringit Aim.: 272 (1696), Mant.: 141 (1700).

Det.: Origanum vulgare L. [?].

78. Palma prunifera foliis Juccae, fructu racemoso
Cerasi-formi, ossiculo duro cinereo, Pisi magnitudine,
Lachrymam Sanguis Draconis dictam fundens Aim.:

277 (1696).

Syn.: Dracone arbore (Clusius, 1576), Draco (Clusius, 1601a,
Draco arbor (Bauhin, 1623; Parkinson, 1640), Draco arbor et
de eius Lacryma, seu sanguine e Draconis cinnabati Veterum
(Bauhin & Cherler, 1650), Palma prunifera foliis Yuccae,
fructu, in Racemis congestis, ceraciformi, duro cinereo pisi
magnitudine: hujus lacryma, Sanguis draconis dicta, Draco
Arbor, Clusi. Belg. Draken-boom (Commelin, 1689), Palma
prunifera foliis Juccae fructu racemoso cerasiformi ossiculo
duro cinereao pisi magnitudine Lachryman sanguis Draconis
dictam sundem (Kiggelaer, 1690).

Det.: * Dracaena draco (L.) L. [a herbarium specimen found
in HS 91: 31 has a label agreeing with Plukenet's name].

79. Papaver corniculatum ru bruin minus ex Insulis
Fortunatis. Mahopola Insulanis dictum Aim.: 279
(1696).

Det.: Papaver rhoeas L. [?].

Com.: The common name 'Mahopola' is reported by
Plukenet. Plants of Papaver spp. are called 'amapola' in
mainland Spain and the Canaries.

21

80. Peucedani folio Plant a Maderensis Aim.: 289
(1696).

Syn.: Planta fruticescens Africana perrara, foliis Peucedani,
floribus conglomeratis Herbae castae Americanae nonnihil
similibus (Breyne, 1678b).

81. Phaseolus teneri folius, fructu coccineo. ex Insulis
Fortunatis a D.D. Uvedal accepimus Aim.: 291 (1696).

Det.: Phaseolus vulgaris L. [?]

82. Polygonum Juncoide Scoparium ex Insulis
Fortunatis Aim.: 303 (1696), Phyt.: t. 311, f. 3 (1694).

Syn.: Alsine alpina junceo folio (Bauhin, 1620, 1623).
Det.: * Plocama pendula Aiton [HS 97: 117].

83. Polygonum ex Insula Palma, Serpylli foliis ad
genicula confertis, argentea coma paleacea Mant.: 154
(1700).

Det.: Polygonum maritimum L. [?]

Com.: Determination based on the description and on the
fact that J. Cuningham collected plants of this species during
his stay in La Palma in the late seventeenth century. A
specimen is in HS 189: 12 in Cuningham's collection from this
island. Plukenet also refers to this species in a phrase name
found in Aim.: 122 (illustrated in Phyt.: t. 277, f. 3); his
description is 'Cruciatae marinae similis, Planta Indiae orien-
talis [. . .] Huic valde similis es Insula Palma nuper est allata
plantula, cui nomem dedi Polygonum Serpylli foliis, ex
adverso binis, sericea coma candicante'. There is no her-
barium specimen for this name.

84. Ranunculus Canariensis hirsutus grumosa radice
Platani sere foliis, flore pallide luteo majore Aim.: 313
(1696).

Syn.: Ranunculus creticus latifolius (Clusius, 1601a), Ranun-
culus asphodeli radice Creticus and also Ranunculus lanugin.
apii folio asphodeli radice (Bauhin, 1623), Ranunculus alter
saxatilis Asphodeli radice (Parkinson, 1640).
Det.: * Ranunculus cortusifolius Willd. [label for the speci-
men of R. cortusifolius is found in HS 97: 144 and quotes the
description given by Plukenet in Aim.: 313].

85. Rubia arborescens asperior Insula rum
Canariensium, foliis ad singula genicula ternis [. . .]
Nettle-Tree a Mercatoribus dicta Aim.: 323 (1696),
Phyt.: t. 311, f. 4 (1694); see Fig. 11.

Det.: * Rubia fruticosa Aiton ssp. fruticosa [HS 97: 171; see
Fig. 12].

86. Saamounae Brasiliensium floribus aemula, Arbor
Sal vitolia sericea, dipetaloides, ex Insulis Fortunatis

Aim.: 326 (1696), Mant.: 164 (1700), Phyt.: t. 313, f. 2
(1694).

Syn.: Teucroides filiculosum foliis laurinis, floribus, galeatis &

labiatis (Sloane, 1696).

Det.: * Teucrium heterophyllum L'Her. [HS 98: 1].

22

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

Fig. 11 Plukenet's illustration of Rubia arborescens asperior Insularum Canariensium, . . . (= Rubia fruticosa Aiton ssp. fruticosa) from his
Phytographia, t. 311, f. 4 (1694).

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

23

/*

wrx

+9 4***

X* 1 / '

/ A?,^ f$**^ffy i> *S?**tr*-*~r

*\ . ' ^ ^ / j?

fr * M j^v

1 4 n &v 4 ***~JK,
' / SY

f/ J? /S '

*"* MT^tfe

eX. fctfy.

>r <^^A^ ' y

nBiiiimiiHiiBmiiiiiiiiiiiiiiiiiiiiiiimiiiiiiiiiiimiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiHiiiii

Fig. 12 Voucher material of Rubia arborescens asperior Insularum Canariensium, . . . (= Rubia fruticosa Aiton ssp. fruticosa) in Plukenet's
herbarium (now part of Herb. Sloane, vol. 97: 171 (BM)).

24

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

Fig. 13 Plukenet's illustration oi Solatium tomentosum Canariense spinosum, fructu Cerasorum forma & magnitudine . . . (= Solanum
vespertilio Aiton) from his Phytographia, t. 316, f. 3 (1694).

87. Salicis folio Canariensis Arbuscula impatiens [. . .] s.
Arbor crepitus; The Snap-Tree nostratibus, & Satterel
Insulanis dicta. Ab Ecbolii Indici s. Adhatodae, cucullatis
floribus Aemula, Hujus ex iisdem locis, parum diversa.
Eademque est, cum Arbuscula Canariensi Salicis aut
pot ins Oleae Sylv. Barbadens. folio & facie, & c. Aim. : 328
(1696), Mant.: 165 (1700).

Syn.: Apancholoa (sorte) sive Herba lymphis infiliens Nier-
amberg. de Exotic fol. 351 [?].

Com.: The use of the common name 'Satterel' is indicated in
this description, but we have been unable to link this common
name with any Canarian species.

88. Salvia auriculata, mucronatis foliis crassis,
lanuginosa, appendicibus aucta Aim.: 329 (1696),
Phyt.: t. 57, f. 3(1691).

Del.: Salvia sp. [HS 97: 184. The phrase-name does not give
any reference to Macaronesia. However, labels from the
herbarium specimen refer both to Phyt.: t. 57 f. 3 and to the
Canary Islands (i.e. 'Sage, salvia trifoliada Canariensis') and
Plukenet's drawing from Phytographia is extremely similar to
this specimen. However, the material belongs to a species
that does not occur in the Canary Islands and it is likely that it
was wrongly assigned there by Plukenet].

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

25

if. 3 $-2. /

/*

* I**'

%

/

miiiiiiriiiiiiiiiiiiiiiiimimiitfiBiiiHiiiiiiiiilllMlllllillllllllllU^

Fig. 14 Voucher material of Solatium tomentosum Canariense spinosum, fructu Cerasorum forma & magnitudine . . . (= Solarium vespertilio
Alton) in Plukenet's herbarium (now part of Herb. Sloane, vol. 98: 57 (BM)).

26

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

Fig. 15 Plukenet's illustration of Tithymalus aizoides lactifluus s. Euphorbia Canariensis, quadrilatera, & quinquelatera Cerei effigie . . . ( -
Euphorbia canariensis L.) from his Phytographia, t. 320, f. 2 (1694).

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

27

. f.

7

. -J, 4-

iiiiiiiiiimiiiiiiiiiiiiiiiiiiiiiimiiiiimiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiuiiiuiiiiii

Fig. 16 Voucher material of 77//z_yma/s aizoides lactifluus 5. Euphorbia Canariensis, quadrilatera, & quinquelatera Cerei effigie . . . (=
Euphorbia canariensis L.) in Plukenet's herbarium (now part of Herb. Sloane, vol. 102: 86 (BM)).

28

89. Salvia sylvestris amplissimis Verbasci foliis,
graveolens, flore albo parvo Canariensis inter
Delineation. [. . .] Arvida Salva Insulanis vulgo. Flos
ejus galea caret. Facie externa cum Stachyde convenit

Aim.: 329 (1696).

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

(Linnaeus, 1738)].

Det.: * Phyllis nobla L. [HS 102: 15. We have also found
sheets in Miller's (from Chelsea Physic Garden) and
Uvedale's herbaria from the late seventeenth and early
eighteenth centuries] .

His.: Leucophae canariensis (L.) Webb & Berth. [W.B.].
Det.: * Sideritis canariensis L. [?]

Com.: There is neither a drawing nor an obvious specimen
for this entry. The description is extremely similar to that of
entry 95 and our determination is based upon that. The
common name 'Arvida Salva' is given by Plukenet, however
Sideritis species are known as 'chahorra' and 'salvia blanca' in
the Canary Islands.

90. Sedum in a jus Canarinum [. . .] pilis ad oras
foliorum hispidis argenteo-lucidis fimbriatum floribus
ex flavo pellescentibus, per ramos numerosissimos
recurvatos eleganti serie dispositis [. . .] Corozone
Celio ab Insulanis dictum Aim.: 340 (1696), Mant.:
169 (1700), Phyt.: t. 314, f. 1 (1694).

His.: Sempervivum canariense L. [Linn.; Gis.; Ten.], Aeo-
nium canariense (L.) Webb & Berth. [W.B.].
Det.: * Aeonium tabulaeforme (Haw.) Webb & Berth, [deter-
mination based on Plukenet's description and illustration.
Nevertheless, one herbarium specimen without a label is
found in HS 101: 201].

Com.: The common name 'Corozone Celio' (perhaps a
derivation of 'corazoncillo') is reported in this description.
The common name 'corazoncillo' is used for Lotus species
whereas plants of Aeonium are known as 'pastel de risco',
'oreja del abad', verol' or 'bejeque'.

91. Sedum maximum villosum ex Insulis Fortunatis La
Frecho de las uvas Insulanis dictum Aim.: 339 (1696).

Syn.: Sedum maximum arborescens, latifolium, flore flavo,
Capitis Bonae Spei, D. ten Rh. (Breyne, 1678a).
Com.: Plukenet gave the common name 'La Frecho de las
uvas' for plants of this species. No obvious Spanish common
name has been found which could be related to that reported
by Plukenet.

92. Sideritis canariensis Mocanes dicta, folio subtus
incano, margine spinulis asperato, calyculis tomentosis

Aim.: 346 (1696), Phyt.: t. 315 f. 4 (1694).

His.: Scutellaria integrifolia L. [Ten.].
Det.: * Forsskaolea angustifolia Retz. [HS 98: 39].
Oth.:HS102: 12.

Com.: The common name 'Mocanes' is given in this descrip-
tion. However the name 'mocan' is used, in the Canary
Islands, for plants of Visnea mocanera L. f. Plants of F.
angustifolia are known as 'ratonera' or 'hierba ratonera'.

93. Sim pi a- nob la Canariensium, Planta oblongis,
amplioribus splendentibus foliis ternis circa caulem
ambientibus, venosis Aim.: 347 (1696).

His.: Phyllis nobla L. [this name was originally published by
Linnaeus (1753). However, reference to Plukenet's phrase-
name was given by Linnaeus only in Hortus cliffortianus

94. Solanum tomentosum Canariense spinosum, fructu
Cerasorum forma & magnitudine [. . .] Hujus fructus
sunt saturate Laccae coloris ex quibus fucum conficiunt
Insulanae Mulieres, quo faciem oblinunt, & ex pallida
rubicundam efficiunt, ut hoc modo Amasiis suis
formosiores reddantur. Bella Donna Canarina spinis
armata; Permenton Insulanis vocatur Aim.: 351
(1696), Phyt.: t. 316, f. 3 (1694); see Fig. 13.

Syn.: Planta spinosa incognita (Bontius, 1658).

His.: Solanum vespertilio Aiton [W.B.].

Det.: * Solanum vespertilio Aiton [HS 98: 57; see Fig. 14].

Oth.:HS88: 122.

Com.: The common name 'Permenton' is given by Plukenet,

this follows closely the names used in the Canary Islands:

'pimentero' and 'pimientero'. This description has an ethno-

botanical interest as it presents a detailed account on the

traditional use of the fruits of this species. Its juice was the

basis for a dye which was used as a facial makeup by the

Canarian women.

95. Stachys amplissimis Verbasci foliis, floribus albis
parvis non galeatis, spica Betonicae, ex Insula
Canarina [. . .] Arvida salva Incolis, & nostratibus
Sage-Tree (i.e.) Salvia arbor nuncupatur Aim.: 356
(1696), Phyt.: t. 322, f. 4(1694).

His.: Sideritis canariensis L. [Linn.; Gis.; Ten.].
Det.: * Sideritis canariensis L. [HS 98: 69; 102: 39].
Oth.:HS102:40.

Com.: The use of the common name 'Arvida salva' is
reported by Plukenet. This description is extremely similar to
that of entry 89 and we believe that both refer to the same
species. For a discussion of the use of the common name by
Plukenet, see entry 89.

96. Tithymalus aizoides lactifluus s. Euphorbia
Canariensis, quadrilatera, & quinquelatera Cerei
effigie, ad angulos per creba intervalla spinis rectis
atronitentibus, Gazellae cornua referentibus, armata

Aim.: 370 (1696), Mant.: 182 (1700), Phyt.: t. 320, f. 2
(1694); see Fig. 15.

Syn.: Quauh cuetz placuitlapilli (Hernandez, 1651), Euphor-
bio similis Sadricalli Indorum (Breyne, 1689), Rangiseri cor-
nua referens planta Zeylanica. Sandricay Zeylan (Hermann,
1689), Tithymalus quadrangularis spinosus S. spinis geminis
aduncis ex eadem sede ortis armatus, succo lacteo acerrimo
turgidus (Kiggelaer, 1690).

His.: Euphorbia canariensis L. [Linn.; Gis.; Ten.; W.B.].
Det.: Euphorbia canariensis L. [HS 102: 86; see Fig. 16].

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

97. 1 it h vina I us dendroides Linariae foliis ex Insula

Canarina Aim.: 369 (1696), Mant.: 181 (1700), Phyt.:
t. 319, f. 5 (1694).

Syn.: Tithy mains perennis & procerior lini folio acuto

(Sloane, 1696).

His.: Othonna tenuissima L. [Gis.], O. linifolia L. f. [Ten.],

Euphorbia regis-jubae Webb & Berth. [W.B.].

Det.: * Euphorbia regis-jubae Webb & Berth, [we have not

found any specimen for this species in Plukenet's herbarium,

this determination is based on the drawing from Phytographia

and on Webb & Berthelot's identification (Webb & Berthe-

lot, 1846-1847)].

29

TAXA FOUND IN PLUKENET'S HERBARIUM
BUT NOT IN HIS WORKS

A study of the whole of Plukenet's herbarium revealed that
there were 34 herbarium specimens for 22 taxa which,
although reported as collected in the Macaronesian area do
not appear to have have been described in Plukenet's works.
Material which fell under this category is listed below. An
entry number is given for each taxon and within each entry
the following headings can be found:

(1) Taxonomic determination of specimens (coded as Det.:).
Endemic taxa are pointed with an asterisk.

(2) Location of specimens in Plukenet's herbarium (coded as
Herb.:). Labels are given in square brackets.

(3) Comments (coded as Com.:). Additional remarks con-
cerning the specimens are included in this heading.

98. Det.: * Apollonias barbujana (Cav.) Bornm.

Herb.: HS 95: 104; 96: 137 ['Laurus canariensis odorata']; 96:
138 ['An tree strawberry of the Canaries']; 100: 167.

99. Det.: Asparagus cf. capensis L.

Herb.: HS 95: 108 ['Asparagus pelroa s. Corruda aculeata ex
Insulis Canarinis'].

Com.: The label of this specimen refers to Phyt.: t. 78, f. 3
and follows the description found in Aim.: 54 (Asparagus
aculeatus, triplice spina surrectus). This species does not occur
in the Canary Island flora. It is worth mentioning that in HS
95: 108 there is another specimen which, though poorly
preserved, resembles the Canarian endemic A. umbellatus
Link.

100. Det.: * Bryonia verrucosa Dryand.
Herb. : HS 95: 200.

101. Det.: Calendula arvensis L.

Herb.: HS 95: 131 ['Calendula sylv. minima Canariensis'].

102. Det.: Centhranthus calcitrapae (L.) Dufr.
Herb.: HS 98: 127 ['Valeriana annua ex Insulis Canarina'].

103. Det.: Chrysanthemum segetum L.

Herb.: HS 95: 162 ['Chrysanthemum from I. Canaries'].

104. Det.: Conyza bonariensis (L.) Cronquist.
Herb.: HS 95: 193 ['From I. Canaries an major [illeg.]' ].

105. Det.: * Echium cf. leucophaeum Sprague &
Hutch.

Herb.: HS 96: 14 ['Echium from I. Canaries'.].

106. Det.: Erica arborea L.

Herb.: HS 100: 26 ['An Erica from the Canaries look in my
former collections you'll find its name'], 100: 27 ['An Erica 2a
Clus. ex Insula Canarina vid autorum'].

107. Det.: Geranium cf. rotundifolium L.

Herb.: HS 96: 53 ['An Geranium fatid. ex Insula Canarina'].

108. Det.: * Gonospermum fruticosum (Buch) Less.
Herb.: HS 96: 195 ['Millefol. Tanaceti fol Canariense'].

109. Det.: Hordeum vulgare L.

Herb.: HS 105: 41 ['Gramen canarium hirsutum'].

110. Det.: Laurus azorica (Seub.) Franco.
Herb.: HS 100: 168.

111. Det.: * Lavandula multifida L. ssp. canariensis
(Mill.) Pit. & Proust.

Herb. : HS 100: 167.

112. Det.: Mentha spp.

Herb.: HS 101: 16 ['Teucrii facie frutice Canariensis floribus
ociquis albis non galeatis Bobart. Of this saith he I observe 3.
varieties suppose this yr. 1 st . The 2 nd hath longer leaves with
ye shape & green ness of Betony and such flowers as this. The
3 d is an uprighter plant more wooly and rounder leav'd, which
I thinke is at Hampton Court. But by Mr Bobarts leave these
3 are specifically distinct. Mentha pilulifera Betonicae forma
ad odore aromatico. vid Almag. Bot. 247'].
Com.: There are six specimens on this page, one of them
identified as Mentha cf . piperita L. , the other five apparently
of a different species.

113. Det.: Mentha longifolia (L.) Huds.

Herb.: HS 96: 187 ['Mentastri spiculi folio longiore candicau-
lis species major ex Insula Canarina'].

114. Det.: Mentha cf. spicata L.

Herb.: HS 96: 189 ['Mentha affigio inodora ex Insula Mad-
era'].

115. Det.: * Pericallis tussilaginis (L'Her.) D. Don.

Herb.: HS 95: 128 ['Cacalia Africana floribus Jacobea pur-
pureis ex insula Canarina'].

30

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

116. Del.: Salvia cf. verbenaca L.

Herb.: HS 96: 100 ['An Hormino Canariensis'].

117. Del.: Sisymbrium cf. erysimoides Desf.
Herb.: HS 96: 18 ['An Eruca Gomerensis'].

118. Del.: * Stachys germanica L. var. canariensis
Font Quer & Svent.

Herb.: HS 98: 69 ['Stachys Canariensis [crossed] Verbasci
folio Canariensis'].

119. Det.: Umbilicus cf. horizontalis (Guss.) DC.

Herb.: HS 95: 200 ['Cotyledon s. Umbilicus radici tuberosa
minor ex Insul. Canar.'].

120. Herb.: HS 89: 68 ['Filicula Canarina elegans'].

Com.: This specimen does not belong to any known species
from the Canary Island flora.

121. Herb.: HS 96: 14 ['Elegans Salvia [illeg.]
Canariensibus Dicta'].

Com.: There is no specimen for this label.

122. Herb.: HS 98: 95 ['An Thymelea from I.
Canaries'].

Com.: The specimen situated close to to this label does not
belong to any species from the Canary Island flora. However
there is another specimen on this page belonging to Daphne
gnidium L. to which this label could refer.

DISCUSSION

With almost 125 plant entries, both Plukenet's herbarium and
his publications can be regarded as the most important
pre-Linnaean account of the Macaronesian flora. The major-
ity of species recorded by this herbalist are the earliest known
references to the plants of this flora. Plukenet's descriptions
of Canarian plants pre-date those of Feuillee (Feuillee, 1724)
which has been traditionally considered as the most complete
pre-Linnaean work for the Canarian flora (Herrera-Pique,
1987). Although it was published later than Plukenet's work,
the approximately 20 species illustrated by this French natu-
ralist are also extremely important as they were the first
known field drawings of Canarian plants made by any Euro-
pean scientist who visited the archipelago (Herrera-Pique,
1987).

In this study we have demonstrated that from Plukenet's
works and collections, the seventeenth century herbalist
clearly had available a considerable amount of information
on the Macaronesian flora. However, subsequent taxonomic
reviews of the region have made hardly any mention of them.
Linnaeus himself referred to only 24 of these phrase-names
and in his own copies of Plukenet's publications there are few
annotations for the Macaronesian species.

Other important works published shortly after Linnaeus
(e.g. Linnaeus filius, 1782; Aiton, 1789; L'Heritier,

1785-1805) do not quote any of Plukenet's names despite
having had access to his publications, and they were based on
the exhaustive plant collections made in the Azores, Canaries
and Madeira between 1776 and 1779 by the collector from
Kew, Francis Masson (Lemmon, 1968). Only L'Heritier
(1788) mentioned Plukenet's polynomial Mentha canariensis
frutescens, foliis [. . .]foliorum longiribus pediculis insidenti-
bus (entry 73) as a synonym of Bystropogon canariense (L.)
L'Her.

More recent publications restricted to the Macaronesian
flora have followed the same pattern. For instance, there are
only two mentions of Plukenet's names in the Flora of the
Canary Islands produced by Pitard & Proust (1908) whereas
in Lowe's Floras of Madeira (Lowe, 1838, 1857-1872) we
have found no reference to any of his polynomials. However,
the Canarian naturalist Viera y Clavijo (1866-1869) men-
tioned, in his Diccionario de historia natural de las Islas
Canarias, twelve of the Plukenet names previously cited as
synonyms by Linnaeus. This poor utilization of Plukenet's
publications by researchers of the Macaronesian flora may
well be due to the fact that his works have no geographical
index, and as E. L. Greene reported on the Almagestum '. . .
in this kind of book, you are helpless to even begin to search,
unless you have some idea of the genus, or a genus, to which
it [a plant] may belong . . .' (Greene, 1983).

It was only in the middle of the nineteenth century that
Webb & Berthelot in their Histoire naturelle des lies Canaries
(Webb & Berthelot, 1836-1850) cited 27 of Plukenet's
phrase-names in the synonymy of some of the taxa described
in their work. Some of these names were not previously
treated by Linnaeus (e.g. Ilex perado Aiton ssp. platyphylla
(Webb & Berth.) Tutin for entry 12, Chamaecytisus prolif-
erus (L. f.) Link for entry 29, Arbutus canariensis Veill. for
entry 18). Furthermore, Webb & Berthelot also looked
critically at Linnaeus' treatment of Plukenet's Macaronesian
names. For instance they found that the polynomial Cytisus
Canariensis, microphyllos, angustifolius, prorsus incanis
[. . .] Esta Insulanis nuncupatur (entry 30) had been incor-
rectly synonymized with Genista canariensis L. by Linnaeus.

Further studies will be needed in order to establish the
taxonomic identity of earlier phrase-names given as syn-
onyms by Plukenet. However, many of them were associated
with plant species from regions other than Macaronesia.
Among them there are taxa from the New World (e.g.
Marggraf, 1648; Hernandez, 1651; Ogilvy, 1671; Plumier,
1693), the Far East (Rheede tot Draakenstein, 1689, 1690)
and the Middle East (Rauwolf, 1707). It is very doubtful
whether these names have any real taxonomic relationship
with those Macaronesian taxa described by Plukenet. Indeed
we could identify what appear to be 'good' synonyms for only
seven of his descriptions (i.e. entries 14, 37, 61, 66, 78, 96 and
97). Although Plukenet's publications have been regarded as
containing one of the most comprehensive reviews of names
from the ancient herbalists (Pulteney, 1790) it seems that
there were many errors in these synonyms. This was noticed
by John Ray, who in one of his letters to Hans Sloane stated
'. . . I have gotten a sight of Dr. Plukenet's 'Almagaestum
bot' [. . .] As far as I am able to judge, he is often out in his
conjectural synonymes . . .' (Henrey, 1975).

In contrast, Plukenet's illustrations of Macaronesian plants
(a selection of which are shown amongst Figs. 3-15) are
remarkable. Although it seems that they were not drawn by
Plukenet himself but by various other artists, the most
important of them being M. van der Gucht (Henrey, 1975),

PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

31

they reveal the skill and observational abilities of Plukenet as
a botanist. This is clearly shown in the drawing of Solanum
vespertilio Aiton (Fig. 13 and entry 94). This endemic from
Tenerife and Gran Canaria is one of the few Solanum species
which has flowers with one of the stamens longer than the
others and the style curved and elongated (Jaeger & Hepper,
1986). This means that flower buds of this species have a
small peak towards their ends. The character is clearly
illustrated in Plukenet's drawing, making its identification as
S. vespertilio unmistakeable.

Most of his references and collections appear to have been
based on material brought to Britain by visitors who stayed in
Macaronesia. It is well known that a strong trade in orchil,
pitch, sugar, wine and cloth was established between the
Canary Islands and Britain during the sixteenth and seven-
teenth centuries (Morales-Lescano, 1973; Fernandez-
Armesto, 1982; Lobo-Cabrera, 1988). This led to the
foundation of trading companies such as the 'Canary Com-
pany' in 1655 (Morales-Lescano, 1965a). Furthermore, other
European visitors visited the islands during this period in
order to study their natural history (e.g. T. Sprat from the
Royal Society in Tenerife in 1667 (Morales-Lescano, 19656),
John Cuningham in La Palma and Hans Sloane in Madeira in
the late seventeenth century (Dandy, 1958)). It is therefore
likely that many of Plukenet's references which concern the
Macaronesian region reflect the trading and scientific links
which existed between Britain and the islands during this
time. Being in close association with the Royal gardens and
with the most important British herbalists, Plukenet could
have had easy access to most of the exotic plant material
which was introduced into Britain during the second half of
the seventeenth century.

The collections and works of Leonard Plukenet provide a
valuable source of information on early accounts of non-
European floras, as already shown for the West Indies
(Howard, 1979). However, it is likely that many of his
references and specimens came originally from sources which
were European in a broad sense. Plukenet's studies in the
Macaronesian region provide a good indication of what
knowledge was had by European naturalists and herbalists
half a century before Linnaeus (1753) published what we now
accept as the first valid binomial names for Macaronesian
plants.

ACKNOWLEDGEMENTS. The authors wish to thank C. Lorenzo-
Millana (Ministerio de Education y Ciencia, Spain) and E. Del
Rio-Hijas (Consejo Superior de Investigaciones Cientificas, Spain)
for the Latin translations from the originals. We are also grateful for
the assistance of staff from the British Library (Main Library and
Manuscript Collections) and from the libraries of the Royal Botanic
Gardens (Kew) and of The Natural History Museum (London). We
are also grateful to the Bayerische Staatsbibliothek (Munich) for
providing us with a microfilm copy of Tenzel's work. Our gratitude
also goes to the Linnean Society of London for permission to study
Linnaeus' annotated copy of Plukenet's works and to J. Perera
(Universidad de La Laguna) for his help in tracing some of the
references. This research was partially supported by a personal grant
(JFO) from Ministerio de Education y Ciencia, Spain: Plan Nacional
de Formation de Personal Investigador en el Extranjero (grant no.
PG88 42044506) and by the financial support of Centre de Investiga-
tion y Tecnologia Agrarias de Canarias.

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PRE-LINNAEAN REFERENCES FOR MACARONESIAN FLORA

33

Tournefort, J. P. 1694. Elemens de botanique 1. Paris.

V allot, A. 1665. Hortus regius. Paris.

Viera y Clavijo, J. 1808. La flora de Canarias. Catdlogo razonado de las plantas

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P.B. Webb & S. Berthelot, Histoire naturelle des lies Canaries 3 (2, sect. 2,
livr. 62). Paris.

1846-1847. Phytographia canariensis (Campanulaceae-
Equisetaceae). In P.B. Webb & S. Berthelot, Histoire naturelle des lies
Canaries 3 (2, sect. 3, livr. 92). Paris.

SYSTEMATIC INDEX

Accepted names for taxonomic determinations are indicated with the symbol #.

Achyranthes argentea Lam. 4

# Achyranthes aspera L. 4
Acrostichum marantae L. 38
Acrostichum velleum L. 38

# Adenocarpus foliolosus (Aiton) DC. 31

# Adiantum reniforme L. 40

Aeonium canariense (L.) Webb & Berth.
90

# Aeonium tabulaeforme (Haw.) Webb &
Berth. 90

# Aizoon canariense L. 55

# Allium spp. 26

# Apollonias barbujana (Cav.) Bornm. 98

# Arbutus canariensis Veill. 18

# Argyranthemum frutescens (L.) Sch. Bip.
ssp. frutescens 20

# Asparagus capensis L. 99

# Asparagus umbellatus Link 99

# Asplenium hemionitis L. 39
Asplenium palmatum Lam. 39

# A triplex glauca L. 19

# Bosea yervamora L. 15

# Bryonia verrucosa Dryand. 100

# Bystropogon canariensis (L.) L'Her. 68,
70,73

# Bystropogon plumosus (L. f.) L'Her. 73,
74

Cacalia kleinia L. 65

# Calendula arvensis L. 101

Callianassa canariensis (L.) Webb &

Berth. 32

Campanula canariensis L. 21

# Canarina canariensis (L.) Vatke 21

# Cedronella canariensis (L.) Webb & Berth.
67,69

# Centhranthus calcitrapae (L.) Dufr. 102

# Chamaecytisus proliferus (L. f.) Link ssp.
proliferus var. proliferus 29

# Cheilanthes catanensis (Cos.) M. P. Fuchs.
38

Chrysanthemum frutescens L. 20
Chrysanthemum indicum L. 67

# Chrysanthemum segetum L. 103

# Convolvulus canariensis L. 26, 50

# Convolvulus fruticulosus Desr. 27

# Conyza bonariensis (L.) Cronquist 104

# Cystopteris fragilis (L.) Bernh. 42
Cytisus proliferus L. f . 29

Danae androgyna (L.) Webb & Berth. 57

# Daphne gnidium L. 122

# Davallia canariensis (L.) J. E. Sm. 41
Digitalis canariensis L. 32

# Dracaena draco (L.) L. 78
Dracocephalum canariense L. 69

# Echium leucopheum Sprague & Hutch. 35,
105

# Erica arborea L. 106

# Erica scoparia L. ssp. platycodon (Webb &
Berth.) Hansen & G. Kunkel 29

# Euphorbia canariensis L. 96
Euphorbia obtusifolia Poir. ssp. regis-jubae
(Webb & Berth.) Maire 97

# Euphorbia regis-jubae Webb & Berth. 97

# F/CMS carica L. 37

# Forsskaolea angustifolia Retz. 92

# Galactites tomentosa Moench 22
Gendarussa hyssopifolia (L.) Webb &
Berth. 34

Genista canariensis L. 31

# Geranium rotundifolium L. 107

# Gonospermum fruticosum (Buch) Less.
108

# //edera /ie/k L. 15

# Hordeum vulgare L. 109

# Hypericum canariense L. 51
Hypericum floribun dum Aiton 51

# Ilex canariensis Poir. 11

# Ilex per ado Aiton ssp. platyphylla (Webb
& Berth. ) Tutin 12

Ilex platyphylla Webb & Berth. 12

# Isoplexis canariensis (L.) Loudon 32, 33

# Jasminum azoricum L. 52, 54

# Juniperus phoenicea L. 28

# Justicia hyssopifolia L. 16, 34

# Kleinia neriifolia Haw. 64, 65

# Laurus azorica (Seub.) Franco 110
Laurus indica L. 59
Lavandula abrotanoides Lam. 62

# Lavandula buchii Webb 62
Lavandula multifida L. 62

# Lavandula multifida L. ssp. canariensis
(W. Mill.) Pit. & Proust 111

# Lethariella canariensis (Ach.) Krog 75
Leucophae canariensis (L.) Webb &
Berth. 89

# Limonium pectinatum (Aiton) Kuntze 63

# Lotus spp. 90

# Lupinus albus L. 66

# Mammea americana L. 13

# Mentha spp. 112
Mentha canariensis L. 73

# A/em/ia longifolia (L.) Huds. 113

# Mentha piper ita L. 112

# Mentha spicata L. 114

# Mesembryanthemum nodiflorum L. 36

# Myrica faya Aiton 17, 23

# Ocotea foetens (Aiton) Baill. 14

# Origanum vulgare L. 77
Othonna linifolia L. 97
Othonna tenuissima L. 97

# Papaver rhoeas L. 79

Pericallis tussilaginis (L'Her.) D. Don 115

# Periploca laevigata Aiton 8, 10

# Persea indica (L.) Spreng. 59

# Phaseolus vulgaris L. 81

# /%// noWfl L. 93

# Plocama pendula Aiton 82

# Polygonum maritimum L. 83
Pyrethrum frutescens (L.) Gaertn. 20

# Ranunculus cortusifolius Willd. 84

# Rubia fruticosa Aiton ssp. fruticosa 85

# Rumex lunaria L. 1
Ruscus androgynus L. 57

# Sa/wfl sp. 88
Salvia africana L. 49

# Salvia canariensis L. 49

# Salvia verbenaca L. 116
Sarothra gentianoides L. 27
Scutellaria integrifolia L. 92

# Semele androgyna (L.) Kunth 57
Sempervivum canariense L. 90

# Sideritis canariensis L. 89, 95

# Sisymbrium erysimoides Desf. 117

# Solanum vespertilio Aiton 94

# Stachys germanica L. var. canariensis Font
Quer&Svent. 118

Statice limonium L. 63

# 7e///ie canariensis (L.) Webb & Berth. 30

# Teucrium heterophyllum L'Her. 86
Trichomanes canariense L. 41

# Umbilicus horizontalis (Guss.) DC. 119

# Usnea articulata (L.) Hoffm. 76

# Viburnum tinus L. ssp. rigidum (Vent.) P.
Silva 53

# Visnea mocanera L. f. 92

Webbia floribunda (Aiton) Webb & Berth.
51

BIBLIOGRAPHY ABBREVIATIONS FOUND IN PLUKENET'S WORKS

Abbreviations for Botanic Gardens and for publications as they are cited in Plukenet's works are given below. Full identification
of these abbreviations can be found in the list of References. Those abbreviations which were not identified are indicated by
'[?]'. Plant entry numbers are also given for each abbreviation.

34

J. FRANCISCO-ORTEGA, A. SANTOS-GUERRA AND C.E. JARVIS

Anguillarae (Anguillara, 1561) 37

Banisteri Cat. Stirp. Virgin. (Banister, 1693)

10,68

Bellonius (Belon, 1553) 25
Bocc. de PI. Sicil. (Boccone, 1674) 4
Bontii (Bontius, 1658) 56, 94
Breyn. Cent. (Breyne, 1678a) 91
Breyn. Fascicul. Rarior. (Breyne, 1678b) 80
Breyn. Prodr. (Breyne, 1689) 96
C. B. P. (Bauhin, 1623) 1, 20, 25, 28, 29, 30,

33, 37, 45, 47, 63, 65, 75, 76, 78, 82, 84
C. B. Phytopin. (Bauhin, 1596) 1

C. B. Prodr. (Bauhin, 1620) 31
Camerar. H. Medic. (Camerarius, 1588) 66
Cat. Jam. (Sloane, 1696) 10, 15, 38, 39, 86, 97
Cat. PI. ad Parad. Bat. append. (Hermann,

1698) 6

Clus. (Clusius, 1576) 78
Clus. (Clusius, 1601) 78
Clus. exot. (Clusius, 1605) 65
Clus. Hispan. (Clusius, 1576) 28
Clus. Hist. (Clusius, 1601) 25, 28, 76, 84
Clusij (Clusius 1601) 39, 63
Cordi. Histor. (Cordus, 1561) 29
Cordi. in Dioscorid. & Hist. (Cordus, 1561)

37

D. Raius Hist. PI. (Ray, 1686) 39
Dalechamp. Hist. Lugd. (Dalechamps, 1586)

29

Dioscorides tradit. [?] 14
Dioscoridis [?] 37

Dodart. Memoir. (Dodart, 1676) 63
Ferrarif. (Ferrarius, 1633) 59
Florent. Schuyl. Cat. H. Leyd. (Schuyl, 1672)

59
Fr. Cupani in Hort. Catholic. (Cupani, 1696)

25,66
Galenus in libr. de Antidotis (Galenus, 1587)

14

Grislisleii, Viridar. Lusit. (Grisley, 1661) 52

H. Amst. (Commelin, 1689) 78

H. Beaum. (Kiggelaer, 1690) 78, 96

H. Eyst. (Besler, 1613) 66

H. Leyd. (Hermann, 1687) 66

H. M. P. 9 (Rheede tot Draakenstein, 1689)

8
H. M. P. 10 (Rheede tot Draakenstein, 1690)

10
H. Malab. Part 9 (Rheede tot Draakenstein,

1689) 32

H. R. P. (Vallot, 1665) 59
Hernand. apuc Recc. (Hernandez, 1651) 46
Hort. Amst. (Commelin, 1689) 61
Hort. Fames. Aldino (Aldino, 1625) 59
Hort. Malab. Part 9 (Rheede tot

Draakestein, 1689) 34
Hort. Malab. Part 10 (Rheede tot

Draakestein, 1690) 67
Hort. Regio Hampton (The Royal Garden at

Hampton Court Palace, on the Thames in

south-west London) 21
J. B. Tom. 1 (Bauhin & Cherler, 1650) 13,

14,30,31,65,75,78
J. B. Tom. 2. (Bauhin & Cherler, 1651) 1, 25,

33,47

Jacobi Sylvij (Manadi & Sylvij, 1598) 14
Johannis Adams in Hort. Comptoniano

(John Adams was gardener to the Duke of

Beaufort at Badminton, Henry Compton

was Bishop of London and his garden was

at Fulham Palace in London (see Dandy,

1958) 46
Lib. 2 de Simplicibus (Manadi & Sylvij, 1598)

14

Lob. Icon. (Lobelius, 1581) 1
Lob. Obs. (Lobelius, 1576) 1
Magnol Botan. Monspel. (Magnol, 1676) 47

Marcgrav. (Marggraf, 1648) 13

Mesue (Manadi & Sylvij, 1598) 14

Mexicana Hernand. apud Recc. (Ray, 1688b)
68

Moris. Hist. (Morison, 1680) 7, 66

Nieramberg. de Exotic [?] 87

Nierembergio (Nieremberg, 1635) 13

Nov. Hisp. apud Reccum (Hernandez, 1651)
69

Nov. Hisp. Terent. apud Reccum (Hernan-
dez, 1651) 21, 59, 96

Nov. Hispaniae Terentij apud Recc. (Her-
nandez, 1651) 65

Ogilv. Americ. (Ogilvy, 1671) 17

P. B. P. (Hermann, 1689) 19, 48, 61, 96

Park. Th. (Parkinson, 1640) 59, 66, 78, 84

Park. Theatr. (Parkinson, 1640) 47

Plinij (Pliny, 1826c) 58, 59

Plinio (Pliny, 1826b) 37

Plinius (Pliny, 1826a) 14

Plumier (Plumier, 1693) 26, 42

Ponae ejusd. Descript. Montis Baldi (Pona,
1601) 1

Prodr. (Bauhin, 1620) 45, 82

Prosp. Alpini de PI. exot. (Alpino, 1627) 50

Raij Hist. PI. (Ray, 1686) 19, 66

Raius Hist. PI. (Ray, 1688a) 47

Rawolf. (Rauwolf, 1707) 28

Rivorum Medicina, Hernand. (Hernandez,
1651)21

Schol. Botan. (Sherard, 1689) 45

Solinus (Solinus, 1958) 14

Turneforti. Elem. Bot. (Tournefort, 1694)
10,49

Turnefort. de opt. Meth. instituend in re
Herber epist. ad Sherard. [?] 67

Virginiensium Ogilv. (Ogilvy, 1671) 23

Vossio, Not, in Pompon. Melam [?] 14

Bull. not. Hist. Mus. Lond. (Bot.) 24(1): 35-48

Issued 23 June 1994

Studies on the lichen genus Sticta (Schreber)
Ach.: II. Typification of taxa from Swartz's
Prodromus of 1788

DAVID J. GALLOWAY

Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD

SYNOPSIS. Of the 18 lichens described from Jamaica by Swartz in his Prodromus (1788), three taxa are referable to
the genus Sticta, viz. Lichen damaecornis, L. laciniatus and L. tomentosus, although Swartz's L. laciniatus is a later
homonym of L. laciniatus Hudson (1762) and is illegitimate. Lichen damaecornis and L. tomentosus are typified
from authentic Swartz material and detailed descriptions and taxonomic notes are given. Swartz material distributed
as L .laciniatus comprises two distinct taxa which are newly described here as Sticta laciniosa and S. swartzii.

INTRODUCTION

Sticta is a widely distributed lichen genus of some 60 species
with a preponderance of taxa found in tropical or subtropical
areas, especially the Caribbean, Central America and the
tropical parts of South America, and the palaeotropics from
Africa and the Indian Ocean Islands to the Pacific. Apart
from the cosmopolitan taxa S. fuliginosa (isidiate) and 5.
limbata (sorediate), found on all major land areas, and the
widespread palaeotropical species 5. sublimbata and 5.
wiegelii, species of Sticta tend to have rather restricted
distributions with both tropical and temperate regions having
well-defined endemic taxa, for example New Zealand (Gallo-
way, 1985), East Africa (Swinscow & Krog, 1988), Juan
Fernandez and southern South America (Galloway, 1994).
Species of Sticta and Lobaria (unlike Pseudocyphellaria) are
particularly strongly represented in the neotropics in com-
parison with numbers of taxa in cool temperate parts of South
America (Galloway & Arvidsson, 1990; Galloway, 1994).
This reflects a basic biogeographical distinction between
these three important Southern Hemisphere genera with
Pseudocyphellaria having pronounced austral affinities (Gal-
loway, 1987, 1988o, 19926) while both Sticta and Lobaria
appear to be predominantly tropical groups (Galloway &
Arvidsson, 1990). Species of Sticta are now recognized as
being important nitrogen producers in both tropical and
temperate forest ecosystems (Green et al., 1980; Galloway,
1988a, 1992a, 19926, 1994). Nitrogen is fixed by cyanobacte-
ria present either as primary photobionts or as internal
cephalodia (James & Henssen, 1976). The taxonomy of Sticta
is still very confused, especially in tropical regions where
speciation in the genus is most marked. A number of early
names in the genus, for example Sticta damaecornis, have
been widely used in the tropics, when it is now known that
several entities are involved. In order to clarify correct use of
names in Sticta, the present paper is one in a series attempting
to define the limits of taxa described in the late eighteenth
century and early nineteenth century.

The Swedish botanist Olof Peter Swartz (1760-1818) was a
student of Linnaeus's son, Carl Linnaeus, completing his
studies in medicine and natural history at the University of

Uppsala in 1783. Twenty-two years old, and furnished with
ample private means, he was keen to travel to distant parts to
study natural history in the tradition of the elder Linnaeus's
'Linnaean apostles' (Stafleu, 1971). Swartz decided on the
West Indies as a suitable area for study, following in the
footsteps of Sir Hans Sloane, Charles Plumier, Patrick
Browne and Nicolaus Joseph Jacquin (Steam, 1980). Choos-
ing the island of Jamaica, he embarked from Sweden on 5
August 1783, and landed at Montego Bay in Jamaica on 5
January 1784 after some time botanizing at Boston, Massa-
chusetts. During his time in Jamaica he collected widely from
the interior of the island and reached the highest summits in
the Blue Mountains (Stearn, 1980). On his return home in
1786 he spent some time in London working on the arrange-
ment and naming of his West Indian collections with the help
of Sir Joseph Banks's herbarium and library and with the
assistance of Jonas Dryander, Banks's Swedish librarian. The
first results of Swartz's West Indian botanical collections
appeared in his book Nova genera et species plantarum seu
Prodromus (Swartz, 1788), a slim but nomenclaturally very
important work (Stearn, 1980; Nicolson & Jarvis, 1990).

Some of the earliest names now recognized in Sticta appear
in Swartz's Prodromus where he described 18 new taxa in the
collective genus Lichen (see Galloway, 1981, 19886), of
which L. damaecornis and L. tomentosus are referable to
Sticta as 5. damaecornis (Sw.) Ach. and 5. tomentosa (Sw.)
Ach. Subsequently, these taxa and the illegitimate Sticta
laciniata (Sw.) Ach., were widely, and often incorrectly,
reported in the literature on tropical lichens. In an attempt to
clarify the confusion which exists in the literature relating to
present distributions of these taxa, they are typified on
authentic Swartz material from Jamaica, descriptions are
given and bibliographic and taxonomic notes are supplied for
each. Original Swartz material distributed as Lichen lacinia-
tus Sw. comprises two distinct taxa; one with a green algal
photobiont which is newly described here as Sticta laciniosa
D.J. Galloway, and one with a cyanobacterial photobiont,
which is newly described as S. swartzii D.J. Galloway.

Authentic Swartz material from Jamaica and reported in
the Prodromus was studied from the following herbaria: BM,
BM-ACH, G, GB, L, PC-MONTAGNE, SBT, UPS, UPS-
ACH, UPS-THUNBERG. More recent material from

The Natural History Museum, 1994

36

D.J. GALLOWAY

Jamaica and from neotropical and palaeotropical regions was
studied from BM collections.

Scanning electron microscopy was performed on air-dried
material coated with gold-palladium on aluminium stubs,
using an Hitachi S-800 microscope. Thin-layer chromatogra-
phy of acetone extracts was carried out according to standard-
ized methods (Culberson, 1972; White & James, 1985).

SYSTEMATIC TREATMENT

1. Sticta damaecornis Ach., Meth. Lich.: 276 (1803). Lichen
damaecornis Sw., Prodr.: 146 (1788). Platisma corn-
udamae Hoffm., Descr. Adumbr. pi. lich. 1: 103, tab.

XXIV, figs 1-7 (1790). Parmelia damaecornis (Sw.)
Eschw. in Martius, Fl. Bras. 1: 213 (1833). Lobaria
damaecornis (Sw.) Trevisan, Lichenotheca veneta exs. 75
(1869). Type: Jamaica, sine loco, Swartz (SET- lectotype
selected here).
Figs 1,2.

Sticta damaecornis f. elongato-laciniata Tuck, in Stizenb.,
Flora, Jena 81: 121 (1895). [Wright, Lichenes Cubae No.
59]; nom. nud.

Thallus 4-8(-10) cm diam., possibly larger, irregularly
spreading, loosely attached, margins free, ascending.
Lobes rather narrow, (1.5-)2-4(-7) mm wide, regularly
dichotomously branching, divergent at apices, discrete, con-

Fig. 1 Lichen damaecornis Sw. Lectotype (SBT). Scale in mm.

STUDIES ON LICHEN GENUS STICTA

37

Fig. 2 Hoffmann's 1794 illustration of Platisma cornudamae. Scale in mm.

38

D.J. GALLOWAY

tiguous to somewhat entangled centrally, plane to distinctly
convex, often markedly canaliculate below, apices blunt,
rounded or shallowly furcate. Margins entire, conspicuously
thickened and ridged below, sinuses smoothly rounded.
Upper surface olivaceous, suffused brown or red-brown at
apices when wet, pale green-grey to olivaceous or buff,
suffused brownish when dry, matt, somewhat coriaceous,
minutely punctate-impressed to irregularly dimpled in parts
to uniformly smooth, pliable, flabby when wet, rather
brittle when dry, minutely maculate (x 10 lens), maculae
visible as a faint, irregular white marbling of upper surface.
Isidia, phyllidia, pseudocyphellae, and soredia absent,
continuous under SEM or with occasional, scattered pores
(Fig. 3 A). Medulla white, K-. Lower surface white or pale
tan to dark brown or brown-black, smooth to irregularly
wrinkled-ridged, glabrous or glabrous only near apices to
tomentose from apices to centre, tomentum sparse to
moderate, pale buff to brown-black, silky to felted-entangled.
Cyphellae common, scattered, conspicuous, rounded, regu-
lar, rather small, 0.3(-0.5) mm diam., margins sharply
defined, conspicuously raised above lower cortex and tomen-
tum, basal membrane white (Fig. 4 A). Thallus
120-200(-240) (Am thick. Upper cortex 20-30 |xm thick, out-
ermost 8-10 |xm pale yellow-brown, of small, closely com-
pacted cells 2-5 jjim diam., inner zone colourless, of larger,
thin- walled, round to irregular cells 6-8.5 |xm diam. Photo-
biont layer dense, continuous, 25-35 |i,m thick, photobiont
green, cells rounded, densely packed, to 4 |jim diam. Medulla
55-145 |xm thick, of loosely interwoven colourless hyphae,
4-5 (xm diam. Lower cortex 20-30 |xm thick, closely similar in
structure to upper cortex, outermost layer of cells pale
red-brown. Tomental hairs pale red-brown to yellow-brown,
to 5.5 jjun diam., and 60-170 (jun long (Fig. 5A).

Apothecia common, marginal or submarginal, rounded,
0.5-1.5 mm diam., subpedicellate, constricted at base,
pedicel 0.2-0.6 mm thick, disc matt, epruinose, shallowly
concave at first, soon becoming plane to subconvex, disc pale
to dark red-brown or blackened, margins persistent,
roughened, paler than disc, exciple below disc pale buff to
dark brownish, roughened to areolate-scabrid, sometimes
with silky whitish to red-brown tomentum. Exciple
112-190 (Jim thick, pale to dark red-brown, of parallel, radiat-
ing, round to oblong, thick-walled cells 8.5-15 |xm diam.
Hypothecium 55-90 ^m thick dark greenish- or olive-brown,
turning olive-black in K and suffusing a distinctive yellow
pigment into thecium and surrounding mounting medium.
Thecium 60-70 |xm tall, pale yellow-brown, becoming pale
yellowish to pale pinkish in K; epithecium 14-20 jxm thick,
dark brown or red-brown, intensely granular, pale red-brown
in K; paraphyses simple, 2.5-3 (xm thick, apices swollen. Asci
cylindrical 70-82(-88) x 14-17(-20) |xm. Ascospores ellip-
soid with pointed ends, pale olive-brown to colourless,
1-3-septate, 25-30.5 x 5.5-8.5 jim.

CHEMISTRY. Nil.

DISTRIBUTION. Jamaica, Cuba (Imshaug, 1957). Palaeotro-
pical and South American material referred to this name (e.g.
Nylander, 1860; Stizenberger, 1895; Zahlbruckner, 1925)
belong in other taxa, for example Sticta dichotoma for
narrow-lobed species from Indian Ocean islands, and 5.
ainoae (Galloway & Pickering, 1990) for collections from
temperate, southern South America.

TYPIFICATION. Original (syntype) Swartz material of Lichen
damaecornis is found in the following herbaria: BM-ACH
(Galloway, 1988ft), BM, G, GB, H-ACH (Vainio, 1915),
PC-MONTAGNE, SET, UPS- THUNBERG [sheet 26188].
Material from Swartz's herbarium (SET sheet 40, Fig. 1) is
selected as lectotype. This agrees with the original description
and the early fine coloured illustration (Hoffmann, 1794: tab.
24, fig. 7) showing the morphological characters which cor-
rectly define this West Indian species (Fig. 2).

OBSERVATIONS. Early accounts of Sticta damaecornis (Hoff-
mann, 1794; Acharius, 1799, 1803, 1810, 1814) follow Swartz
(1788, 1811) in giving the West Indies as habitat for the
species, and Acharius (1814) included two varieties, wiegelii
and canariensis, which are today referred to the species Sticta
wiegelii (Acharius) Vainio and S. canariensis (Florke) Delise.
The former occurs widely in both tropical and temperate
habitats (Galloway, 1994), the latter in Macaronesia, Spain,
Portugal, France, Great Britain, Ireland and Norway (Purvis
et al., 1992). Hooker (1822) recorded it from the neotropics,
while Delise (1825) cited it from America, Jamaica and
Reunion and described a similar though distinct species, 5.
dichotoma Delise, from Mauritius and Reunion. Fee (1837)
recognized 5. damaecornis as being distinct from Delise's 5.
dichotoma while in contrast, Nylander (1860), who recorded
5. damaecornis as a widespread tropical species, recognized
several varieties viz., var. sinuosa, var. macrophylla, var.
caperata and var. dichotoma. Tuckerman (1882) like
Nylander, also had a wide and obviously heterogeneous
concept of the species, including in it Swartz's Lichen lacinia-
tus.

A more restricted distribution for S. damaecornis is
accepted here (see above) with other tropical and temperate
taxa being referred to other taxa (see Galloway & Pickering,
1990). Sticta damaecornis is characterized by rather narrow,
regularly dichotmously branching lobes which are diver-
gent at the apices, noticeably thickened-ridged at the margins
below and commonly distinctly canaliculate below. It has a
white medulla; a green photobiont; sparse to moderate
tomentum on the lower surface which may be pale buff to
brown-black; conspicuous, scattered, small cyphellae with
sharply defined margins projecting above the tomentum;
apothecia are marginal or submarginal, the disc red-brown to
black, the exciple roughened to areolate-scabrid and some-
times whitish tomentose.

SPECIMENS EXAMINED. Jamaica: sine loco, Mr Wiles (BM);
Ibid., Hart, June 1886 (BM); Mt Diablo, H.N. Ridley (BM);
steep ridge on flanks of Blue Mountains in headwaters of
Mabess River, 1460 m, 16 December 1988, P.J.Bellingham
1/13: 863600 (BM) [epiphytic at c. 1 m on stem of Smilax
balbisiana under tall forest canopy]; Grand Ridge of the Blue
Mountains E. of John Crow Peak, 1600 m, 8 January 1989,
P.J.Bellingham 1/13: 856600 (BM) [on a fallen rotten branch
Mull Ridge forest]; Grand Ridge of the Blue Mountains
between Morce's Gap and John Crow Peak, 1600 m, 10
February 1989, P.J.Bellingham 1/13: s.n. (BM) [from the
trunk of Haenianthus incrassatus at 2 m height in montane
rainforest]; immediately N. of the Grand Ridge of the Blue
Mountains between John Crow Peak and Morce's Gap, 1580
m, 23 March 1989, P.J.Bellingham 1/13: 856601 (BM) [epi-
phytic at 1.5 m on trunk of Eugenia virgultosa in montane
rainforest]; steep ridge flanks at headwaters of Mabess River,
NW of Belle Vue Peak on the Grand Ridge of the Blue

STUDIES ON LICHEN GENUS STICTA

39

Fig. 3 SEM of upper cortex. A. S. damaecornis (SET). B. 5. laciniosa (BM). C. 5. swartzii (BM). D. 5. tomentosa (SET). All x 2000.

Mountains, 1710 m, 13 June 1990, P.J.Bellmgham 1/13:
867597 (BM) [epiphytic at 0.3 m on trunk of Clethra occiden-
talis in montane rainforest]; steep ridge flanks at headwaters
of Mabess River N. of the Grand Ridge of the Blue Moun-
tains between Morce's Gap and John Crow Peak, 1440 m, 23

August 1990, P.J.Bellmgham 1/13: 857602 (BM) [on the stem
of a large Marcgravia brownei at 0.5 m under tall montane
rainforest on steep, bluffed slopes]; Portland, near Hardwar
Gap, near the Portland-St Andrew Line, 17-27 December
1968, W.L. & C.F. Culberson 13,299 [A. Vezda, Lich.Sel-

40

D.J. GALLOWAY

Fig. 4 SEM of cyphellae. A. 5. damaecornis (SET) x 150. B. 5. laclniosa (BM) x 250. C. 5. swartzii (BM) x 250. D. 5. tomentosa (SET)
x 150.

.Exs. 863] (BM); Portland, Abraham's Ridge, 2000-3000 ft,
17 December 1973, B.D. Morley & C. Whitefoord 588 (BM).
Cuba: sine loco, Wright 59, 60, 61 [Lichenes Cubae] (BM);
Sierra Mae stra, cerca Pico Bayamesa, S. del poblado Pino

del Agua, 1440 m, 1 December 1978, T. Pocs 9067 (BM);
Sierra Maestra, Estribo de Turquino, 1600-1700 m, 20 April
1979, T. Pocs 9092 (BM); Sierra Maestra, 1300 m, 20 April
1979, T. Pocs 9087 (BM); Sierra de la Gran Piedra, Pico

STUDIES ON LICHEN GENUS STICTA

41

Fig. 5 SEM of tomental hairs. A. S. damaecornis (SET) x 2000. B. S. laciniosa (BM) x 2000. C. S. swartzii (BM) x 1500. D. S. tomentosa
(SET) x 2000.

Mogota, 900-1000 m, 26 May 1979, T. Pocs 9123 [A. Vezda,
Lich.Sel.Exs. 1680] (BM).

2. Sticta laciniosa D.J. Galloway, sp. nov.

Thallus viridis vel olivaceous foliaceus, laciniosus 5-8(-15)
cm latus, lacinii irregulariter divisa subdichotomae vel trunca-
tae, (2-)5-15(-30) mm latae, margis integerrimis, supra lae-
vigatae non faveolatae; medulla niveis vel dilute stramineis,

42

K + rubra; subtus dense tomentosus, cyphellis numerosis
profunde excavatis; apothecia 0.5-3(-6) mm lata, marginalia
et aliquando supra lacinias, excipulo scabrido; sporae 8:
nae, dilute olivaceus, 1-3-septatae, (22-)25-33(-36) x
5.5-8.5 u,m.

Typus: Jamaica, Parish of Portland, Grand Ridge of the Blue
Mountains between John Crow Peak and Morce's Gap,
1805'N 7639'W, c. 1610 m, on the trunk of a Lyonia
octandra (Sw.) Griseb., (Ericaceae) at 0.5 m height in
montane rainforest, 2 April 1989, P.J. Bellingham (BM-
holotype). Fig. 6.

Lichen laciniatus Sw., Prodr.: 147 (1788) nom. illegit. (Art.
64.1) [Note 1]. Platisma laciniatum (Sw.) Hoffm., Descript.
Adumbr. pi. lich. 3: 14 (1801). nomen sed non planta. Sticta
laciniata (Sw.) Ach., Meth. Lich.: 279 (1803). Lobaria lacini-
ata (Sw.) Trevisan, Lichenotheca veneta exs. 75 (1869). [Note
2]. Type: Jamaica, sine loco, Swartz (SBT-lectotype selected
here).

NOTE 1. Sticta laciniata Ach.

Lichen laciniatus (Swartz, 1788) on which Acharius's Sticta
laciniata is based is a later homonym of Lichen laciniatus
Hudson (Hudson, 1762: 449) and is accordingly illegitimate
(Art. 64.1). Material of Lichen laciniatus in Swartz's her-
barium (SET) comprises two taxa, a green photobiont species
with entire margins and a K+ red medullary reaction, which
is here described as Sticta laciniosa, and a cyanobacterial
photobiont species, with delicately phyllidiate margins which
is described below as S. swartzii. Material of this latter taxon
was discussed and figured by Hoffmann (1801) as Platysma
laciniata (Sw.) Hoffm., based on the illegitimate L. laciniatus
Sw.

NOTE 2. Lobaria laciniata (Sw.) Trevisan

Hudson's Lichen laciniatus (Hudson, 1762: 449) is an earlier,

D.J. GALLOWAY

but largely forgotten, name for Scopoli's Lichen amplissimus
(Scopoli, 1772: 386), the basionym for the well-known lichen
Lobaria amplissima (Scop.) Forssell, and is cited as a syn-
onym of Lobaria amplissima in several treatments (e.g.
Crombie, 1894; Zahlbruckner, 1925). Vainio (1899) recog-
nized this when he made the combination Lobaria laciniata
(Hudson) Vainio, but failed to recognize that his new combi-
nation was a later homonym of Lobaria laciniata (Sw.)
Trevisan (Trevisan, 1869) and therefore illegitimate. Thus,
Hudson's Lichen laciniatus becomes unavailable for use in
Lobaria and does not take precedence over Lobaria amplis-
sima.

Thallus irregularly spreading, 5-8(-15) cm diam., loosely to
closely attached from margins to centre. Lobes broadly
laciniate, subdichotomously to irregularly branched,
branches discrete and somewhat truncate at margins,
becoming subimbricate centrally (2-)5-15(-30) mm wide,
thick, coriaceous. Margins entire, unevenly sinuate or
truncate, slightly thickened below, occasionally to com-
monly furnished with projecting, short black tufts of tomen-
tum. Upper surface lettuce green to olive green, occasionally
suffused brownish at margins when wet, pale olivaceous or
pale glaucous-greyish or yellowish or brownish in parts
when dry, mainly plane or subconvex to minutely, irregularly
and shallowly pitted or wrinkled, not faveolate or punctate-
impressed, matt, without isidia, maculae, phyllidia,
pseudocyphellae or soredia, continuous under SEM, rarely
with occasional, scattered pores (Fig. 3B). Medulla whitish to
pale yellowish, K+ red. Lower surface smooth or minutely
wrinkled especially at margins, pale brown at margins, black
centrally or black and shining from margins to centre in older
lobes, glabrous in a narrow, marginal zone, uniformly
densely tomentose from margins to centre, except for young
lobe tips, tomentum dark brown to black, thick, entangled,
shaggy or woolly. Cyphellae common, scattered, round or

Fig. 6 Sticta laciniosa. Holotype (BM). Scale in mm.

STUDIES ON LICHEN GENUS ST1CTA

43

subirregular, 0.1-0.5(-1.2) mm diam., deeply excavate, often
sunk in dark tomentum, margins thin, sharply defined (Fig.
4B), concolorous with lower cortex, pit membrane yellowish
to pale ochre. Thallus 200-450(-550) jxm thick. Upper cortex
40-55 (Jirn thick, colourless, of round to irregular isodia-
metric cells, 2.5-8.5 |xm diam. Photobiont layer 40-55 |xm
thick, photobiont green, cells rounded, 3.5-5.5 Jim diam.
Medulla 150-300 |xm thick, colourless in upper parts, pale
red-brown near lower cortex, hyphae loosely interwoven to
5.5 n-m diam. Lower cortex 40-65 jxm thick, dark red-brown,
cells round to irregular, thick-walled, 2.5-11 |xm diam.
Tomental hairs 5-8.5 |xm diam., dark red-brown to 180 |xm
long, in clustered fascicles (Fig. 5B).

Apothecia prominent, mainly marginal and submarginal,
occasionally laminal, 0.5-3(-6) mm diam., subpedicellate,
round to subirregular, concave at first becoming plane or
subconvex at maturity, disc orange-brown to dark red-brown,

rarely blackened, glossy especially when young, epruinose,
smooth to matt and minutely roughened-papillate at matu-
rity. Proper exciple swollen, conspicuous, persistent,
obscuring disc when young, slightly darker than disc, con-
spicuously verrucose-areolate, margins slightly raised above
surface of disc, without projecting marginal hairs. Exciple
150-230 (jtm thick, pale red-brown in outer parts, colour-
less internally, of parallel, radiating, round to irregular
thick-walled cells 8-22 (Jim diam. Hypothecium 50-65 |xm
thick, densely interwoven, olive brownish to red-brown,
unchanged in K. Thecium 160-200 u.m tall, pale yellow-
brown to dark brown; epithecium 14-20 jxm thick, dark
brownish to red-brown, unchanged in K; paraphyses simple,
2-3 (Jim thick, apices swollen to 5 (xm diam. Asci cylindrical
to clavate-cylindrical (80-)88-106 x 16-19.5 |xm. Ascospores
elongate-ellipsoid, apices pointed, pale olive-brown, 1-3-
septate, (22-)25-33(-36) x 5.5-8.5 |xm. Pycnidia common,

Fig. 7 Lichen laciniatus Sw. Lectotype, right-hand specimen (SET); left-hand specimen is S.swartzii. Scale in mm.

44

D.J. GALLOWAY

scattered, often crowded, ostiole punctate, 0.1 mm diam.,
dark brown-black, often with noticeably swollen margin
concolorous with thallus.

CHEMISTRY. Medulla K+ red: containing a complex mixture
of terpenoids, pigments and other lichen substances sepa-
rated by TLC. The presence of acetone-soluble secondary
compounds is extremely unusual in species of Sticta, although
gyrophoric acid, congyrophoric acid and an unidentified,
fast-running compound (Rf class 7) are known from the
Asian species 5. nylanderiana Zahlbr., S. platyphylloides Nyl.
(lacking gyrophoric acid) and S. praetextata (Rasanen)
Awasthi (Joshi & Awasthi, 1982; Chen, 1993). The above
chemical pattern in S. laciniosa is presently under investiga-
tion.

DISTRIBUTION. Jamaica (all records seen except one); also in
Colombia (one nineteenth century collection only, no
recently collected material was seen).

OBSERVATIONS. Original material of Lichen laciniatus Sw. is
found in the following herbaria: H-ACH (Vainio, 1915),
SBT, UPS-ACH [S.L. 232], UPS, UPS-THUNBERG (sheet
26193 pr.p.). Material from Swartz's herbarium in Stockholm
(SBT sheet 38, right-hand specimen, Fig. 7) is selected as
lectotype since it accords with the original description, the
illustration published by Swartz (1811: pi. 7) and with the
most recent correct usages of the name (Malme, 1899;
Vainio, 1915) which refer to the characteristic K-f red
reaction of the medulla. As is mentioned above, the name
Sticta laciniata (Sw.) Ach. is illegitimate and cannot now be
used for this characteristic species which is accordingly here
described as S. laciniosa. The left-hand specimen in SBT
sheet 38 is a different species of Sticta having a cyanobacterial
photobiont, and delicate coralloid, marginal isidia. This
taxon was illustrated in Hoffmann (1801: tab. LXV, 3) and
material of it is present also in GB as Lichen laciniatus
(Arvidsson, 1989). This taxon is described below as 5.
swartzii.

Sticta laciniosa is characterized by laciniate, somewhat
truncate lobes rather variable in width with entire margins
and having a green photobiont, a mainly smooth upper
surface which is not faveolate or punctate impressed, a white
to very pale yellowish medulla (K + red), prominent, mar-
ginal and laminal ascomata, a thickly tomentose lower sur-
face, the brown-black tomentum often projecting beyond the
lobe margins, and scattered, deeply excavate cyphellae sunk
in the tomentum and having sharply defined, thin margins
and a pigmented basal membrane.

SPECIMENS EXAMINED. Jamaica: sine loco, Purdie (BM);
Ibid., Mr Wiles (BM); Ibid., June 1886, Hart (BM); immedi-
ately N. of the Grand Ridge of the Blue Mountains between
Morce's Gap and John Crow Peak, on steep slopes at the
head of the Mabess River catchment, 1580 m, 14 January
1989, P.J.Bellingham 1/13: 856601 (BM) [common over a
small damp, shaded rock face under montane rainforest];
Grand Ridge of the Blue Mountains between John Crow
Peak and Morce's Gap, 1600 m, 3 February 1989, P.J.Bel-
lingham 1/13: 855600 (BM) [on trunk of Hedyosmum arbore-
scens at 1.5 m height, under predominantly Clethra
occidentalis canopy]; Grand Ridge of the Blue Mountains
between John Crow Peak and Morce's Gap, 1600 m, 10
February 1989, P.J.Bellingham 1/13: s.n. (BM) [from trunk
of Haenianthus incrassatus at 1 m height, in montane rainfor-

est; from trunk of Hedyosmum arborescens at 2 m height, in
montane rainforest]; Grand Ridge of the Blue Mountains
between John Crow Peak and Morce's Gap, 1610 m, 2 April
1989, P.J.Bellingham 1/13: 854602 (BM) [on trunk of Lyonia
octandra at 0.5 m height in montane rainforest]; immediately
S. of the summit of Blue Mountain Peak, 2240 m, 4 Septem-
ber 1990, P.J.Bellingham 1/18: 945549 (BM) [on trunk of
Vaccinium meridionale at 1 m height in elfin forest. Colom-
bia: [as N. Granada] sine loco, Mrs Blagbourne (BM).

3. Sticta swartzii D.J. Galloway, sp. nov.

Thallus cinereus foliaceus, laciniosus, 4-7. 5 (-10) cm latus;
lacinii irregulariter divisa, 2-5(-10) mm latae, margis isidiatis
vel phyllidiatis vel ciliatis; supra nitida, laevigatae vel faveola-
tae; medulla niveis K-; photobion Nostocaceis; subtus
tomentosus, margine glabrae, cyphellae profunde excavatae;
apothecia 0.5-1.0(-1.5) mm lata, marginalia et aliquando
supra lacinias, excipulo pallido, minute scabrido; sporae 8:
nae, incolores, 1-3-septatae, ellipsoideae, apices acutae,
(28-)33-36 (-42) x 8.5-12(-14) jjun.

Typus: Jamaica, Parish of St Andrew, Grand Ridge of the
Blue Mountains, west of the summit of St John Peak, 1805'N
7639'W, c. 1910 m, epiphytic at 0.5 m on the trunk of
Eugenia alpina (Sw.) Willd. (Myrtaceae) in upper montane
rainforest, 29 April 1990, P.J. Bellingham (BM-holotype).
Fig. 8.

Thallus 4-7.5(-10) cm diam., irregularly spreading or clus-
tered in partial rosettes, loosely attached centrally, margins
free. Lobes irregularly laciniate, 2-5(-10) mm wide, subdi-
chotomously to irregularly branched, free, discrete at
margins, entangled- imbricate centrally, rather thin and
papery, fragile. Margins slightly thickened and delicately
ridged below, entire in parts and then with prominent
whitish to black or brown projecting cilia, 0.2-0.5 mm long,
to ragged, lacerate-isidiate-phyllidiate. Upper surface dark
slate blue or blue-black when wet, pale to dark grey, here and
there suffused red-brown or brown when dry, mainly plane or
subconvex, matt or shining, smooth in places or irregularly
shallowly ridged, occasionally dimpled to regularly
punctate-impressed towards margins, isidiate, maculate,
phyllidiate, without soredia or pseudocyphellae, continu-
ous under SEM (Fig. 3C). Maculae minute, white, (use x 10
lens), with scattered larger photobiont-free areas appearing
as irregular pale buff to whitish blotches. Isidia marginal,
0.5-2 mm tall, terete, fingerlike to coralloid, becoming
flattened-phyllidiate. Phyllidia marginal, 0.5-2 mm tall, con-
stricted at base, dorsiventral, lanceolate at first to ragged-
subcoralloid, best developed in older parts of thallus.
Medulla white, K-. Lower surface minutely and irregularly
wrinkled at margins, ridged to faveclate centrally, pale
whitish or greyish at margins to buff or brown centrally,
tomentose, tomentum rather variable, from thin and
arachnoid to thick and woolly, often well-developed centrally
with margins glabrous, or continuous from margins to
centre, pale buff to dark brown. Cyphellae scattered, round,
rarely elongate, 0.1-0.5(-1.00) mm diam., deeply excavate,
margins very thin, sharply defined, concolorous with lower
surface, basal membrane white or creamish, not pigmented
(Fig. 4C). Thallus 55-135(-170) |xm thick. Upper cortex
20-30(-35) (Jim thick, colourless, of 3-4 rows of round to
irregular, thick-walled cells, 5.5-16 |xm diam., cells close to

STUDIES ON LICHEN GENUS STICTA

45

iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiifiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiiii

Fig. 8 Sticta swartzil. Holotype (BM). Scale in mm.

photobiont layer much larger than cells of outermost layer.
Photobiont layer 20-45 jxm thick, dense, continuous, photo-
biont INostoc, cells rounded, 5.5-9 fjun diam., clustered in
packets. Medulla 35-70 (xm thick, almost lacking in young
lobes, hyphae colourless, loosely interwoven, 3-5 u,m diam.
Lower cortex 14-25 |xm thick, colourless, 1-2 rows of round
to irregular, thick-walled cells 5.5-22 JJLITI diam. Tomental
hairs 5.5-8.5 |xm diam., colourless, to 170 u,m long, single or
in fascicles (Fig. 5C).

Apothecia rather rare, marginal or submarginal,
0.5-1.0(-1.5) mm diam., subpedicellate, constricted at base,
round, very shallowly concave at first soon becoming plane,
disc red-brown to dark brown, epruinose, matt, slightly
roughened. Proper exciple pale pinkish brown, markedly
paler than disc, translucent when wet, minutely corrugate-
scabrid, without projecting marginal hairs. Exciple
110-225 u,m thick, pale red-brown in outermost 15-20 u,m,
remainder colourless, of parallel, radiating, round to irregu-
lar, thick- walled cells, 11-28 |xm diam., largest diameter cells
towards hypothecium. Hypothecium 55-115 u,m thick,
opaque, upper 20-35 |xm red-brown, paler in K, remainder
colourless. Thecium 135-170 (xm tall, colourless; epithecium
10-17 |xm thick, red-brown, paler in K, contiguous with
hypothecium at margins of fruit; paraphyses simple, 2-3 (Jim
thick, swollen to 5 u.m at apices. Asci clavate-cylindrical
70-110 x 16-25 |xm. Ascospores elongate-ellipsoid, apices
pointed, colourless, 1-3-septate, (28-)30-36(-42) x 8.5-12
(-14) (Jim. Pycnidia not seen.

CHEMISTRY. TLC nil.

DISTRIBUTION. Jamaica.

OBSERVATIONS. Original Swartz material from Jamaica hav-

ing a cyanobacterial photobiont, delicate marginal coralloid
isidia and a K- medulla and labelled Lichen laciniatus is
found in the following herbaria: GB, UPS-THUNBERG
(sheet 26193 pr.p.). This taxon was also illustrated in Hoff-
mann (1801: tab. LXV, 3). As indicated above (see under
discussion of 5. laciniosa) this is a good independent taxon
which is here described as 5. swartzii.

Sticta swartzii is characterized by rather narrow, irregularly
laciniate, thin, papery lobes with coralloid isidiate to phyllidi-
ate margins and often fine, projecting cilia. It has a shining
upper surface which is irregularly punctate-impressed; a
white medulla which is K-; occasional, small, marginal or
submarginal apothecia with prominent pale, glabrous mar-
gins; and a variably tomentose lower surface with scattered,
round, deeply excavate cyphellae with a white or creamish
basal membrane.

SPECIMENS EXAMINED. Jamaica: sine loco, Swartz (GB,
UPS-THUNBERG 26193 pr.p.); headwaters catchment of
the Mabess River, N. of the Grand Ridge of the Blue
Mountains, Parish of Portland, c. 1480 m, 1 May 1989,
P.J.Bellingham 1/13: 858601 (BM) [Occasional on a shaded
rock face (shale) on a steep ridge under montane rainforest] ;
Grand Ridge of the Blue Mountains W. of the summit of Sir
John Peak, Parish of St Andrew, c. 1910 m, 29 April 1990,
P.J. Bellingham 1/13: 877603 (BM) [On the trunk of Cyrilla
racemiflora in upper montane rainforest]; Grand Ridge of the
Blue Mountains immediately W. of Belle Vue Peak, Parish of
St Andrew, c. 1740 m, 12 May 1990, P.J.Bellingham 1/13:
867597 (BM) [Epiphytic at 1 m on the trunk of Cyathea
pubescens in montane rainforest]; NE flanks of Sir John
Peak, at a small headwater gully of the Spanish River, Parish
of Portland, c. 1840 m, 9 September 1990, P.J.Bellingham
1/13: 88160 (BM) [Epiphytic at 1 m height on the trunk of a

46

D.J. GALLOWAY

Cyathea pubescens in deep shade at the head of a moist gully
in montane rainforest]; steep ridge flanks at headwaters of
the Green River, W. of High Peak, Parish of St Thomas, c.
1780 m, 18 May 1991, P.J.Bdlingham 1/13: 885594 (BM)
[Epiphytic at 0.5 m height on the trunk of Podocarpus urbanii
in tall montane rainforest].

4. Sticta tomentosa (Sw.) Ach., Meth. Lich.: 279 (1803).
Lichen tomentosus Sw., Prodr.: 147 (1788). Lobaria
tomentosa (Sw.) Rauschel, Nomendat. Bot. ed 3: 330
(1797). Stictina tomentosa (Sw.) Nyl., Syn. meth. lich.
1(2): 343 (1860). Dystictina tomentosa (Sw.) Clem., Gen.

fung.: 175 (1909). Type: Jamaica, sine loco, Swartz (SBT-
lectotype selected here).
Fig. 9.

Sticta bicolor Taylor in Lond. J.Bot.6: 183 (1847). Type:
Brazil, Organ Mountains, [near summit, March 1841],
Gardner 1001 (BM- lectotype selected here).

Thallus 10-50(-85) mm diam., orbicular, rosette-forming,
loosely attached centrally, margins free. Lobes rather broad,
rounded, 5-10(-20) mm diam., discrete at margins or
shallowly imbricate, shallowly or occasionally deeply incised,

Fig. 9 Lichen tomentosus Sw. Lectotype (SET). Scale in mm.

STUDIES ON LICHEN GENUS STICTA

47

rarely from margins to centre. Margins entire, not thickened
below, wavy, slightly crisped and subascendent above, occa-
sionally to commonly with minute, silky, white, glistening,
projecting tufts of tomentum. Upper surface dark blue-black
or glaucous blue-grey when wet, pale blue-grey when dry,
smooth, undulate, occasionally but erratically punctate-
impressed, matt or rarely glossy in parts, never velvety-
pilose, tomentose or scabrid, rather thin and papery in
texture, brittle when dry, flabby, pliable when wet, maculate,
without isidia, pseudocyphellae or soredia, continuous
under SEM or with occasional scattered pores (Fig. 3D).
Maculae white, minute, scattered (x 10 lens). Medulla white,
K-. Lower surface white at margins, pale tan to brownish
centrally, glabrous in a wide marginal zone, matt, minutely
wrinkled, tomentum white, rarely pale brownish centrally,
silky, sparse to densely felted. Cyphellae common, scat-
tered, round to subirregular, 0.1-0.8(-1.5) mm diam., mar-
gins very narrow, sharply defined, rising abruptly and
vertically from lower surface (Fig. 4D), central cavity white,
deeply concave. Thallus 140-225 |xm thick. Upper cortex
14-20 (Jim thick, colourless, of two layers of thick-walled
cells, wall 2.5-3 |xm thick, uppermost layer of smaller cells
2.5-5 (Jim diam., inner cells larger, 8.5-12 (Jim diam. Photo-
biont layer continuous, 40-55 |xm thick, photobiont cyano-
bacterial INostoc, cells in clusters in a colourless gelatinous
matrix, cells 3-5.5 |xm diam. Medulla 70-130 fjim thick, of
loosely woven, colourless hyphae to 4 jim diam. Lower
cortex a single layer of thick-walled, rectangular, colourless
cells, 14-20 u,m tall and 8-11 (Jim thick, wall to 3 |xm thick.
Tomental hairs colourless, single or in clusters, to 4 |xm
diam., 35-135 u,m long (Fig. 5D).

Apothecia common, richly developed and often clustered at
margins, rare or absent centrally, rounded, sessile, con-
stricted at base to very shortly pedicellate, insertion of disc
showing a marked concavity on lower surface,
0.1-1.5(-2) mm diam., disc plane to subconvex, matt, pale to
dark red-brown when dry, epruinose, pale brown, opaque
when wet. Proper exciple persistent when dry, excluded when
wet, white to pale buff or creamish, smooth to minutely
crenulate with occasional projecting silky white hairs at
margins of disc, especially noticeable in young fruits. Exciple
55-125 |xm thick, outermost 30 |xm dilute yellow-brown,
remainder colourless, of round to irregular, isodiametric
cells 8-22 |xm diam. Hypothecium 40-55 (Jim thick, densely
interwoven, upper 22-28 u,m pale red-brown, remainder
colourless, unchanged in K. Thecium 80-90 (Jtm tall, colour-
less; epithecium 5-9(-13) |xm thick, pale olive-brown to red-
brown, unchanged in K; paraphyses distinctly septate,
constricted at septa and appearing long-moniliform,
3.5-8 |xm diam., swollen and tinged olive-brown at apices.
Asci cylindrical, (72-)80-92 x 11-17 |jun. Ascospores colour-
less, long-ellipsoid, apices pointed, straight or curved,
3-septate, 27.5-33.5(- 36) x 5.5-8.5 jxm.

CHEMISTRY. Nil.

DISTRIBUTION. Neotropics and palaeotropics. Jamaica
(Imshaug, 1957), Mexico (Imshaug, 1956ft), Panama
(Imshaug 1956a), Colombia, Venezuela, Peru, Brazil
(Malme, 1934). East Africa including Kenya, Tanzania,
Uganda (Swinscow & Krog, 1988), St Helena (Leighton,
1871), South Africa, Madagascar, Hawaii (Magnusson &
Zahlbruckner, 1943; Magnusson, 1956).

TYPIFICATION. Original (syntype) Swartz material of Lichen
tomentosus is found in the following herbaria: G, GB, L
[910,213-1824], SBT, UPS, UPS-THUNBERG [sheet
26202]. Material from Swartz's own herbarium (SBT sheet
44) is selected as lectotype (Fig. 9) since it accords with the
original description and closely resembles the coloured illus-
tration later provided by Swartz (1811: pi. IX). Material in
BM-ACH labelled Sticta tomentosa by Acharius (Galloway,
19886) is a mixture of two taxa neither of which appears to be
Sticta tomentosa.

OBSERVATIONS. In commenting on his species Sticta bicolor,
Thomas Taylor (1847) gives the following notes which are a
good description of the morphology of 5. tomentosa;
'. . .Thallus 4 inches wide, lobes scarcely one quarter of an
inch broad, the central parts of an ash-grey, the extreme of a
chestnut brown, but little deepened by moisture. The thick
dark grey scabrous pubescence of the inferior surface of the
thallus reappears on the backs of the apothecia. The smooth
surface of the thallus and the crowded marginal sessile
apothecia readily distinguish this species from S. sylvatica
Ach.'.

SPECIMENS EXAMINED. Jamaica: sine loco, Mr Wiles (BM);
sine loco, on branches and twigs of trees, Feb.-March 1905,
Miss C.E. Cummings [Lich. Exs., G.K.Merrill 193] (BM);
headwaters of catchment of the Mabess River, N. of the
Grand Ridge of the Blue Mountains, 1340 m, 8 May 1989,
P.J.Bellingham 1/13: 860602 (BM) [from trunk of Myrcian-
thes fragrans on a rock bluff in montane rainforest]; steep
slopes at the headwaters of the Mabess River N. of the Grand
Ridge of the Blue Mountains between Morce's Gap and John
Crow Peak, 1420 m, 5 July 1990, P.J.Bellingham 1/13: 859601
(BM) [Epiphytic at 0.5 m height on stems of Picea weddellii in
a steep rubbly channel at the edge of an old landslide]; steep
ridge at headwaters of Mabess River, N. of the Grand Ridge
of the Blue Mountains between Morce's Gap and John Crow
Peak, 1440 m, 23 August 1990, P.J.Bellingham 1/13: 857602
(BM). Mexico: La Cima, 3050 m, 14 July 1908, C.G.Pringle S
19,184 (BM); Amecameca, 14 September 1908, C.G.Pringle
S 19,225 (BM); Hills Patzcuaro, November 1891, C.G.Prin-
gle S 22,511 (BM). Panama: Chiriqui, between Los Planes de
Hornito and Fortuna Lake. Trail to Zarzo, 1200 m, 8 March

1985, R.Hampshire & C.Whitefoord 335 (BM). Colombia:
Rio Magdalena, Mr J. Weir (BM); Cali-Dagua Road, after
Bitaco turning, 1000 m, 17 December 1967, R.M. Garrett 36
(BM); sine loco., ex Herb. Lindig 2521 (BM). Venezuela:
Sierra de Sto Domingo, 1 August 1958, R.W.G. Dennis
1935b (BM). Peru: In declivibus Andium peruvianarum. pr.
Sachapata, Sept. 1854, W.Lechler 3124 (BM). Brazil: Organ
Mountains, Gardner (BM); sine loco, Mr Weir 61 (BM). St
Helena: High Peak National Forest, 600 m, 17 December

1986, A. B. Barlow (BM). Tanzania: Arusha Distr., Mt Meru,
south side, 2000 m, February 1974, T.D.V. Swinscow T16/5
(BM). Kenya: Meru Distr., Mt Kenya, east side, Themwe,
2100 m, February 1974, T.D.V. Swinscow 3K 16/8 (BM); Mt
Kenya, 2 km NW of Irangi Forest Station, 2000 m, February
1974, T.D.V. Swinscow K48/15 (BM); Mt Kenya, near Castle
Forest Station, 1900 m, February 1974, T.D.V. Swinscow K
49/6 (BM). Uganda: Kigezi District, Kinkizi County, 1600 m,
December 1971, T.D.V. Swinscow 3U 56/4 (BM). South
Africa: Cape Province [Kaffraria], Barziya (?), Rev. R. Baur
(BM). Madagascar: sine loco, Barron (BM). Hawaii: Mauii,
Puu Kukui, Mount Kaulawelewele, 295 m, 25 April 1970,

48

D.J. GALLOWAY

A.C. Jermy s.n. (BM) [on soft sandstones or as epiphytes in
Metrosideros rainforest with Pomantia arborea as a dominant
shrub].

ACKNOWLEDGEMENTS. I am grateful to Dr Lars Arvidsson (Gote-
borg), Mr Lars E. Kers and Prof. Bengt Jonsell (Bergianska
Tradgarden, Stockholm) and Dr Roland Moberg (Uppsala) for the
loan of original Swartz material; to Dr Peter Bellingham (Botany
School, Cambridge University) for access to his lichen collections
from the Blue Mountains, Jamaica; and to Mr Phil Crabbe, Phil
Hurst and Pat Hart (BM) for expert photographic assistance. I thank
my colleague Prof. Per Magnus J0rgensen (Bergen) for his helpful
discussions and advice on nomenclatural matters, and assistance with
the Latin diagnoses.

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Bull. not. Hist. Mus. Land. (Bot.) 24(1): 49-90

Issued 23 June 1994

Seaweeds of the western coast of tropical
Africa and adjacent islands: a critical
assessment. IV. Rhodophyta (Florideae) 4.
Genera L-O

DAVID M. JOHN, GEORGE W. LAWSON, JAMES H. PRICE

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD

WILLEM F. PRUD'HOMME VAN REINE

Research Institute Rijks herbarium! Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The
Netherlands

WILLIAM J. WOELKERLING

School of Botany, La Trobe University, Bundoora, Victoria 3083, Australia

CONTENTS

Introduction 49

Species list 51

Numerical list of references 80

References . . 82

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 coast 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 recombination is made, Nothogenia magnified (Pilger) J.H. Price.

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, 1988, 1992; 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 L-O
and constituent species are listed in alphabetical order.

Each main entry consists of:

(i) The major bold heading, representing the currently-
accepted name and authorities.

(ii) Subsidiary italicized 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 in 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 taxon level; only when clarification was
required for comprehension have changes been made in
subhead 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 generalized 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, generalized distri-
bution statements are included verbatim since it is not always
clear for precisely which countries within the area they
establish records.

References are presented in two ways:
(a) as a numbered list (p. 80) to allow inclusion of manuscript
and expedition sources, and (b) as a list of full references in
alphabetical order. Numbers within parentheses after the
geographical names refer to corresponding numbers in the
references. References relevant only in previous parts of this
series are omitted here and numbers are therefore not fully
standardized between the present and past parts.

The Natural History Museum, 1994

50

D.M. JOHNETAL

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,*
Bioko [=Macias Nguema Biyogo, Fernando Poo]; 18, Principe; 19, Sao Tome; 20,* 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, Annobon [=Pagalu]. 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.

* Nos 17 (Bioko) and 20 (Spanish Guinea, = Rio Muni) on the original map (part I) are now jointly administered as Equatorial Guinea.
Bioko is 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 (FLORIDEAE) OF TROPICAL AFRICA

51

(iv) Additional qualifying notes appear below whole entries or
individual parts of entries to which they specifically refer. In
these notes, references containing species records consist of
authors' names, followed by the reference number in the
terminal list and, where appropriate, the relevant page num-
bers after a colon. Other references 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 italicized headings and
any other discarded combinations that require reference are
included as cross-referencing entries to the currently-
accepted names in the overall list. Without extended field and
herbarium studies, the treatments presented here are essen-
tially preliminary. Critical updating of the overall text is kept
firmly in mind for the whole series and we would appreciate
notification of any detected errors and omissions from any of
the parts.

SPECIES LIST

Lasiothalia sp.

Cote d'lvoire (288).
Liberia (129; 350; 586).

Note. Very tentative identification, with reservations, by Dr E.
Wollaston of specimens from Harper, Cape Palmas, Liberia; not
identifiable to species. A small epiphyte possibly easily overlooked or
mistaken for Callithamnion, which it closely resembles (129). A few
filaments have also been collected at nearby Tabou in Cote d'lvoire.
Probably more widespread than suggested by records.

Laurencia

For a set of treatments of certain specific groups and subge-
neric relationships across Laurencia from the western and
central Pacific through the Pacific USA coasts and over to
UK, see Saito (1982). Early studies by the same author exist
on the genus from Japan (Saito, 1967); Hawaii, the Philip-
pines and adjacent areas (Saito, 1969a); Pacific North
America (Saito, 1969ft); and southern Australia (Saito &
Womersley, 1974). Saito (1982: 306) believes that '. . . we
should not rearrange the species of Laurencia until the
characteristics of many other species from other areas of the
world have been clarified'. Work on the taxonomy of this
genus in the Canary Islands, through DNA studies, is in
progress (Gil-Rodriguez and others). See also the short
review by Gil-Rodriguez & Haroun (658).

Laurencia brachyclados Pilger

Annobon (139 ;350 ;456 ;457 ;496 ;535 ;563 ;586) .
Ascension (475).

Note. Steentoft (535) commented: '. . . specimen of L. brachycla-
dos Pilger (Mildbraed 6719 from Annobon) in Hb B0rgesen would
seem to represent the basal parts of a young plant of [L. obtusa] var.
rigidula, but it is so minute, and at such variance with the published
description (Pilger, 1920: 6) that it would seem better to leave it
alone for the time being'. Mildbraed specimen 6719 bears little
resemblance to Pilger's (456) description. Steentoft (535) also sug-
gested that L. brachycladus might possibly be a dwarf or immature
form of L. perforata, based solely on its description. Examination of
the isotype in Herb. Agardh (No. 36616) seemed to confirm this

assertion, but Yamada (563), examining the L. brachyclados holo-
type, stated that it was not L. perforata. Much earlier De Toni (139:
371) had commented: 'Mea sententia haec species videtur forsan cum
Laurencia perforata Mont, comparanda' (This species perhaps com-
pares with Laurencia perforata Mont.). Cribb (112: 166-7) consid-
ered that L. brachyclados and L. pygmaea Weber-van Bosse '. . .
may be the same species' whereas as Pilger (456: 6) commented [L.
brachyclados]: 'Die neue Art gehort in die Verwandschaft van L.
scoparia J. Ag. [q.v.]; sie ist durch ihr sprossform ausgezeichnet'.

Laurencia brongniartii J. Agardh

Canaries (658).

Ghana (299;350;376;377;586;590).

'. . . Probably widespread in tropical seas. . .' (350;586).

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

[As Fucus pinnatifidus Linnaeus]

Ghana (271).

Note. The tentative attribution of Hornemann's (271) record
given above for Ghana is explained under the entry for Laurencia
pinnatifida, which L. brongniartii superficially resembles. See also
entry for Laurencia concinna Montagne. It has been suggested by
John & Lawson (590) that this species may sometimes be mistaken
for Laurencia pinnatifida or L. undulata Yamada, so reports of these
species from Sao Tome (93) and Senegal (122) respectively may
require reinvestigation.

Laurencia caespitosa Lamouroux

See Laurencia hybrida (DeCandolle) Lenormand ex Duby
and Laurencia canariensis Montagne in Kiitzing.

Laurencia canariensis Montagne in Kiitzing

Canaries (25;26;27;109;133;318;323;407).
'warm Atlantic. . .'(410).
[As Laurencia caespitosa Lamouroux]
Canaries (401; 407).

Note. Cotton's (109) text indicated that this is a replacement
name for material that, in 1841, Montagne had considered as
Laurencia caespitosa Lamouroux. The latter is normally considered
to be a synonym, if correctly used, of L. hybrida (q.v.). This calls into
question the status of this taxon in that records need careful analysis
against L. hybrida. B0rgesen (71: 68-69) and Dangeard (119: 182)
very firmly placed L. canariensis Montagne in Kiitzing in synonymy
with L. hybrida. In 1841 Montagne (401) had commented under his
Laurencia caespitosa entry: '. . . Je doute beaucoup de la legitimite
specifique de cette Algue, que la plupart des phycologues reunissent
peut-etre avec raison a la precedente. Nos echantillons sont assez
fidelement represented dans la figure citee de Gmelin.' 'la prece-
dente' in this case was Laurencia pinnatifida Lamouroux. J. Agardh
(25: 769; 26: 769) indicated that Montagne (401: 154) had originally
considered his species as conspecific with Laurencia hybrida but later
came to believe them to be separate. This depends on the conspeci-
fity or separateness of Laurencia caespitosa and L. hybrida. In 1876
J. Agardh (27: 662) placed this taxon in his 'Species inquirendae'.

Laurencia chondrioides B0rgesen

Canaries (598;663).

[As Chondriopsis dasyphylla Woodward]

Canaries (439).

Note. The placement of Chondriopsis dasyphylla under Chondria
dasyphylla in Price et al. (1986) is incorrect.

Laurencia complanata (Suhr) Kiitzing
See notes to Laurencia concinna Montagne.

52

D.M. JOHNETAL

Laurencia concinna Montagne

Note. The question of whether Laurencia brongniartii J. Agardh
should be recorded under that name or L. concinna remains open
although arbitrarily decided as the former. Yamada (563) considered
Laurencia concinna Montagne as synonymous with Laurencia
brongniartii J. Agardh. Papenfuss (1943: 91), by contrast, considered
L. concinna to be different from L. brongniartii and L. complanata
(Suhr) Kutzing. Cribb (113: 114-5) followed Yamada (563) in
synonymizing the two taxa; he used the name L. brongniartii J.
Agardh (1841) which seems to antedate L. concinna. Earlier, Cribb
(112: 162-163) had followed Papenfuss (1943: 91) in maintaining the
taxa as distinct since the type specimen of L. brongniartii '. . . is
hardly complanate in the dried condition. . .'. Wynne (1986a) in his
tropical/subtropical western Atlantic check-list, has accepted L.
concinna as synonymous with L. brongniartii.

Laurencia corallopsis (Montagne) Howe

Canaries (633;658).

Cape Verde Islands (Leiden Herbarium, det. M.C. Gil-
Rodriguez and R.J. Haroun).
Salvage Islands (R.J. Haroun, in litt. 1990).
[As L. corrallopsis Howe]
Canaries (686).

[As Laurencia (grex) corallopsis (Montagne) Howe]
Canaries (634).

Laurencia cruciata Harvey

St. Helena (142;260;391;655).

Note. Dickie (142) regarded this species as very close to Lauren-
cia obtusata, and this comment is repeated by Mellis (391) and
Hemsley (260).

Laurencia densa (P. Dangeard) J. Feldmann

See Chondria densa P. Dangeard and Laurencia microcladia
Kutzing.

Laurencia elata (C. Agardh) Harvey
See notes to Laurencia flexuosa Kutzing.

Laurencia filiformis (C. Agardh) Montagne
See note under Laurencia scoparia J. Agardh.

Laurencia flexilis Setchell

Canaries (658).

Salvage Islands (Leiden Herbarium, det. M.C. Gil-Rodriguez
and R.J. Haroun).

Laurencia flexuosa Kutzing

[As Laurencia flexuosa J. Agardh]
Mauritanie (349;516).

Note. Jaasund (279: 62), under the name Laurencia elata (C.
Agardh) Harvey, indicated that Yamada (563: 241, pi. 26, 27)
included L. flexuosa Kutzing and L. luxurians (Harvey) J. Agardh
within L. elata, the distribution for which was cited as Australia,
Tasmania and South Africa, Jaasund then adding East Africa (Tan-
zania).

Laurencia galtsoffii Howe

Gabon (294;350;586).

Ghana (288;350;586;590).

Liberia (129;288;350;586).

'. . .in tropical parts of the Atlantic and Pacific Oceans. . .'

(350;586).

Tropical Africa (N. Gambia - Congo river)' (598).
[As Laurencia galstoffi Howe]
Gabon (294).

[As Laurencia cf. galstoffii Howe]
Cape Verde Islands (652).

Laurencia hybrida (DeCandolle) Lenormand ex Duby

Canaries (8;16;33;38B;38D;71;118;128A;191;227;229; 230;
237;263;375;489;517;598;633;634;658).

Cape Verde Islands (38B;38D;598).

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

'. . . Atlantico, desde el sur de Inglaterra a Canarias' (517).

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

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

[As Laurencia caespitosa Lamouroux]

Canaries (3;38;44;221;254;305;401).

Cape Verde Islands (38;408;596).

'. . . De 1'Angleterre aux Canaries' (38).

'. . . D'Angleterre aux Canaries' (89).

[As Laurencia caespitosa Lamouroux var. subsimplex Mon-
tagne]

Cape Verde Islands (38;408;597).

[As Laurencia hybrida DeCandolle forma]

Canaries (387).
Note. B0rgesen (71: 68-69) very firmly placed Canaries Laurencia

caespitosa and L. canariensis in synonymy with L. hybrida. See also

the entry for Laurencia canariensis Montagne in Kutzing.

Laurencia intermedia Yamada

Cape Verde Islands (652;683).

Cote d'lvoire (350;586).

Ghana (295;350;586;590;695).

Liberia (129;295;350;586).

'. . . does not extend from the Gulf of Guinea into Senegal'

(487).
'. . . in tropical parts of the Atlantic and Pacific Oceans'

(350;586).
'. . . probably widespread in many warm temperate and

tropical seas' (590).

'. . . Tropical Africa (N. Gambia- Congo river)' (598).
[As Laurencia papillosa (Forsskal) Greville]
Ghana (153;338;537).
'warm Atlantic' (78).

Note. For clinal morphological variation between L. papillosa<
>L. intermedia< >L. paniculata, see the notes to L. paniculata.
Laurencia tropica Yamada, L. flexilis Setchell and (so far as treat-
ment of some areas of the Indian Ocean are concerned) L. intermedia
Yamada are very similar. Yamada (563: 234), quoted also in Jaasund
(279: 61), commented that more specimens becoming available from
different localities could well result in his L. tropica being reduced to
synonymy with L. flexilis Setchell. Tanzanian plants clearly identifi-
able with B0rgesen's Indian Ocean material that he described as L.
flexilis, correspond to the detailed description provided by Saito
(1967: 39-45) for L. intermedia Yamada. In the end, Jaasund (279:
62) remained uncertain about the conspecificity of Tanzanian (and
Indian Ocean) L. flexilis and L. intermedia, whilst using the latter

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

53

name and generally being inclined to believe that only one taxon was
involved.

Laurencia implicata J. Agardh

Canaries (Leiden Herbarium, collected during trip of Helgo-
land Research Vessel 'Heincke').

[As L. intricata Lamouroux]

Cape Verde Islands (652).

Sao Tome (350;535;586).

Senegal (5 1;59).

?Sierra Leone (30;350;586).

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

'. . . Subtropical Africa [Senegal (N. of Gambia); Maurita-
nia; Former W. Sahara]. . .' (598).

'. . . Tropical Africa (N. Gambia - Congo river)' (598).
Note. See notes to Laurencia majuscula and L. obtusa. Laurencia

implicata (as L. intricata) has been sometimes considered to be a

variety of L. obtusa (e.g. by Yamada, 563). Lawson & John (350:

586) suggest that Aleem's report of this species from Sierra Leone,

growing on wave exposed shores as 'tufty cushions' with Centroceras

and Gelidium, may have been a misidentification for L. tenera. Often

referred to as L. intricata, but the correct name is L. implicata

according to Silva et al. (1987).

Laurencia intricata Lamouroux
See L. implicata J. Agardh.

Laurencia lata Howe & Taylor

Senegal (59;399).

'Subtropical Africa [Senegal < > of Gambia); Mauritania;

Former W. Sahara]' (598).

Note. Bodard & Mollion (59) referred certain of their specimens
with dorsiventral symmetry to this species.

Laurencia luxurians (Harvey) J. Agardh
See the notes to Laurencia flexuosa Kiitzing.

Laurencia majuscula (Harvey) Lucas

Canaries (633;647;658;686).

Cameroun (350;586).

Cape Verde Islands (652;683).

Gabon (294;350;586).

Gambia (296;350;586).

Ghana (178;299;300;350;376;377;586;590;654;695).

'. . . Probably pan tropical. . .' (350;586).

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

Mauritanie (Leiden Herbarium, det. M.C. Gil-Rodriguez

and R.J. Haroun).
Salvage Islands (Leiden Herbarium, det. M.C. Gil-Rodriguez

and R.J. Haroun).

Note. This plant was once (257) considered to be a variety of
Laurencia obtusa, since the two are almost identical anatomically.
Morphologically it resembles L. implicata but is readily separated by
its palisade-like cortical cells.

Laurencia microcladia Kiitzing

Canaries (686).
Cape Verde Islands (686).

Mauritanie (Leiden Herbarium, det. M.C. Gil-Rodriguez
and R.J. Haroun).

Senegal (54;59;529).

'Atlantique tropicale' (529).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara] (598).
'warmer parts of Atlantic Ocean' (97).
[As L. obtusa (Hudson) Lamouroux]
Canaries (439 pro parte).

Note. See the entry for Chondria densa P. Dangeard, where
additional information is presented. Bodard (54), when referring to
L. microcladia, placed in brackets '( = L. densa J. Feldmann,
= Chondria densa Dangeard)' implying that these entities are conspe-
cific. No further information is given to justify the implication.
Prud'homme van Reine et al. (663) have re-investigated Piccone's
material (439) and considered that some plants referred to him as L.
obtusa should be more correctly attributed to L. microcladia.

Laurencia minuta sine auctorum

Canaries (647).
Note. Cited without authority in a comparative table.

Laurencia natalensis Kylin

See notes on Laurencia obtusa (Hudson) Lamouroux.

Laurencia nidifica J. Agardh

Cape Verde Islands (652).

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

Ghana (288;350;586;590).

Liberia (129;295;350;586).

St. Helena (655).

'. . .in pluribus calidioribus oceanis formae steriles consimi-

les adsunt, quae an invicem specie differant parum

constat. . .' (27).
'. . . probably widespread in warm temperate and tropical

seas. . .' (350;586).
'. . . Tropical Africa (N. Gambia- Congo river). . .' (598).

Note. This taxon has always presented difficulties of determina-
tion due, in part, to sparse material. Doubts recorded in many of the
above references relate to that.

Laurencia obtusa (Hudson) Lamouroux

Annobon (456;457;535).
Cameroun (337;350;535;537;586).
Canaries

(8;16;38B;38C;38D;71;128A;191;216;226;227;237;252;

379 ;392 ;401 ;439 ;489 ;490 ;499 ;5 17 ;535 ;546;547 ;555 ;556 ;

557A;584;598;633;634;647;658;662;668;684).
Cape Verde Islands

(38B;38C;38D;252;408;499;535;555;556;598).
Gambia (296;350;586).
Ghana (BM Herbarium, Foote 1949).
Mauritanie (38B;38C;38D;252;349;535;555;556).
Principe (93;350;535;586).
Salvage Islands

(38B;38C;38D;215;216;231;375;555;556;598).
Sao Tome (93;251;265;350;535;586).
Senegal (38B;38C;38D;122;529;535;542;555;556).
Sierra Leone (30;350;586).

'African and American coasts; Canary Islands. . .' (177).
'. . . an den atlantischen Kiisten von Grossbritannien bis zu

den Canarischen Inseln. . .' (501).
'. . . Atlantic coasts from Britain to the Canary Islands. . .'

(269).

54

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

'. . . Atlantique tropical et tempere' (542).

'. . . In mari atlantico et ejus sinubus ab ins. britannicis usque

ad Brasiliam et Cap. Bon. Spei. . .' (318).
'. . .in oceano atlantico ... an ubique eadem?' (27).
'Nordwestafrika' (499).
'Pantropical' (529).

'. . . toutes les mers chaudes. . .' (190).
'. . . Subtropical Africa [Senegal (N. of Gambia); Maurita-
nia; Former W. Sahara]. . .' (598).

'. . . Tropical Africa (N. Gambia- Congo river). . .' (598).
'warmer parts of Atlantic Ocean' (375).
'Warmer parts of the Atlantic Ocean. . .' (62;71).
'Westafrika' (499).
'. . . widespread from boreal-antiboreal to tropical seas'

(350;586).

[As Laurencia obtusa Lamouroux]
Canaries (38;44;89;439;547).
Cape Verde Islands (38;145;528).
'. . . Atlantic [Ocean] . . . north and south . . . temperate

and tropical latitudes. . .' (254).
'. . . De la Grande Bretagne aux Canaries. . .' (38;89).
[As Laurencia obtusa Hudson]
Cape Verde Islands (150).
Sao Tome (263;264).

'Atlantic and Pacific, temperate and subtropical' (143).
'Warmer parts of the Atlantic' (144).
[As Laurencia obtusa (Hudson) Lamouroux var. gracilis

Harvey]
Canaries (242).

[As Laurencia obtusa Lamouroux var. gracilis Kiitzing]
Canaries (439).
[As Laurencia obtusa (Hudson) Lamouroux var. natalensis

(Kylin)]

Sao Tome (535).
[As Laurencia obtusa (Hudson) Lamouroux var. rigidula

Grunow]
Annobon (535).
Sao Tome (535).
[As Laurencia obtusa (Hudson) Lamouroux var. gelatinosa

B0rgesen]
Canaries (71).

[As Laurencia hybrida (De Candolle) Lenormand ex Duby]
Canaries (439 pro parte).

Note. Prud'homme van Reine et al. (663) have re-investigated
Piccone's material (439) and considerd that some plants referred to
him as L. hybrida should more correctly be attributed to L. obtusa.
[As Laurencia papillosa Forsskal and var. gracilis Kiitzing]
Canaries (439).

Note. See note for Laurencia papillosa. [As Laurencia grex.
[presumably 'prox.'] obtusa]
Canaries (232B).

Note. For comment on the status of varieties sometimes recog-
nized in L. obtusa and on the features distinguishing L. obtusa/L.
implicata (the latter considered a variety of the former by Yamada,
563), see Lawson & John, 350: 340). See also the note under
Laurencia majuscula. According to Steentoft (535), the Sao Tome
plants are close to two varieties - var. natalensis (Kylin) B0rgesen
which is more delicate, smaller, and more irregularly branched
than the type, and var. rigidula Grunow, also smaller, more rigid,
with more dense erect branches than the type. These minor
differences are not a good basis for varietal recognition where
morphological plasticity is high. Piccone (439: 45) indicated that
he had material of this very polymorphic species that was very
similar to varieties recognized under the names of gracilis and

D.M. JOHNETAL

gelatinosa. See notes under Laurencia papillosa, L. brachyclados
and L. viridis for probable records of L. obtusa.

Laurencia paniculata sine auctorum

See under L. patentiramea (Montagne) Kiitzing.

Laurencia papillosa (C. Agardh) Greville

[As L. papillosa (Forsskal) Greville]

Ascension Island (474;475).

Annobon (456;457).

Cameroun(337;350;454;484;500;586).

Canaries (128A;227;584;658).

Cape Verde Islands (38;150;191;598;683;686).

Mauritanie (624).

'. . . in oceano Atlantico ad littora calidiora Africae. . .'

(133).

'. . .in oceano atlantico calidiori ad littora Africae. . .' (26).
'. . . toutes les mers tropicales' (190).
'. . . Tropical Africa (N.Gambia- Congo river). . .'(598).
'Warmer parts of the Atlantic Ocean. . .' (62).
[As Laurencia papillosa J. Agardh var. thyrsoides]
Cape Verde Islands (38).

[As Laurencia papillosa Forsskal var. thyrsoides}
Cape Verde Islands (150).
[As Laurencia papillosa (Forsskal) J. Agardh]
'. . .in caldiore atlantico. . .' (27).
[As Laurencia papillosa Greville]
'Warm Atlantic' (410).

Note. Yamada (563), who actually described Laurencia interme-
dia, regarded L. papillosa and L. paniculata as probably representing
'. . . the extreme forms of one very variable species in which L.
intermedia may be included. . .'. Piccone (439: 44-45) indicated that
he had only one very small specimen from the Canaries. B0rgesen
reported (71: 68) that Forti had allowed him to see the small
fragment, stating '. . . It was so small that I did not feel inclined to
make an anatomical examination of it, but according to its appear-
ance and colour, more reddish than L. papillosa, it can most
probably be referred to Laurencia obtusa'' . The Mauritanian record
(624) is given with a query. See also under L. obtusa.

Laurencia patentiramea (Montagne) Kiitzing

[As L. paniculata (C. Agardh) Kiitzing]

Canaries (38C;38D;598).

Cape Verde Islands (38C;38D;44;598).

Senegal (37;450;451).

[As Laurencia paniculata (C. Agardh) J. Agardh]

Canaries (658).

Senegal (529).

[As Laurencia paniculata J. Agardh]

Mauritanie (624).

Note. Audiffred (38C) indicated his record as new to the Canary
Islands. Lawson & John (350, 586) repeated Yamada's (563) state-
ment of opinion that L. patentiramea (as L. papillosa) and 'L.
paniculata J. Agardh' may represent '. . . the extreme forms of one
very variable species in which L. intermedia may be included'.
Askenasy (37: 47) indicated in a footnote that the specimens of
Laurencia were very fragmentary and that Bornet, who had deter-
mined them, had given most with doubt. Given as Laurencia
paniculata! (Agardh) Kiitzing by Piccone (450, 451), the Piccone
(451) record was based on the Naumann (Gazelle) collection deter-
mined by Askenasy. See also the 'Laurencia sp. A' from Senegal
recorded by Sourie (529) and also note under Laurencia papillosa.
For the correct name of this taxon, see Silva et al. (1987: 67, 68).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA
Laurencia perforata Montagne

See under Laurencia tenerrima (Clemente) Cremades &
Perez-Cirera.

Laurencia pinnatifida (Hudson) Lamouroux

Annobon (456).

Canaries (13;38D;226;227;229;230;232B;237;252;253-

375;379;401;489;517;584;598;633;658).
Cape Verde Islands (Leiden Herbarium, det. M.C. Gil-
Rodriguez and R.J. Haroun).
?Ghana (350;586).

Mauritanie(38D;252;253;344;349;567).
Salvage Islands (38B;38D;231;375;598).
Sao Tome (93;251;295;350;586;590).
Senegal (38D;530?).
Western Sahara (38D;349).

'. . . Atlantico Oriental (Inglaterra- Mauritania). . .' (253).
'. . . Atlantique (de Norvege a la Mauritanie). . .' (33).
'. . . Norwegen bis Mauritanien' (567).
'. . . widespread in boreal-antiboreal seas and less common in

tropical seas. . .' (350;586).
'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Tropical Africa (N. Gambia - Congo River)' (598).
[As Laurencia pinnatifida Lamouroux]
Canaries (44;214;268).
SaoTome(261;263).
'. . . Atlantique depuis les cotes anglaises jusqu'en

Mauritanie. . .' (222).
'. . . west coast of Africa. . .' (268).
[As Laurencia pinnatifida (Gmelin) Lamouroux]
Annobon (457;535).

Canaries (16;42;71;191;230;236;387;392;499;535;546;556).
Mauritanie (122;529;535;556).
Salvage Islands (38B;556;556A).
Sao Tome (93;535).
Senegal (529;535;556).
Western Sahara (556).
'Afrikanischen Kiiste von Marokko und den atlantischen

Inseln bis zum Kap' (239).

'. . . Faeroes southwards to the Canary Islands. . .' (71).
'. . . most eastern Atlantic coasts south to Senegal' (535).
'Nordwestafrika' (499).
'Westafrika' (499).

[As Laurencia pinnatifida (Turner) Lamouroux]
'. . . De la Grande-Bretagne aux Canaries. . .' (89).
[As Fucus pinnatifidus Linnaeus]
Ghana (271).

Note. Sao Tome plants were regarded with some doubt by
Lawson & John (350, 586) since dredged from llm; elsewhere this
species is intertidal or shallow subtidal. Records are not confirmable
since neither the Newton (1881) plant (see 535) nor the Carpine (93)
plant (Huve, pers. comm. to DMJ) are traceable. The plant superfi-
cially resembles L. brongniartii and might be readly mistaken for it.
Sourie (529) expressed doubt, noting Dangeard's opinion that Dakar
plants approached more closely L. undulata (q.v.). The nomencla-
tural equivalence of Fucus pinnatifidus and Laurencia pinnatifida
may be in error, although L. pinnatifida is found elsewhere in the
check-list area. If drift material was involved in the original P.E. Isert
collections, the flattened species L. brongniartii, frequent in deep
waters off the Ghana coast, may have been the plant in question. The
original Isert specimens, if not destroyed in the 1807 fire at Copen-
hagen Herbarium (C), require examination for correct attribution.
This need is emphasized by the failure to re-record specimens during

55

the intensive work of recent decades. The record for Senegal (530) is
given with a query.

Laurencia platycephala Kiitzing
Canaries (634?;658).

Laurencia poiteaui (Lamouroux) Howe

[As L. papillosa (Forsskal) Greville]

Canaries (439).

[As L. pinnatifida (Hudson) Lamouroux]

Canaries (439).

[As L. poitei (Lamouroux) Howe]

Sao Tome (350;535;586).

Senegal (59).

'. . . probably widespread in warm temperate and tropical

seas. . .' (350;586).
'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Tropical Africa (N. Gambia - Congo River)' (598).
[As Laurencia tuberculosa Agardh]
Sao Tome (251 ;265).
[As Gracilaria poitei Lamouroux]
SaoTome(251;263;264).
[As Gracilaria poitei (Lamouroux) Agardh.]
Sao Tome (265).

Note. Prud'homme van Reine et al. (663) have re-investigated
Piccone's material (439) and consider that plants referred to as L.
pinnatifida and L. papillosa are correctly attributed to L. poiteaui.
[As L. paniculata (C. Agardh) Kiitzing]
Canaries (38C;38D;598).
Cape Verde Islands (38C;38D;44;598).
Senegal (37;450;451).

[As Laurencia paniculata (C. Agardh) J. Agardh]
Senegal (529).

[As Laurencia paniculata J. Agardh]
Mauritanie (624).

Note. Audiffred (38C) indicated his record as new to the Canary
Islands. Lawson & John (350, 586) repeated Yamada's (563) state-
ment of opinion that: 'L. patentiramea (as L. papillosa) and 'L.
paniculata J. Agardh' may represent '. . .the extreme forms of one
very variable species in which L. intermedia may be included.
Askenasy (37: 47) indicated in a footnote that the specimens of
Laurencia were very fragmentary and that Bornet, who had deter-
mined them, had given most with doubt. Given as Laurencia
paniculata? (Agardh) Kutzing by Piccone (450,451); the Piccone
(451) record was based on the Naumann (Gazelle) collection deter-
mined by Askenasy. See also the 'Laurencia sp. A' from Senegal
recorded by Sourie (529) and also note under Laurencia papillosa.
For the correct name of this taxon, see Silva et al. (1987: 68).

Laurencia poitei (Lamouroux) Howe
See L. poiteaui (Lamouroux) Howe.

Laurencia pygmaea Weber-van Bosse
See the notes to Laurencia brachyclados Pilger.

Laurencia scoparia J. Agardh

Senegal (47;52;54;59).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Note. Amongst the varied statements of name and authorities
throughout Bodard & Mollion (59), that employed in table HID (pp.

56

219-220) is: 'L. scoparia (Lamour.) Howe'. Laurencia scoparia sensu
J. Agardh is very similar in external morphology to L. flagellifera J.
Agardh (26: 747); the latter differs in having elongate and palisade-
like epidermal layer cells and lenticular thickenings in the medulla
cell walls. According to Wynne (1986a), the correct name is L.
filiformis (C. Agardh) Montagne.

Laurencia senegalensis Bodard

Senegal (399).

Note. This is probably simply a name of convenience, carelessly
allowed to pass into the publication. It seems neither to have been
described anywhere in print by Bodard, nor used elsewhere by
Bodard, Mollion, or anyone else.

Laurencia tenera Tseng

Canaries (647; 658).

Cape Verde Islands (Leiden Herbarium, det. M.C. Gil-
Rodriguez and R.J. Haroun).

Cote d'lvoire (287;295;350;586).

Gambia (296;350;586).

Ghana (288;297;350;491;586;590).

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

Mauritanie (Leiden Herbarium, det. M.C. Gil-Rodriguez
and R.J. Haroun).

Sierra Leone (295;350;586).

St. Helena (655).

Togo (288;293;350;586;590).

'. . . probably pantropical. . .' (350;586;590).

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

Laurencia tenerrima (Clemente) Cremades &
Perez-Cirera

[As Laurencia perforata Montagne]

Canaries (27;38;44;89;97;128A;133;141A;323;351;390;

401;407;439;563;598;662).
Cape Verde Islands (38;598).
Gabon (350; 586).
Gambia (296;350;586).
Mauritanie (624).
Salvage Islands (38B;598).
Sao Tome (93;251;265;350;586).
'. . . in maribus caldioribus in arena et inter minores Algas

repens. . .' (27).

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

(350;586).

[As Laurencia perforata (Bory) Montagne]
Annobon (456).
Canaries (2;5;8;13;16;38B;38C;71;89;112;191;226;227;229;

253;318;323;375;379;490;583;658;686).
Cape Verde Islands (38B;38C;100;123;183;191).
Gabon (250).

[As Laurencia cf . perforata (Bory) Montagne]
Cape Verde Islands (652).
Salvage Islands (38B;38C;375).
Sao Tome (251 ;265).
'. . . extendida por los mares templados y tropicales. . .'

(253).
[As Laurencia perforata (Bory) Montagne in Barker Webb &

Berthelot]
Annobon (535).
Gabon (535).

D.M. JOHNETAL

Gambia (535).

Sao Tome (535).

Canaries (535;633;634).

[As Laurencia perporata Montagne]

Canaries (25;26).

[As Laurencia cf. perforata (Bory) Montagne]

Cape Verde Islands (652).

Note. See comments concerning this species in notes under L.
brachyclados Pilger.

Laurencia tuberculosa J . Agardh

SaoTome(251;265).

Note. See L. poiteaui and remarks of Steentoft (535). The identity
of specimens clearly requires confirmation.

Laurencia undulata Yamada

Senegal (55;122?;529;590).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Note. Despite recording his material as L. pinnatifida, Sourie
(529: 116) indicated Dangeard's (122) opinion that the Dakar plants
approached L. undulata Yamada. There is also the possibility of
confusion with Laurencia brongniartii J. Agardh (q.v.).

Laurencia viridis Gil-Rodriguez & Haroun

Canaries (647).

Cape Verde Islands (647).

Salvage Islands (647).

Note. A common alga in Macaronesia, before its description
usually misidentified as L. obtusa.

Laurencia spp.

Cameroun (337;344;537).

Canaries (5;8;13;71;226;229;232B;237;281;301;351;379;490;

633; 658).

Cape Verde Islands (100;183;351;411).
Cote d'lvoire (287).
Ghana (42A;297;299;300;335;336;338;344;376;377;487;491;

537;567).

Guinea-Bissau (529).
Guinee (529).
Liberia (287).
Mauritanie (349;537).
Namibia (164).
St. Helena (533).
Salvage Islands (38B).

Senegal (59;123;344;399;411;529;530;531;537;542).
West Africa (290;344;479).

Note. Several of these references represent generalized state-
ments secondarily based on more specific or similarly generalized
data published elsewhere. Some of the records cover more than one
undetermined species within Laurencia; examples are numbers 100
and 183 (Cape Verde Islands - spp. I and II); 490, 633, 529 and 530
(Guinee and Senegal - 2 species, one identified as 'L. sp. A'). Sourie
(529: 119) also stated that one of his species resembles 'L. paniculata
(C. Ag.) J. Ag.'. Amongst the records are some regarded by
publishing author(s) as new species (albeit often with doubt, e.g. 164:
437); sterile material sometimes prevented further critical work
(38B), occasionally subsequently resolved and reflected in specific
entries.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

Lejolisia mediterranea Bornet

Canaries (665).

Leptofauchea brasiliensis Joly

Canaries (664).

Leptofauchea rhodymenioides Taylor

Cape Verde Islands (Prud'homme van Reine, collected on
CANCAP VII expedition).

Leptophytum Adey (1966), nomen inquirendum

The status and disposition of the genus Leptophytum Adey
(669: 323) is unresolved. The generic type collection [i.e. the
type of L. laeve (Stroemfelt) Adey] is missing and thus
application of the name Leptophytum lacks the nomencla-
tural foundation essential for stability (Woelkerling, 1988:
2-3, 217-218). In addition, Woelkerling & Irvine (1986a:
76-77) questioned whether Leptophytum should be recog-
nized as a distinct genus, noting the difficulties outlined by
Adey (669: 28) in ascribing species to Leptophytum vs
Phymatolithon. Nevertheless, Chamberlain (1990: 198) felt
that 'As an interim statement, therefore, I consider that
Leptophytum should be accepted as a distinct genus, and that
it and L. laeve should be interpreted according to Adey's
(1966) concepts until further data are available'. The present
account follows Woelkerling (1988), where Leptophytum is
not recognized as a distinct genus but rather is treated as a
nomen inquirendum and thus a genus requiring further
evaluation. Two species ascribed to Leptophytum have been
recorded from the region under consideration.

Leptophytum bisporum (Foslie) W. Adey

See Phymatolithon bisporum (Foslie) Afonso-Carrillo.

Note. This species was originally described as Lithothamnion
bisporum Foslie (19066: 18) based on material from Puerto Orotava,
Tenerife, Canary Islands. Subsequently, Adey (669: 30) transferred
the species to Leptophytum and then Afonso-Carrillo (11: 134)
placed it in Phymatolithon. Data on the holotype are provided by
Woelkerling (678: 39).

Leptophytum bornetii (Foslie) W. Adey
See Lithothamnion bornetii Foslie.

Leptosiphonia schousboei (Thuret in Bornet & Thuret)
Kylin

Canaries (684).

Leptosiphonia sp.

Senegal (59).

Note. Occurs in the Bodard & Mollion (59) text only in the
terminal table HID (Dredging along the coast of the '. . . Sud de la
petite cote . . . Senegal').

Liagora

A complex genus on which much work remains to be done.
Abbott's recent (656,688) studies of type material relevant to
J. Agardh's (1896) and Lamouroux's (1812) works on the
genus indicated much duplication of names among subgeneric

57

taxa and misappreciation of 'species' limits, apart from mis-
determinations and omission of many previously published
taxa. She was still moved to comment that, of taxa known by
1896, 'many remain little-known to this day'. All of which
makes usage of names in the list area require some reserva-
tion and confirmation.

Liagora albicans Lamouroux

[As Liagora decussata Montagne]

Ascension (475).

Canaries (38;68;139;390;408;439;598).

Cape Verde Islands (25;38;131;191;318;407;408;410;423;528;

551;564;597;598;688).

Note. Kutzing's (318: 538) statement 'Ad insulae St. Vincentii
oras rejecta' presumably refers to the Cape Verde Islands. J.
Agardh's (25: 429) description was quoted direct from Kvitzing (318)
'. . . ad insularum St. Vincentii'. The Latin description was also given
in full in Webb (551) where he referred to 'Montag. MS'. Both that
and Montagne's (1849: 64) description were published in the same
year, with Montagne published in January 1849 and Webb in
November-December 1849. The most recent review of the genus
(656: 308; 688: 1 19) concluded that L. decussata is a later synonym of
L. albicans Lamouroux.

Liagora canariensis B0rgesen

Canaries (2;3;8;13;16;38B;38D;68;191;226;227;235;236;237;

303;351;375;379;489;598;634).
Salvage Islands (38B;38D;598).
[As Liagora fragilis Zanardini var.]
Canaries (439 pro parte).

Note. Piccone (439) gave this (p. 55) in his summary list of species
recorded by Liebetruth and Bolle. L. fragilis was apparently a
Liebetruth record. One of us (PVR) has examined the specimens on
which this record is based and found them to be referrable to two
species, L. canariensis and L. distenta.

General note. Acuna Gonzales (2) stated: '. . . en nuestro archip-
ielago [Islas Canarias] tambien existen algunas [especies] que son
endemicas, como . . . Liagora canariensis. . .'. With substantiable
records from elsewhere, this statement is clearly in error. Levring
(375) has also recorded the species from Cabo Girao, Funchal and
Deserte Grande, both Madeira group, and observed (375: 54): '. . .
no doubt closely related to L. valida and it may be difficult to tell
them apart. . .'. This comment is probably based on Feldmann's
(191: 414) own comment that L. canariensis is near to L. valida, a
pantropical species present on both sides of the Atlantic. According
to Abbott (656: 309, 312 et seq.) L. valida Harvey is itself a synonym
of L. fragilis Zanardini.

Liagora ceranoides Lamouroux

Ascension (474).

Canaries (38B; 38D; 68; 72; 128A; 191; 226; 227; 303; 375;

556; 564; 584; 598; 651).
Cape Verde Islands (688).

Salvage Islands (38B;38D;231;375;556;556A;598).
'Warm Atlantic' (410).
[As Liagora pulverulenta Agardh]
Canaries (547).

Note. Lamouroux's (331: 239) original record (repeated in Lam-
ouroux, 332) is given as '. . . Sur les cotes de 1'ile St. Thomas. Ded.
Weber.'. This is presumably the West Indian island, not that in the
Gulf of Guinea. See also the discussion by Abbott (688).

Liagora complanata C. Agardh

See Liagora distenta (Mertens in Roth) Lamouroux.

58

D.M. JOHNETAL

Liagora corymbosa B0rgesen
See Liagora farinosa Lamouroux.

Liagora decussata Montagne
See Liagora albicans Lamouroux.

Liagora distenta (Mertens in Roth) Lamouroux

Canaries (13;38;38B;38D;68;191;226;227;229; 303;375;439;

489;490;517;556;584;598;634;648;688).
Salvage Islands (38B;38D;215;231;375;556;598).
'. . . Atlantico de Cadiz a Canarias. . .' (517).
'. . . Atlantique (de Cadix aux Canaries)' (188).
'. . . warmeren atlantischen Ocean. . .' (502).
[As Liagora complanata Agardh]
Salvage Islands (381;439;452).
[As Liagora distenta (Mertens in Roth) C. Agardh]
Canaries (392).
Cape Verde Islands (683).

[As Liagora distenta J. Agardh var. complanata J. Agardh]
'Warm Atlantic' (410).
[As Liagora fragilis Zanardini var.]
Canaries (439 pro parte).
See note under L. canariensis B0rgesen.
[As Liagora ramellosa Sender ex Kiitzing]
Canaries (319).

[As Liagora ramellosa Sender in Kiitzing]
Canaries ('De Cadiz aux Canaries. . .') (89).

Note. Reported by Weisscher (556) from the Salvage Islands
solely on the basis of the Gil-Rodriguez et al. (231) record; not found
by CANCAP Expeditions. Bornet himself (89: 105) indicated 'Le
Liagora ramellosa ne me parait pas specifiquement distinct du L.
distenta'. See comment under L. distenta concerning the possible
identity of the Salvage Island record. For discussion of type material,
see Abbott (688).

Liagora elongata Zanardini
See Liagora farinosa Lamouroux.

Liagora farinosa Lamouroux

Canaries (1;16;18;68;80;191;227;229;230;372;416;564).

Cape Verde Islands (652;683;688).

?Prfncipe (350;586).

?Sao Tome (350;586).

'Seems to occur in all warmer seas' (68).

'. . . widespread in warm temperate and tropical seas'

(350;586).

[As Liagora corymbosa B0rgesen]
Salvage Islands (231).
[As Liagora elongata Zanardini]
Canaries (67;246;390;439;547).
[As Liagora megagyna B0rgesen]
?Principe (535).
?Sao Tome (535).

Note. B0rgesen (68: 59-62) examined Lamouroux's specimens
from the Red Sea and confirmed the common identity of L. farinosa,
L. elongata, L. cheyneana and L. corymbosa as a rather variable but
characteristic plant. Abbott (656: 308 et seq.) has confirmed B0rges-
en's conclusion. Steentoft (535) listed L. megagyna records from Sao
Tome and Principe. The latter record was based on exsiccata
specimens collected by F. Newton, who never visited Principe
(Steentoft, pers. comm.) so possibly there was a mistake on the label.

After re-examining Newton's collection, Lawson & John (350)
suggest the identity of the plants to be L. farinosa rather than L.
megagyna; see also the latter entry. The most recent critical treat-
ment of morphology and nomenclature in the species here main-
tained as L. farinosa Lamouroux is that begun by Abbott in 1984 (1)
and carried further in 1990 (656: 308; 688: 122). In 1984 she
commented that L. farinosa '. . . has a number of features about it
that make recognition of the species easy. . .'. She went on to detail
these, adding afterwards 'Nevertheless, the taxon has been given at
least 1 1 specific names . . . and as L. farinosa was designated as the
type specimen of Ganonema Fan & Wang (1974). It was segregated
from Liagora principally on the relationship of the carpogonial
branch to its supporting branch and its location'. Most species of
Liagora, including the type species L. viscida, show carpogonial
branches that are accessory to an established vegetative branching
pattern. Fan & Wang (1974) established that in L. farinosa, the
carpogonial branches were borne only on secondary filaments, or
tertiary filaments of a cortical cluster. Abbott (1) noted that the
location of the carpogonial branches correlated strongly with the
season or age of the plant, varying in position/location/bearing
branchlet type; hence, since this is the feature by which the genus
Ganonema is recognized, it is too unstable a character for generic
distinction. The latter genus (Ganonema farinosa (Lamouroux) Fan
& Wang and Ganonema pinnetiramosa (Yamada) Fan & Wang) was
therefore reduced by Abbott (1) to synonymy with Liagora farinosa
Lamouroux. See also Abbott (688).

Liagora fragilis Zanardini

[As L. valida Harvey]

Canaries (38C;598).

Cape Verde Islands (38C;100;183;191;423;598;652).

'. . . warmer parts of the Atlantic and Pacific (Abbott,

1945). . .' (416).

Note. See also under L. distenta (Mertens in Roth) Lamouroux
and L. canariensis B0rgesen.

Liagora gymnarthron B0rgesen

Canaries (38B;68;88;191;565;598).
[As L. gymnorthron B0rgesen]
Canaries (227).

[As Liagora cf. gymnarthron B0rgesen]
Salvage Islands (38B).

Note. Feldmann (191: 414) indicated that L. gymnarthron appears
to approach L. decussata Montagne (Antilles). Audiffred & Weiss-
cher's Salvage Island record (38B) is expressed with doubt since their
plant differed from Borgesen's description in having regularly
dichotomous branches dispersed as well as alternate/decussate on the
main axes. According to Abbott (pers. comm. to Prud'homme van
Reine) this is probably L. distenta. See also the entry for L. albicans
Lamouroux.

Liagora megagyna B0rgesen

Principe (535).
Sao Tome (535).

Note. Steentoft (535: 121) indicated that only a single plant in
poor condition was available to her. This was the Newton specimen
from Principe; it is therefore not clear why she gave only Sao Tome
as the African distribution, but see L. farinosa entry. Steentoft (535)
indicated that her determination must be regarded as uncertain.
Liagora farinosa (q.v.) is suggested by Lawson & John (350; 586) as
the most likely identity for the specimen.

Liagora perforata

An erroneous statement in Audiffred & Weisscher (38B: 19)
regarding a 'host' of Champia parvula (C. Agardh) Harvey

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

59

on the island of Selvagem Grande. Presumably it correctly
relates to Laurencia perforata Montagne, which is listed in
the body of the text by the same authors.

Liagora pulverulenta C. Agardh

See Liagora ceranoides Lamouroux.

Note. Yamada (564: 20-22) considered there to be two groups in
the taxon L. ceranoides, rather easily distinguishable from each other
on general habit. Since these had hitherto been called pulverulenta J.
Agardh and leprosa J. Agardh he made new combinations at formal
level: L. ceranoides a pulverulenta (Agardh) Yamada and L. cera-
noides p leprosa (J. Agardh) Yamada. Abbott (656: 308) has
confirmed the common identity of L. ceranoides Lamouroux and L.
pulverulenta Agardh, suggested by B0rgesen (68: 58).

Liagora ramellosa Sender ex/in Kiitzing
See Liagora distenta (Mertens) Agardh.

Liagora tetrasporifera B0rgesen

Canaries (2;13;16;38B;38C;38D;57;68;88;188;191;214A;226;

227;229;327;351;490;556;564;598).
Salvage Islands (38B;38C;38D;556;598).
[As L. viscida (Forsskal) C. Agardh]
Canaries (439; 547).

Note. Under the entry for L. viscida Piccone (439: 34) com-
mented that he had only few specimens, differing somewhat from the
typical form. One of us (PVR) has checked the Piccone specimens
under this name and found them to be attributable to L. tet-
rasporifera. See also the entries for Liagora viscida (Forsskal) C.
Agardh. There is some reason to believe that confusion has existed
for this area in attribution of viscida as epithet to specimens likely to
represent tetrasporifera. In the absence of definitive means of revi-
sion, we have not attempted to resolve the situation but have
maintained the authors' naming patterns.

Liagora valida Harvey

See Liagora fragilis Zanardini.

Liagora viscida (Forsskal) C. Agardh

Canaries (38D;216;227;499;584;598).

St. Helena (259;401;655).

'Macaronesia' (656).

'. . . ad oras Europae et Africae; ad ins. St. Thomae. . .'

(318).

'. . .In den warmeren Teilen des Atlantischen Ozeans' (499).
'. . . in oceano Atlantico caldiore. . ; ad insulam Sancti

Thomae. . .' (131).

'. . . littus occidentale (Press); ad insulam St. Thomae' (25).
'Nordwestafrika' (499).
'. . . warmeren atlantischen Ocean. . .' (502).
[As Liagora viscida Agardh S. laxa Kiitzing]
?Sao Tome (318).
[As Liagora viscida C. Agardh]
St. Helena (260).
'Warmer Atlantic' (410).
[As Liagora viscida Forsskal]
Canaries (142).
St. Helena (142;391).

Note. Available evidence tends to suggest that references above
to 'St. Thomae' refer to the West Indian island and not to the African
island of that name. Confusion has been caused by the form of
citation in Kutzing (318: 538), quoted in detail here. Data are

included for completeness. Problems of determination, referred to in
the note to Liagora tetrasporifera B0rgesen (q.v.), have been wors-
ened by the characteristic referred to by Feldmann (188: 271) '. . .
cette algue est assez polymorphic. . .'.

Liagora spp.

Canaries (5;38C;89;117;118;229;247;301;302;304;490;567).
Cape Verde Islands (552).

Note. Weber-van Bosse (552) identified material as Liagora,
possibly a new species, and Bornet (89) indicated he detected six
species amongst Schousboe's material. Audiffred (38C: 179) made
the interesting point 'Even Isabella Abbott did not recognise this
species'!

Lictoria taxiformis (Delile) J. Agardh
See Asparagopsis taxiformis (Delile) Trevisan.

Litholepis

Based on studies of relevant type collections, Woelkerling
(1986) concluded that Litholepis Foslie (203: 5) was a hetero-
typic synonym of Titanoderma Na'geli (1858: 532). Subse-
quently, Campbell & Woelkerling (1990) subsumed
Titanoderma into Lithophyllum Philippi (1837: 387), and
Woelkerling & Campbell (1992: 81) concluded that the type
species of Litholepis, L. caspica (Foslie) Foslie, was a hetero-
typic synonym of Lithophyllum pustulatum (Lamouroux)
Foslie. These conclusions are followed in this paper. Two
species ascribed to Litholepis have been recorded from the
region under consideration.

Litholepis mediterranea Foslie

Cape Verde Islands (366;368;598).

Note. The status and disposition of this species are uncertain, and
records from the Cape Verde Islands require confirmation once a
detailed study of the holotype (see 678: 147) has been undertaken.
The holotype was collected at Banyuls sur Mer, France. Feldmann
(188: 317) referred the species to Fosliella (now considered a
heterotypic synonym of Hydrolithon; Penrose & Chamberlain, 1993)
while Adey (669: 15) referred the species with some doubt to
Lithoporella. Both Hydrolithon and Lithoporella belong to the
subfamily Mastophoroideae (Woelkerling, 1988: 115) whereas the
type of Litholepis belongs to the subfamily Lithophylloideae (Woelk-
erling, 1988: 92).

Litholepis sauvageaui Foslie

See Lithoporella sauvageaui (Foslie) Adey.

Lithophyllum Philippi

The concept of Lithophyllum adopted in this paper follows
Woelkerling & Campbell (1992: 17). According to this con-
cept, the following characters collectively delimit Lithophyl-
lum from other genera of Corallinaceae: 1) thallus
nongeniculate; 2) crustose portions of thallus with a dorsi ven-
tral internal organization; 3) haustoria absent; 4) cells of
contiguous vegetative filaments commonly joined by second-
ary pit-connections; 5) fusions between vegetative cells
absent or very rare; 6) tetrasporangial/bisporangial concep-
tacles uniporate; and 7) tetrasporangia/bisporangia lacking
apical plugs. The earlier taxonomic history of Lithophyllum is
summarized by Woelkerling (19836), who also provided
accounts of the original collections of the four species that

60

D.M. JOHNETAL

Philippi (1837) included in the genus. Hyperantherella Hey-
drich (1900: 316) and Crodelia Heydrich (1911: 12) are
homotypic synonyms of Lithophyllum (see Woelkerling,
1988: 99-100). Heterotypic synonyms include Dermatolithon
Foslie (682: 11) (a homotypic synonym of Titanoderma; see
below and also Woelkerling et al., 1985 and Campbell &
Woelkerling, 1990), Litholepis Foslie (203: 5) (see Woelker-
ling, 1986: 260; Woelkerling & Campbell, 1992: 81), Perisper-
mon Heydrich (1900: 316) (see Woelkerling, 1991),
Pseudolithophyllum Lemoine (1913a: 45) (see Woelkerling,
1988: 103), Stichospora Heydrich (1900: 316) (see Woelker-
ling, 1983a: 184; Woelkerling, 1988: 102), and Titanoderma
Nageli (1858: 532) (see Campbell & Woelkerling, 1990).
Chamberlain (1991: 13, 23-24) and Chamberlain et al. (1991:
164-165) proposed that Titanoderma be maintained as a
genus distinct from Lithophyllum, but Woelkerling & Camp-
bell (1992: 17-18) concluded that characters suggested by
Chamberlain (1991) and Chamberlain et al. (1991) to be
diagnostic of Titanoderma (a predominance of primigenous
palisade cells and the occurrence of at least some bistratose
margin) were too variable in southern Australian collections
to be used reliably in generic delimitation.

Lithophyllum absimile Foslie & Howe in Foslie
See under Spongites wildpretii Afonso-Carrillo.

Lithophyllum accretum (Foslie & Howe) Lemoine

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

Lithophyllum accretum (Foslie & Howe) Lemoine
f . canariensis Foslie

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

Lithophyllum accretum (Foslie & Howe) Lemoine var.
canariense (Foslie) Lemoine

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

Lithophyllum aequinoctiale Foslie

See Porolithon aequinoctiale (Foslie) Foslie.

Lithophyllum africanum Foslie

See Porolithon africanum (Foslie) Foslie.

Lithophyllum amplexifrons (Harvey) Heydrich

See Pneophyllum amplexifrons (Harvey) Chamberlain &
Norris.

Lithophyllum aninae Foslie

Cape Verde Islands (6;100;101;136;139;207;210;212;366;597;

598;678).

Note. Foslie (207: 28) based Lithophyllum aninae on a single
collection from Sao Vincente, Cape Verde Islands. Adey (669: 4)
referred the holotype to Lithophyllum without comment. The holo-
type (678: 27), however, has not been studied in detail in a modern
context, and thus the status and disposition of the species are

uncertain, as is the identification of specimens from the West African
region.

Lithophyllum applicatum Lemoine in B0rgesen

See Neogoniolithon hirtum (Lemoine in B0rgesen) Afonso-
Carrillo.

Lithophyllum bisporum Foslie

See Phymatolithon bisporum (Foslie) Afonso-Carrillo.

Lithophyllum byssoides (Lamarck) Foslie.

Mauritanie (356;359;360).

[As Goniolithon byssoides (Lamarck) Foslie]

Mauritanie (354).

Note. This species was originally described as Nullipora byssoides
Lamarck (1801: 374) and was based on material depicted by Seba
(1758: pi. 116, fig. 7). Subsequently, the species has been placed in
Millepora (Lamarck, 1816: 203), Lithothamnion (Philippi, 1837:
388), Spongites (Kutzing, 1869: 35), Goniolithon (Foslie, 1898a: 5),
Lithophyllum (682: 20), and Titanoderma (Chamberlain & Woelker-
ling in Woelkerling, 1988: 260). Woelkerling (1983a: 177-180, figs
12-16) outlined the nomenclatural history of the species, neotypified
it with a Philippi collection and provided an account of that material.
Later, Woelkerling (1988: 216, 217, 260) concluded that the species
belonged to Titanoderma sensu Woelkerling et al. 1985 (see also
Woelkerling, 1988). Following Campbell & Woelkerling (1990) and
Woelkerling & Campbell (1992), Titanoderma is regarded here to be
a heterotypic synonym of Lithophyllum, and thus Nullipora byssoides
is dealt with as a Lithophyllum. Specimens on which the records from
Mauritanie are based now need to be re-examined to determine
whether they are conspecific with the neotype of L. byssoides.

Lithophyllum calcareum (Pallas) Areschoug

See Phymatolithon calcareum (Pallas) Adey & McKibbin.

Lithophyllum canariensis Foslie

See Mesophyllum canariensis (Foslie) Lemoine.

Lithophyllum capense Rosanoff

Cape Verde Islands (38;598).

'. . . Afrique meridionale. . .' (38).

[As Lithothamnium capense (Rosanoff) Foslie]

Canaries (354).

Note. Rosanoff (1866: 86) based this species on material from
southern Africa that Hohenacker had distributed as Alg. mar. sice.
no. 236. Type material of this species has not been studied in detail in
a modern context, and thus the status and disposition of the species
are uncertain as is the identification of specimens from the West
African region.

Lithophyllum caribaeum Foslie

See Neogoniolithon caribaeum (Foslie) Adey.

Lithophyllum corallinae (P. & H. Crouan) Heydrich

Mauritanie (349).

[As Dermatolithon corallinae (P. & H. Crouan) Foslie]

Salvage Islands (38B;231;375).

[As Melobesia corallinae Solms-Laubach]

Mauritanie (59;252).

Senegal (252).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

[As Melobesia corallinae Crouan]
Canaries (439)

Note. Detailed accounts of this species, including information on
the lectotype, are provided by Chamberlain (1991: 66-69, figs
208-224, as Titanoderma) and Woelkerling & Campbell (1992:
41-56, figs 22-32). All specimens on which published records for the
West African region are based need to be checked to determine
whether they are conspecific with the lectotype. Lithophyllum coral-
linae also was referred to by Price et al. (1986: 86) under Dermatoli-
thon in a previous part of this critical assessment series.

Lithophyllum cristatum Meneghini f . crassa (Lloyd)
Hauck

See Tenarea tortuosa (Espoer) Lemoine.

Lithophyllum crouanii Foslie

Canaries (598).

[As Lithophyllium crouani]

Canaries (235).

Note. The lectotype material (678: 68) of Lithophyllum crouanii
was collected at Berwick-on-Tweed, England and has been examined
in detail by Chamberlain et al. (1988), who also provided an account
of the species in the British Isles. Records of this species from the
Canary Islands require confirmation, especially in view of the misi-
dentifications uncovered by Chamberlain et al. (1988).

Lithophyllum cystoseirae (Hauck) Heydrich.

Annobon (455).

[As Dermatolithon cystoseirae (Hauck) Huve]

Canaries (227;582)

Mauritanie (262;349;367;368).

[As Dermatolithon cystoseirae (Hauck) Foslie]

Mauritanie (367).

[As Dermatolithon cystoseirae (Hauck) Foslie f. saxicola

Huve]

Canaries (368).
Mauritanie (368).
[As Dermatolithon cystoseirae (Hauck) Foslie var. saxicola

Huve]

Canaries (368).
Mauritanie (368).

'. . . Atlantique (de 1'Angleterre a la Mauritanie). . .' (33)
[As Melobesia cystoseirae Hauck]
Annobon (139)

[As Dermatolithon papillosum (Zanardini) Foslie var. cys-
toseirae (Hauck) Lemoine]
Annobon (350)
'. . . Gulf of Guinea region. . .' (350;367)

Note. This species was originally described as Melobesia cystosei-
rae (Hauck, 1883: 266, pi. 3, figs 1, 2, 6) and was based on specimens
from the Adriatic Sea. Studies of type material by Huve (272) and
Athanasiadis (1989) have confirmed that this species belongs to
Lithophyllum as delimited by Campbell & Woelkerling (1990) and
Woelkerling & Campbell (1992), but a thorough study of reproduc-
tive anatomy needs to be undertaken before the relationships of it to
others in the genus can be properly determined. Once such a study is
completed, all specimens on which published records for the West
African region are based need to be re-examined to determine
whether they are conspecific with the type. Lithophyllum cystoseirae
was also referred to by Price et al. (1986: 86) under Dermatolithon in
a previous part of this series.

61

Lithophyllum daedaleum Foslie & Howe

Canaries (666).

Note. Foslie & Howe (1906: 133) based this species on material
from Puerto Rico. The holotype (678: 70) has not been studied in
detail in a modern context, and thus the status and disposition of this
species are uncertain, as is the identification of the material from the
Canary Islands.

Lithophyllum decussatum (Ellis & Solander) Philippi
f . pianist-ilia Foslie

Canaries (211).

Note. Foslie (211: 22) based Lithophyllum decussatum f. planis-
cula on material from Morocco (see 678: 173) that previously had
been referred to Lithophyllum expansum. Foslie (211: 23) also
suggested that 'En lignende form foreligger ogsaa fra de Kanariske
0er' but Foslie did not definitely refer the Canary Islands material to
f. planiscula. No such specimen is filed in the Foslie herbarium under
Lithophyllum decussatum (6: 44), and until the relevant specimen is
located and examined in a modern context, this record must be
regarded as questionable.

Lithophyllum duckeri Woelkerling

Ascension (541).

[As Lithothamnion crassum Philippi]

Ascension (541).

Canaries (547;598).

St Helena (142;260;391).

Note. Lithophyllum duckeri is a nom. nov. for Lithothamnion
crassum Philippi. An account of the lectotype and the nomenclatural
history of Lithophyllum duckeri are provided by Woelkerling (1983a:
180-184, figs 17-22), who showed that the type of Lithothamnion
crassum belonged to Lithophyllum. Upon transfer into Lithophyl-
lum, the new specific epithet duckeri was required because Rosanoff
(1866: 93) had used the combination Lithophyllum crassum for
another species. Woelkerling (1983a) noted that the relationships of
L. duckeri to other species of Lithophyllum was uncertain. The
lectotype was collected from the west coast of Sicily. Subsequent
studies of southern Australian species of Lithophyllum (Woelkerling
& Campbell, 1992) have shown that characters relating to tetraspo-
rangial conceptacle roof anatomy are important in species delimita-
tion, and because these were not considered by Woelkerling (1983a),
further studies of the type of L. duckeri are required before its status
and relationships can be fully determined. Once such studies are
completed, all specimens on which records from Ascension, the
Canaries, and St. Helena are based must be checked to determine
whether they are conspecific with the lectotype of L. duckeri.

Lithophyllum esperi (Lemoine in B0rgesen) South &
Tittley

[As Pseudolithophyllum esperi Lemoine]

Canaries (70;191 ;227;362;363;499;582;598).

Cape Verde Islands (366;598).

'. . . Golfe de Guinee. . .' (366;586).

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

Note. Lemoine (363: 63) based this species on four collections
from Puerto Orotava, Tenerife, Canary Islands but did not designate
a type. South & Tittley (1986: 43) transferred the species into
Lithophyllum, but the original collections of Lemoine, from which a
lectotype must be chosen, have not been studied in detail in a modern
context, and thus the status and disposition of this species are
uncertain, as is the identification of other specimens from the West
African region.

62

D.M. JOHNETAL

Lithophyllum expansum Philippi

See Mesophyllum lichenoides (Ellis) Lemoine.

Lithophyllum geometricum Lemoine

Canaries (191;362;363;687)

[As Dermatolithon geometricum (Lemoine) Dawson]

Canaries (227;366;367;368;369;582;674).

Cape Verde Islands (366;367;368;369;582).

'. . . Atlantique african. . .' (369).

[As Lithophyllum sp. (geometricum^)]

Canaries (230).

[As Lythophyllum sp. (geometricum'?) Lemoine]

Canaries (230).

Note. Lemoine (363: 47) based this species on two collections
from Puerto Orotava, Tenerife, Canary Islands but did not designate
a type. The species subsequently has been transferred to Goniolithon
Setchell & Mason (1943: 89), Dermatolithon (674: 273) and Titano-
derma (Price et al., 1986: 86). The type material apparently has not
been re-examined in detail in a modern context and a lectotype has
not been designated; thus the status and disposition of the species are
uncertain, as is the identification of other specimens from the West
African region. Lithophyllum geometricum also was referred to by
Price et al. (1986: 86) under Dermatolithon in a previous part of this
series.

Lithophyllum gracile Foslie

Cape Verde Islands (6;136;139;;207;210;212;366;597;598).
'. . . Africa occidental' (136).

Note. A detailed study of the holotype collection (678: 108),
which comes from Sao Vincente, Cape Verde Islands, has not been
undertaken in a modern context, and thus the status and disposition
of the species are uncertain, as is the identification of specimens from
the West African region. Adey (669: 5) referred the holotype to
Lithophyllum without comment.

Lithophyllum hapalidioides (P. & H. Crouan) Foslie
See Lithophyllum pustulatum (Lamouroux) Foslie.

Lithophyllum hauckii (Rothpletz) Lemoine

See Neogoniolithon mamillosum (Foslie) Setchell & Mason

Lithophyllum hirtum Lemoine in B0rgesen

See Neogoniolithon hirtum (Lemoine in B0rgesen) Afonso-
Carrillo.

Lithophyllum illitus Lemoine in B0rgesen

See Neogoniolithon illitus (Lemoine in B0rgesen) Afonso-
Carrillo.

Lithophyllum incrustans Philippi

Canaries (3;6;70;109;134;227;252;354;356;359;493;582;584;

598;633).

Cape Verde Islands (100;366;375;598).
Mauritanie (6 ;248 ;252 ;349 ;356 ;359 ;63 1) .
Western Sahara (252;349;631).
'. . . Atlantique: depuis les cotes anglaises jusqu'en

Mauritanie. . .' (222).
'. . . Atlantique (des Feroes a la Mauritanie, lies du Cap

Vert). . .' (33).

'. . . Atlantico norte (hasta Marruecos y Mauritania)' (517).
'. . . Atlantique nord (jusqu'au Maroc et a la Mauritanie)'

(188).

'. . . cotes . . . africaine de 1'Atlantique. . .' (357).
'. . . si commune sur les cotes atlantique a maree basse. . .'

(359).
'subtropical Africa (N. of Gambia); Mauritania; former W.

Sahara' (598).

[As Lithophyllum incrunstans[sic\] Philippi]
Canaries (253).
Cape Verde Islands (253).

'. . . Atlantico Oriental (Inglaterra - Cabo Verde). . .'(253).
[As Lithothamnion incrustans (Philippi) Foslie]
Canaries (547).

[As Lithothamnion ponderosum Foslie]
Sao Tome (197;265).

Note. Records of this species from the West African region
require confirmation. An account of the holotype is provided by
Woelkerling (19836: 313-317, figs 15-22), but unfortunately, details
of tetrasporangial conceptacle roof anatomy, now known (Woelker-
ling & Campbell, 1992) to be important in species delimitation within
Lithophyllum, were not included and further study of the type is
required. The holotype was collected along the west coast of Sicily.

Lithophyllum irregulare (Foslie) Huve ex Steentoft

Canaries (535).

Sao Tome (535).

[As Lithophyllum irregularis Foslie]

Canaries (227).

[As Lithothamnium irregulare Foslie]

Sao Tome (206;350;586).

'. . . in warm temperate and tropical parts of the eastern

Atlantic Ocean' (350;586).
[As Lithothamnion irregulare Foslie]
Canaries (582).

'Gulf of Guinea' (6;135;139;206;212;582).
'. . . vestkysten af Afrika' (209).
[As Pseudolithophyllum irregulare (Foslie) Adey]
Canaries (598).

Tropical Africa (N. Gambia-Congo river)' (598).
[As Tenera irregularis (Foslie) Lemoine]
Sao Tome (70).

Note. This species was originally described as Lithothamnion
irregulare (206: 6), based on material from Sao Tome. The species
subsequently was transferred to Tenarea (363: 56), then to Lithophyl-
lum (see below), and then with some doubt to Pseudolithophyllum
(669: 13). The combination Lithophyllum irregulare was first effected
by H. Huve (1957: 138), but because Huve did not cite the basionym
(Lithothamnion irregulare, 206: 6), her combination is invalid
(Article 33.2 in the International code of botanical nomenclature; see
Greuter, 1988). Steentoft (535: 128) subsequently validated the
combination which is correctly cited as either Lithophyllum irregulare
(Foslie) H. Huve ex Steentoft or Lithophyllum irregulare (Foslie)
Steentoft. The holotype collection (678: 130) was examined by
Steentoft (535: 128), but there is no detailed account of the holotype
in a modern context, and thus the status and disposition of this
species is uncertain. Retention here of the species in Lithophyllum is
purely arbitrary as it is unknown whether the type possesses second-
ary pit-connections (characteristic of Lithophyllum) or cell fusions
(implied by placement in Pseudolithophyllum sensu Adey, 1970).
Further data on the name Pseudolithophyllum are provided by
Woelkerling (1988: 103). All specimens on which published records
for the West African region are based need to be checked to
determine whether they are conspecific with the holotype.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

Lithophyllum kaiseri (Heydrich) Heydrich

Annobon (586).

[As Lithophyllum Kaiseri Heydrich]

Annobon (367;368;455).

[As Lithophyllum kotschyanum Unger]

Annobon (139;397;455;457).

Note. This species originally was described as Lithothamnion
kaiseri (Heydrich, 1897a: 64) and is based on material from El Tor,
the Red Sea. Subsequently, Heydrich (1897&: 412) transferred the
species to Lithophyllum. Heydrich did not designate a type, and
Heydrich's main herbarium has apparently been destroyed (Stafleu
& Cowan, 1979: 187; Hiepko, 1987: 230). However, syntype material
occurs in TRH (678: 132). Because a lectotype has not been
designated and studied in detail in a modern context, the status and
disposition of the species are uncertain, as are the records from the
West African region. According to Lemoine (368: 6), plants from
Annobon identified by Pilger (455: 419) as Lithophyllum kotschy-
anum Unger are really Lithophyllum kaiseri. Once L. kaiseri has
been lectotypified and studied in detail, Pilger's plants need to be
re-examined to determine whether they are conspecific with the
lectotype of L. kaiseri.

Lithophyllum kotschyanum Unger

See Lithophyllum kaiseri (Heydrich) Heydrich.

Lithophyllum leptothalloideum Pilger

Annobon (455;457).

[As Pseudolithophyllum leptothalloideum (Pilger) De Toni]

Annobon (139;350;586).

Note. A detailed study of the type material (whereabouts uncer-
tain; see Stafleu & Cowan, 1983: 265), collected at Annobon, has not
been undertaken in a modern context, and thus the status and
disposition of this species are uncertain.

Lithophyllum lobatum Lemoine in B0rgesen

Canaries (70;227;362;365;366;685;687).

Cape Verde Islands (366).

Mauritanie (349;366).

Senegal (529).

'Lusitano- Africano-Mediterraneen' (529) .

[As Pseudolithophyllum lobatum (Lemoine) Verlaque &

Boudouresque]
Canaries (582;598).
Cape Verde Islands (598).

'N.W. Africa and Atlantic Islands' [translation] (582).
'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).
[As Lithophyllum expansum Philippi]
Canaries (21 1;493).

Note. Lemoine (363) states that: 'les echantillons determines sous
le nom [Lithophyllum expansum} par M. Foslie, et cites par M.
Sauvageau [493: 185] dans la localite de Puerto Orotava sont en
realite L. lobatum'.

[As Mesophyllum lobatum Lemoine]
Canaries (191).

Note. Lemoine (362: 40) based this species on four collections
from Puerto Orotava, Tenerife, Canary Islands, but did not specify a
type. As far as known, a lectotype has not been designated and there
has been no detailed study of the protologue collections in a modern
context. Thus the status and disposition of the species are uncertain,
as is the identification of specimens from the West African region.
According to Lemoine (362), the cited entries for Lithophyllum
expansum involve misidentified specimens. Feldmann (192: 414) used
the binomial Mesophyllum lobatum Lemoine instead of Lithophyl-

63

lum lobatum Lemoine in B0rgesen. It is not clear whether this is an
error or whether Feldmann had intended to transfer lobatum to
Mesophyllum; the binomial probably should be cited as Mesophyllum
lobatum (Lemoine) Lemoine ex J. Feldmann.

Lithophyllum marlothii Heydrich

Sao Tome (6;265).

[As Lithophyllum Marlothii Heydrich, forma]

Sao Tome (251).

Note. Heydrich (18976: 61) originally described this species as
Lithothamnion marlothii but soon (Heydrich, 1897a: 410) transferred
it to Lithophyllum. Heydrich based the species on collections from
several localities in South Africa but did not designate a type.
Heydrich's main herbarium apparently has been destroyed (Stafleu
& Cowan, 1979: 187; Hiepko, 1987: 230), but syntype material of L.
marlothii occurs in TRH (678: 145). Because a lectotype has not been
designated and studied in detail in a modern context, the status and
disposition of the species are uncertain, as are the records from the
West African region.

Lithophyllum mildbraedii Pilger

Annobon (455;457;500).

Bioko (500).

Cameroun (500).

[As Pseudolithophyllum mildbraedii (Pilger) De Toni]

Annobon (139).

Bioko (350).

Cameroun (350).

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

'. . . only known from the eastern parts of the tropical

Atlantic' (350; 386).

Note. A detailed study of the type material (whereabouts uncer-
tain; see Stafleu & Cowan, 1983: 265), collected at Annobon, has not
been undertaken in a modern context, and thus the status and
disposition of this species are uncertain as are records from the West
African region.

Lithophyllum oligocarpum Foslie

See Porolithon onkodes (Heydrich) Foslie.

Lithophyllum orbiculatum (Foslie) Foslie

Salvage Islands (38B;231;375).

'Norway to Morocco: W. Mediterannean' (649).

[As Pseudolithophyllum orbiculatum (Foslie) Lemoine]

Salvage Islands (598).

Note. A detailed study of the lectotype (678: 164), which comes
from Kristiansund, Norway, has been undertaken by Chamberlain et
al. (1991) who also noted confusions resulting from specimens being
misidentified as Lithophyllum orbiculatum. Chamberlain et al. (1991:
161-162) did not confirm the occurrence of this species in the West
African region, and all specimens on which published records from
the area are based need to be checked to determine whether they are
conspecific with the lectotype.

Lithophyllum orotavicum Foslie

See Neogoniolithon orotavicum (Foslie) Lemoine.

Lithophyllum papillosum (Zanardini ex Hauck) Foslie

Canaries (188;191;359;362;363;365).

[As Dermatolithon papillosum (Zanardini) Foslie]

Mauritanie (248).

[As Goniolithon papillosum (Zanardini) Foslie]

64

D.M. JOHN ET AL

Canaries (18;582).

[As Titanoderma papillosum (Zanardini) Price, John & Law-
son]

Annobon (586).
'. . . Gulf of Guinea region. . .' (586).

Note. This species was originally described as Lithothamnion
papillosum Zanardini ex Hauck (1883: 272, pi. 2, fig. 4) and has been
lectotypified (272: 224) with a collection from Susak Island in the
Adriatic Sea. The lectotype has been examined by Huve (272) and by
Woelkerling (1988: 217-218). Further studies of the lectotype are
required, however, to elucidate the reproductive anatomy of the
species, to determine its status and disposition, and to determine the
status and disposition of Goniolithon Foslie (1898a: 5), which is
typified by G. papillosum (see Woelkerling, 1988: 217-218). Once
these matters have been resolved, all specimens from the West
African region ascribed to this species need to be checked to
determine whether they are conspecific with the lectotype (see also
582: 25). Lithophyllum papillosum also was referred to by Price et al.
(1986: 86) under Dermatolithon in a previous part of this series.

Lithophyllum polycephalum Foslie

Canaries (6;212;493).

Cape Verde Islands (6;201;210;212;597).

[As Lithophyllum (Dermatolithon) polycephalum (Foslie)

Foslie]

Canaries (191 ;362;363).
Cape Verde Islands (363;674).
[As Dermatolithon polycephalum (Foslie) Foslie]
Canaries (227;366).
Cape Verde Islands (139;366).

[As Goniolithon polycephalum (Foslie) Afonso-Carrillo]
Canaries (11 ;582).
Cape Verde Islands (582).

Note. Foslie (201) based this species on a collection from Sao
Vincente, Cape Verde Islands. Although the holotype (678: 174) has
been examined by Afonso-Carrillo (11: 139) and by Woelkerling &
Campbell (1992: 22), a detailed study in a modern context has not
been undertaken, and thus the status and disposition of this species
are uncertain, as is the identification of specimens from the West
African region. Lithophyllum polycephalum was also referred to by
Price et al. (1986: 86) under Dermatolithon in a previous part of this
critical assessment series.

Lithophyllum polyclonum Foslie

[As Dermatolithon polyclonum (Foslie) Foslie]
Mauritanie (349).

Note. Foslie (201: 18) based this species on a collection from the
West Indies. The holotype (678: 175) has not been studied in detail in
a modern context, and thus the status and disposition of this species
are uncertain, as is the identification of the material from Mauritanie.
Lithophyllum polydonium also was referred to by Price et al. (1986:
86) under Dermatolithon in a previous part of this series.

Lithophyllum ponderosum Foslie

See Lithothamnion ponderosum Foslie.

Lithophyllum proboscideum (Foslie) Heydrich
See Porolithon africanum (Foslie) Foslie, and 678: 176.

Lithophyllum pustulatum (Lamouroux) Foslie

Canaries (363)

[As Lithophyllum (Dermatolithon) pustulatum (Lamouroux)
Foslie]

Canaries (109;354;356;359).

Mauritanie (359).

'Cape Blanc, Senegal' (356).

[As Dermatolithon pustulatum (Lamouroux) Foslie]

Canaries (227;235;390;499;584).

Cape Verde Islands (366;499) .

'Nordwestafrika' (499).

[As Melobesia (Dermatolithon) pustulatum]

Canaries (6).

[As Melobesia pustulatum Lamouroux]

Canaries (441 ;444;547).

Cape Verde Islands (26;38;145;408).

'Du Nord de la Grande-Bretagne aux Canaries' (38;89).

[As Lithophyllum pustulatum Lamouroux f . australis Foslie]

Canaries (202;678).

[As Dermatolithon hapalidioides (P. & H. Crouan) Foslie]

Canaries (227;361;375;582).

Salvage Islands (38B;375).

[As Dermatolithon hapalidioides (P. & H. Crouan) Foslie f.

confinis (P. & H. Crouan) Foslie]
'. . . Atlantique (. . .Canaries. . .)' (33).
[As Lithophyllum hapalidioides (P. & H. Crouan) Foslie]
Canaries (188;191;356;362;363).
[As Lithophyllum hapalidioides (P. & H. Crouan) Foslie var.

confinis (P. & H. Crouan) Lemoine]
Canaries (363).

[As Dermatolithon nepalidioides (P. & H. Crouan) Foslie]
Salvage Islands (231).

Note. This species was originally described as Melobesia pustulata
(331: 315). Woelkerling et al. (1985) lectotypified the species with a
Lamouroux collection from France and provided a detailed account
of the material. Additional data on the lectotype are provided by
Chamberlain (1991, as Titanoderma ) and Woelkerling & Campbell
(1992). These workers examined the types of Melobesia confinis P. &
H. Crouan and M. hapalidioides P. & H. Crouan and concluded that
they are heterotypic synonyms of Melobesia pustulata Lamouroux.
Chamberlain (1991, as Titanoderma) recognized four distinct variet-
ies from the British Isles, whereas Woelkerling & Campbell (1992:
90-94) found so many intermediate specimens in southern Australia
that it seemed to them neither desirable nor advantageous to
recognize distinct varieties. Specimens on which all published records
of Lithophyllum pustulatum from the West African region are based
need to be checked to determine whether they are conspecific with
the lectotype (also see 576: 25). Foslie (202: 117) based Lithophyllum
pustulatum f. australis on specimens from a number of localities
including the Canary Islands. Woelkerling (678: 35) designated the
Canary Islands specimen as lectotype and provided additional com-
ments. The lectotype, however, has not been studied in detail in a
modern context, and thus the status and disposition of this taxon are
uncertain. Lithophyllum pustulatum was also referred to by Price et
al. (1986: 86) under Dermatolithon. Entries in this earlier part of the
series for Dermatolithon confinis, D. hapalidioides and D. nepalidio-
ides also pertain to Lithophyllum pustulatum.

Lithophyllum racemus (Lamarck) Foslie

St. Helena (541).

Note. This species was originally described as Millepora racemus
(Lamarck, 1816: 203) and is based on material from 'les mers de la
Guiane?' collected by Turgot. Turgot's material (whereabouts uncer-
tain) has not been studied in detail in a modern context, and thus the
status and disposition of this species is uncertain, as is the identifica-
tion of the material from St. Helena. Additional notes on the name
Lithophyllum racemus appear below in the entry for Mesophyllum
brachycladum.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

65

Lithophyllum retusum (Foslie) Foslie

Cape Verde Islands (683).

Ghana (6;211;350;535;586).

Sao Tome (6;134;197;198;211;212;350;535;586).

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

[As L. retusum Foslie, forma]

SaoTome(251;265).

Note. Foslie (197: 15) first described this species as Lithotham-
nion retusum, but later (681: 9) transferred it to Goniolithon Foslie
and then (682: 18) to Lithophyllum (see also 669: 5). The species is
based on a single collection from Sao Tome (Henriques no. 24,
collected by M011er; see 197: 15). The holotype (678: 189), however,
has not been examined in detail in a modern context and thus the
status and disposition of the species are uncertain, as are records
from the West African region. According to Steentoft (535), the
Hariot (251: 164) and Henriques (265: 166) records of Lithophyllum
retusum Foslie forma do not represent the same taxon. Several
collections referred to by Steentoft (535) are represented in TRH (6:
43).

Lithophyllum simile Foslie

Sao Tome (6;211;212;350;535;586).

'. . . in tropical parts of the eastern Atlantic Ocean'

(350;586).
Tropical Africa (N. Gambia - Congo river)' (598).

Note. Foslie (211: 30) based Lithophyllum simile on a single
collection from Sao Tome. A detailed study of the holotype (678:
201), however, has not been undertaken in a modern context, and
thus the status and disposition of the species are uncertain, as is the
identification of specimens from the West African region. Adey (669:
6) referred the holotype to Lithophyllum without comment.

Lithophyllum solutum (Foslie) Lemoine
See Lithothamnion solutum (Foslie) Lemoine.

Lithophyllum subtenellum (Foslie) Foslie

Sao Tome (6;190;251;265;350;586).

'. . . widespread in warm temperate and tropical parts of the

eastern Atlantic Ocean' (350;586).
[As Lithothamnium subtenellum Foslie]
Mauritanie (359).
Sao Tome (356;535).

Tropical Africa (N. Gambia - Congo river)' (598).
[As Lithothamnium subtenellum (Foslie) Lemoine]
Sao Tome (188).

Note. A detailed study of the lectotype collection (678: 215; see
also 669: 6), which comes from Guethary, France, has not been
undertaken in a modern context, and thus the status and disposition
of the species are uncertain, as is the identification of specimens from
the West African region.

Lithophyllum tortuosum (Esper) Huve
See Tenarea tortuosa (Esper) Lemoine.

Lithophyllum vickersiae Lemoine in B0rgesen

Canaries (70;227;362;366;375;499;556;687).

Cape Verde Islands (366;556).

Senegal (366).

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

[As Lithophyllum incrustans]

Canaries (70;493;547).

[As Lithophyllum cf. vickersiae Lemoine]
Salvage Islands (38B;556).

Note. Determination based on external features only.
[As Lithothamnium vickersiae Lemoine]
Canaries (100;191).
[As Pseudolithophyllum vickersiae (Lemoine in B0rgesen)

Afonso-Carrillo]
Canaries (11;18;582).
Cape Verde Islands (598).
'southern limit in the Gulf of Guinea' [translation fom the

Spanish] (582).
'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Note. Based on studies of the original Canary Islands collections
of Lemoine (362: 42), Afonso-Carrillo (11: 139) decided that this
species belonged to Pseudolithophyllum sensu Lemoine (1913). Sub-
sequently, however, Woelkerling (1988: 103) concluded that Pseudo-
lithophyllum sensu Lemoine (1913) is a heterotypic synonym of
Lithophyllum, and thus L. vickersae is retained here in Lithophyl-
lum. Afonso-Carrillo (11) did not designate a lectotype from amongst
the seven collections cited in the protologue (362: 42), and he did not
present a detailed account of the type material. Thus the status and
disposition of the species are uncertain, as is the identification of
specimens from the West African region. According to Lemoine
(362: 42), some plants of L. vickersae had earlier been associated
with L. incrustans (see listings above).

Lithophyllum zostericolum

See Pneophyllum amplexifrons .

Lithophyllum spp.

Cameroun (454).

Canaries (212;229;230;301 ;302;304).

Cape Verde Islands (652;683).

Namibia (348).

[As Lythophyllum sp.]

Canaries (229;230).

[As Dermatolithon sp.]

Canaries (253;314;582).

Note. Price et al. (1986: 86) also record Dermatolithon sp. in a
previous part of this critical assessment series but provide no refer-
ences.

Lithoporella Foslie

The concept of Lithoporella adopted in this paper follows
Woelkerling (1988: 124-128). Historical data on the genus
are summarized by Turner & Woelkerling (1982a, b), who
also give an account of the lectotype species, L. melobesio-
ides (Foslie) Foslie. A revised key to the genera of Masto-
phoroideae, including Lithoporella, is provided by Penrose &
Chamberlain (1993: 303). According to Turner & Woelker-
ling (19826: 233) and Woelkerling (1988: 128), uncertainties
surrounds the delimitation and circumscription of most spe-
cies in the genus.

Lithoporella atlantica (Foslie) Foslie

See note to Lithoporella melobesioides (Foslie) Foslie.

66

D.M. JOHNETAL

Lithoporella conjuncta (Foslie) Foslie

Cape Verde Islands (139;366;597).

Mauritanie (139;21 1 ;349;366;597).

Senegal (597)

[As Lithoporella (Eulithoporella) conjuncta Foslie]

'Atlantique africain' (371).

[As Mastophora (Lithoporella) conjuncta Foslie]

Cape Verde Islands (6;136;207).

Mauritanie (6;136;207;210;212).

Note. This species was originally described as Mastophora con-
juncta Foslie (207: 30) and is based on collections from Cap Blanc,
West Africa and Sao Vincente, Cape Verde Islands. Subsequently,
Foslie (211: 59) transferred the species to Lithoporella, and Adey
(669: 15) lectotypified it with the Cap Blanc collection. A detailed
study of the lectotype (678: 61) has not been undertaken in a modern
context, and thus the status and disposition of the species are
uncertain, as is the identification of specimens from the West African
region.

Lithoporella melobesioides (Foslie) Foslie

Cape Verde Islands (366;598).

Mauritanie (366).

'Pantropical' (366).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]. . .' (598).

Note. The lectotype collection (678: 148), from South Nilandu,
Maldive Islands, has been studied in detail by Turner & Woelkerling
(1982fl, b), who also provided an account of the species in southern
Australia. All records of this species from the West African region
require confirmation. Errors associated with the typification of L.
melobesioides are indicated by Woelkerling (678: 148).

Lithoporella sauvageaui (Foslie) Adey

Canaries (18;582;598).

Cape Verde Islands (582;598).

[As Fosliella (Litholepis) sauvageaui (Foslie)]

Canaries (191).

[As Litholepis sauvageaui Foslie]

Canaries (6;139;203;204;227;366;368;493;678;687).

Cape Verde Islands (366;368).

[As Melobesia (Litholepis) sauvageaui Foslie]

Canaries (70;359;362).

Cape Verde Islands (100;633).

[As Melobesia sauvageaui Foslie]

Canaries (235).

Note. This species was originally described as Litholepis sau-
vageauii Foslie (203: 6) and is based on a single collection (678: 195)
from Puerto Orotava, Tenerife, Canary Islands. Subsequently, the
species was transferred to Melobesia (363: 66) and then to Litho-
porella (669: 15). There has been no detailed study of the holotype in
a modern context, and thus the status and disposition of the species
are uncertain, as is the identification of specimens from the West
African region.

Lithothamnion Heydrich, 1897a: 412, nom. cons.; non
Lithothamnium Philippi, 1837: 387.

The concept of Lithothamnion adopted in this paper follows
Woelkerling (1988: 169). The taxonomic history of
Lithothamnion Heydrich and of Lithothamnium Philippi are
summarized by Woelkerling (1983a), who proposed (Woelk-
erling, 19856) the conservation of Lithothamnion Heydrich
with L. muelleri Lenormand ex Rosanoff as type species. This
proposal has been approved and incorporated into the Inter-

national code of botanical nomenclature (Greuter , 1988: 116).
Further data on the nomenclature and infrageneric classifica-
tion are summarized by Woelkerling (1988: 173-175) who
also noted that no world monograph of the genus has been
published and that species concepts are poorly known. In the
literature, the spellings Lithothamnion and Lithothamnium
are both widespread. As the correct orthography is now
Lithothamnion, all entries are made under that spelling.

Lithothamnion amplexifrons (Harvey) Lemoine
See Lithophyllum amplexifrons (Harvey) Foslie.

Lithothamnion angolense Romanes

Angola (139;370;677).

Note. Lithothamnion angolense is based on fossil material from
four localities (677: 584). A detailed study of the type material
(whereabouts uncertain) has not been undertaken in a modern
context, and thus the status and disposition of this species are
uncertain.

Lithothamnion antarcticum (Hooker f . & Harvey)
Heydrich

See note to Mesophyllum ectocarpon (Foslie) Adey.

Lithothamnion bisporum Foslie

See Phymatolithon bisporum (Foslie) Afonso-Carrillo and
notes under Lithothamnion hispanum Foslie ex Gonzalez
Henriques.

Lithothamnion bornetii Foslie

Canaries (227; 598).

Note. This species originally was described as Lithothamnion
bornetii Foslie (18986: 9). Subsequently, Adey (669: 30) transferred
the species to Leptophytum (see comments on Leptophytum under
the entry for that genus), but then Adey & Adey (1973: 347, as
Lithothamnion ) indicated that the species was either a Leptophytum
or a Phymatolithon and that it occurred in France, Spain, and the
British Isles. Parke & Dixon (1976: 534) then placed the species in
Phymatolithon while South & Tittley (1986: 44) made the combina-
tion Phymatolithon bornetii (Foslie) Foslie. Foslie never placed this
species in Phymatolithon, and the binomial Phymatolithon bornetii
coined by both Parke & Dixon (1976) and South & Tittley (1986) is
invalid because a full reference to the basionym was not made in
accordance Article 33.2 of the International code of botanical nomen-
clature (see Greuter, 1988). Chamberlain (1990, as Leptophytum}
provided an account of British Isles material and of an isotype from
France in PC. Chamberlain (1990) also reported that some specimens
from France identified as 'bornetii' by Lemoine were misidentified
plants of Phymatolithon purpureum (P. & H. Crouan) Woelkerling &
L. Irvine, and she was unable to confirm the French and Spanish
records of Adey & Adey (1973). Information on the holotype is
provided by Chamberlain (1990) and Woelkerling (678: 41). Records
of this species from the Canary Islands require confirmation, and the
generic placement of the species requires further evaluation once the
status of Leptophytum as a genus is clarified.

Lithothamnion brachycladum Foslie

See Mesophyllum brachycladum (Foslie) Adey.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA
Lithothamnion brassica-floridum (Harvey) Areschoug

See Neogoniolithon brassica-florida (Harvey) Setchell &
Mason.

Lithothamnion calcareum (Pallus) Areschoug

See Phymatolithon calcareum (Pallus) Adey & McKibbin.

Lithothamnion californicum Foslie

See note on Phymatolithon tenuissimum (Foslie) Adey.

Lithothamnion canariense Foslie

See Mesophyllum canariense (Foslie) Lemoine.

Lithothamnion capense (Rosanoff) Foslie
See Lithophyllum capense Rosanoff.

Lithothamnion corallioides P. & H. Crouan

Canaries (15;18;582;648).
Cape Verde Islands (582).

Note. P. & H. Crouan (1867: 151, pi. 20, gen. 133, figs 8-10)
based Lithothamnion corallioides on material from Brest, France.
The type material has not been studied in detail in a modern context
and thus the status and disposition of this species are uncertain, as is
the identification of specimens from the West African region.

Lithothamnion corticiformis (Kiitzing) Foslie
See Melobesia membranacea Kiitzing.

Lithothamnion crassum Philippi

See Lithophyllum duckeri Woelkerling.

Lithothamnion crispatum Hauck

Mauritanie (356).

[As Lithothamnium Philippii Foslie = L. crispatum Hauck]

Mauritanie (354).

Note. Hauck (1878: 289) based Lithothamnion crispatum on
specimens from Rovigno in the Adriatic Sea. Subsequently, Hauck
(1883: 270) transferred the species to Lithophyllum. Foslie (1898a)
then transferred the species to Archaeolithothamnion, but later
(Foslie, 1904: 13) reclassified it as Lithothamnion philippii f. crispata
(Hauck) Foslie. There has been no detailed study of the type material
in a modern context, and thus the status and disposition of the species
is uncertain, as noted by Athanasiadis (668: 41). Once such a study is
undertaken, the specimens from Mauritanie need to be checked to
determine whether they are conspecific with the type.

Lithothamnion ectocarpon Foslie

See Mesophyllum ectocarpon (Foslie) Adey.

Lithothamnion erubescens Foslie

See Mesophyllum erubescens (Foslie) Lemoine.

Lithothamnion floridanum Foslie

See Mesophyllum floridanum (Foslie) Adey.

67

Lithothamnion fruticulosum (Kiitzing) Foslie
See Spongites fruticulosum Kiitzing.

Lithothamnion hispanum Foslie ex Gonzalez
Henriques

Canaries (598).

[As Lithothamnion hispanum Foslie]

Canaries (235).

Note. Gonzalez Henriques (235) listed this entity from the Canary
Islands and attributed the specific epithet to Foslie. As far as can be
determined, however, neither Foslie (see 678) nor other earlier
authors have described a species with the specific epithet hispanum,
and thus the epithet appears to be newly coined by Gonzalez
Henriques (235). The account of Gonzalez Henriques (235) lacks a
description, and consequently, Lithothamnion hispanum must be
considered a nomen nudum. It is possible that Gonzalez Henriques
(235) was referring to Lithothamnion bisporum (treated in this series
under Phymatolithon), a species based on type material from the
Canary Islands (678: 39), but this cannot be confirmed.

Lithothamnion incrustans Philippi
See Lithophyllum incrustans Philippi.

Lithothamnion indicum Foslie

Mauritanie (6).

Note. A detailed study of the lectotype collection (678: 125),
which comes from Corner Inlet, Victoria, Australia, is presently
being undertaken in conjunction with monographic studies of non-
geniculate corallines in southern Australia. Until that study is com-
pleted, the status and disposition of the species will remain uncertain.
Once the study of the lectotype is complete, the specimen on which
the record from Mauritanie is based needs to be checked to deter-
mine whether it is conspecific with the lectotype.

Lithothamnion irregulare Foslie

See Lithophyllum irregulare (Foslie) Huve ex Steentoft.

Lithothamnion lenormandii (Areschoug) Foslie
See Phymatolithon lenormandii (Areschoug) Adey.

Lithothamnion lichenoides Heydrich

See Mesophyllum lichenoides (Ellis) Lemoine.

Lithothamnion mamillare (Harvey) Areschoug

See Neogoniolithon mamillare (Harvey) Setchell & Mason.

Lithothamnion mamillosum Hauck

See Neogoniolithon mamillosum (Hauck) Setchell & Mason.

Lithothamnion membranaceum (Esper) Foslie
See Melobesia membranacea (Esper) Lamouroux.

Lithothamnion onkodes Heydrich

See Spongites onkodes (Heydrich) Penrose & Woelkerling
and the entry for Porolithon.

68

D.M. JOHNETAL

Lithothamnion orbiculatum Foslie

See Lithophyllum orbiculatum (Foslie) Foslie.

Lithothamnion philippi Foslie

Canaries (6;204;227;356;363;493;594;598).

Mauritanie (6;354;359).

Senegal (248).

'Subtropical Africa [Senegal (N. of Gambia); Mauritanie;

Former W. Sahara]' (598).
[As Mesophyllum philippi) (Foslie) Adey]
Canaries (18).

Note. The complex nomenclatural history of this species is
explained by Woelkerling (678: 171), who designated a new lectotype
from the Gulf of Naples, Italy. The lectotype, however, has not been
studied in detail in a modern context, and thus the status and
disposition of this species are uncertain, as is the identification of
specimens from the West African region. Afonso-Carrillo et al. (576:
25, as Mesophyllum ) commented that previous references to the
taxon from the Canaries are doubtful and require confirmation.

Lithothamnion polymorphum auctorum

See entries for Phymatolithon purpureum (P. & H. Crouan)
Woelkerling & L. Irvine and Phymatolithon calcareum (Pal-
las) Adey & McKibbin.

Lithothamnion ponderosum Foslie

Sao Tome (197;265).

Note. Foslie (197: 15) based this species on a specified collection
from Sao Tome. Subsequently Foslie (211: 42) considered
Lithothamnion ponderosum to be conspecific with Lithophyllum
africanum Foslie (199: 3) but incorrectly adopted the 1900 epithet
africanum rather than the 1897 epithet ponderosum. Steentoft (535:
128, under Lithophyllum africanum f. intermedia}, provides some
additional comments on the Sao Tome material, but the holotype
(678: 175) has not been studied in detail in a modern context and thus
the status and disposition of the species are uncertain.

Lithothamnion racemus (Lamarck) Areschoug

See notes under Mesophyllum brachydadum (Foslie) Adey.

Lithothamnion solutum (Foslie) Foslie

Cape Verde Islands (6;210).

Mauritanie (6;252;356).

[As Lithophyllum solutum (Foslie) Lemoine]

Mauritanie (248;359;360;556).

Salvage Islands (38B;556).

[As Mesophyllum solutum (Foslie) Lemoine]

Cape Verde Islands (366;598).

Mauritanie (349;366).

Salvage Islands (598).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Note. A detailed study of the lectotype collection (678: 203, as
Lithothamnion fruticulosum f. soluta Foslie) has not been undertaken
in a modern context and thus the status and disposition of this taxon
are uncertain, as is the identification of specimens from the West
African region. This taxon was originally described as Lithothamnion
fruticulosum f. soluta Foslie (1904: 7). Subsequently it was raised to
species status [as Lithothamnion solutum (Foslie) Foslie (204: 14)],
then transferred to Lithophyllum solutum (Foslie) Lemoine (356:
13), and later transferred to Mesophyllum solutum (Foslie) Lemoine
(366: 238; see also 349: 115). The binomial Mesophyllum solutum is

invalid because the combination was proposed without a full, direct
reference to its basionym (Article 33.2 of the International Code of
Botanical Nomenclature; see Greuter, 1988).

Lithothamnion sonderi Hauck

Canaries (177;191;227;362;365;375;582;584;598;631;649).
'. . . cotes atlantique d'Europe (de la Norvege aux Canaries)'

(188).

Note. A detailed study of the type collection, which comes from
Helgoland, Germany has been undertaken by Chamberlain (631),
who also provides data on the species in the British Isles. Specimens
on which published records for the West African region are based
need to be checked to determine whether they are conspecific with
the type.

Lithothamnion subtenellum Foslie

See Lithophyllum subtenellum (Foslie) Foslie.

Lithothamnion tenuissimum Foslie

See Phymatolithon tenuissimum (Foslie) Adey.

Lithothamnion vickersiae Lemoine

See Lithophyllum vickersiae Lemoine in B0rgesen.

Lithothamnion spp.

Angola (312).

Cape Verde Islands (114;145;366).
Ghana (335;350;487;586).
Namibia (348;437;438).
Western Sahara (349;476).

Lithothamnium

In the literature, the spellings Lithothamnion and Lithotham-
nium both are widespread. As the correct orthography is now
Lithothamnion, all entries are made under that spelling.

Lomentaria articulata (Hudson) Lyngbye

Canaries (2; 3; 8; 16; 38D; 70; 71; 191; 214; 226; 227; 229;

235; 375; 379; 392; 489; 490; 499; 517; 584; 598; 633).
Cameroun(239;350;454;500;586;591).
Salvage Islands (38B;38D;231;598).
Western Sahara (349).

'. . . Atlantique (de la Norvege au Rio de Oro). . .' (33).
'. . . Atlantique: depuis les cotes anglaises jusqu'au Rio de

Oro. . .' (222).
'A Boreal-Atlantic alga' (2).

'. . . Atlantique Nord (des Feroe aux Canaries). . .' (189).
'. . . Atlantischer Ozean von den Faroer an siidwarts bis

Westindien und Westafrika. . .' (499).
'. . . Atlantico norte (de Faeroes a Canarias)' (517).
'. . . Faeroes southwards to Morocco and the Canary

islands. . .' (70).
'. . . Nordwestafrika' (499).
'. . . Norway (Nordland) to Cameroun; Canary Isles. . .'

(273).

'. . . Tropical Africa (N. Gambia- Congo river)' (598).
'. . . West Kuste Afrikas' (239).
'. . . widespread from boreal-antiboreal to tropical seas'

(350;586).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

[As Chylodadia articulata Greville]

Canaries (439).

'. . . den atlantischen - Cap Inseln. . .' (239).

[As Lomentaria articulata Lyngbye]

Canaries (44;547).

[As Lomentaria articulata (Lightfoot) Lyngbye]

Canaries (401).

Note. Yarish et al. (591: 217-218) have commented upon the
significance of the ill-supported Cameroun statements.

Lomentaria bailey ana (Harvey) Farlow

Canaries (598;667).

Mauritanie (Cap Blanc) (516; 565 A).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).
[As Lomentaria bayleyana (Harvey) Farlow]
Mauritanie (349).

Note. In view of Bodard & Mollion's (59) records from Senegal, it
is possible that these records (or even the whole taxon) should be
transferred to (or referred to by the name of) Lomentaria uncinata
Meneghini. See notes under Lomentaria uncinata. Cribb (113: 71-72)
indicated the dichotomy of opinion that exists for the relationship of
L. baileyana and L. uncinata, some authors considering them conspe-
cific, others distinct.

Lomentaria exigua De Notaris
See Chylodadia reflexa Lenormand.

Lomentaria fastigiata Kiitzing

Sao Tome (132;323).

Note. Doubt expressed by De Toni (132) related to the possibility
that this was the western Atlantic island, not that in the Gulf of
Guinea. Included for completeness, since the situation is unclear.

Lomentaria fir ma (J. Agardh) Berthold

Senegal (59;399).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).
[As Lomentaria firma (J. Agardh) Kylin]
Senegal (531).

Note. See Lomentaria sp.l. Bodard & Mollion (59: 200) com-
mented as follows: '. . . Les deux Lomentaria sont plus originaux et
leur determination a ete delicate. Pour le premier nous avons trouve
dans 1'herbier Thuret un Chondria provenant de 1'ile de Goree
(Herbier Le Prieur) sans determination mais groupe avec le Lomen-
taria firma Berthold (Chylodadia firma J. Ag.). Cette espece pre"-
sente de nombreux synonymes, elle correspond au L. firma Kiitz.
Tab. Phyc. XV, t. 78 et vraisemblablement au Chondrothamnion
irregulare Kiitz. Tab. Phyc. XV, t. 82, decrite de la Mediterranee,
elle se retrouve au Maroc dans 1'herbier Schousboe, des echantillons
Zanardini sont egalement de la meme espece et proches des notres,
c'est done une espece africano-mediterraneene . . . les deux Lomen-
taria sont irreguliers au Senegal'.

Lomentaria impudica Montagne

See Catenella impudica (Montagne) J. Agardh.

Lomentaria kaliformis sine auctorum

See Chylodadia verticillata (Lightfoot) Batters.

69
Lomentaria linearis (Zanardini) Zanardini

Canary Islands (Prud'homme van Reine, unpublished; col-
lected during trip of Helgoland research vessel 'Heincke' to
the Canaries).

Lomentaria ovalis (Hudson) J. Agardh
See Gastrodonium ovatum (Hudson) Papenfuss.

Lomentaria parvula C. Agardh

See Champia parvula (C. Agardh) Harvey.

Lomentaria patens Kiitzing

Namibia (128).

Note. By straightforward nomenclatural synonymy, this record of
a single plant would probably have represented Chylodadia verticil-
lata (Lightfoot) Bliding (q.v.). However, from this named location,
the presence of that species as currently conceived is not credible. In
regard to a similar report by the same authors from South Africa,
Seagrief (570) has commented '? synonym of Chylodadia capensis
Harvey. . .' and the same comments could even more realistically be
made here for Namibia.

Lomentaria pygmaea auctorum

See Gastrodonium reflexum (Chauvin) Kiitzing.

Lomentaria reflexa Chauvin

See Gastrodonium reflexum (Chauvin) Kiitzing.

Lomentaria subdichotoma Ercegovic

Canaries (648).

Lomentaria uncinata Meneghini in Zanardini

Senegal (59).

'. . . cotes occidentales d'Afrique et aux Canaries. . .' (184).
Note. See Lomentaria sp. 2. Bodard & Mollion (59: 200) com-
mented as follows: '. . . Les deux Lomentaria sont plus originaux et
leur determination a ete delicate. . . La seconde est Lomentaria
uncinata Menegh. (1840) dont le synonyme principal est L. baileyana
. . . Farlow dont les varietes S et - sont identiques a 1'espece L.
uncinata. Cette espece peut-etre considered comme une tropicale
atlantique. Ces deux Lomentaria sont irreguliers au Senegal'. See
also notes under L. baileyana. Boudouresque et al. (1984: 45)
incorrectly cited the authorities for this taxon as (Meneghini ex
Kiitzing) Farlow.

Lomentaria uvaria (Wulfen) Duby

See Botryodadia botryoides (Wulfen) J. Feldmann.

Lomentaria sp.

Senegal (529).

Note. Sourie (529: 108) recorded it in his list as 'Lomentaria n. sp.
Feldm.' and in his notes (p. 116) as 'encore non decrit par J.
Feldmann'.

Lomentaria sp.l.

Senegal (55).
Note. See notes under Lomentaria firma (J. Agardh) Berthold.

70

Lomentaria sp.2.

Senegal (55).
Note. See Lomentaria uncinata Meneghini.

Lophocladia trichoclados (C. Agardh) Schmitz

Annobon (139;292;456;457).

Canaries ( 128 A ; 139 ;292 ;633 ;634 ;648 ;662) .

Ghana (290;292;299;300;350;376;377;491 ;586).

Principe (139;350;586).

Sierra Leone (30;350;586).

'. . . widespread in warm temperate and tropical seas..'

(292?;350;586).
[As Lophocladia trichoclados (Mertens in C. Agardh)

Schmitz]

Canaries (71;227;303;598;648).
Cape Verde Islands (598).
Principe (93).
Sao Tome (93).

Tropical Africa (N. Gambia - Congo river)' (598).
[As Lophocladia trichoclados (Mertens) Schmitz]
Canaries (13;38D;191;226;375).
[As Dasya trichoclados J. Agardh]
Canaries (439;547).

Lophosiphonia adhaerens Pilger

Annobon (139 ;295 ;350 ;456 ;457 ;586) .
?Sierra Leone (295).
St. Helena (655).
Note. See Lophosiphonia spp. for possible Sierra Leone records.

Lophosiphonia cristata Falkenberg

Ascension Island (474).

Canaries (38B;634;657).

Salvage Islands (38B;598;657).

[As Lophosiphonia cf. cristata Falkenberg]

Canaries (38C).

Note. Audiffred (38C: 179-180) indicated that his specimens did
not have the usual hooked apices of branchlets associated with L.
cristata, nor did the rhizoids remain in open connection with the
pericentrals. He debated the wide variation in reported numbers of
pericentral cells in Lophosiphonia (4-18 according both to species
and author); his own specimens had 9 pericentrals. The difficulties in
distinguishing between sterile Lophosiphonia and Polysiphonia were
indicated.

Lophosiphonia obscura (C. Agardh) Falkenberg in
Engler & Prantl

See under Lophosiphonia reptabunda (Suhr) Kylin.

Lophosiphonia reptabunda (Suhr in Kiitzing) Kylin

Angola (352).

Ascension (475).

Bioko (346;350;586).

Cameroun (350;586).

Canaries (38B;38C;38D;54;110;556;598;662;684).

Cape Verde Island (598).

Cote d'lvoire (350;586).

Gabon (350;586).

Ghana (288;350;586).

Salvage Island (38B;38C;38D;556;598).

D.M. JOHNETAL

Sierra Leone (350;586).

St. Helena (655).

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

'. . .in warm temperate and tropical parts of the Atlantic. . .'

(350;586).

'. . .on many rocky shores of West Africa. . .' (347 A).
Tropical Africa (N. Gambia - Congo river)' (598).
'. . . warmer parts of the Atlantic Ocean. . .' (71).
[As Lophosiphonia reptabunda (Kiitzing) Kylin]
Angola (298).
Gabon (295).
Sierra Leone (295).
[As Lophosiphonia reptabunda Kylin]
Canaries (489;517).

[As Lophosiphonia reptabunda (Suhr in Kiitzing) Cribb]
Canaries (110).

[As Lophosiphonia obscura J. Agardh]
'. . . cotes occidentale d'Afrique et aux Canaries. . .' (184).
[As Lophosiphonia obscura (C. Agardh) Falkenberg]
Cameroun (337;537).
Canaries (108;227;375;482;584).
Cape Verde Islands (38).
Ghana (153;338;487).
'. . . in den warmeren Teilen des Atlantischen Ozeans. . .'

(501).

'. . . warmer parts of the Atlantic Ocean' (62).
[As Lophosiphonia obscura Falkenberg]
Canaries (190).

[As Lophosiphonia obscura auct., Howe]
Canaries (71;191;310).

'. . . in warmer parts of the Atlantic Ocean. . .' (71).
[As Lophosiphonia sp.]
Ascension Island (474).
Sierra Leone (339).
[As Polysiphonia obscura J. Agardh]
Cape Verde Islands (38;145).

'. . . au sud de 1'Angleterre aux Canaries. . .' (38;89).
[As Polysiphonia reptabunda Suhr]
'Ad oras africanas' (321).
'. . . In mari atlantico subtropico ad Alga parasitica. . .'

(318).

Note. Erroneously recombined by Jaasund (280: 411) as Lopho-
siphonia reptabunda (Suhr) nov. comb, who seemed to be unaware of
the fact that both Cribb and Kylin had carried out such recombina-
tion independently in 1956. Of these it appears that Kylin (281)
antedated Cribb (110) who discussed (p. 140-141) its nomenclature.
Kapraun et al. (310: 890-891) discussed the validity of records by
B0rgesen (71) from the Canaries, considering that the material
concerned could not represent Lophosiphonia obscura and is prob-
ably a species of Polysiphonia. It would appear that Lophosiphonia
obscura as such occurs north of the area considered here.

Lophosiphonia scopulorum (Harvey) Womersley

Canaries (633;634;648).
Salvage Islands (38B; 556).
Note. See also Polysiphonia scopulorum Harvey.

Lophosiphonia v ilium (J. Agardh) Setchell & Gardner

See Polysiphonia scopulorum Harvey var. villum (J. Agardh)
Hollenberg.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

71

Lophosiphonia spp.

Ascension (474) (probably in the main referable to L. rept-

abunda).
Cameroun (460).
Cape Verde Island (652).
Ghana (299;376;377).
Senegal (529).
Sierra Leone (295;339;350;468;586).

Note. The Lophosiphonia material referred to in 339 and 460 may
well be L. reptabunda but the specimens cannot now be traced. The
species referred to in 350 and 586 closely resembles the little-known
L. adhaerens Pilger (456).

Lophura episcopalis (Montagne) Kutzing
See Halopitys incurvus (Hudson) Batters.

Lythophyllum calcareum (Pallas) Areschoug

See Phymatolithon calcareum (Pallas) Adey & McKibbin.

Lythophyllum sp. (geometricum?) Lemoine

See Lithophyllum geometricum Lemoine and Dermatolithon
geometricum (Lemoine in B0rgesen) Price, John & Lawson.

Lythophyllum sp.

See Lithophyllum sp.

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

Canaries (598;633;634;657).

'. . . from northern Russia south to Portugal and from

Morocco south possibly to Rio de Oro, Mauritania. . .'

(245;657).

[As Gigartina stellata (Stackhouse in Withering) Batters]
Canaries (13;227).
Mauritanie (349).
Western Sahara (349).

'. . . Atlantique (de 1'Arctique au Rio de Oro)' (33).
'. . . Atlantique: depuis la Scandinavie jusqu'au Rio de

Oro. . .' (222).
'Circumboreale,. . . de 1'Arctique aux Cotes Atlantique

d'Europe et d'Afrique jusqu'a la Mauritanie. . .' (173)

Note. The distributional statement in 657 is reproduced directly
from Guiry et al. (245). Afonso-Carrillo et al. (657: 289) present the
first report from the Canaries of what is probably the Petrocelis
cruenta phase (the sporophyte) of M. stellatus; unfortunately, the
material was sterile.

Mastophora Decaisne

The concept of Mastophora adopted in this paper follows
Woelkerling (1988: 129-133) who provides nomenclatural
and systematic data on the genus. Historical data on the
genus are summarized by Turner & Woelkerling (1982a, b),
who also give an account of the lectotype species, M.
licheniformls Decaisne [a heterotypic synonym of M. rosea
(C. Agardh) Setchell]. A revised key to the genera of
Mastophoroideae, including Mastophora, is provided by Pen-
rose & Chamberlain (1993: 303). According to Turner &
Woelkerling (I982b: 233) and Woelkerling (1988: 133),

uncertainties surrounds the delimitation and circumscription
of most species in the genus.

Mastophora conjuncta Foslie

See Lithoporella conjuncta (Foslie) Foslie.

Mastophora melobesioides Foslie

See Lithoporella melobesioides (Foslie) Foslie.

Melobesieae

Certain records have been omitted on the grounds of impon-
derability. An example is the organism(s) referred to by
Viero y Clavijo (548) under the name Bellaria Lapidea
Canariensia. Martin Aguado (386), in analysing the treat-
ments of algae by Viero y Clavijo (548), concluded that
melobesioids were being referred to in this entry, otherwise
headed 'Confites', but that it was not possible to be more
precise than 'subfamilia Melobesieae'.

Melobesia Lamouroux

The concept of Melobesia adopted in this paper follows
Woelkerling (1988: 186-191), who also provides information
on the nomenclatural and taxonomic history of the genus.
Many species that at some stage were ascribed to Melobesia
are now placed in other genera.

Melobesia amplexifrons Harvey
See Pneophyllum amplexifrons.

Melobesia brassica-florida Harvey

See note to Neogoniolithon brassica-florida (Harvey) Setchell
& Mason.

Melobesia callithamnioides sensu Falkenberg

See references for Melobesia callithamnioides under Fosliella
farinosa (Lamouroux) Howe.

Note. Chamberlain (1983: 351-352) provided a detailed historical
account of Melobesia callithamnioides sensu Falkenberg and con-
cluded that Melobesia callithamnioides sensu Falkenberg was conspe-
cific with Fosliella farinosa (Lamouroux) Howe [=Hydrolithon
farlnosum (Lamouroux) Penrose & Chamberlain]; see Penrose &
Chamberlain, 1993], where she treated it as a distinct variety under
the name Fosliella farinosa f. callithamnioides (Foslie) Chamberlain.
Chamberlain (1983: 352), however, was unable to confirm this
placement by comparative examination of type material and indeed
did not designate a lectotype. Until a lectotype is designated and
studied in detail in a modern context, the status and disposition of
Fosliella farinosa f. callithamnioides (Foslie) Chamberlain will remain
uncertain, as will records of this taxon from the West African region.

Melobesia confervicola (Kutzing) Foslie

See Pneophyllum confervicolum (Kutzing) Chamberlain.

Melobesia confinis P. & H. Crouan

See Lithophyllum pustulatum (Lamouroux) Foslie.

72

D.M. JOHNETAL

Melobesia corallinae P. &. H. Crouan

See Lithophyllum corallinae (P. & H. Crouan) Heydrich.

Melobesia corticiformis Kiitzing

See Melobesia membranacea (Esper) Lamouroux.

Melobesia cystoseirae Hauck

See Lithophyllum cystoseirae (Hauck) Heydrich.

Melobesia farinacea Lamouroux

See Fosliella farinosa (Lamouroux) Howe.

Note. Based on a comparative study of relevant type and other
collections, Penrose & Chamberlain (1993) have concluded that the
genus Fosliella, which is typified by F. farinosa, is a heterotypic
synonym of Hydrolithon, and they have transferred Lamouroux's
species to that genus, as Hydrolithon farinosum (Lamouroux) Pen-
rose & Chamberlain.

Melobesia farinosa Lamouroux

See Fosliella farinosa (Lamouroux) Howe and note under
Melobesia farinacea Lamouroux.

Melobesia lejolisii Rosanoff

See Pneophyllum lejolisii (Rosanoff) Y. Chamberlain.

Melobesia mamillaris Harvey

See Neogoniolithon mamillare (Hauck) Setchell & Mason.

Melobesia membranacea (Esper) Lamouroux

Canaries ( 191 ;227 ;232B ;439 ;441 ;582 ;584 ;598 ;633 ;634) .

Cape Verde Islands (408;598).

Mauritanie (252;349;439;441).

Salvage Islands (598).

'. . . regions temperees et chaudes de 1'Atlantique. . .' (188).

'Subtropical Africa [Senegal(N. of Gambia); Mauritania;

Former W. Sahara]' (598).
[As Melobesia membranacea (Esper) Foslie]
Cape Verde Islands (408).
[As Melobesia membranacea Lamouroux]
Cape Verde Islands (38).
Congo (249;250).

[As Melobesia corticiformis Kiitzing]
Canaries (547).
Cape Verde Islands (41;42).
[As Epilithon membranaceum (Esper) Heydrich]
Angola (541).

Canaries (70;3 14 ;354;356 ;359 ;362 ;363 ;5 17 ;624) .
Cape Verde Islands (366).
Congo (356).
Mauritanie (356;359).
Salvage Islands (38B;231;375).
Senegal (356).
'. . . optima verio in mari atlantico ab oris Norvegiae usque

ad promontorium Cap b. spei. . .' (25).
'. . . regions temperees et chaudes de 1'Atlantique. . .' (355).
[As Lithothamnion membranaceum (Esper) Foslie]
Cape Verde Islands (252).
Mauritanie (252).

Senegal (252).

'. . .in oceano Atlantico ab oris Norvegiae usque ad promon-
torium Capitis Bonae Spei Africae australis. . .' (134).
'Nordwestafrika' (499).

Note. This species originally was described as Corallina mem-
branacea Esper (1806: Corallina Tab. XII). Esper (1806) did not
indicate the locality from which his material came, but subsequently
(1830: 363, as Melobesia) stated that the species occurred on the
western shores of France and probably in other regions. Lamouroux
(1812: 186) transferred the species to his new genus Melobesia, and as
noted by Mason (1953: 319), Chamberlain (1985: 673) and Woelker-
ling (1988: 189), M. membranacea (Esper) Lamouroux must be
considered the type species of Melobesia. The species also has been
placed in Epilithon (Heydrich, 1897a: 408) and in Lithothamnion
(681: 7). Chamberlain (1985) neotypified the species with a Lamour-
oux specimen (CN); accounts of the neotype are provided by
Chamberlain (1985) and Wilks & Woelkerling (1991), the latter in
conjunction with a monographic account of southern Australian
species of Melobesia. Specimens on which published records for the
West African region are based now need to be checked to determine
whether they are conspecific with the neotype. Chamberlain (1983:
300, 306) concluded from studies of the type of Melobesia corticifor-
mis Kiitzing that it was a heterotypic synonym of Melobesia mem-
branacea. References to Melobesia corticiformis above are based on
comments of Lemoine (363: 29) who indicated that these specimens
were misidentified plants of M. membranacea (as Epilithon ).

Melobesia minutula Foslie

See Pneophyllum confervicolum (Kiitzing) Y. Chamberlain.

Melobesia pustulata Lamouroux

See Lithophyllum pustulatum (Lamouroux) Foslie.

Melobesia sauvageauii Foslie

See Lithoporella sauvageaui (Foslie) Adey.

Melobesia solmsiana Falkenberg

See references for Melobesia solmsiana under Fosliella fari-
nosa (Lamouroux) Howe.

Note. Chamberlain (1983: 351-352) concluded that Melobesia
solmsiana Falkenberg was conspecific with Fosliella farinosa (Lam-
ouroux) Howe [= Hydrolithon farinosum (Lamouroux) Penrose &
Chamberlain; see Penrose & Chamberlain, 1993] and treated it as a
distinct form under the name Fosliella farinosa f. callithamnioides
(Foslie) Chamberlain. Chamberlain (1983: 352), however, was
unable confirm this placement by comparative examination of type
material and indeed did not designate a lectotype. Until a lectotype is
chosen and studied in detail in a modern context, the status and
disposition of Melobesia solmsiana Falkenberg and Fosliella farinosa
f. callithamnioides (Foslie) Chamberlain will remain uncertain, as will
records of this taxon from the West African region.

Melobesia solmsii Bornet ex Lemoine

Canaries (356).

Note. Melobesia solmsii is an herbarium name of Bornet that was
first published by Lemoine (354: LXIV), who suggested that Bornet's
plants were the same as plants identified by Falkenberg as Melobesia
callithamnioides . Subsequently, Lemoine (356: 26) recorded Melobe-
sia solmsii from various regions, including the Canary Islands. The
Canary Islands record was listed in Price et al. (1986: 92) under their
entry for Fosliella farinosa. The nomenclatural history of Melobesia
callithamnioides sensu Falkenberg (1879) has been summarized by
Chamberlain (1983: 352), who noted that Falkenberg (179: 109)
subsequently described his material as a new species, Melobesia

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

73

solmsiana. Given these facts, Melobesia solmsii Bornet ex Lemoine
must be interpreted as a homotypic synonym of Melobesia solmsiana
Falkenberg. Chamberlain (1983: 351-352) concluded that Melobesia
solmsiana Falkenberg was conspecific with Fosliella farinosa (Lam-
ouroux) Howe [=Hydrolithon farinosum (Lamouroux) Penrose &
Chamberlain); see Penrose & Chamberlain, 1993] and treated it as a
distinct variety under the name Fosliella farinosa L callithamnioides
(Foslie) Chamberlain. Chamberlain (1983: 352), however, was
unable to confirm this placement by comparative examination of type
material and indeed did not designate a lectotype. Until a lectotype is
chosen and studied in detail in a modern context, the status and
disposition of Melobesia solmsiana Falkenberg (including M. solmsii
Bornet ex Lemoine) and Fosliella farinosa f. callithamnioides (Foslie)
Chamberlain will remain uncertain, as will records of this taxon from
the West African region.

Melobesia stictaeformis Areschoug

Salvage Islands (439).

Note. Areschoug (1852: 517) based this species on a collection
from the Mediterranean. A detailed study of the type (presumably in
LD), however, has not been undertaken in a modern context, and
thus the status and disposition of this species are uncertain, as is the
record from the Salvage Islands.

Meredithia microphylla (J.Agardh) J.Agardh

Canaries (227;598;661).

Cape Verde Islands (598).

[As Callymenia microphylla J. Agardh]

Canaries (70; 177).

[As Callymenia (Meredithia) microphylla J. Agardh]

Canaries (493).

[As Callymenia reniformis (Turner) J.Agardh]

Canaries (547).

[As Kallymenia microphylla J. Agardh]

Canaries (191 ;273;392;584).

Cape Verde Islands (37).

Note. For background data on this species, see Guiry & Maggs
(661). See note under Kallymenia microphylla J. Agardh in Price et
al. (1992).

Meristiella echinocarpa (Areschoug) Cheney &
Gabrielson

[As Meristotheca! decumbens Grunow in Piccone]

Cape Verde Islands (439).

[As Rhabdonia decumbens (Grunow) Grunow in Askenasy]

Cape Verde Islands (37)

[As Mychodea schrammii P. & H. Crouan]

Cape Verde Islands (38;450;451).

Note. Prud'homme van Reine et al (663) have re-investigated the
Macaronesian algae studied by Piccone (see 439,450,451) and
Grunow (see 37,38) and concluded that Askenasy had in error
identified both Meristiella echinocarpa (from the Cape Verde Islands)
and Meristotheca ? decumbens (from Madeira, the Canaries and also
from the Cape Verde Islands) as Mychodea schrammii. The speci-
mens collected by Naumann (450, 451) have not been studied in
detail by Prud'homme van Reine et al. (663).

Meristiella schrammii (P. & H. Crouan) Cheney &
Gabrielson

See Meristiella echinocarpa (Areschoug) Cheney & Gabriel-
son.

Meristotheca coacta Okamura

Ghana (299).

Note. See under Halichrysis depressa (J. Agardh) Schousboe in
Bornet for the validity of this record based on sterile plants.

Meristotheca ? decumbens Grunow in Piccone

Canaries (439)

Cape Verde Islands (439;450;451).

Note. See note under Meristiella echinocarpa (Areschoug)
Cheney & Gabrielson.

Meristotheca duchassaingii J. Agardh

See Halymenia duchassaingii (J.Agardh) Kylin.

Meristotheca senegalense J.Feldmann [mscr.]

Senegal (529;531).

[As Meristotheca senegalensis J. Feldmann]

Senegal (59;398).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).
[As Sarcodia ceylanica Harvey]
Senegal (122).

Note. According to Mollion (398) this is a manuscript name. Prior
to this, Sourie (529: 116) had noted '. . . espece nommee, mais
encore non decrite par J. Feldmann'.

Meristotheca sp.

Gabon (294).
Senegal (529).

Note. For comments on validity of Gabon record, see note to
Halichrysis depressa (J. Agardh) Schousboe in Bornet.

Mesophyllum

The concept of Mesophyllum adopted in this paper follows
Woelkerling & Harvey (1993), who provide an updated
description of the genus taking into account new data on
spermatangial conceptacles (Woelkerling & Harvey, 1992).
Additional nomenclatural and systematic data on Mesophyl-
lum are provided by Woelkerling & Irvine (19866) and
Woelkerling (1988: 191-196)

Mesophyllum applicatum Lemoine

See Neogoniolithon hirtum (Lemoine in B0rgesen) Afonso-
Carrillo.

Mesophyllum brachycladum (Foslie) Adey

Canaries (598).

Principe (350;535;586).

St. Helena (535).

'. . . in warm temperate and tropical parts of the eastern

Atlantic Ocean. . .' (350;586).
Tropical Africa (N. Gambia - Congo river)' (598).
[As Lithothamnion brachycladum Foslie]
Angola (541 ;634).
Canaries (6;134;198;354;356;634)
Principe (134;363).
St. Helena (6;134;198;212;634;655).
'. . . West Coast of Africa. . .' (208).
'. . . probably nearly all along the west coast of Africa' (198).

74

D.M. JOHNETAL

[As Lithothamnion racemus (Lamouroux) Foslie]

Canaries (547).

St. Helena (142;391).

Note. This species originally was described as Lithothamnion
brachycladium (198: 3) and is based on material from St. Helena.
Subsequently, Adey (669: 22) transferred the species to Mesophyl-
lum, expressing some doubt about this placement. The holotype
(678: 42-43) has not been examined in detail in a modern context,
and thus the status and disposition of the species are uncertain, as is
the identification of specimens from the West African region. In the
protologue, Foslie (198: 3) suggested that the holotype of Lithotham-
nion brachydadum is probably the same specimen that Dickie (142:
179) referred to Lithothamnion racemus, and thus the same specimen
on which the record of Mellis (391: 382) is based. Although the
Dickie and Mellis records are included under brachydadum, Foslie's
suggestion requires confirmation. Lemoine (354: LIV) noted that a
Canary Islands specimen in the Thuret herbarium under the name
Lithothamnion branchydadium had been determined by Foslie as
Lithophyllum racemus.

Mesophyllum canariense (Foslie) Lemoine in B0rgesen

Canaries (191 ;362 ;363 ;375 ;489 ;493 ;535 ;582 ;598) .

Sao Tome (350;535;586).

'Gulf of Guinea' (582).

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

'. . . widespread in warm temperate and tropical seas. . .'

(350;586).

[As Lithophyllum marlothii Heydrich]
SaoTome(6;251;265).

[As Lithophyllum Marlothii Heydrich forma Foslie]
Sao Tome (251).

[As Lithothamnion canariense Foslie]
Canaries (6 ; 139 ;205 ;212 ;493 ;660) .
[As Lithothamnion canaricae Foslie]
Canaries (387).

[As Mesophyllum canariense (Foslie) Lemoine]
Canaries (191;227;361;687).
[As Mesophyllum canariensis (Foslie) Lemoine]
Canaries (229).
[As Mesophyllum canariense (Foslie) Lemoine var. fasciata

Lemoine]
Canaries (363).
[As Mesophyllum canariense (Foslie) Lemoine var. difformis

Lemoine]
Canaries (363).

[As Lythophyllum canariensis ?]
Canaries (237).

Note. This species originally was described as Lithothamnion
canariense (205: 17) and is based on material from Puerto Orotava,
Tenerife, Canary Islands. Lemoine (363: 31) subsequently trans-
ferred the species into Mesophyllum. The holotype (678: 46) has
recently been studied in a modern context by Reyes & Afonso-
Carrillo (687), who state that it certainly belongs to the genus
Mesophyllum as delineated by Woelkerling & Irvine (19866). The
identification of specimens from much of the West African region
remains uncertain. Both Mesophyllum canariense var. difformis
Lemoine (363: 33) and Mesophyllum canariense var. fasciata Lem-
oine (363: 31) are based on single collections from the Canary
Islands, but the holotypes have not been examined in detail in a
modern context, and thus the status and disposition of these varieties
are also uncertain. According to Steentoft (535: 130), specimens
from Sao Tome referred to Lithophyllum marlothii by Hariot (251:
164) and Henriques (265: 166) are considered by Lemoine to belong
to Mesophyllum canariense. Similarly, Adey & Lebednik (6: 19)
indicate that one specimen listed under Lithophyllum marlothii really
belongs to Mesophyllum canariense. These specimens need to be

re-examined to determine whether they are conspecific with the
holotype of Mesophyllum canariense. Gonzalez (237: Table II) lists a
Lythophyllum canariensis from the Canary Islands without further
taxonomic or nomenclatural comments. It is presumed here that
Gonzalez is referring to Mesophyllum canariense. As far as can be
determined, the binomial Lithophyllum canariensis has never been
validly published.

Mesophyllum ectocarpon (Foslie) Adey

Canaries (17;598;634).

Cape Verde Islands (366;597;598).

Mauritanie (349;597).

Senegal (366).

'Makaronesische Inseln, Nordwest und West-Afrika' (597).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

former W. Sahara]' (598).
[As Lithothamnion ectocarpon Foslie]
Canaries (136;191;227;362;363;366).
Cape Verde Islands (6;136;139;207;210;362;363;366).
Mauritanie (6;139;207;210;212;362;363;366).
'Vestkysten af Afrika: Kap Blanco . . . og St. Vincent' (207).
Senegal (6).

Note. This species was originally described as Lithothamnion
ectocarpon Foslie (207: 11) based on specimens from the Cape Verde
Islands and Cap Blanc, Africa. Adey (in 6: 83) lectotypified the
species with the Cap Blanc collection and subsequently (669: 23)
transferred it to Mesophyllum with some question. The lectotype
(678: 82) has not been studied in detail in a modern context, and thus
the status and disposition of the species are uncertain, as is the
identification of specimens from the West African region.

Mesophyllum erubescens (Foslie) Lemoine

Canaries (366).

Cape Verde Islands (100;101;366;598).

'Pan tropical' (366).

Note. This species was originally described as Lithothamnion
erubescens (198: 9) and is based on material from Brazil. Lemoine
(361: 252) transferred the species into Mesophyllum without com-
ment. The holotype (678: 85) has not been studied in detail in a
modern context and thus the status and disposition of the species are
uncertain, as is the identification of specimens from the West African
region. Afonso-Carrillo et al. (582) commented that despite previous
references to the taxon from the Canary Islands, its presence there is
doubtful and requires confirmation.

Mesophyllum floridanum (Foslie) Adey

Sao Tome (350;586).

[As Lithothamnion floridanum Foslie]

Sao Tome (6;139;204;535).

Note. This species was originally described as Lithothamnion
floridanum (204: 11) and was based on material from Florida. In the
protologue, Foslie (204: 12) also mentions two sterile specimens from
Sao Tome which resemble Lithothamnion floridanum but cannot be
definitely identified to species. The holotype (678: 96) has not been
studied in detail in a modern context and thus the status and
disposition of the species are uncertain, as is the identification of
specimens from Sao Tome.

Mesophyllum lichenoides (Ellis) Lemoine

Canaries (188;191;227;232B;361;363;517;546;582;584;598;

633;666).
Mauritanie (649).
'. . . Atlantico (de Gran Bretana a Marruecos y Canarias)'

(517).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

'. . . Atlantique (de 1'Angleterre a la Mauritanie, Canaries)'

(33).

'. . . Atlantique jusqu'au Rio de Oro. . .' (222).
[As Mesophyllum lichenoides (Linnaeus) Lemoine]
Mauritanie (349).
Western Sahara (349).
[As Lithophyllum lichenoides Philippi]
Canaries (70;527).

[As Lithophyllum expansum Philippi]
Canaries (356;448;584).
Salvage Islands (89).
'Cap Blanc' (356).

[As Lithophyllum expansum Philippi f. involvens Vinassa]
Mauritanie (211).
[As Lithophyllum expansum Philippi f. strictaeformis

(Areschoug) Foslie]
Salvage Islands (215).

[As Lithophyllum expansum Philippi f . exigua Foslie]
'Cap Blanc' (252).

[As Pseudolithophyllum expansum (Philippi) Lemoine]
Canaries (18;70;177;221;227;598).
Mauritanie (70;177;188;221;248;349;359;365).
Salvage Islands (38B;598)
'Subtropical Africa [Senegal (N. of Gambia); Mauritanie;

Former W. Sahara]' (598).
'. . . existe aussi sur les cotes atlantique d'Espagne et sur les

cotes d'Afrique jusqu'an Senegal . . .' (365).

Note. This species originally was described as Corallium
lichenoides (Ellis, 1768: 407) and was probably based on material
from Cornwall, England. The subsequent taxonomic and nomencla-
tural history of Mesophyllum lichenoides is outlined by Woelkerling
& Irvine (19866), who neotypified the species with plants from West
Looe, Cornwall, England and provided a detailed account of the
neotype material and the species. Additional data on M. lichenoides
are provided by Woelkerling & Harvey (1992, 1993) in conjunction
with studies of southern Australian species of Mesophyllum. Speci-
mens on which published records for the West African region are
based now need to be checked to determine whether they are
conspecific with the neotype. The lectotype of Lithophyllum expan-
sum Philippi has been examined in detail by Woelkerling (19836:
307-313) who concluded that the lectotype belonged to Mesophyllum
as delimited by Johansen (1976, 1981) and Cabioch (1972) and that
circumstantial evidence suggested that the lectotype was conspecific
with Mesophyllum lichenoides. Woelkerling (19836: 312) also noted
that Philippi's epithet expansum has been widely misapplied to plants
referable to Lithophyllum. Thus all specimens from the West African
region identified as Lithophyllum expansum or Pseudolithophyllum
expansum need to be re-examined to determine whether they belong
to the Lithophylloideae or the Melobesioideae and then to determine
their true generic and specific identity. It is likely that many of these
plants belong to Lithophyllum rather than Mesophyllum. This also
applies to specimens identified as Lithophyllum expansum Philippi f .
exigua Foslie (197: 3), Lithophyllum expansum Philippi f. involvens
Vinassa (1892: 59), and Lithophyllum expansum Philippi f. strictae-
formis (Areschoug) Foslie (199: 18) [Basionym: Melobesia strictae-
formis Areschoug, 1852: 517]. The types of these formae also need to
be re-examined in a modern context to determine their status and
disposition. Data on the types of Lithophyllum expansum Philippi f.
exigua Foslie and Melobesia stictaeformis Areschoug are provided by
Woelkerling (678: 88, 207); the whereabouts of the type of Litho-
phyllum expansum Philippi f. involvens Vinassa is uncertain. See
note under entry for Lithophyllum lobatum concerning Sauvageau's
(493) and Foslie's (211) record of 'Lithopyllum expansum' from the
Canaries.

75

Mesophyllum lobatum Lemoine
See Lithophyllum lobatum Lemoine in B0rgesen.

Mesophyllum philippii (Foslie) Lemoine
See Lithothamnion philippii Foslie.

Mesophyllum solutum (Foslie) Lemoine, nominum
invalidum

See Lithothamnion solutum (Foslie) Foslie.

Mesophyllum sp.

Canaries (229;237).
Cape Verde Islands (366).

Note. Lemoine (366: 236) indicated: '. . . deux especes indeter-
minees, un Lithothamnion et un Mesophyllum ce dernier appartient
peut-etre a une espece non decrite des Canaries'.

Mesothamnion caribaeum B0rgesen

See Pleonosporium caribaeum (B0rgesen) R. Norris.

Microcladia capensis (Kutzing) Papenfuss
See the note to Ceramium capense Kutzing.

Microcladia glandulosa (Solander ex Turner) Greville

Gambia (296).

Senegal (133).

Senegambia (296).

'. . .ad litus Senegambiae. . .' (318).

[As Microcladia glandulosa Greville]

Senegal (283).

[As Microcladia tenuis Kutzing]

'Gabon, Guinea' (149).

'Embouchure de la riviere de Gabon, Guinee' (321).

Microcladia tenuis Kutzing

See Microcladia glandulosa (Solander ex Turner) Greville.

Micropeuce mucronata (Harvey) Kylin

Ghana (350;376;377;586).

'. . . probably widespread in many warm temperate and

tropical seas' (350;586).
'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).
Tropical Africa (N. Gambia - Congo river)' (598).

Monospora furcellata auctorum
See Griffithsia arachnoides C. Agardh.

Monospora pedicillata Solier in Castagne

See Monosporus pedicillatus (J. E. Smith) Solier in Castagne.

76

D.M. JOHNETAL

Monosporus pedicillatus (I.E. Smith) Solier in
Castagne

Canaries (71;633;668;684).

[As Monospora pedicellata Solier]

Canaries (547).

[As Corynospora pedicellata (J.E. Smith) J. Agardh]

Canaries ( 1 3 ;38D ;227 ;375 ;547 ;556;584) .

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

[As Neomonospora pedicillata (Smith) G. Feldmann & Mes-

lin]

Canaries (191).

'. . . Atlantique (de 1'Angleterre aux Canaries). . .' (33).
'. . . Atlantique nord: de la Grande-Bretagne jusqu'aux

Canaries. . .' (222).
'. . . Atlantique nord, de 1'Angleterre aux Canaries. . .'

(190).
'. . . Atlantique nord, de Grande Bretagne aux Canaries. . .'

(196).
[As Neomonospora furcellata (J. Agardh) G. Feldmann &

Meslin]
Canaries (191).

Murray el la periclados (C. Agardh) Schmitz

Angola (298;347A;352;458;466).

Bioko (346;350;586).

Cameroun(337;350;460;463;535;537;586).

Congo (535).

Gabon (458;471).

Gambia (296;350;586).

Ghana ( 153 ;336;337 ;338 ;342 ;344 ;350;458 ;460 ;463 ;464 ;466 ;

473;535;586).
Guinee (350;460;535;586).
Sao Tome (350;458;466;470;473;535;586).
Sierra Leone (30;295;336;339;344;350;374;378;458;460;464;

466 ;468 ;470;473 ;535 ;586) .

'. . . present in many parts of West Africa. . .' (347 A).
Tropical Africa (N. Gambia - Congo river)' (598).
'West Africa' (290).
'. . . widespread in warm temperate and tropical seas. . .'

(350;586).

Note. Post's (458) Angola record is based on her redetermination
of Welwitsch no. 60 (BM), from Cabo Lombo, Loanda, under the
manuscript name of Ceramium arachnoideum Welwitsch (q.v.). The
record (460) from Cameroun does not relate to Post's comments
(458: 79) in redetermining 'Bostrychia periclados' as Bostrychia
moritziana (Sonder ex Kiitzing) J. Agardh (q.v.), rather than Mur-
ray >ella periclados. Records from Gabon are said (458; 471: 150) to be
from 'siisswasser' but are included for completeness.

Murrayella sp.

Sierra Leone (460).

Mychodea schrammii P. & H. Crouan

See Meristiella echinocarpa (Areschoug) Cheney & Gabriel-
son.

Myriogramme costata P. Dangeard

Senegal (89;122;221).

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

Note. See Nitophyllum dentatum for doubt concerning the separa-
tion of the genus Myriogramme from Nitophyllum.

Myriogramme dentata (Schousboe) nomen nudum

Senegal (529).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

Note. Combination has not been validly proposed. The relation-
ship between this taxon and Nitophyllum dentatum Bornet (q.v.)
requires further investigation.

Myriogramme minuta Kylin
Canaries (633;634;648).

Myriogramme sp.

Senegal (55;56).

Naccaria corymbosa J. Agardh

Salvage Islands (38B;598).

Note. See Searles & Leister (1980) concerning former known
geographical distribution only in North America.

Naccaria sp.

Canaries (227).

Nemalion amoenum (Pilger) B0rgesen

Cameroun (80;281;350;433;586).
Tropical Africa (Gambia - Congo river)' (598).
[As Dermonema amoenum Pilger]
Cameroun (139;433;454).

Note. See discussion in B0rgesen (80: 26-27) for placing Der-
monema in the genus Nemalion which he does with some doubt,
stating (p. 27) 'final decision as to its real place can of course not be
taken before its female organs are found, nevertheless I think that at
present its right place is in the genus Nemalion' '.

Nemalion helminthoides (Velley in Withering) Batters

Canaries ( 18 ;226 ;227 ;232B ;253 ;583 ;584;598 ;633 ;634) .

Western Sahara (349;476;659).

'. . . Atlantico norte de Inglaterra y Norvega a

Marruecos. . .' (517).

'. . . Atlantico (Norvega -Canaries). . .' (253).
'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W.Sahara] '(598).

[As Nemalion multifidum (Weber & Mohr) J. Agardh]
Canaries (254;305).
[As Nemalion heminthoides]
Canaries (237).
[As Nemalion lubricum Duby]
'. . . Atlantischer Ozean: Nordwest-africanische Kiiste. . .'

(499).

Nemalion lubricum Duby

See Nemalion helminthoides (Velley) Batters.

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA
N einal ion multifidum (Weber & Mohr) J. Agardh

'. . . in oceano atlantico a littore Norvegiae usque ad Canar-

ias(Webb.). . .'(25).

Note. Feldmann (188) considers the possibility that this species is
a mere growth form of N. helminthoides (Velley) Batters.

Nemalion sp.

Canaries (70).

Nemastoma canariensis (Kiitzing) J. Agardh

Canaries (13;38C;70;89;134;191;227;232B;315;540;598;662).

[As Gymnophloea canariensis Kiitzing]

Canaries (24;318).

[As Halymenia capensis Montagne]

Canaries (401).

[As Nemastoma canariensis J. Agardh]

Canaries (547).

[As Nemastoma canariensis (Kiitzing) Montagne]

Canaries (139;390;407).

[As Nemastoma canariense (Kiitzing) Montagne]

Canaries (633;634).

Note. The comments under Nemastoma confusum probably also
apply equally to this species.

Nemastoma confusum Kraft & D. John

Cape Verde Islands (652;683).

Ghana (299;315;350;376;586).

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

'. . . tropical West Africa' (350;586).

Note. Athanasiadis (630) has reassessed the genus Nemastoma
based on a study of the type species (N. dichotomum) and proposed a
new generic circumscription. Many species traditionally placed in
Nemastoma cannot now be accommodated. Anthanasiadis comments
thus (p. 31) 'Others, like Nemastoma confusum Kraft et John (1976),
which should apparently be excluded, necessitate a better knowledge
of the related genera in order to be transferred to the taxon where
they actually belong'.

Nemastoma multifida J. Agardh

See Platoma cydocolpum (Montagne) Schmitz.

Nemastoma sp.

Cape Verde Islands (652).

Neogoniolithon

The concept of Neogoniolithon adopted in this paper follows
Penrose (1992), who studied the type species, N. fosliei, and
used the following combination of features to delimit Neogo-
niolithon from other genera of the subfamily Mastophoroi-
deae: 1) thallus non-endophytic and lacking haustoria; 2)
thallus lacking a vertical layer of filaments composed of
palisade cells; 3) spermatangia simple and borne on the floor
and roof of male conceptacle chambers; and 4) gorrimoblast
filaments arising dorsally from fusion cells. Tetrasporangial
conceptacle ontogeny in Neogoniolithon (Penrose, 1992: 342)
is similar to that in Spongites (Penrose, 1991). A revised key
to the genera of Mastophoroideae r including Neogoniolithon,
is provided by Penrose & Chamberlain (1993: 303). Addi-
tional nomenclatural and systematic data on Neogoniolithon
are provided by Woelkerling (1988: 139-145).

77

Neogoniolithon absimile (Foslie & Howe) Cabioch
See Spongites wildpretii Afonso-Carrillo.

Neogoniolithon accretum (Foslie & Howe) Setchell &
Mason

Canaries (227;582).

[As Neogoniolithon accretum Foslie & Howe f. canariense

(Foslie) De Toni]
Canaries (139;205;212;382;493).
[As Lithophyllum accretum (Foslie & Howe) Lemoine]
Canaries (71;362;363;493).

[As Lithophyllum accretum (Foslie & Howe) Lemoine]
Canaries (71;362;363;493).
[As Lithophyllum accretum (Foslie & Howe) Lemoine f.

canariensis Foslie]
Canaries (205).
[As Lithophyllum accretum (Foslie & Howe) Lemoine var.

canariense (Foslie) Lemoine]
Canaries ( 191 ;363).

Note. This species originally was described as Goniolithon accre-
tum (Foslie & Howe, 1906: 131) and is based on material from Sands
Key, Florida. Subsequently, the species was transferred to Lithophyl-
lum (357: 159) and then to Neogoniolithon (Setchell & Mason, 1943:
90). The holotype (678: 16) has not been studied in detail in a modern
context, and thus the status and disposition of this species are
uncertain, as is the identification of specimens from the West African
region. Foslie (205: 19) also described Lithophyllum accretum f.
canariensis Foslie based on material from Puerto Orotava, Tenerife,
Canary Islands. This form apparently has not been transferred into
Neogoniolithon .

Neogoniolithon brassica-florida (Harvey) Setchell &
Mason

[As Lithothamnion brassica-florida (Harvey) Areschoug]

St. Helena (142;260;391).

[As Lithothamnion brassica-florida Harvey var.]

St. Helena (391).

Note. This species originally was described as Melobesia brassica-
florida Harvey (1849: 110) and is based on material from Algoa Bay,
South Africa. Subsequently, the species was transferred to
Lithothamnion (Areschoug, 1852: 523), then to Goniolithon (681: 9)
and then to Neogoniolithon (Setchell & Mason, 1943: 91). The
lectotype (678: 43) has been studied in detail by Woelkerling et al.
(1993), who confirmed that this species belonged to Neogoniolithon
as circumscribed by Penrose (1992). Woelkerling, Penrose & Cham-
berlain (1993) also concluded that N. fosliei (Heydrich) Setchell &
Mason, the type species of Neogoniolithon, was a heterotypic syn-
onym of N. brassica-florida. The specimens from St. Helena now
need to be re-examined to determine whether they are conspecific
with the lectotype of N. brassica-florida.

Neogoniolithon caribaeum (Foslie) Adey

Canaries (227;582;598).

Cape Verde Islands (598).

[As Lithophyllum caribaeum Foslie]

Canaries (71;191;362;363;366).

Cape Verde Islands (366).

[As Lithophyllum (?) cariboeum Foslie]

Cape Verde Islands (100).

Note. This species was originally described as Lithophyllum
decipiens Foslie (204: 18) based on material from the US Virgin
Islands. Subsequently, Foslie (206: 22) raised it to species status as
Lithophyllum caribaeum, and Adey (669: 8) transferred it to Neogo-

78

niolithon. There has seen no detailed study of the lectotype collection
(678: 48) in a modern context, and thus the status and disposition of
this species are uncertain, as is the identification of specimens from
the West African region.

Neogoniolithon hirtum (Lemoine in B0rgesen)
Afonso-Carrillo

Canaries (11;582;598;633;634).

[As Lithophyllum application Lemoine]

Canaries (227;362;363).

[As Mesophyllum applicatum Lemoine]

Canaries (191 ;687).

[As Lithophyllum hirtum Lemoine in B0rgesen]

Canaries (227;362;363;687).

[As Tenarea adhaerens Lemoine in B0rgesen]

Canaries (191;227;362;363).

Note. Based on studies of the original Canary Islands collections
of Lemoine, Afonso-Carrillo (11: 131) concluded that Lithophyllum
hirtum Lemoine in B0rgesen (363: 37) belonged to Neogoniolithon
(sensu Cabioch, 1972) and that Lithophyllum applicatum Lemoine in
B0rgesen (363: 38) and Tenarea adhaerens Lemoine in B0rgesen
(363: 59) were heterotypic synonyms. These conclusions need to be
re-evaluated in light of the revised concept of Neogoniolithon pre-
sented by Penrose (1992). Afonso-Carrillo (11) did not present
detailed accounts of the type collections, and it is not clear whether
the species involved possess the features now considered diagnostic
of Neogoniolithon. Lemoine (363: 37, 38) based Lithophyllum hirtum
on two collections and L. applicatum on five collections, but neither
she nor Afonso-Carrillo (11) specified types. Until lectotypes are
chosen and examined in detail in a modern context, the status and
disposition of these species will remain uncertain, as will the identifi-
cation of specimens from the West African region. Feldmann (191:
413) used the binomial Mesophyllum applicatum Lemoine instead of
Lithophyllum applicatum Lemoine. It is not clear whether this is an
error or whether Feldmann had intended to transfer applicatum to
Mesophyllum; the binomial probably should be cited as Mesophyllum
applicatum (Lemoine) Lemoine ex J. Feldmann.

Neogoniolithon illitus (Lemoine in B0rgesen)
Afonso-Carrillo

Canaries (11 ;582;598).

Cape Verde Islands (582;598).

[As Lithophyllum illitus Lemoine in B0rgesen]

Canaries (191 ;227;362;363;366;687).

Cape Verde Islands (100;366;645).

Note. Based on studies of the original Canary Islands collections
of Lemoine, Afonso-Carrillo (11: 131) concluded that Lithophyllum
illitus Lemoine in B0rgesen (363: 37) belonged to Neogoniolithon.
This conclusion needs to be re-evaluated in light of the revised
concept of Neogoniolithon presented by Penrose (1992). Afonso-
Carrillo (11) did not present detailed accounts of Lemoine's original
collections, and it is not clear whether the species possesses the
features now considered diagnostic of Neogoniolithon. Lemoine
(363: 54) based this species on 14 collections obtained from several
localities, but neither she nor Afonso-Carrillo (11) specified a type.
Until a lectotype is chosen and examined in detail, the status and
disposition of this species will remain uncertain, as will records from
the West African region. The record in Feldmann (183: 1071) is given
with a query.

Neogoniolithon mamillare (Harvey) Setchell & Mason

Annobon (366).
Bioko (350;586).
Cameroun (350;586).
Cape Verde Islands (366;598).

D.M. JOHNETAL

St. Helena (655).

Tantropical' (350;366;586).

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

[As Goniolithon mamillare (Harvey) Foslie]

Annobon (397;455;457;500).

Bioko (500).

Cameroun (500).

'. . .ad littora Africae ad Caput Verde. . .' (134).

[As Goniolithon mamillaris]

Cape Verde Islands (BM Herbarium, Coralline Catalogue).

[As Lithothamnion brassica-florida Harvey]

St. Helena (142;259;391).

[As Lithothamnion mamillare Harvey]

Cape Verde Islands (145).

[As Lithothamnion mamillare (Harvey) Areschoug]

Cape Verde Islands (38).

[As Melobesia mamillaris Harvey]

Cape Verde Islands (254;366).

[As Porolithon mamillare (Harvey) Foslie]

Annobon (139).

Cape Verde Islands (139;357).

Senegal (357).

Note. This species was originally described as Melobesia mamill-
aris Harvey (1849: 109). Subsequently, the species has been trans-
ferred to Lithothamnion (Areschoug, 1852: 521), Goniolithon (681:
9), Porolithon (357: 177), and Neogoniolithon (Setchell & Mason,
1943: 91). Harvey based the species on material from Brazil, Tierra
del Fuego, South Africa and the Cape Verde Islands, but did not
specify a type. Printz (212: pi. 47, legend to fig. 15), however,
lectotypified the species with a Brazilian specimen. According to
Woelkerling (678: 144) and Porter (1987: 200), this specimen appears
to be missing both from TRH and TCD, but TRH contains two
syntype fragments collected at Bahia by C. Darwin. Until the
lectotype can be located (or a new neotype designated) and studied in
detail in a modern context, the status and disposition of this species
are uncertain, as is the identification of specimens from the West
African region.

Neogoniolithon mamillosum (Hauck) Setchell &
Mason

Cape Verde Islands (366;597;598).

Mauritanie (349;366).

Senegal (597).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W.Sahara]' (598).
[As Goniolithon mamillare (Harvey) Foslie]
Annobon (397;454;455;500).
Bioko (500).
Cameroun (500).
[As Goniolithon mamillosum (Hauck) Foslie f. microcarpa

Foslie]

Cape Verde Islands (139;210;597).
Mauritanie (139;597).

Note. Goniolithon mamillosum f. microcarpa Foslie (207: 24) has
been lectotypified (678: 149) with a collection from Sao Vincente,
Cape Verde Islands, but this collection has not been studied in a
modern context and thus the status and disposition of f. microcarpa
are uncertain.

[As Goniolithon mamillosum (Hauck) Foslie]

Ascension (6).

Cape Verde Islands (6).

Mauritanie (6).

St. Helena (6).

Senegal (6).

RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

[As Lithophyllum hauckil (Rothpletz) Lemoine]
Mauritanie (188;349;359)

Note. There has been no detailed study of the type of Neogonioli-
thon mamillosum, and thus the status and disposition of the species
are uncertain, as is the identification of specimens from the West
African region. The basionym of Neogoniolithon mamillosum
(Hauck) Setchell & Mason, Lithothamnion mamillosum Hauck
(1883: 272) is a later homonym of Lithothamnion mamillosum
Gembel, and this led Rothpletz (1891: 304) and then Foslie (1895:
580) to independently coin the new name Lithothamnion hauckii
Rothpletz for Hauck's taxon (see 678: 115). In accordance with
Articles 55.1 and 64.1 of the International code for botanical nomen-
clature (see Greuter, 1988), Hauck's epithet mamillosum is correct
when used in the combination Neogoniolithon mamillosum (Hauck)
Setchell & Mason, but because of homonymy must become hauckii
when the species is placed in Lithothamnion. The type of both
Neogoniolithon mamillosum and Lithothamnion hauckii is the same.
In the protologue, Hauck (1883) depicted several specimens and
indicated that the material came from the Adriatic without indicating
specific localities. A lectotype for the species needs to be selected
from amongst extant syntypes, several of which are in TRH (678:
144-145).

Neogoniolithon orotavicum (Foslie) Lemoine

Canaries (368;583;598;633;634).

Cape Verde Islands (366;368;598).

'. . . Golfe de Guinee' (368).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).
[As Neogoniolithon orotavicum Foslie]
Canaries (227;366).
Cape Verde Islands (366).
Senegal (366).
[As Neogoniolithon orotavicum (Foslie) Lemoine ex Afonso-

Carrillo]

Canaries (11;253).
Senegal (253).
'. . . Atlantico Oriental (Archipelagos macaronesicas y costas

del Senegal). . .' (253).

[As Neogoniolithon orotavicum (Foslie) Afonso-Carrillo]
Canaries (17;582).

[As Lithophyllum orotavicum Foslie]
Canaries (2;70;191;362).

Note. This species originally was described as Goniolithon oro-
tavicum Foslie (205: 20) and is based on a single collection (see 678:
165) from Puerto Orotava, Tenerife, Canary Islands. Subsequently,
Lemoine (363: 49) transferred the species to Lithophyllum, and then
(366: 236, 238) to Neogoniolithon. Lemoine's 1964 (366) combination
Neogoniolithon orotavicum does not conform to Article 33.2 of the
International Code of Botanical Nomenclature with respect to
basionym citation and thus is invalid. Subsequently, however, Lem-
oine (368: 12) properly validated the binomial, an action apparently
unknown to Afonso-Carrillo (11: 133-134). Although the holotype
has been examined by Adey (669: 9) and Afonso-Carrillo (11:
133-134), neither provided a detailed morphological/anatomical
account of the material. Thus the status and disposition of this species
in relation to Penrose's (1992) circumscription of Neogoniolithon are
uncertain, as is the identification of specimens from the West African
region.

Neogoniolithon spp.

Canaries (10).

Cape Verde Islands (366).

Sao Tome (93;350;535).

79

Neomonospora furcellata (J. Agardh) G. Feldmann &
Meslin

See Griff ithsia arachnoides C. Agardh.

Neomonospora pedicellata (Smith) G. Feldmann &
Meslin

See Monosporus pedicellat us (Smith) Solier & Castagne.

Neomonospora sp.

Senegal (59).

Neuroglossum binderianum Kiitzing
Namibia (348).

Nitophyllum dentatum Bornet

Senegal (89; 132).

Note. Bornet (89: 293-294) quoted the description direct from
Schousboe, but the latter's name was Areolaria dentata Schousboe,
Tanger. Rarissime reperi inter Algas ad Dar Hamra circa Tingin
lectas mense augusto . . . petris calcareis vel Lithophytis innatum'.
Dangeard (118), in an addendum concerning Myriogramme costata
and Nitophyllum dentatum, comments on the close similiarity of these
two taxa. His Myriogramme had numerous teeth, close together and
a polystromatic frond centrally. In Bornet's description of N. dentata,
the teeth are only few and sparse, and the frond distromatic centrally.
See note under Myriogramme dentata Schousboe.

Nitophyllum fissum (Greville) J. Agardh
See Hymenema venosa (Linnaeus) Kylin.

Nitophyllum laceratum (Gmelin) Greville

'. . . in oceano Atlantico ab oris Angliae usque ad

Senegalliam. . .' (25;132).

Note. It is likely that this taxon is attributable to Cryptopleura
ramosa (Hudson) Kylin ex Newton.

Nitophyllum punctatum (Stackhouse) Greville

Canaries ( 13 ;226 ;227 ;584 ;598 ;633 ;634 ;662 ;684) .

Mauritanie (624).

'Nordwestafrika' (499).

[As Nitophyllum punctatum (Stackhouse) Greville var. ocel-

lata J. Agardh ]
'. . . Atlantic Ocean (. . . African. . .coasts, Canary Islands)'

(177).

Nitophyllum uncinatum (Turner) J. Agardh

See Acrosorium uncinatum (J. Agardh) Kylin (now A. venu-
losum (Zanardini) Kylin).

Nitophyllum venosum Harvey

See Hymenena venosa (Linnaeus) Kylin.

Nitophyllum sp.

Angola (41 ;352).
Senegal (59;122;529).
[As Nitophyllum sp. A]

80

Guinea-Bissau (529).
Senegal (529).

Note. Dangeard (122) and Sourie (529) suggest that this species
was 'voisin de N. punctatum (Stackh.) Greville'.

Nothogenia erinacea (Turner) Parkinson

Namibia (32A;36B).

[As Chaetangium erinaceum (Turner) Papenfuss]

Namibia (348;438).

[As Chaetangium ornatum (Linnaeus) Kutzing]

Namibia (128;160;348;453;500).

Note. According to Seagrief (570) Chaetangium dlchotomum
Kutzing (1869), Chaetangium hystrix (C. Agardh) Kutzing (1869) and
Chaetangium ornatum (L.) Kutzing are all synonyms of C. erina-

Nothogenia magnified (Pilger) J.H. Price comb. nov.

Basionym: Chaetangium magnificum Pilger in Hedwigia 48:

181-182, fig. C 1-3 (1908).
[As Chaetangium magnificum Pilger]
Namibia (139;348;429;453;500).
[As Chaetangium magnificum (Linnaeus) Kutzing]
Namibia (160).

[As Chaetangium ornatum (Linnaeus) Kutzing]
Namibia (429).

Note. The recombination is made here as there seems good
reason to doubt the conspecificity of Chaetangium magnificum with
that previously known often as Chaetangium ornatum (Linnaeus)
Kutzing [now Nothogenia erinacea (Turner) Parkinson, q.v.]. Papen-
fuss (429: 11) considered Pilger's (453) 'Chaetangium magnificum' a
synonym under 'Chaetangium ornatum'.

Nothogenia ovalis (Suhr) Parkinson

Namibia (32 A).

[As Chaetangium ovale (Suhr) Papenfuss]
Namibia (348;437).

'. . . Luderitz [S.W.A.] to Storms River mouth [S.A.]. . .
more common on the west coast. . .' (523).

D.M. JOHN ET AL

Olivia ustulata (Montagne) Montagne
See Caulacanthus ustulatus (Turner) Kutzing.

Ophidocladus simpliciusculus (P. & H. Crouan)
Falkenberg

Canaries (71;191;227;488;598).

Mauritanie (349).

Namibia (36B).

Western Sahara (349).

'. . . Atlantique (de la Bretagne aux Canaries). . .' (33).

'Subtropical Africa [Senegal (N. of Gambia); Mauritania;

Former W. Sahara]' (598).

[As Polysiphonia (?) simpliciuscula Crouan frat.]
Canaries (547).

Orcasia pulla Simons

See Streblocladia comptoclada (Montagne) Falkenberg.

Osmundaria volubilis (Linnaeus) R.E. Norris

[As Vidalia volubilis (Linnaeus) J. Agardh]
Canaries (439).

Note. This striking and unmistakable species has only been found
during the research trip of the Italian vessel 'Corsaro' (439). During
that trip many collections were also made in the Mediterranean, an
area where Osmundaria volubilis is common. It cannot be discounted
that a labelling error might have occurred, so accounting for the
record.

ACKNOWLEDGEMENTS. We acknowledge with thanks assistance from
many sources, of which the principal are: (i) Department of Botany,
The Natural History Museum, London, for provision of the neces-
sary research facilities to continue the series of papers of which the
present is part; (ii) editorial expertise of Dr R.J. Huxley and Marian
West for detecting inaccuracies in the submitted text; (iii) a grant (for
GWL) towards this work from The Systematics Association.

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RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA

81

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482 Reinbold, 1907.
484 Richardson, 1969.
487 Round, 1981.
488Saenger, 1971.

489 Santos Guerra, 1972.

490 Santos Guerra et al., 1970.

491 Sanusi, 1980.
493 Sauvageau, 1912.
496 Schmidt, 1924.

499 Schmidt, 1931.

500 Schmidt & Gerloff, 1957.

501 Schmitz & Falkenberg, 1897.

502 Schmitz & Hauptfleisch, 1896-97.

516 Seoane-Camba, 1960.

517 Seoane-Camba, 1965.

82

523 Simons, 1974.

527 Solms-Laubach, 1881.

528 Sender, 1852.

529 Sourie, 1954a.

530 Sourie, 1954ft.

531 Sourie, 1954c.
533 Southward, 1958.
535 Steentoft, 1967.

537 Stephenson & Stephenson, 1972.
540 Taylor, 1960.
541Tittleyetal., 1984.
542Trochain, 1940.
546Varoetal., 1979.

547 Vickers, 1897.

548 Viera y Clavijo, 1866.

551 Webb, 1849.

552 Weber-van Bosse, 1899.

555 Weisscher, 1982.

556 Weisscher, 1983.
556A Weisscher et al., 1982.

557 Weisscher etal., 1957.
557A Wffle, 1890-1891.

563 Yamada, 1931.

564 Yamada, 1938.

565 Yamada, 1941.
565AYarishetal., 1985.
567Luning, 1985.

570 Seagrief, 1984.

576 Afonso-Carrillo, 1985.

582 Afonso-Carrillo et al., 1985.

583 Haroun Tabraue et al., 1985.

584 Ribera Siguan et al., 1985.
586 Lawson & John, 1987.

590 John & Lawson (unpublished).

591 Yarish etal., 1986.
594 Lawson, 19546

596 Bailey & Harvey, 1862.

597 Prud'homme van Reine & Lobin, 1986.

598 Prud'homme van Reine (in litt., 10/4/
1987).

624 Marcot-Coqueugniot, 1991.

625 Prud'homme van Reine & van den Hoek,
1988.

630 Athanasiadis, 1988.

631 Chamberlain, 1992.

633 Pinedo etal., 1992.

634 Elejabeitia etal., 1992.

645 Lemoine, 1935 (cf. 100).

646 Sanson etal., 1991.

647 Gil-Rodriguez & Haroun, 1992.

648 Ballesteros et al., 1992.

649 Irvine & Chamberlain, 1994.

651 Brodie & Norris, 1992.

652 Otero-Schmitt & Sanjuan, 1992.

653 Prud'homme van Reine & van den Hoek,
1990.

D.M. JOHN ET AL

654 John & Lawson, 1991.

655 Lawson etal., 1993.

656 Abbott, 1990a.

657 Afonso-Carrillo et al., 1992.

658 Gil-Rodriguez & Haroun, 1993.

659 Hoek, 1982.

660 Masaki, 1968.

661 Guiry & Maggs, 1984.

662 Viera-Rodriquez et al., 1987ft.

663 Prud'homme van Reine et al., 1994.

664 Haroun etal., 1993.

665 Morales Ayala & Viera Rodriquez, 1990.

666 Viera-Rodriquez, 1985, thesis.

667 Viera-Rodriquez et al., 1987a.

668 Athanasiadis, 1987.

669 Adey, 1966.
674Dawson, 1955.

677 Romanes, 1916.

678 Woelkerling, 1993.

681 Foslie, 1898c.

682 Foslie, 1900d.

683 Otero-Schmitt, 1993.

684 Kristiansen et al., 1993.

685 Medina & Haroun, 1993.

686 Haroun & Prud'homme van Reine, 1993.

687 Reyes & Afonso-Carrillo, 1993.

688 Abbott, 1990ft.

695 Hardy & Seku, 1993.

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90

D.M. JOHNETAL

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Bull. nat. Hist. Mus. Land. (Bot.) 24(1): 91-99

Issued 23 June 1994

Studies on the Cretan flora 3. Additions to the
flora of Karpathos

NICHOLAS J. TURLAND

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD

LANCE CHILTON

22 River View, Ordsall, Retford, Nottinghamshire DN22 7UL

SYNOPSIS. Floristic notes are provided for 47 taxa, of which 42 were recorded by the authors in April 1992 as new to
the South Aegean island of Karpathos. These were first published in Flora of the Cretan area: annotated checklist &
atlas (Turland, Chilton & Press, 1993), but without comprehensive details of collections or observations, which are
instead provided here. Of the 42 taxa, 36 are new to the whole Karpathos island group (Karpathos, Kasos, Saria and
adjacent islets). Of these, Ranunculus cupreus Boiss. & Heldr. was previously considered endemic to Crete,
Bellevalia trifoliata (Ten.) Kunth is confirmed as present in the Cretan area (the above islands together with Crete
itself), Scilla bifolia L. is new to the Cretan area, and Asyneuma giganteum (Boiss.) Bornm. is new to Europe
(previously known only from the East Aegean islands). Details are also provided confirming the occurrence of five
taxa which were formerly known in the Karpathos island group only from old (pre-1930) records.

INTRODUCTION

The island of Karpathos lies in the South Aegean region some
75 km to the north-east of Crete. At 48 km long and up to 12
km wide (305 km 2 ), it is the largest component of the
Karpathos island group (Fig. 1), which also includes Kasos
(18 km x 6 km) and Saria (9 km x 6 km). This archipelago,
together with Crete and its satellite islands, comprise the
floristic territory of the Cretan area ('Cr') as defined in Flora
Europaea (Tutin et al., 1964-1980) and the Med-Checklist
(Greuter, Burdet & Long, 1984-1989).

Karpathos lies on a north-south axis and reaches 1215 m on the
calcareous mass of Mount Kalilimni , which is located more or less
in the centre of the island. On the eastern side, at c. 700 m, is the
upland plain of Lastos, much of which was formerly under
cultivation, while on the western side are vertical cliffs and steep
rocky slopes above woods of Pinus brutia Ten. The area north of
Kalilimni consists of a steep-sided ridge with several summits
over 500 m, more pinewoods (largely burnt) and extensive
exposure of schistose rock. In the northernmost part of Karpa-
thos are located the villages of Avlona, Diafani and Olimbos,
which lacked vehicular access until 1975, and where traditional
cultivation of cereals on narrow hillside terraces is still practised.
The region south of Kalilimni is less precipitous topographically
and contains a large proportion of the island's cultivated land. On
the eastern coast is Pigadia, the principal population centre (also
known as Karpathos). The southernmost part of the island is
low-lying and more or less flat, with some cultivated fields, but
mainly large expanses of very dry, exposed calcareous phrygana.

Flora of the Cretan area: annotated checklist & atlas
(Turland, Chilton & Press, 1993) was published as a first
product of the European Plant Information Centre (EPIC) at
The Natural History Museum. In this checklist, the total
number of native vascular plant species recorded from the
island group is given as 905, of which 10 (1.11%) are endemic
and 32 (3.54%) endemic to the Cretan area. There are also 26
introduced species.

The Natural History Museum, 1994

Of the taxa dealt with in the present paper, 42 were
recorded as additions to the flora of Karpathos in Turland et
al. (1993), as a result of a field trip to the island by the present
authors in April 1992. Comprehensive details of collections
or observations were not included in the checklist because of
its concise format, and are instead provided here. During the
field trip, the present authors also confirmed the occurrence
of five taxa which were formerly known in the Karpathos
island group only from old (pre-1930) records. They too are
listed here together with full details.

Taxonomy and nomenclature follow Flora of the Cretan
area. Page references to the text and distribution maps
contained in that work are provided for each taxon, using the
acronym 1 FCA\ The sequence of localities of collections or
observations runs from north to south. Transliteration of
Greek place names follows Turland (1992), which in turn
follows official Greek usage. The aim is to provide the closest
possible phonetic equivalent to the Greek pronunciation
without resorting to unwieldy letter combinations. In order to
avoid repetitiveness in citing the records, the collectors
'Chilton & Turland' are abbreviated to 'C. & TV and 'obser-
vation' to 'obs.' All collections have been lodged at The
Natural History Museum (BM).

FLORISTIC NOTES

Pteridophyta

Aspleniaceae

Asplenium ruta-muraria L. subsp. ruta-muraria (new
to the Karpathos island group - FCA: 31, 200)

92

N.J. TURLAND AND L. CHILTON

Fig. 1 The Karpathos island group, showing localities mentioned in the text, and the mapping grid used by Turland et al. (1993). The basic,
unadjusted square size is 8.25 km x 8.25 km.

CRETAN FLORA 3. ADDITIONS TO KARPATHOS

93

W. of summit of Mt Kalilimni, 1150 m, crevices of W. -facing
calcareous cliff, 9 April 1992, C. & T. 307 (living material, no
longer extant).

This small north temperate fern also occurs in the three main
mountain massifs of Crete (Lefka Ori, Psiloritis and Dikti).

Spermatophyta
Angiospermae
Dicotyledones
Apocynaceae

Vinca major L. subsp. major (new to the Karpathos
island group - FCA: 36, 206)

Aperi village, 300 m, side of road, 7 April 1992, C. & T. obs.;
2 km NNW of Piles, 250 m, abandoned cultivated ground, 6
April 1992, C. & T. obs.

This evergreen subshrub is cultivated for ornament in Crete
and occasionally becomes naturalized. It occurs as a native
plant in western and central parts of the Mediterranean
region.

Araliaceae

an old record from Mount Kalilimni cited by Rechinger
(1943a: 455, as Lithospermum arvense var. sibthorpianum
(Griseb.) Halacsy). L. sibthorpianum is related to L. arvense
and L. incrassatwn Heldr. ex Guss. (Buglossoides arvensis
subsp. gasparrinii (Heldr. ex Guss.) R. Fern.) These two
species are unknown in the Karpathos island group, although
they do occur in Crete together with L. sibthorpianum.

Cactaceae

Opuntia ficus-barbarica A. Berger (new to the
Karpathos island group - FCA: 41, 212)

O. ficus-indica sensu Flora Europaea

Road entrance to Olimbos village, 350 m, field-margins, 7
April 1992, C. & T. obs.; Aperi village, 300 m, 7 April 1992,
C. & T. obs.; 1.5 km W. of Menetes, 350 m, near side of
road, 10 April 1992, C. & T. obs.

Opuntia ficus-barbarica was introduced to the Mediterranean
region from tropical America. It is planted in Crete and also
naturalized in cultivated and waste areas, usually near human
habitation.

Campanulaceae

Hedera helix L. (new to the Karpathos island group -
FCA: 37, 206)

Piles village, 300 m, wall at side of road, 6 April 1992, C. & T.
obs.

Hedera helix is widespread in Crete as subsp. helix (with
blackish fruits) and is a native plant in woodland and on cliffs.
The species also occurs on walls in Cretan towns and villages,
where it may sometimes be planted, either as subsp. helix or
subsp. poetarum Nyman (with orange-yellow fruits). In Kar-
pathos it was observed only at the cited locality - a man-made
habitat - which suggests that its native status on the island is
doubtful. In the absence of fruits, the plants could not be
identified to subspecific rank. Outside the Cretan area the
species is distributed in temperate parts of Europe and Asia.

Boraginaceae

Lithospermum sibthorpianum Griseb. (confirmation
for the Karpathos island group - FCA: 40)

Buglossoides arvensis subsp. sibthorpiana (Griseb.) R. Fern.

5.5 km S. of Olimbos, cultivated area at head of valley
running to NNW, 200 m, among cereal crop on cultivated
terrace, 8 April 1992, C. & T. 291; SE side of summit of Mt
Kalilimni, 1150 m, soil-patch in calcareous phrygana, 9 April
1992, C. & T. 304; E. slope of Mt Kalilimni, 900 m, small
open area in dry stream-bed surrounded by garigue of
Phlomis floccosa D. Don and Sarcopoterium spinosum (L.)
Spach, 5 April 1992, C. & T. 247; 1.5 km E. of Lastos plain,
700 m, bare soil at side of track, 9 April 1992, C. & T. obs.

This annual species was given for Karpathos by Greuter et al.
(1983: 49, as Lithospermum arvense L.) purely on the basis of

Asyneuma giganteum (Boiss.) Bornm. (new to Europe
-FC4:42, 213)

W. of summit of Mt Kalilimni, 1150 m, crevices of W.-facing
calcareous cliff, 9 April 1992, C. & T. 311 (living material, in
hort. Turland).

Asyneuma giganteum is a monocarpic chasmophyte which
occupies a very isolated taxonomic position within the genus
(Damboldt, 1978: 71). Its discovery in Karpathos extends the
known distribution into Europe, the previously known popu-
lations being on the East Aegean islands of Rodos (Dam-
boldt, loc. cit.) and Halki (Carlstrom, 1987: 97, 223). The
population on Mount Kalilimni appears to be extremely small
and localized: up to 10 plants were found on one cliff only, all
of them diminutive (up to 5 cm high), with only a few leaves
and no fertile parts present, or indeed any dead remains of
those of the previous growing season. The single plant
collected has subsequently grown well in cultivation in Brit-
ain. By autumn 1992, it had produced a rosette of numerous
leaves and, although still sterile, was clearly identifiable with
a fertile specimen collected in Rodos (In monte Prophet Elias
(Monte Prof eta) prope Salakos, in saxosis calc. Turka', c.
600 m, 2 July 1935, Rechinger 8508 (BM!)).

Two other taxa have a similar bi-regional endemic distribu-
tion spanning the phytogeographical divide between Europe
and Asia: Astragalus austro-aegaeus Rech. f. (Leguminosae)
is known only from Kasos, Karpathos and Rodos, while
Ophrys umbilicata subsp. rhodia H. Baumann & Kiinkele
(Orchidaceae) is known only from Karpathos and Rodos.

94

N.J. TURLAND AND L. CHILTON

Caryophyllaceae

('crust in in brachypetalum Pers. (new to the Karpathos
island group - FCA: 45, 218)

Lastos plain, 700 m, fallow field, 5 April 1992, C. & T. 241.

This small annual species occurs in the higher mountains of
Crete, where two subspecies are known: the endemic subsp.
doerfleri (Halacsy ex Hayek) P. D. Sell & Whitehead, appar-
ently restricted to Mount Kedros, and the more widespread
subsp. roeseri (Boiss. & Heldr.) Nyman. Unfortunately it was
not possible to identify the Karpathos collection to subspe-
cific rank and it is recorded, for the time being, as the species
sensu lato. Cerastium brachypetalum is widely distributed in
western, central and southern Europe where several subspe-
cies have been recognised.

Sagina apetala Ard. (new to Karpathos - FCA: 49,

223)

1 km NW of Pigadia, sea-level, damp bare sandy mud at edge
of Phragmites bed, 11 April 1992, C. & T. 339.

This is another diminutive annual with a widespread Euro-
pean distribution which includes Crete. The species was first
recorded from the Karpathos island group by Greuter et al.
(1983: 50), from Kasos.

Sagina maritima Don (new to Karpathos - FCA: 49,

223)

0.5 km W. of Pigadia, 5 m, damp bare sandy ground by track,
4 April 1992, C. & T. 226.

Sagina maritima is morphologically similar to the preceding
species, but differs in having leaves muticous or very shortly
mucronate, rather than distinctly aristate. It occurs along the
coasts of most of Europe, including Crete. In common with
Sagina apetala, it was first recorded from the Karpathos
island group by Greuter et al. (1983: 50), from Kasos.

Stellar iu media (L.) Vill. (confirmation for the
Karpathos island group - FCA: 53, 228)

Diafani village, sea-level, 8 April 1992, C. & T. obs.; road
entrance to Olimbos village, 350 m, 7 April 1992, C. & T.
obs.; 1.5 km SW of Spoa, 300 m, foot of NW-facing calcare-
ous cliff, 7 April 1992, C. & T. obs.; Aperi village, 300 m, 7
April 1992, C. & T. obs.; immediately NW of Piles, 300 m,
bare stony ground of dry stream-bed by road, 6 April 1992,
C. & T. 260; immediately W. of Menetes, by church, 350 m,
grassy area, 10 April 1992, C. & T. obs.; Menetes village, 350
m, 10 April 1992, C. & T. obs.

This taxon is the species sensu stricto (i.e. subsp. media sensu
Flora Europaed), which is distributed throughout Europe.
The species was given for Karpathos by Greuter et al. (1983:
51) purely on the basis of an old record from Volada cited by
Rechinger (1943a: 148). Jalas & Suominen, in Atlas Florae
Europaeae (1983: 72, map 820), map Stellaria media sensu
lato from Karpathos (i.e. including 5. cupaniana (Jord. &
Fourr.) Beg., 5. neglecta Weihe and S. pallida (Dumort)
Pire), using a symbol indicating the existence of a post-1930
record, but they do not provide a separate map for the species
sensu stricto.

Chenopodiaceae

Beta vulgaris subsp. maritima (L.) Arcang. (new to
the Karpathos island group - FCA: 54, 230)

Diafani, sea-level, cliff immediately above seashore, 8 April
1992, C. & T. obs.; coast WSW of Piles, 5 m, margin of
abandoned field above beach, 7 April 1992, C. & T. 272.

At the cited localities, the plants appeared to be referable to
the native subsp. maritima, rather than subsp. vulgaris, which
occurs as a casual escape from cultivation in both Crete and
Karpathos. Subsp. maritima occurs along the coasts of west-
ern and southern Europe (including Crete), Asia and North
Africa, as well as on saline soils inland.

Compositae (Asteraceae)

Bellis sylvestris Cirillo (new to the Karpathos island
group -FCA: 59, 237)

Lower E. slopes of Mt Kalilimni above Lastos plain, 750 m,
garigue, 5 April 1992, C. & T. obs.; along track from Lastos
plain to E. coast, 700 m, garigue, 9 April 1992, C. & T. obs.;
upper end of Volada village, 470 m, garigue, 5 April 1992, C.
& T. obs.; W. of Othos, on road to Stes, 450 m, phrygana, 6
April 1992, C. & T. 249; immediately W. of Menetes, by
church, 350 m, grassy area, 10 April 1992, C. & T. obs.

Numerous flowering individuals of this perennial southern
European species were observed in Karpathos - enough to
suggest that spring is at least part of its main flowering period
there. In Crete, the species seems to flower mainly in the
autumn, and only rarely in spring.

Centaurea calcitrapa L. subsp. calcitrapa (new to the
Karpathos island group - FCA: 61, 240)

0.5 km SE of Olimbos, 350 m, bare ground at side of road, 7
April 1992, C. & T. obs.

This biennial plant occurs as a native in southern-central
Europe and the Mediterranean region, and is naturalized
elsewhere in western and central Europe. It is found in Crete,
where it usually grows in weedy habitats similar to the
Karpathos locality.

Rhagadiolus edulis Gaertn. (new to the Karpathos
island group - FCA: 71, 253)

Valley 4 km N. of Mt Kimaras, between semi-deserted village
and gorge, 100 m, grove of Pinus brutia Ten. on valley-floor,
growing with Aristolochia cretica Lam., Cyclamen creticum
(Dorfl.) Hildebr., Scaligeria napiformis (Spreng.) Grande
and Theligonum cynocrambe L., 8 April 1992, C. & T. 298; 1
km SE of Lastos plain, 650 m, rocky ground sheltered by a
shrub on margin of abandoned field, 9 April 1992, C. & T.
301; 1 km SE of Menetes, 280 m, foot of E. -facing calcareous
cliff, 10 April 1992, C. & T. obs.

Rhagadiolus edulis has not always been separated from R.
stellatus (L.) Gaertn. sensu lato, which is distributed from
southern Europe eastwards to Iran and was already known to
occur in Kasos and Karpathos (Rechinger, 19430: 676; Greu-
ter et al., 1983: 54). R. edulis was not given for the Karpathos
island group by Rechinger (op. cit.: 677), and was not treated

CRETAN FLORA 3. ADDITIONS TO KARPATHOS

95

separately by Greuter et al. (loc. cit.) It is characterized by its
lyrately divided leaves and glabrous phyllaries and is prob-
ably only worthy of separation at varietal rank, if at all.
Turland et al. (1993) do not treat it separately, except in their
distribution maps. R. stellatus sensu stricto, with dentate to
incised leaves and pubescent or minutely setulose phyllaries,
also occurs in Karpathos, and both taxa occur more or less
sympatrically in Crete. R. edulis has a tendency to grow in
more humid habitats than R. stellatus in the Cretan area.

Tyrimnus leucographus (L.) Cass. (new to the
Karpathos island group - FCA: 73, 257)

0.25 km SW of Avlona, 300 m, cultivated terraces on
calcareous substratum, 8 April 1992, C. & T. obs.; immedi-
ately N. of Olimbos, 300 m, field, 8 April 1992, C. & T. obs.;
E. side of Mt Kimaras, on Spoa-Olimbos road, 350 m, rocky
ground at side of road, 7 April 1992, C. & T. 276.

Old records of this Mediterranean species from Kasos and
Karpathos (Rechinger, 1943a: 655) were considered errone-
ous and were referred to Carduus argentatus L. by Greuter et
al. (1983: 52). The latter species was also observed by the
authors on the terraces at Avlona.

Crassulaceae

Crassula alata (Viv.) A. Berger (new to the Karpathos
island group - FCA: 76, 259)

Beach 1 km NW of Pigadia, sea-level, sandy ground at top of
beach, 11 April 1992, C. & T. 335 - det. J.R. Akeroyd.

This minute annual species is currently known in Europe only
from the Aegean region (Crete, Karpathos and the
Kiklades). Otherwise, it occurs in North Africa and south-
western Asia. It was first recorded from Crete by Greuter &
Raus (1981: 276), and has not always been distinguished
there from the very similar Crassula tillaea Lest.-Garl. (Greu-
ter, Malthas & Risse, 1984: 275-276).

Cruciferae (Brassicaceae)

Eruca sativa Mill, (new to the Karpathos island group

-FCA: 82, 268)

E. vesicaria subsp. sativa (Mill.) Thell.

1.5 km NE of Lefkos, 100 m, 6 April 1992, C. & T. obs.; 0.5
km SW of Volada, 500 m, 6 April 1992, C. & T. obs.; 2.25 km
NNW of Piles, 250 m, 6 April 1992, C. & T. obs.; Pigadia
town, 15 m, 4 April 1992, C. & T. obs.

Eruca sativa is an annual species widely distributed in the
Mediterranean region. It occurs in Crete, where it grows in
weed communities on cultivated ground.

Teesdalia coronopifolia (J. P. Bergeret) Thell. (new to
the Karpathos island group - FCA: 86, 272)

Lastos plain, 700 m, fallow field, 5 April 1992, C. & T. 240.

This mainly southern European and North African annual is
also known from western Crete, where it has been recorded
under similar ecological conditions at an altitude of 1050 m on

the Omalos plain in the Lefka Ori massif (Rechinger, 1943ft:
79).

Dipsacaceae

Knautia integrifolia (L.) Bertol. (new to the
Karpathos island group - FCA: 86, 273)

Along track from Lastos plain to E. coast, 700 m, 9 April
1992, C. & T. obs.; immediately NE of Stes, 450 m, weed
community in olive grove, 6 April 1992, C. & T. obs.; 1.5 km
W. of Menetes, 350 m, 10 April 1992, C. & T. obs.

Knautia integrifolia is an annual species represented in Crete
by subsp. mimica (Borbas) Greuter, which otherwise occurs
in Albania and Greece, and on the East Aegean island of
Rodos by subsp. urvillei (Coult.) Greuter, which otherwise
occurs from Turkey-in-Europe to Palestine. The precise
identity of the populations in Karpathos has yet to be
ascertained; they could belong to either of these taxa, or
possibly to subsp. integrifolia, which is distributed mainly in
the western and central Mediterranean region, extending
eastwards to Bulgaria and Greece.

Euphorbiaceae

Euphorbia helioscopia L. (new to Karpathos - FCA:

89, 277)

0.25 km SW of Avlona, 300 m, cultivated terraces, 8 April
1992, C. & T. obs.; ravine immediately SW of Diafani, 50 m,
8 April 1992, C. & T. obs.; 5.5 km S. of Olimbos, cultivated
area at head of valley running to NNW, 200 m, 8 April 1992,
C. & T. obs.; 1.5 km W. of Menetes, 350 m, 10 April 1992, C.
& T. obs. ; 1.5 km N. of Airport, 20 m, margin of fallow field,
10 April 1992, C. & T. 324.

This annual species occurs almost throughout Europe
(including Crete), North Africa and Asia, and is widely
introduced elsewhere. The species was first recorded from
the Karpathos island group by Greuter et al. (1983: 57), from
Kasos.

Geraniaceae

Erodium moschatum (L.) L'Her. (new to the
Karpathos island group - FCA: 92, 281)

Olimbos village, 350 m, 7 April 1992, C. & T. obs.; gorge 4.5
km N. of Mt Kimaras, 100 m, 8 April 1992, C. & T. obs.;
semi-deserted village 3.5 km N. of Mt Kimaras, 150 m, 8
April 1992, C. & T. obs.; 1.5 km SW of Spoa, 300 m, foot of
NW-facing calcareous cliff, 7 April 1992, C. & T. obs.;
immediately NE of Stes, 450 m, 6 April 1992, C. & T. obs.;
Pigadia town, 15 m, 4 April 1992, C. & T. obs.; immediately
W. of Menetes, by church, 350 m, 10 April 1992, C. & T. obs.

Erodium moschatum is an annual or biennial species widely
distributed in Europe (including Crete), North Africa and
Asia, and introduced elsewhere.

96

Geranium tuberosum L. subsp. tuberosum (new to the
Karpathos island group - FCA: 93, 282)

Immediately NE of Stes, 450 m, weed community at side of
track, 6 April 1992, C. & T. 253.

This tuberous perennial plant is distributed in southern
Europe, North Africa and south-western Asia. It occurs in
Crete, where it is characteristic of areas in which traditional
methods of cultivation are still practised. The plants typically
grow in cultivated and fallow fields, as for example in the
fields of the Lasithi plain in the Dikti massif in eastern Crete.

Guttiferae (Hypericaceae)

Hypericum perforatum L. (new to the Karpathos
island group - FCA: 94, 284)

0.25 km SW of Avlona, 300 m, growing through calcareous
rocks of dry-stone wall, 8 April 1992, C. & T. 285; 0.5 km E.
of Pigadia, 11 April 1992, C. & T. obs.; between Menetes and
Arkasa, growing through calcareous rocks of dry-stone wall,
10 April 1992, C. & T. obs.

Hypericum perforatum occurs almost throughout Europe
(including Crete), as well as in North Africa and Asia. The
plants seen in Karpathos were sterile, but they can neverthe-
less be referred to this species with confidence.

Labial ae (Lamiaceae)

Melissa officinalis subsp. altissima (Sm.) Arcang.
(confirmation for the Karpathos island group - FCA :
96, 286)

Immediately NE of Stes, 450 m, heap of stones in partly
cultivated area, 6 April 1992, C. & T. obs.; 2 km NNW of
Piles, 250 m, near stream, 6 April 1992, C. & T. obs.

Melissa officinalis was given for Karpathos by Greuter et al.
(1983: 58) purely on the basis of an old record from Othos
cited by Rechinger (19430: 522). Subsp. altissima occurs in
southern Europe (including Crete), North Africa and south-
western Asia.

Rosmarinus officinalis L. (new to the Karpathos island
group -FCA: 99, 289)

0.5 km W. of Volada, by track to Lastos plain, 500 m, one
plant at top of bank by track, C. & T. obs. ; near Piles, 6 April
1992, C. & T. obs.

This largely Mediterranean shrub is occasionally cultivated
for ornament or culinary use in both Crete and Karpathos,
and it is very likely that the few plants seen at the cited
localities are deliberately planted individuals rather than
native or naturalized. The native status of the species in Crete
is similarly doubtful.

Stachys arvensis (L.) L. (new to the Karpathos island
group -FCA: 100,292)

2 km SE of Aperi, 250 m, 5 April 1992, C. & T. obs.
Stachys arvensis is found in Europe, Africa, south-western

N.J. TURLAND AND L. CHILTON

Asia and America. It occurs in Crete, where it grows among
rocks and in stony places.

Leguminosae (Fabaceae)

Lathy rus annuus L. (new to the Karpathos island
group -FCA: 106,299)

5.5 km S. of Olimbos, cultivated area at head of valley
running to NNW, 200 m, among cereal crop on cultivated
terrace, 8 April 1992, C. & T. 292; immediately NE of Stes,
450 m, weed community at side of track, 6 April 1992, C. &
T. 251; 1 km NW of Pigadia, sea-level, annual community on
damp ground at edge of Phragmites bed, growing with
Bromus spp., Hordeum leporinum Link, Medicago spp. and
Vicia villosa subsp. varia (Host) Corb., 11 April 1992, C. &
T. 338.

This annual species is distributed from the Mediterranean
region eastwards to central Asia. It occurs in Crete, where its
usual habitats are much the same as those in Karpathos.

Lens culinaris Medik. (confirmation for the Karpathos
island group - FCA: 107)

5.5 km S. of Olimbos, cultivated area at head of valley
running to NNW, 200 m, 8 April 1992, C. & T. obs. ; 1 km NE
of Lefkos, 40 m, fallow field, 6 April 1992, C. & T. 264.

Lens culinaris is an annual species widely cultivated in
Europe, Asia and elsewhere for its edible seeds. The plant
was given as cultivated in Karpathos by Greuter et al. (1983:
60) purely on the basis of an old record cited by Ciferri (1944:
61, as Ervum lens L.) The plants at both localities cited here
are probably no more than relics of cultivation, although
those at Lefkos may have become locally naturalized, since
the field appeared to have been lying fallow for some years.

Vicia faba L. (new to the Karpathos island group -
FCA: 115)

Lastos plain, 700 m, fields, 5 April 1992, C. & T. obs.; Aperi
village, 300 m, 7 April 1992, C. & T. obs.; 1.5 km W. of
Menetes, 350 m, 10 April 1992, C. & T. obs.; 1.5 km N. of
Airport, 20 m, fields, 10 April 1992, C. & T. obs.

Vicia faba is cultivated for its edible seeds in both Crete and
Karpathos. In Crete, the species also occurs as a casual relic
or escape from cultivation, and the records from Karpathos
most probably represent such occurrences.

Malvaceae

Lavatera cretica L. (new to the Karpathos island
group -FCA: 118,317)

Road entrance to Olimbos village, 350 m, 7 April 1992, C. &
T. obs.; Pigadia town, 15 m, waste area, 11 April 1992, C. &
T. 343.

This annual or biennial species is distributed from western
Europe through the Mediterranean region to Arabia. It
occurs in Crete, where it is most often found in cultivated and
waste areas.

CRETAN FLORA 3. ADDITIONS TO KARPATHOS

Plantaginaceae

97

Scrophulariaceae

Plantago amplexicaulis Cav. subsp. amplexicaulis (new
to the Karpathos island group - FCA: 124, 323)

E. edge of Pigadia town, 20 m, waste area at side of road, 11
April 1992, C. & T. 344.

This annual plant is found in southern Spain, southern Italy,
the South Aegean region (including eastern Crete), North
Africa and Cyprus.

Linaria chalepensis (L.) Mill, (new to the Karpathos
island group - FCA: 142, 348)

5.5 km S. of Olimbos, cultivated area at head of valley
running to NNW, 200 m, 8 April 1992, C. & T. obs.

Linaria chalepensis is an annual species found in the Mediter-
ranean region and south-western Asia. It occurs in Crete,
where it grows as a weed of cultivated areas.

Ranunculaceae

Ranunculus cupreus Boiss. & Heldr. (new to the
Karpathos island group - FCA: 132, 334)

Summit area of Mt Kalilimni, 1200 m, soil-pockets in calcare-
ous rocky slope, 9 April 1992, C. & T. 306; E. side of Mt
Kalilimni, 1000 m, crevices and soil-pockets in calcareous
rock-face exposed to N., 5 April 1992, C. & T. 244.

Ranunculus cupreus is a tuberous perennial species which,
prior to its discovery on Mount Kalilimni in Karpathos, was
considered endemic to Crete, where it occurs sporadically
from sea-level up to 2200 m. The plants are frequent on
Kalilimni above c. 1000 m. The basal leaves are hairy and the
undersides of the honey-leaves sometimes copper-tinted -
both diagnostic features of R. cupreus, and relevant in
distinguishing this species from R. subhomophyllus (Halacsy)
Vierh., which has also been recorded from Mount Kalilimni
(Rechinger, 1943a: 190; Strid, 1986: 220). It seems probable
that R. cupreus in Karpathos has previously been misidenti-
fied as R. subhomophyllus: the authors observed large num-
bers of plants of the former but none of the latter during their
field trip. Indeed, the accuracy of all records of R. subho-
mophyllus from Karpathos should not be taken for granted.
The same could be said of records from Mount Afendis
Kavousi in eastern Crete (Rechinger, 1943&: 74; Strid, loc.
cit.): this mountain is the type locality of R. cupreus, which
still occurs there in large numbers (clearly agreeing with the
Kalilimni material), whereas R. subhomophyllus has not yet
been found there by either of the authors.

Resedaceae

Scrophularia peregrina L. (new to Karpathos - FCA:

143, 349)

Gorge 4.5 km N. of Mt Kimaras, 100 m, 8 April 1992, C. & T.
obs.; semi-deserted village 3.5 km N. of Mt Kimaras, 150 m,
8 April 1992, C. & T. obs.; Aperi village, 300 m, 7 April
1992, C. & T. obs.; immediately NW of Piles, 300 m, heap of
soil and stones by dry stream-bed by road, 6 April 1992, C. &
T. 259; 0.5 km E. of Pigadia, 11 April 1992, C. & T. obs.;
immediately W. of Menetes, by church, 350 m, 10 April 1992,
C. & T. obs.

This annual species is found mainly in the Mediterranean
region, including Crete, where it grows in rocky places and
weed communities mainly in the west of the island. In the
Karpathos island group, it is known from an old record from
Kasos cited by Rechinger (1943a: 477), and was recorded as
new to Saria by Greuter et al. (1983: 68). It appears to have
been hitherto overlooked in Karpathos, however, in spite of
its moderately frequent occurrence there.

Umbelliferae (Apiaceae)

Smyrnium olusatrum L. (new to the Karpathos island
group -FCA-. 153,363)

Aperi village, 300 m, dry-stone wall at side of road, 7 April
1992, C. & T. 267; Piles village, 300 m, 6 April 1992, C. & T.
obs.; Menetes village, 350 m, 10 April 1992, C. & T. obs.

Smyrnium olusatrum is a biennial species distributed mainly
in the Mediterranean region. It occurs in Crete, where it is
often found in or around villages.

Reseda luteola L. (confirmation for the Karpathos
island group - FCA: 134, 336)

2.5 km N. of Mt Kimaras, 200 m, hard bare ground on track,
8 April 1992, C. & T. 296; 2 km NW of Pigadia, 10 m,
roadside bank, 4 April 1992, C. & T. obs.

This biennial species was given for Karpathos by Greuter et
al. (1983: 66) purely on the basis of an old record from the
Olimbos area cited by Rechinger (1943a: 245). It occurs in
Europe (including Crete) and North Africa eastwards to
central Asia.

Monocotyledones

Agavaceae

Agave americana L. (new to the Karpathos island
group -FCA: 157,368)

Diafani village, 8 April 1992, C. & T. obs.

This large succulent perennial was introduced to the Mediter-
ranean region from Mexico. It is cultivated in Crete and is
possibly also naturalized on roadsides and near villages. The
exact status of the population at Diafani is uncertain.

98
Cyperaceae

Carex illegitima Ces. (new to the Karpathos island
group -FCA: 160,372)

2.5 km N. of Mt Kimaras, 200 m, steep slope of stream gully
in area of burnt Pinus brutia woodland, forming clumps
amongst regenerating vegetation of Arbutus unedo L., Pista-
cia lentiscus L. and Cistus on schistose substratum, 8 April
1992, C. & T. 295; along track from Lastos plain to E. coast,
700 m, in field layer of very open Pinus brutia woodland, 9
April 1992, C. & T. obs.

This mainly eastern Mediterranean sedge was considered
doubtfully present in the Cretan area until recorded by
Turland (1992: 163) from the lower western slopes of Mount
Afendis Kavousi in eastern Crete where, as in Karpathos, it
grows in Pinus brutia woodland.

Gramineae (Poaceae)

Bromus diandrus Roth (new to the Karpathos island
group -FCA: 165,378)

Immediately NE of Stes, 450 m, weed community at side of
track, 6 April 1992, C. & T. 250; 1 km NW of Pigadia,
sea-level, annual community on damp ground at edge of
Phragmites bed, 11 April 1992, C. & T. obs.; Pigadia town,
15 m, 4 April 1992, C. & T. obs.; immediately W. of
Menetes, by church, 350 m, 10 April 1992, C. & T. obs.

Bromus diandrus is an annual grass found in southern
Europe, North Africa and south-western Asia. It occurs in
Crete, growing in a variety of disturbed habitats.

Bromus hordeaceus subsp. divaricatus (Bonnier &
Layens) Kerguelen (new to the Karpathos island
group -FCA: 166,379)

B. hordeaceus subsp. molliformis (Lloyd ex Godr.) Maire &
Weiller

1.5 km N. of airport, 20 m, fallow field, 10 April 1992, C. &
T. 323 - del. M.H. Martin (University of Bristol).

This annual Mediterranean grass also occurs in Crete, where
it grows in marshes, damp grassland and sandy alluvial soils,
as well as fallow fields.

Horde um spontaneum K. Koch (new to the Karpathos
island group - FCA: 171, 385)

Immediately W. of Menetes, by church, 350 m, 10 April
1992, C. & T. obs.; immediately E. of Menetes, 340 m, waste
ground at side of road, 10 April 1992, C. & T. 317; 1.5 km N.
of Airport, 20 m, side of road, 10 April 1992, C. & T. obs.

Greuter, Maltha's & Risse (1985: 36) suggested that this
annual grass is 'actively spreading as a roadside weed in
Greece and on many Greek islands'. The same authors (loc.
cit.) considered it to be fairly recently introduced to Crete,
and now naturalized there. Elsewhere, it is known from
Libya, Egypt and south-western and central Asia.

N.J. TURLAND AND L. CHILTON

Imperata cylindrica (L.) Raeusch. (new to the
Karpathos island group - FCA: 111, 386)

1.5 km SW of Piles, 20 m, wet ditch at side of road, 6 April
1992, C. & T. obs.; 1 km NW of Pigadia, sea-level, damp
ground at edge of Phragmites bed, 11 April 1992, C. & T.
340.

This rhizomatous perennial grass is distributed from southern
Europe to central Asia, as well as in the Old World tropics
and subtropics and Chile. In Crete, it occurs in damp sandy
places, ditches and moist grassland near the sea.

Lamarckia aurea (L.) Moench (new to Karpathos -
FCA: 171,386)

Semi-deserted village 3.5 km N. of Mt Kimaras, 150 m, fallow
cultivation terrace, 8 April 1992, C. & T. 300; S. edge of
Pigadia town, 25 m, weed community in olive groves, 11
April 1992, C. & T. obs.

Lamarckia aurea is an annual grass found from the Mediter-
ranean region (including Crete) eastwards to Pakistan. It was
first recorded from the Karpathos island group by Greuter et
al. (1983: 72), from Saria.

Iridaceae

Romulea ramiflora Ten. susbp. ramiflora (new to
Karpathos - FCA: 179, 396)

Lastos plain, 700 m, fallow field, growing with Romulea
bulbocodium (L.) Sebast. & Mauri, 9 April 1992, C. & T.
302.

This geophyte is distributed in the Mediterranean region,
including Crete, and was first recorded from the Karpathos
island group by Greuter et al. (1983: 74), from Kasos.

Liliaceae

Bellevalia trifoliata (Ten.) Kunth (new to the
Karpathos island group, and confirmation for the
Cretan area - FCA: 182, 401)

Lastos plain, 700 m, scrub, 5 April 1992, C. & T. obs.; 1 km
SE of Lastos plain, 650 m, near margin of abandoned field, 5
April 1992, C. & T. 237; area immediately NE of Stes,
450-470 m, several populations in olive groves, margin of
scrub and abandoned fields, 6 April 1992, C. & T. obs.;
immediately W. of Menetes, by church, 350 m, abandoned
terraces, 10 April 1992, C. & T. obs.

This Mediterranean bulbous species is given as doubtfully
present in the Cretan area by Heywood (1980: 45), presum-
ably on the basis of an old record from Crete cited by
Rechinger (1943a: 728) and doubted by Greuter (1974: 161).
The new records from Karpathos confirm the presence of the
species in the Cretan area. Furthermore, the authors have
recently received confirmation of the occurrence of the
species in Crete itself (Apokoronos: S. of Vrises, by road to
Hora Sfakion, beneath olive tree in stony field, 10 April 1993,
/. Palmer colour transparency!).

Greuter (1974: 161) also casts doubt upon old records of
Bellevalia dubia (Guss.) Rchb. cited by Rechinger (I943a:

CRETAN FLORA 3. ADDITIONS TO KARPATHOS

728) from Kasos. This species has a distribution similar to
that of B. trifoliata and is likewise given by Heywood (1980:
45) as doubtfully present in the Cretan area. Its occurrence
there has recently been confirmed by a record from the
Korikos peninsula in north-western Crete (Strasser, 1988: 6,
22), although its presence in Kasos remains doubtful. It is
possible that the records from that island are in fact referable
to B. trifoliata.

Scilla bifolia L. (new to the Cretan area - FCA: 185,
406)

W. of summit of Mt Kalilimni, 1100 m, head of valley
beneath W. -facing cliffs, among low open scrub of Acer
sempervirens L., 9 April 1992, C. & T. 308; immediately S. of
col S. of summit of Mt Kalilimni, 1150 m, in protection of
spiny shrub beneath NW-facing calcareous cliffs, 5 April
1992, C. & T. 246.

This small bulbous species is widely distributed in central and
southern Europe and south-western Asia. It occurs through-
out most of Greece and on several of the Aegean islands
(Andersson, 1991: 695). However, before its discovery in
Karpathos, it was known in the South Aegean region only on
the island of Rodos, where it has been recorded at altitudes of
1000-1200 m on Mount Attaviros south-south-east of Embo-
nas by Carlstrom (1987: 119, 260), as Scilla 'longistyla' Speta,
a mis-print of S. longistylosa Speta, which is included within
S. bifolia by Mordak (1984: 216). The plants occur in large
numbers at the first cited locality on Kalilimni, and it is
intriguing that this conspicuous and attractive species has
been overlooked there for so long. In the high mountains of
Crete (Lefka Ori, Kedros, Psiloritis, Dikti and Afendis
Kavousi massifs) Scilla bifolia is unknown, even under eco-
logical conditions analogous to the Kalilimni locality.
Instead, there occurs the endemic Scilla nana (Schult. &
Schult. f.) Speta (= Chionodoxa nana (Schult. & Schult. f.)
Boiss. & Heldr., including C. cretica Boiss. & Heldr. and 5.
albescens Speta).

Typhaceae

Typha domingensis (Pers.) Steud. (new to the
Karpathos island group - FCA: 195, 416)

2.5 km N. of Mt Kimaras, 200 m, wet stream-bed in area of
burnt Pinus brutia woodland, 8 April 1992, C. & T. obs.

Typha domingensis is a hygrophilous species distributed in
the Mediterranean region, Asia, tropical Africa and tropical
America. It occurs in Crete, growing in watery places mainly
in the west of the island.

99

ACKNOWLEDGEMENTS. We should firstly like to express our gratitude
for support from the Park Fund, at The Natural History Museum,
which provided all the expenses of the field trip to Karpathos. We
would also like to thank Dr J.R. Akeroyd, for determining Crassula
alata, Dr M.H. Martin (University of Bristol), for determining
Bromus hordeaceus subsp. divaricatus, Mrs I. Palmer, for communi-
cating her Cretan record of Bdlevalia trifoliata, and all the local
people in Karpathos, particularly George Philippidis of Pigadia, who
helped to make the trip both trouble-free and enjoyable.

REFERENCES

Andersson, I. A. 1991. Scilla L. In A. Strid & Kit Tan (Eds), Mountain flora of

Greece 2: 694-697. Edinburgh.
Carlstrom, A. 1987. A survey of the flora and phytogeography ofRodhos, Simi,

Tilos and the Marmaris peninsula (SE Greece, SW Turkey). Lund.
Ciferri, R. 1944. Flora e vegetazione delle isole italiane dell'Egeo. Atti 1st. hot.

Univ. Lab. crittogam. Pavia Suppl., V, A.
Damboldt, J. 1978. Asyneuma Griseb. & Schenk. In P. H. Davis (Ed.), Flora

of Turkey and the East Aegean Islands 6: 65-81. Edinburgh.
Greater, W. 1974. Floristic report on the Cretan area. Mems Soc. broteriana

24: 131-171.

- Burdet, H. & Long, G. (Eds) 1984-1989. Med-Checklist. A critical
inventory of vascular plants of the circum-mediterranean countries. 1, 3-4.
Geneve.

Malthas, U. & Risse, H. 1984. Additions to the flora of Crete, 1973-1983
(1984) - II. Willdenowia 14: 269-297.

1985. Additions to the flora of Crete, 1973-1983 (1984) - III.
Willdenowia 15: 23-60.

Pleger, R. & Raus, T. 1983. The vascular flora of the Karpathos island

group (Dodecanesos, Greece). A preliminary checklist. Willdenowia 13:
43-78.

- & Raus, T. (Eds) 1981. Med-Checklist notulae 4. Willdenowia 11:
271-280.

Heywood, V. H. 1980. Bellevalia Lapeyr. In T. G. Tutin. V. H. Heywood, N.

A. Surges et al. (Eds), Flora Europaea 5: 44-45. Cambridge.
Jalas, J. & Suominen, J. (Eds) 1983. Atlas Florae Europaeae 6. Helsinki.
Mordak, E. V. 1984. Scilla L. In P. H. Davis (Ed.), Flora of Turkey and the

East Aegean Islands 8: 214-224. Edinburgh.
Rechinger, K. H. 1943a. Flora aegaea. Denkschr. Akad. Wiss. Wien 105(1).

19436. Neue Beitrage zur Flora von Kreta. Denkschr. Akad. Wiss. Wien
105(2,1).

Strasser, W. 1988. West-Kreta. Botanische Studien 1987. Steffisburg. Privately

published by the author.
Strid, A. 1986. Ranunculus L. In A. Strid (Ed.), Mountain flora of Greece 1:

210-225. Cambridge.
Turland, N. J. 1992. Studies on the Cretan flora 1. Floristic notes. Bull. Br.

Mus. nat. Hist. (Hot.) 22: 159-164.

Chilton, L. & Press, J. R. 1993. Flora of the Cretan area: annotated
checklist & atlas. London.

Tutin, T. G., Heywood, V. H., Burges, N. A. et al. (Eds) 1964-1980. Flora
Europaea 1-5. Cambridge.

Bulletin of The Natural History Museum

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Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete

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Volume 17 Volume 21

No. 1 Studies in Pseudocyphellaria (lichens) 1. The New No. 1 Historical and taxonomic studies in the genus

Zealand species. D.J. Galloway. 1988. Pp. 1-267, 124 Titanoderma (Rhodophyta, Corallinales) in the British

figs. 51.00 Isles. Y.M. Chamberlain. 1991. Pp. 1-80, 247 figs.

Volume 18

No. 1 An illustrated catalogue of the type specimens in the
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No. 3 Seaweeds of the western coast of tropical Africa and

adjacent islands: a critical assessment. IV. Phodophyta
(Florideae) 2. Genera G.J.H. Price, D.M. John & G.W.
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No. 4 Some Cretaceous and Palaeogene Trinacria (diatom)
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plates. 9.10

No. 5 A monograph of Dryopteris (Pteridophyta:

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Volume 19

A new species of Maytenus (Celastraceae) in Ethiopia.
Sebsebe Demissew. 1989. Pp. 1-3, 1 fig.
Central American Araliaceae - a precursory study for the
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A revision of the Solarium nitidum group (section Holo-
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I Six new species of Solatium sect. Geminata from South

America. S. Knapp. 1989. Pp. 103-112, 8 figs.
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Revision of Piper (Piperaceae) in the New World 1.
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Volume 20

No. 1 Studies in the genus Hypericum L. (Guttiferae) 8.

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No. 2 The marine algal flora of Namibia: its distributions and
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169-192, 13 figs.

Revision of Piper (Piperaceae) in the New World. 2. The
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fig-

A taxonomic study of the species referred to the

ascomycete genus Leptorhaphis . B. Aguirre-Hudson.

1991. Pp. 85-192, 76 figs.

The typification and identification of Calymperes

crassilimbatum Renauld & Cardot (Musci:

Calymperaceae). L.T. Ellis. 1991. Pp. 193-194, 1 fig.

Volume 22

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No. 2 Palynological evidence for the generic delimitation of

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Lira-Saade & J. Caballero. 1992. Pp. 109-121.

Seaweeds of the western coast of tropical Africa and

adjacent islands: a critical assessment. IV. Rhodophyta

(Florideae) 3. Genera H-K. J.H. Price, D.M. John &

G.W. Lawson. 1992. pp. 123-146.

Two new species of Solanum section Geminata

(Solanaceae) from Cerro del Torra in western Colombia.

S. Knapp. 1992. Pp. 147-152.

Fissidens ceylonensis Dozy & Molkenb. (Musci:

Fissidentaceae) and some allied taxa from southern

India. L.T. Ellis. 1992. Pp. 153-156, 2 figs.

New species of Piper (Piperaceae) from central America.

M. Tebbs. 1992. Pp. 157-158.

Studies on the Cretan flora 1. Floristic notes. N.J.

Turland. 1992. Pp. 159-164.

Studies on the Cretan flora 2. The Dianthus juniperinus

complex (Caryophyllaceae). N.J. Turland. 1992.

Pp. 165-169. 37.50

Volume 23

No. 1 Revision of Piper (Piperaceae) in the New World 3. The
taxonomy of Piper sections Lepianthes and Radula. M.C.
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Mounting techniques for the preservation and analysis of
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No. 2 New taxa of Gentiana (Gentianaceae) from Western

China and; the Himalayan region. T.-N. Ho and S.-W.

Liu. 1993. Pp. 55-60. 2 figs.

New combinations, names and taxonomic notes on

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Studies in Hypericum: validation of new names. N.K.B.

Robson. 1993. Pp. 67-70.

Generic monograph of the Asteraceae-Anthemideae. K.

Bremer and C.J. Humphries. 1993. Pp. 71-177. 12

figs. 37.50

CONTENTS

1 Pre-Linnaean references for the Macaronesian flora found in Leonard Plukenet's works and
collections

J. Francisco-Ortega, A. Santos-Guerra and C.E. Jarvis
35 Studies on the lichen genus Sticta (Schreber) Ach.: II. Typification of taxa from Swartz's

Prodromusof 1788

D.J. Galloway
49 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment.

IV. Rhodophyta (Florideae) 4. Genera L-O

D.M. John, G.W. Lawson, J.H. Price, W.F. Prud'homme van Reine and W.J. Woelkerling
91 Studies on the Cretan flora 3. Additions to the flora of Karpathos

N. J. Turland and L. Chilton

atural History Must

BOTANY SERIES

Vol. 24, No. 1, June 1994