ISSN 0968-044(
Bulletin of The Natural History Museum
Botany Series
THE NATURAL HISTORY MUSEUM
VOLUME 28 NUMBER 2 26 NOVEMBER 1998 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum
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The Natural History Museum, 1998 Botany Series
ISSN 0968-0446 Vol. 28, No. 2, pp. 67-150
The Natural History Museum Cromwell Road London SW7 5BD Issued 26 November 1998
Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Hull. nut. Hist. Mux. Loud. (Bot.) 28(2): 67-1 13 HISTORY MUSE A revision of Brillantaisia (Acanthaceae 19 NOV 1998 PRESENTED GENERALLY KAREN S1DWELL 1 Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD
CONTENTS
Introduction 68
Taxonomic history 68 Morphology 69 Habit 69 Indumentum 71
Cystoliths 71 Stem 71 Wood anatomy 71 Leaves 71 Inflorescence 74 Bracts and bractlets 74 Calyx 74 Corolla 74 Androecium 76 Pollen 76 Gynoecium 76 Fruit 76 Seeds 76
Concepts of species and higher taxa 76 The species of Brillantaisia 78 Relations of the species of Brillantaisia 1: formulation of the data matrix 78 Choice of outgroup 78 Choice of characters 78 The data matrices 78
Relations of the species of Brillantaisia 2: analysis of the data matrix 78 Results 79 Taxonomic account 83 Key to fertile specimens of Brillantaisia 83 Nomen dubium 105
Species transferred to Hygrophila 106 Species transferred to Eremomastax 106 References 106 Exsiccatae 107
Systematic index 1 13
SYNOPSIS. BrillantaisiaP. Beauv. is a genus of mainly woody herbs and subshrubs from tropical Africa and Madagascar. In this revision, a brief taxonomic history of Brillantaisia is presented, the morphological variation of the genus is described, and concepts of species and higher taxa are discussed. The Phylogenetic Species Concept of Nelson & Platnick is adopted as the criterion for species delimitation whilst hennigian monophyly is the criterion used for recognition of higher taxa. Twelve species of Brillantaisia are delimited. The relations between the twelve species of Brillantaisia and eleven outgroup species representing the genera, Duosperma, Dyschoriste, Eremomastax, Hygrophila, Mellera and Mimulopsis, are investigated using cladistic methodology and a phylogeny is presented. The results of cladistic analysis show that Brillantaisia and six species ofHygrophila both belong within a monophyletic group based on a suite of pollen characters. Brillantaisia is a monophyletic genus based on
the laterally compressed upper corolla lip, membranous hinge at the apex of the corolla tube and winged petiole. Hygrophila is paraphyletic, and until further studies are undertaken on that genus, the sister group to Brillantaisia remains unresolved. Within Brillantaisia, section Stenanthium of Lindau is monophyletic; however, subgeneric taxa are not formally recognized in this treatment. A taxonomic account of the twelve species of Brillantaisia is provided with illustrations, distribution maps and a key. The main taxonomic changes are that B. stenopteris is new to science; B. riparia is raised from a variety ofB. pubescens to species level; B. grottanellii is recognized as distinct from B. madagascariensis; B. kirungae, B. ulugurica and B. cicatricosa are considered conspecific and B. owariensis is circumscribed as a single widespread and variable species.
The Natural History Museum. 1998
.0 68 K. SIDWELL
on a cultivated Hooker INTRODUCTION species could be seen single specimen. therefore united all the species recognized by Nees (1847) and owariensis P. Bentham ( 1 849) under the earliest name, Brillantaisia The aim of this paper is to present a revision of Brillantaisia with a Beauv.' . key for identification of species, addressing the following questions: Since that time, new of Brillantaisia have been 1 many species in . 1 . How many species are there Brillantaisia described (see Sidwell, 1 997 for detailed discussion). Gustav Lindau 2. Is the genus Brillantaisia monophyletic? was by far the most prolific describer of new species in the genus. 3. Are the subgeneric groups within Brillantaisia proposed by Between 1893 and 1904 Lindau published descriptions of 25 new Lindau (1895) monophyletic? species of Brillantaisia. The first, largest and possibly most import- 4. Is Hygrophila the sister group to Brillantaisia! ant of these publications was Acanthaceae Africanae I (Lindau, 5. Is Hygrophila monophyletic? 1893), in which, ten new species of Brillantaisia were described. Brillantaisia P. Beauv. is a genus of mainly woody herbs and Characters used by Lindau at the species level included habit (herb subshrubs from tropical Africa and Madagascar. Plants of Brillant- or shrub), leaf shape (ovate or obovate), leaf base shape (cordate or aisia generally have ovate leaves with a winged petiole, an open not cordate), leaf apex shape (acuminate or not), cystoliths (present paniculate inflorescence of bilabiate purple flowers and capsules or absent), whole plant indumentum (pubescent or glabrous), sepal containing numerous seeds, each held on a hooked retinacula. At the length (equal or unequal) and number of ovules. By the time of his outset of this study, there was no consensus concerning the delimi- account of Acanthaceae in Engler and Prantl's Die Natiirlichen tation of Brillantaisia, species boundaries within the genus or Pflanzenfamilien (Lindau, 1895a) Lindau had published fourteen species relationships. The genus was thought to contain between 13 names in Brillantaisia, and was the first and only worker to delimit (Vollesen, pers. comm.) and 40 (Mabberley, 1987) species. The subgeneric taxa within the genus. Lindau subdivided Brillantaisia monospecific genus Plaesianthera (C.B. Clarke) Livera from Sri into two sections: three species with spicate inflorescences were Lanka had recently been transferred to Brillantaisia as an Asian included in section Stenanthium Lindau, and nine species with representative of the genus by Cramer (1991); however, this genus paniculate inflorescences were mentioned in section Euryanthium has been shown to be best placed within Hygrophila R. Br. (Sidwell, Lindau. Section Euryanthium was further subdivided on leaf shape in press a) and is not discussed further in this paper. Prior to characters into Group A with lanceolate leaves (containing B. land- undertaking cladistic analyses, Brillantaisia is hypothesized as mono- folia Lindau) and Group B with non-lanceolate leaves (containing phyletic (sensu Hennig, 1950, 1979), based on the presence of two all other species with a paniculate inflorescence). posterior stamens, two staminodes and a laterally compressed upper Brillantaisia is now generally considered to be more closely sister corolla lip; the genus Hygrophila is hypothesized as the group related to the genus Hygrophila R. Br. than to other members of the to Brillantaisia based on previous classifications of Nees ( 1 847), Acanthaceae because both genera possess a bilabiate corolla and Bentham (1876), Lindau (1895fl) and Bremekamp (1953, 1965), numerous seeds. However, historically, the position of Brillantaisia and the monophyly of the taxon Hygrophileae containing only within the Acanthaceae has been somewhat confused. Nees von Brillantaisia and Hygrophila, is provisionally accepted on the pres- 1 first to of Esenbeck ( 847) was the produce a worldwide monograph ence of four aperturate pollen (Scotland, \992a, b, 1993; Furness, the Acanthaceae. He refined his work on Asian Acanthaceae (Nees, 1994) and a unique ndhF sequence (Scotland et al., 1995). 1 832), changing the rank of some taxa, and placed both Brillantaisia (as Belantheria and Leucoraphis) and Hygrophila in the subfamily Taxonomic history Echmatacantheae (possessing retinacula), tribe Hygrophileae (co- Brillantaisia was first described in 1 8 1 8 by Ambroise Marie Palisot rolla 5-parted; stamens 2 or 4; seeds numerous; fruits explosive), de Beauvois from a collection made near Agathon in Benin, West Subtribe 1 . This was the first grouping ofBrillantaisia andHygrophila Africa (Palisot de Beauvois, 1818). A single species, Brillantaisia in the same taxon. However, in the same year Lindley (1847) listed owariensis P. Beauv., was described as having a four angled stem, Brillantaisia as of unknown affinity and noted that three eminent - violet flowers in a panicle and ovate-lanceolate toothed leaves with botanists working on the Acanthaceae Brown, Nees and Meisner - an acute tip and a winged petiole. Thirty years later, Nees von were unable to agree on subdivision of the family. Lindley (1847:
Esenbeck ( 1 847) overlooked the original publication ofBrillantaisia 679) noted that 'there are few natural Orders which now demand, in and described two new genera of Acanthaceae from West Africa: so eminent a degree, a searching investigation as that ofAcanthads'.
1 a Belantheria Nees and Leucoraphis Nees. Bentham (1849) recog- Bentham ( 876) subdivided the Acanthaceae into five tribes using nized the oversight of Nees and transferred the two species of combination of characters (corolla aestivation, ovule number, seed Leucoraphis to Brillantaisia as B. lamium (Nees) Benth. and B. shape and retinacula type). Brillantaisia andHygrophila were placed vogeliana (Nees) Benth. and synonomized Belantheria belvisiana in tribe Ruellieae Benth. (aestivation contorted; seeds 2-many per Nees with Brillantaisia owariensis. In 1853, Hooker addressed the locule; seeds laterally compressed; retinacula hooked), subtribe already problematic issue of species delimitation in Brillantaisia. Hygrophileae (corolla two-lipped; filaments laterally connate).
He agreed with Bentham (1849) that the three species described by Baillon ( 1 89 1 ) recognized six series in Acanthaceae and considered Nees (1847) should be placed within Brillantaisia; however, after Brillantaisia sufficiently distinct to be placed in the monogeneric observation of a newly introduced garden plant from Sierra Leone series Brillantaisiees characterized by the presence of two fertile (Lindley, 1853), Hooker (1853) concluded that the separation of posterior stamens, a strongly bilabiate corolla and many seeds per those species on leaf margin and leaf shape characters was invalid as capsule. the character variation previously used to distinguish between the The research of Lindau (1895a) placed Brillantaisia and
' There remains some confusion here as Hooker did not explicitly state that Belantheria Nees and Leucoraphis Nees (separated on the presence or absence of staminodes) should be united under one species of Brillantaisia. Typographic error increases the confusion as two species of Bentham (1849), Brillantaisia lamium and Brillantaisia vogeliana, are mis-named apparently by Hooker (1853). as 'Belantheria lamium' and 'Belantheria vogeliana', two combinations which had never been formally published prior to that work. The of lumping Brillantaisia lamium, B. vogeliana and B. owariensis by Hooker is not upheld in this work. Although leaf margin characters are very variable within the genus, there are other discrete morphological characters supporting the recognition of these three species as distinct. REVISION OF BR1LLANTA1S1A 69
Hygrophila in the Acanthoideae-Contortae-Hygrophileae having structures using a mm ruler or calipers which measure to 0.1 mm. 'rippenpollen' with four equatorial apertures. Bremekamp (1965) Finer measurements were made using an 8 mm 0.1 graticule in a also placed Brillantaisia with Hygrophila, in Acanthoideae- Leica Wild M8 dissecting microscope. Floral dissections were made Ruellieae-Hygrophilineae, a group with bilabiate flowers in cymes, after placing a flower into cold water and heating slowly until the stamens united into a single group, style not held in place by a row water almost boiled. This usually took about 30 seconds and was or bundles of trichomes and subglobose, banded, usually four- sufficient to soften the delicate corolla. Boiling caused the corolla to aperturate pollen. As mentioned above, recent studies confirmed the disintegrate. Photographs were taken using a Nikon F-801s camera monophyletic nature of the group consisting of these two genera with FP4 black and white film and a Microtech MF80 fibre optic based on a pollen type not found elsewhere in Acanthaceae (Scot- light source. Pollen preparations were acetolysed using a modifica- land, 1993) and a unique ndhF sequence (Scotland et al., 1995). tion of Erdtman (1960) (see Sidwell, 1997). Light micrographs of Since the time of Nees von Esenbeck (1847) no monographic pollen grains were taken using a Dialux 20EB photomicroscope. work has been undertaken for Brillantaisia and the prolific descrip- Scanning electron micrographs were taken at The Natural History tion of species by Lindau (1893; 1894; 1895a, b, c, d; 1896; 1897; Museum, London. Stubs were coated in gold-palladium 80:20% and 1898; 1904) served to exacerbate difficulties in understanding the viewed using an Hitachi S-800 SEM microscope with a working group. Several regional floras (Heine, 1966; Benoist, 1967; distance of 5-10 mm. Permanent slides of Brillantaisia wood were Champluvier, 1985; Vollesen, in prep, a & b) have addressed the prepared according to Gourlay (pers. comm.): the stem was boiled problem of the number of species within Brillantaisia in Gabon, for an hour, cut in transverse, tangential longitudinal and radial Madagascar, Rwanda and East Africa respectively. The authors of longitudinal sections on a sliding microtome and stained in safranin. these works disagree in particular on the number of species in The sections were passed through a dehydration series, washed three several morphologically distinct groups within Brillantaisia. Tax- times in alcohol and mounted in Canada balsam. onomists working on Brillantaisia have tended to split it into poorly defined groups and formally name many slightly differing forms of Habit the great morphological variation within the genus. Many new species ofBrillantaisia have been described, often from little material Brillantaisia species range in habit from delicate herbaceous and based on characters that are highly variable within the genus. Brillantaisia land/alia Lindau, approximately 50 cm high, to the Identification of specimens has been inconsistent and a study cover- large woody species B. kirungae Lindau which reaches 7 metres tall. ing the entire range of geographical and morphological variation of Within this range, Brillantaisia species are mainly erect, perennial the genus is essential to clarify this confusion. herbs around 1 .5 metres tall which often become woody at the base of the stem. Brillantaisia lamium, B. vogeliana and B. mada- gascariensis T. Anderson ex Lindau, for example, are all erect, perennial herbs that produce adventitious roots at the lower nodes or MORPHOLOGY along branches that touch the ground, enabling them to spread vegetatively. The terms herb, shrub and tree have been loosely Initially I considered it impossible to separate description of the applied by collectors to describe the habit of Brillantaisia species. morphological variation of Brillantaisia from discussion of the For example, B. madagascariensis ranges in habit from a 'soft transformation that information into succulent herb' 607 1 or a herb with of comparative morphological (Mooney , S) 'perennial creeping data (characters and character states) for phylogenetic analysis. and ascending stems' (Friis & Vollesen 613, C) to a 'shrub to 2m tall' However, morphological data are analysed at two different levels in (Kibuwa 5191, UPS). According to the model of plant architecture, this study. Characters and character states of individual specimens described by Weberling (1989) after Troll (1964), plants can be are analysed when delimiting species of Brillantaisia, and charac- divided into four distinct and clearly defined zones: main flores- ters and character states of taxa are analysed when forming hypotheses cence; enrichment zone; inhibition zone and innovation zone. This of relations among those species. In order to emphasize the filtering model was used by Manktelow (1996) in her monograph of of data undertaken when transforming observation and description Phaulopsis and can be applied to species of Brillantaisia (Fig. 1). into comparative data, a descriptive approach to discussion of Figure 1 illustrates the architecture of Brillantaisia from the apex characters is presented prior to discussion of character coding for to the base of the plants as follows. The inflorescence is divided here cladistic analysis. A purely descriptive morphological section pro- into two distinct and more precisely defined parts: the terminal vides an overview of the genus, directly comparable with many thyrse with lateral dichasial branches forms the main florescence published monographic studies and some flora accounts for the and flowering lateral branches (coflorescences) which 'repeat the Acanthaceae, recent examples of which include studies on Justicia structure of the main axis to some extent' (Weberling, 1989: 225) (Graham, 1988), Justicia sect. Harnieria (Hedren, 1989), Justicia form the enrichment zone. These two parts are separated from one sect.Ansellia (Ensermu, 1990), Ruellia (Ezcurra, 1993), Strobilanthes another by the basal internode (bi on Fig. 1), and the whole flower- (Wood, 1995), Dicliptera (Balkwill et al., 1996) and Phaulopsis ing region is termed a synflorescence. Below the enrichment zone, (Manktelow, 1996). Brief discussion of morphological variation is where the axillary buds do not develop into coflorescences, is the given here and species that exhibit the range of variation of a inhibition zone, and below that, the region of vegetative growth is is considered the particular organ are used as examples. Herbarium specimens rel- termed the innovation zone. The innovation zone evant to the discussion are referenced by collector's name, number region of production of new shoots after flowering in perennial B. B. owariensis and B. and herbarium acronym (e.g. Kibuwa 5191, UPS). plants. Brillantaisia lamium, vogeliana, The data were obtained through observation of herbarium speci- madagascariensis all exhibit the type of growth pattern described mens from BM*, BR*, C, DSM*, E*, ETH, FHO*, G, HBG, K*, above and illustrated in Figure lA.Two species, B. lancifolia and/?. LISC, M, MHU, MO*, NY, P*, PRE, S, SCA*, SRGH, W, WAG*, debilis Burkill have a main florescence and lack the enrichment zone YA* and Z (abbreviations follow Holmgren et al., 1990). Visits have (Fig. IB). In the annual species, B. pubescens T. Anderson ex Oliv. been made to the herbaria marked with an asterisk. Gross morpho- inhibition of the lateral shoots does not occur, lateral coflorescences to the base of the and the inhibition and innovation zones logical features were measured on mature vegetative and reproductive occur plant 70 K. SIDWELL
Inhibition Innovation zone ' ' zone
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are absent (Fig. 1C). In two species ofBrillantaisia, the main axis of Wood anatomy the plant continues to grow vegetatively above the flowering region. A review of the wood anatomy of 38 species of Acanthaceae was This phenomenon is termed proliferation. Troll (1964) considered written by Carlquist & Zona (1988). No species of Brillantaisia or the type of vegetative growth seen in B. oligantha Milne-Redh. and Hygrophila were included in his survey which concluded that 'Woods occasionally in B. riparia (Vollesen & Brummitt) Sidwell to be late of Acanthaceae are characterized by relatively narrow vessels with proliferation which 'consists of the return of the inflorescence apex and alternate lateral wall to a simple perforation plates pitting, septate vegetative condition' (Weberling 1989: 257; Fig. ID). The libriform vasicentric axial robust fibers, scanty parenchyma, rays both Brillantaisia kirungae has been described as a 'very large, multiseriate and uniseriate, erect ray cells abundant in rays ... straggling, bushy herb' (Ball 62, SRGH), a 'robust subshrub' (Bridson numerous small crystals or in cells . . . and nonstoried & Lovett 544, MO) or even a 'small tree' (Thomas 3790, MO; White cystoliths ray structure' (Carlquist & Zona, 1988: 201). The stems of plants of 13908, FHO). White's description of B. kirungae includes bole Brillantaisia often become lignified at the base, particularly in B. measurements which clearly indicate that the plant is a tree, 4 m high owariensis and B. kirungae. Data on woodiness are incomplete, with a trunk 1 .4 m tall and 1 2 cm in diameter at 70 cm height (White, however, as the lower stem and woody rootstock are very rarely 1984); however, such accurate measurement and application of collected on herbarium specimens. One specimen (Poulsen 523, terminology is unusual. I was unable to study this species in the field FHO) included a collection of the lignified lower stem of B. kirungae and the precise pattern of architecture of B. kirungae remains 20 mm in diameter. This of stem showed a clear central with uncertain. piece pith wood 7 mm thick and bark 1 mm thick. Transverse sections (Fig. 2A, Indumentum B) show the vessels, which are relatively broad for the family, to be usually solitary and arranged radially with columns of parenchyma The variation of trichomes on different of Acanthaceae has organs forming medullary rays between the regions of vessels. The vessels been studied in some detail by previous researchers Ahmad, (e.g. have simple perforation plates and minute pitting on the outer walls 1 976, 1 978). Singh & Jain ( 1 975) studied the structure and ontogeny (Fig. 2F). The fibres are long, thin-walled, septate cells (Fig. 2C, E, of trichomes of Acanthaceae and classified them into in forty types F) between which are regions multiseriate rays characterized by a two major groups: glandular and eglandular. ofBrillantaisia Species predominance of erect ray cells (Fig. 2C, D). Cystoliths are visible in both and trichomes. possess glandular eglandular Eglandular ray cells but cannot be seen in the sections shown here. trichomes on Brillantaisia are of three types: simple filiform trichomes, three to nine cells long, tapering towards the apex; short Leaves uniseriate trichomes, two cells long, also tapering to the apex and The leaves ofBrillantaisia are characteristically ovate with a unicellular trichomes with a rounded apex that is often purple. winged petiole. However, they vary enormously in size, shape, margin, the Glandular trichomes are stalked with a multicelled, capitate head extent of the petiole wing and the distribution and density of (Selvaraj & Subramanian, 1 983) and a two to many celled stalk. The trichomes on both surfaces. Leaf shape ranges from linear-lanceo- number of cells in the head and stalk of glandular hairs varies late to ovate The be or continuously between organs and among individuals ofBrillantaisia. broadly (Fig. 3). petiole may unwinged, winged to the base. The margin is often irregular, sometimes having Detailed discussion of indumentum types on the different parts of dentate, crenate and serrate teeth and be toothed. Brillantaisia plants is presented in Sidwell (1997). may secondarily The leaf texture ranges from chartaceous in B. vogeliana to slightly Cystoliths coriaceous in B. owariensis and B. kirungae. The venation is eucamptodromus with tertiary veins becoming perpendicular to the Calcium carbonate cystoliths are present in most members of midrib toward the apex of the leaf. In B. kirungae, the leaves often Acanthaceae and were extensively documented in the last century by dry black. Some Brillantaisia species can be recognized to species Hobein (1884). Recent studies by Inamdar et al. (1990) indicated level when sterile. For example, B. lamium has perhaps the most that variation of cystolith morphology may be a useful taxonomic characteristic leaves of all the species, with a usually cordate leaf character above the generic level in Acanthaceae. The cystoliths of base, an entire margin and an unwinged petiole (Fig. 3A). Brillantaisia are solitary, elongated and pointed at one end. All Brillantaisia land/alia has lanceolate, entire leaves (Fig. 31). Plants vegetative parts, bracts, bractlets and calyces are covered in this type of B. madagascariensis, B. debilis, B. oligantha, B. pubescens and of cystolith. The size of cystoliths in Brillantaisia varies B. riparia all possess entire-margined, ovate leaves with a cuneate intraspecifically and even within a single organ of one specimen. leaf base narrowly tapering into a winged petiole (Fig. 3B, F, G, H, The with toothed leaves are Stem L). species very variable and relatively difficult to distinguish from one another. Leaves of B. vogeliana can In this work, the stem is treated as the axis of the plant below the be easily recognized by the presence of irregular teeth on the petiole main florescence and basal internode as shown in Figure 1 and is wing, particularly where it widens into the lamina base (Fig. 3C),
to the 1 is equivalent 'hypotagma' of Weberling ( 989: 230). The stem and by the rather open tertiary venation with unclear quaternary square towards the apex, longitudinally grooved on opposite sides veins. Leaves of Brillantaisia kirungae are also characteristically and more rounded towards the base of the plant. The corners of the grossly, irregularly toothed (Fig. 3E) and have prominent horizontal upper stem are usually slightly curved but may be sharply angled and and vertical tertiary and quaternary veins giving a very close squared winged. The stem is slightly swollen at or just below the nodes in all pattern. The leaves of B. owariensis exhibit a large range of leaf size, species. Below the basal internode the stem varies in diameter from shape and margin type (e.g. Fig. 3K). The leaves often have promi- 1-2 mm in Brillantaisia debilis to c. 10 mm in B. kirungae. In B. nent, parallel tertiary veins running between the secondary veins, madagascariensis, B. vogeliana and B. lamium adventitious roots with the quaternary veins not clear. Unfortunately, herbarium speci- may be produced at the lower nodes. The epidermal stem surface is mens tend not to show the extent of variation of leaf size in the genus. covered in cystoliths, oriented parallel to the stem. Multicellular Brillantaisia kirungae and some forms of B. owariensis have very trichomes which may be glandular or eglandular are present on the large lower leaves which have rarely been collected as they do not upper stem and are particularly dense at the nodes. readily fit into a plant press. 72 K. SIDWELL
D /I I'M ti
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Fig. 2 Wood anatomy of Brillantaisia kirungae (Poulsen et al. 523, FHO, C, K). A. TS x 50; B. TS x 250; C. TLS x 50; D. TLS x 250; E. RLS x 50; F. RLS x 250. REVISION OF BR1LLANTAISIA 73
Fig. 3 Leaf shape variation in Brillantaisia. A. Brillantaisia lamium (Figuieredo & Arriegas 32, LISC); B. B. debilis (Louis 2161. WAG); C. B. vogeliana (Manktelow et al. 89, UPS); D. B. lamium (1FI 32657, MO); E. B. kirungae (Friis & Vollesen 58, C); F. B. oligantha (Brass 17042, MO); G. B. madagascariensis (Loveridge 71, MO); H. B. pubescens (Vollesen 2392, UPS); I. B. lancifolia (Talbot 200, BM); J. B. grottanellii (Ensermu & Zerihun 619, ETH); K. B. owariensis (Barbosa 1812, LISC); L. B. riparia (Robyns 151, BR). 74 K. SIDWELL
Inflorescence again. Indeterminate main florescences may also have lower coflorescences as seen in B. riparia (Fig. 4F). Brillantaisia inflorescences have been described in general terms The development of a general terminology for Acanthaceae inflo- as 'panicules' (Heine, 1966: 83), 'panicules terminales amples ou rescences to help clarify higher level homology assessment is spicifomes, formes de cymes laches ou contractees' (Benoist, considered essential, but beyond the boundaries of this study as it 1967: 28) or 'large open or contracted panicles, rarely in spiciform will require detailed ontogenetic, anatomical investigations across a racemes' (Vollesen, in prep. a). As with many genera of wide range of taxa. Acanthaceae, it is often difficult to pinpoint a precise division between the vegetative and floral parts of Brillantaisia plants and Bracts and bractlets inflorescence structure and variation cannot be described by The bracts of Brillantaisia are paired at the nodes of the main simply using the broad conventions of botanical terminology such florescence axis. In most species of Brillantaisia the bracts are as panicle, raceme, spike or cyme. Although several workers have foliose, becoming smaller towards the apex of the inflorescence. The studied the morphology of Acanthaceae inflorescences in detail, bracts are often caducous, for example, B. owariensis, has large leaf- for example in the genera Pseuderanthemum, Ruellia, Barleria like bracts present in the young developing inflorescence which fall and Lepidagathis (Sell, 1969fl), Anisotes (Baden, 1981), Justicia soon after the inflorescence The bracts of B. kirungae are (Graham, 1988), Duvernoia and Adhatoda (Manning & Getliffe- opens. large, rounded at the apex, amplexicaul and pubescent with short Norris, 1985), Ruellia (Ezcurra, 1993), Phaulopsis (Manktelow, trichomes. Amplexicaul bracts are not seen in other species of 1996) and Dicliptera (Balkwill, 1996) and discussed the problem Brillantaisia. Hedren (1989: 21 ) used the term bracteole to refer to of appropriate descriptive terminology, there has been no attempt 'small, narrow bracts', differentiating them from 'leaf-like bracts'. to reach a consensus regarding the application of terms to the His is considered as 'bracteole' inflorescences of Acanthaceae. Several of these studies (Baden, terminology imprecise specifically refers to 'a leaflike a flower in an inflorescence 1981; Manning & Getliffe Norris, 1985; Graham, 1988) illustrated organ subtending that is itself subtended by a bract' (Blackmore & Tootill, 1984: 48) and discussed modifications of different inflorescence types from and does not refer to the size or of the The bractlets of an 'ancestral' compound dichasium via 'elaboration', 'reduction' shape organ. Brillantaisia do not subtend the flowers, but occur at the dichoto- and 'condensation' (Graham, 1988: 555). Hedren's (1989: 20) mous branching of the dichasial florescence. The term bractlet is terminology for Justicia sect. Harnieria attempted to 'find a sim- used (as by Daniel, 1984) in preference to the term bracteole and ple and consistent terminology for supposedly homologous refers to the second, third or higher order leaf-like structures at the structures'. However, his terms were different from those previ- nodes of the lateral branches of the main florescence or ously used for Justicia or other Acanthaceae and have not been coflorescences. The bractlets of most species of Brillantaisia are generally applied at that level. The lack of generally applicable linear, sessile, and rounded at the apex. Plants of Brillantaisia terminology and the lack of parallel usage of available terminol- pubescens have petiolate, rounded bractlets which are ogy has meant that the easiest solution for each botanist working shortly per- sistent and characteristic of the on Acanthaceae inflorescences has been to develop his or her own highly species. clear, explicit terminology for a particular group of plants. Brillantaisia inflorescences are described below in light of both Calyx important general works on inflorescence morphology (Rickett The calyx of Brillantaisia has five linear or spathulate, pubescent 1944, 1955; Troll, 1964; Sell, 1969a; Weberling, 1989) and the sepals that are united at the base. The upper adaxial sepal is usually studies of Acanthaceae mentioned above. larger than the other four although B. lamium and some forms of B. The model of the habit of Brillantaisia species described above owariensis have five subequal sepals. In B. kirungae, the upper sepal (Fig. 1 A) is used as a starting point for understanding inflorescence may be several times wider and longer than lateral sepals. Plants of architecture. The habit model divides the inflorescence into a termi- B. pubescens, B. riparia and B. vogeliana have clearly spathulate nal, main florescence and a lower enrichment zone of lateral sepals. During fruit maturation, the calyx may lengthen significantly. coflorescences, the whole flowering region of a plant being termed In B. pubescens, the largest sepal is as long as the fruit. In B. a synflorescence (Fig. 4A-F). The basic 'unit' of Acanthaceae owariensis the upper sepal is occasionally much longer than the fruit. synflorescences is the dichasium, described by Rickett (1944: 216) as 'a cluster formed a by dichotomy beneath a terminal flower' Corolla which 'in its simplest form consists of three flowers' (Fig. 4A). In Brillantaisia synflorescences, the basic dichasium structure is re- The corolla of Brillantaisia has left contort aestivation (Scotland et peated and modified so that in most species the amount of dichasial al., 1994) common to all members of the Contortae sensu Lindau and monochasial branching increases, more or less regularly, down (1895a). The two lipped corolla of Brillantaisia is highly character- the synflorescence from the apex (Fig. 4A & B). Following the istic of the genus. The corolla tube is cylindrical, the upper lip is
terminology of Weberling (1989: fig. 109) the main florescence of hooded and strongly laterally compressed, the lower lip is usually Brillantaisia is usually a regular, dichasially branching thyrse or reflexed with three lobes at the apex. Between the base of the lower double thyrse (Fig. 4A). Below the main florescence, lateral lip and the corolla tube is a region of membranous folded tissue coflorescences may be present (Fig. 4C). Plants of B. which acts as a hinge allowing the corolla lobes to pivot on the madagascariensis and B. grottanellii Pichi-Sermoli have very short corolla tube presumably facilitating pollination. Corolla morphol- lateral inflorescence branches, hence the spicate florescence on ogy varies in the length and width of the tube, the ratio of the tube to which Lindau based his sectional the division of Brillantaisia men- lobes, the shape of the lower lip, and the pubescence of the petals tioned above (Fig. 4D).The apex of the main florescence may return (see Figs 8-16). Corollas range in colour from deep purple to pink to indeterminate vegetative growth as in B. oligantha (Fig. 4E). In with white forms found occasionally in B. vogeliana, B. lamium and this case, the lateral dichasia are produced in the axil of a leaf rather B. madagascariensis. The West African white-flowered B. than a bract and the question of whether the whole structure is the madagascariensis was previously recognized, on the basis of this main florescence or a synflorescence of lateral dichasia arises once character alone, as a separate species. REVISION OF BRILLANTAISIA 75
Determinate apex
Leaf B
Indeterminate apex
Leaf D
Fig. 4 Synflorescence structure of Brillantaisia species. A. basic structure of main florescence: determinate at the apex, with lateral branching once or twice dichasial; bracts on main axis foliose becoming smaller towards the apex, smaller foliose bractlets on lateral branches; B. main florescence with lateral branching both dichasial and monochasial; C. main florescence with lower lateral coflorescence repeating the structure of the main florescence, the whole forming a synflorescence; D. main florescence with reduced lateral branches forming a spike with large persistent bracts; E. main florescence with indeterminate apex; F. main florescence with indeterminate apex and lower lateral coflorescence. 76 K. SIDWELL
Androecium Fruit
Species of Brillantaisia have two stamens and two staminodes. It is The fruit of Brillantaisia are dry, elastically dehiscent capsules with the two posterior stamens in Brillantaisia that are fertile, unlike retinacula (Figs 8-16), characteristic of many members of the other Acanthaceae with two stamens in which the anterior pair Acanthaceae (Sell, 19696). The capusles are erect on the lateral develop (e.g. Fig. 12C).This was mentioned by Lindau (1895a: 278) synflorescence branches, each valve has a longitudinal groove from who stated 'Die Reductionen imAndroecium gehen stets so vor sich, base to apex and within each valve, the seeds are held on two rows dass die hinteren Stb. zuerst zu Staminodien werden oder ganz of dry, comparatively large, hook-shaped retinacula (Fig. 14A & E). verschwinden, nur bei Brillantaisia finden sich ausnahmsweise die The retinacula is a hardened outgrowth from the funicle of the beiden vorderen Stb. zu Staminodien umgebildet'. The fertile an- developing embryo. The capsules of Brillantaisia are many seeded thers are symmetrical, sagittate, dorsifixed and dehiscent through (Figs 8-16), compared to the capsules of most Acanthaceae which long lateral slits. The filaments are flattened towards the base, usually contain only 2 or 4 seeds. Often one or two ovules at the narrowing towards the anther and are almost as long as the corolla narrowed base or apex of the fruit do not develop into fully mature
lobes and held in the upper hooded lip. The staminodes usually seeds. The fruit vary between species in size and the number of seeds extend beyond the corolla tube and have a small flattened vestigial per capsule and pubescence, for example: the fruit of B. pubescens anther at the apex. Infl. pubescens (Fig. 8E), B. riparia (Fig. 9B) and are the smallest in the genus (Fig. 8F); plants of B. vogeliana have B. oligantha (Fig. 10B) the staminodes are minute or absent. small fruit with up to 30 seeds per valve (Fig. 1 3A & C); plants of B. kirungae have large glandular pubescent fruit with 8-10 seeds per Pollen valve (Fig. 12E) and the fruit of B. debilis and B. lamium are glabrous (Figs 14A & 16C). The pollen grains of Acanthaceae are very variable and have been used extensively in taxonomic research on the family (Radlkofer, Seeds 1883; Lindau, 1895a; Bremekamp, 1944; Raj, 1961; De, 1960; Immelman, 1989; Scotland, 1991, 1992a, b, 1993; Furness, 1994). Seed surface characters have been shown to be taxonomically useful The pollen of Brillantaisia is usually oblate spheroidal but ranges at the species level in several groups of Acanthaceae including from suboblate to spheroidal (terms according to Punt et al., 1994). Hypoestes (Balkwill & Getliffe-Norris, 1985) and Peristrophe
1 et in the Lindau ( 895a) used the term rippenpollen to describe the usually (Balkwill al., 1986), and recognition of infrageneric taxa in oval-ellipsoidal grains with pseudocolpi and equatorial apertures Justicia (Immelman, 1990; Lester & Ezcurra, 1991) and found in Brillantaisia. Pollen grains of Brillantaisia are four- Siphonoglossa (Immelman, 1990). The production of mucilage aperturate (see Fig. 5 and plates in Furness, 1994). The apertures are (myxospermy) is common in the Acanthaceae. The mucilage was compound colporate structures consisting of an elongated colpus considered by Kippist (1845) to come from pores in the ends of the with a circular endoaperturate pore in the centre. Between the four seed trichomes. Grubert (1974) said the mucilage was produced by apertures are pseudocolpi, similar to the colpi but lacking an the seed coat. endoaperture. The number of apertures on a pollen grain occasion- The seeds ofBrillantaisia were first described in detail by Schaffnit ally varies and may range from four to six within a single sample. In (1906). The seeds are flattened, asymmetrical, kidney-shaped and one pollen sample of B. owariensis studied by Furness (1994) all the covered with long, hygroscopic trichomes which are adpressed pollen grains had five apertures, but this character has not been when the seed is dry. When wet, the seeds produce mucilage and the consistently found in other samples of the same species or elsewhere hygropscopic trichomes rapidly expand. Trichomes of seeds of all in the genus. Scanning electron micrographs (Fig. 5) clearly show Brillantaisia species are along, thin, single cell tapering towards and thai Brillantaisia pollen has raised bands of tectum with pseudocolpi rounded at the apex with regular annular and occasionally spiral between these bands. The collumellae are fused into a distinct thickening from base to apex. Similar trichomes are common to all bireticulate tectum in most species of Brillantaisia forming a pri- species of Brillantaisia andHygrophila (Schaffnit, 1906), and were mary reticulum with a smaller reticulate secondary tectum in the also reported in Ruellia by Kippist (1845). lumen (Fig. 5E). Brillantaisia madagascariensis has very character- istic pollen with irregularly clavate tectum ornamentation (Fig. 5A-C). Brillantaisia is similar to grottanellii morphologically very CONCEPTS OF SPECIES AND HIGHER TAXA B. madagascariensis, however, investigation of pollen morphology showed this species to lack any tectal ornamentation (Fig. 5D-F). The of the and methods used These data combined with gross morphological data, support recog- importance explicitly stating concepts for of taxa in botanical was nition of B. grottanellii as a distinct species. recognition monography highlighted by Luckow (1995) and McDade (1995), among others, and has been Gynoecium discussed in relation to a revision of Brillantaisia in an earlier paper (Sidwell, in press b). Implicit taxonomic convention that the per- The of Brillantaisia is 1-2 mm in diameter ovary species cylindrical, sonal opinion of an expert botanist is sufficient to delimit taxa was and to 6 mm and at the to the base of the up long pointed apex, tapering deemed unnacceptable for an unbiased scientific study of style (e.g. Fig. 9D). The sits on an annular disc and has ovary parietal Brillantaisia. The literature on species concepts was carefully re- with two rows of placentation ovules on each placental axis. viewed and evaluated and a brief outline of the conclusions of that Brillantaisia species have between twelve and ovules in contrast sixty paper is presented below. Above the level of species, relationships of to most other Acanthaceae which have four ovules usually just per higher taxa are hierarchical and all groups are recognized on the ovary. The is linear and with a small flattened at style slightly region single criterion of Hennigian monophyly. Species are thought of as the apex forming the end 12D). of stigma (Fig. Many species different from higher taxa and are delimited independently from Acanthaceae and related families have a bifid whereas simple stigma considerations of monophyly and prior to any cladistic analysis. the second lobe of the stigma of Brillantaisia is reduced to a barely They are 'consistent with cladistic theory but independent of con- visible raised 'tooth' at the base of the receptive stigmatic surface. straints of autapomorphy' (Nixon & Wheeler, 1990: 213) and can be REVISION OF BRILLANTAISIA 77
x 1 .5 and B. A-C B. pollen (Gutzwiler 841): A. SEM polar view K; Fig. 5 The pollen of Brillantaisia madagascariensis gmttanellii. madagascariensis of view x 1 K with clavate on the tectum. D-F the smaller pollen B. SEM close up of tectum x 7 K; C. Light micrograph, equatorial 'spines' x 7 F. view x 1 K, with D. view x 1 .5 K; E. SEM close up of tectum K; Light micrograph, equatorial B. gmttanellii (Ensermu et al. 821 ): SEM subpolar an even, non-spiny bireticulate tectum. 78 K. SIDWELL
and influences the results 'recognized by either unique characters or unique combinations of vation and analysis greatly [of phylogeny little '.Thiele characters' (Luckow, 1995: 595). Nelson & Platnick (1981) noted reconstruction], but has nevertheless received attention described the method of of the fact that 'species' are only represented by the samples available (1993: 275) converting description into data for as a filter that to a particular biologist and that in practice 'those samples that a morphology comparative analysis the of variation and the of biologist can distinguish, and tell others how to distinguish (diag- 'operates between discovery recording matrix. Details of of the filter are nose), are called species' (Nelson & Platnick, 1981: 11). However, that variation in the data operation is The main issues to be considered in they go on to point out that a pattern based species concept often obscure...'. coding - are the of continu- incomplete without an element of process that of self perpetuation morphological data for cladistic analysis coding - versus discrete characters Pimentel & 1987; and state that '. . . species are simply the smallest detected samples ous (e.g. Riggins, of self-perpetuating organisms that have unique sets of characters' Cranston & Humphries, 1988;Chappill, 1989;Batemanetal., 1992; the treatment of multistate versus data (Nelson & Platnick, 1981: 12). Gift & Stevens, 1997), binary (compare Pimentel & Riggins, 1987; Pleijel, 1995; Wilkinson, ordered versus unordered characters Mickevich & The species of Brillantaisia 1995), (see Lipscomb, 1991: 127; Scotland, 1992c: 16; Quicke, 1993: 25 for Twelve of Brillantaisia were All twelve species species recognized. illustrations of different character state transformation series), and could be recognized on the presence of unique sets of morphological missing entries in a data matrix (Doyle & Donoghue, 1986; Nixon & characters; however, two kinds of are present. Seven species group Davis, 1991; Platnick, 1991; Maddison, 1993). of these species are monothetic, possessing a single character diag- In this study, 'ordinary character coding is based on the notion of nostic for that and five polythetic, lacking any species group species character- state transformation series, in which states of the same character. As well as within the definition of single diagnostic falling character are seen as transformations of one another, and the whole a defined by the possession of a unique set of polythetic group, assemblage of mutually transformable states forms a character' (de characters, Brillantaisia owariensis could also be considered a Pinna, 1996: 10). In this paper, the term binary refers only to widespread and variable 'oc/z/o-species' (Sidwell, 1997). The term presence/absence characters and not to two-state characters, for was first coined by White (1962: 79) to describe three oc/z/o-species example 2 or 4 stamens. Multistate characters are consequently of which are all and species Diospyros (Ebenaceae) widespread taken to be all characters with two or more states. This definition 'have very complicated variation patterns'. White wrote 'Some of clarifies the difference between binary and two-state characters their variation is closely correlated with geography, but most of it is where the may mean two completely different things: either is the not, so that the pattern is of the checkerboard type discussed by Mayr absence of character state 1 (binary), or is the presence of a (1942: 37) but more complicated than any of his examples. Such different character state homologous to 1 (two-state/multistate). species cannot be satisfactorily subdivided and could conveniently Only discrete characters were selected for cladistic analysis. Con- be distinguished from monotypic and polytypic species by calling tinuously variable characters were excluded in this study because of them oc/z/o-species' (White, 1962: 79). Without further extensive subjectivity in coding character states (Pimentel & Riggins, 1987; and detailed field study of numerous populations of B. owariensis, Gift & Stevens, 1997) and because these data inflate tree length recognition of subspecific taxa remained informal. (Cranston & Humphries, 1988). All assumptions regarding the direction of transformation series were considered inappropriate prior to analysis (Hauser & Presch, 1991) and characters were RELATIONS OF THE SPECIES OF treated as unordered throughout. BRILLANTAISIA I: FORMULATION OF THE DATA MATRIX The data matrices
In preliminary analyses, discussed in Sidwell (1997), 48 characters were scored for 23 taxa forming a baseline data matrix from which Choice of outgroup all further analyses were derived. Missing data were present in 19/48 Multiple outgroups were chosen. Species were selected both from characters and in 19/23 taxa either because it was inapplicable to within the tribe Hygrophileae (Hygrophila species) and outside the score a character for a particular taxon, or because a taxon was tribe Hygrophileae. Within the tribe Hygrophileae, six species of polymorphic for a particular character. In the analysis presented Hygrophila were selected. These species represent the morphologi- here, the baseline data matrix was transformed to remove both cal and geographical variation of the genus, they are well delimited inapplicable and polymorphic missing data from the analyses. Three with no problems regarding species concept and they are repre- groups of characters were receded to remove inapplicable missing sented by plenty of herbarium material at K and BM. Outside the data following Maddison (1993) and seven taxa, polymorphic for at tribe Hygrophileae, single species of the five genera considered least one character, were divided into monomorphic subunits fol- most closely related to the Hygrophileae by Vollesen (pers. comm.) lowing Nixon & Davis (1991). The 32 characters used in these
were included in the analysis. analyses are presented in Table 1 . The data matrix of 32 terminal units scored for the 32 characters is presented in Table 2. Choice of characters
Stevens (1991), Wilkinson (1995) and Gift & Stevens (1997) all noted that individual scientists take very different approaches to RELATIONS OF THE SPECIES OF character choice and character state delimitation and unfortunately, BRILLANTAISIA 2: ANALYSIS OF THE DATA rarely explicitly state the criteria used at this stage of their research. MATRIX Stevens (1984: 395) stated that 'the common meeting place of the systematist and morphologist is the analysis of similarity and its Data were analysed using the mh*bb* command of HENNIG86 conversion into hypotheses of homology' and recently Pleijel (1995: version 1.5(Farris, 1988). Tree Gardener version 2.2 (Ramos, 1997) 309) wrote that 'character coding represents the link between obser- was used to edit matrices, run HENNIG86 via a windows shell and to REVISION OF BR1LLANTAISIA 79
Table 1 List of 32 characters, numbered 0-3 1 to fit with conventions of HENNIG86 (Farris, 1989), scored for cladistic analysis of 32 taxa of Acanthaceae listed in Table 2.
No. Character description
0. Inhibition and/or innovation zones below synflorescence: absent (0), present (1)
1 . Leaf blade shape: broadly ovate to ovate 1:1-2 (0), linear-lanceolate ( 1 ) 2. Leaf margin: entire or subentire (0), strongly rather irregularly toothed (1) 3. Leaf pubescence: glabrous (0), pubescent (1) 4. Petiole: absent (0), present unwinged (1), present entire or subentire wing (2), present clearly toothed wing (3) 5. Proliferation zone: absent (0), present (1) 6. Main florescence shape: spicate (0), paniculate (1)
7. Enrichment zone of axillary/lateral inflorescences, absent (0), present (1) 8. Bract shape: broadly ovate/ovate/linear (0), obovate/rounded (1) 9. Bract base: not clasping rachis (0), clasping rachis (1) 10. Bract pubescence: eglandular (0), glandular (1) 11. Sepal length: equal/subequal (0), unequal (1) 12. Sepal shape: linear (0), spathulate (1) 13. Sepal fusion: free (0), fused (1)
14. Sepal width: upper sepal slightly wider than the lower four (0), upper sepal highly developed, usually twice as wide as the lateral sepals (1)
15. Corolla shape: not bilabiate (0), bilabiate hooded but not laterally compressed (1), bilabiate hooded and laterally compressed (2),
16. Corolla hinge: absent (0), present (1)
17. Upper corolla lip pubescence: subglabrous (0), glandular pubescent (1)
18. Stiff trichomes on lower corolla lip: absent (0), present (1) 19. Stamen arrangement: four fertile stamens (0), two posterior stamens with staminodes ca 1mm long (1), two posterior stamens with staminodes >2mm long (2), two anterior stamens with no staminodes
20. Anther spur: absent (0), present (1) 21. Number of colporate apertures in pollen: three (0), four (1) 22. Colpus size: long (0), short/very short (1) 23. Paired aperture position: equatorial (0), opposite pairs at uneven height (1) 24. Atrium at pore: absent (0), present (1)
25. Pollen shape: spheroidal (0), prolate or subprolate (1) 26. Collumellae shape: not branched at base (0), branched at base (1) 27. Ornamentation: single reticulum (0), double reticulum (1) 28. Spines on pollen: absent (0), present (1) 29. Style: not persistent in fruit (0), persistent in fruit (1) 30. Number of ovules in the fruit (both locules): less than ten (0), greater than 10(1)
1 3 1 . Fruit trichomes: absent (0), present ( )
view trees. CLADOS (Nixon, 1992) was used to look at character same in both trees. Although the consensus tree (Fig. 7) is accepted distribution on the HENNIG86 trees. Two types of character transfor- as a working hypothesis of the phylogeny ofBrillantaisia, the results mations were plotted onto the trees using CLADOS: either as unique are discussed here with reference to the characters shown on the tree occurrences (synapomorphies) in black or white or convergence or in Figure 6. The results show that Brillantaisia and six species of within a the tribe parallelism (homoplasy) in grey. The values of tree length, consist- Hygrophila both belong monophyletic group, ency index (CI) and retention index (RI) are provided. All characters Hygrophileae, based on a suite of pollen characters [21(1); 22(0); are treated throughout as unweighted. 23(1); 24(0); 25(0); 26(0) &27(1)] and an entire to subentire leaf margin [2(0)] (this character reverses further up the tree [2( 1 )] in the B. B. owariensis and B. The Results species kirungae, vogeliana). outgroup to the Brillantaisia-Hygrophila clade is a group with non-bilabiate corollas the Eremomastax The analysis produced 122 equally parsimonious trees, 87 steps [15(0)] containing species speciosa, Mellera lobulata and solmsii. The long, CI=0.42 and RI=0.76.Tree 25/1 22 is presented in Figure 6 and Mimulopsis ((Brillantaisia- clade is the strict consensus of these trees is presented in Figure 7. Apart Hygrophila) (Eremomastax (Mellera-Mimulopsis))) characterized the of inflorescences from a few collapsed nodes, the consensus tree (Fig. 7) is very by presence paniculate [6(1)] and which are free or fused at the base similar to the tree given in Figure 6, and the major clades are the sepals only slightly [13(0)]. 80 K. SIDWELL
Table 2 Data matrix of 32 terminal units derived from receding polymorphic missing data scored for the 32 morphologi-
cal characters presented in Table 1 .
Taxon Character number (M 5-9 10-14 15-19 20-24 25-29 30/31
Duosperma crenatum (Lindau) P.G. Meyer REVISION OF BRILLANTAIS1A 81
4 Du.crenatum 1 10202831 IHHH} 1110 Dy.verticillaris 18 E.speciosa 1 15 1 HHh 20 | Me.lobulata 1011 I 1 19 1 1 M.solmsii
1 6 -HH6 13 //. auricula!n 1 o H. laevis
3 7 1118 H. linearis 1931 2 1121222324252627 rUH ~ 3 ff H.thwaitesii-a -0-f-HH-iHHHH011010001 HhHH8 27 I H.thwaitesii-b
H.didynama 1030 -0-1- 1 1 H.pilosa
12 0~i~~ B.pubescens 1
B.riparia-a 1718 23 -HH 1 1 1 B.riparia-d
5. riparia-b
B.riparia-c
1 4 1516 HI0221I B.oligantha-a
B.oligantha-b 12 -IH 10 B. stenopteris-a
B.stenopteris-b
B.grotanellii-a 6 1229 HHH 1 B.grotanellii-b 1 1 1 1028 B. madagascariensis 1 4 9 14 19 B.kirungae 1 1 1
I B.owariensis-a 12 -IH B.owariensis-b 10 10 B. vogeliana 1
B.lamium
31 B.debilis 3 7
1 4 i B.lancifolia-a -HH 3 1 Mh- B.lancifoliab K. SIDWELL 82
Du.crenatum
Dy.vertidllaris
E.spedosa
Me.lobulata
M.solmsil - H.auriculata
H.laevis
H.linearis
H.thwaitesii-a
H.thwaitesii-b
H.didynama
H.pilosa
B.oligantha-a
B.oligantha-b
B.stenopteris-a
B.stenopteris-b
B.pubescens
B.riparia-a
B.riparia-b
B.riparia-c
B.riparia-d
B.owariensis-a - B.o\variensis-b
B.kirungae
B.vogeliana
B.grotanellii-a
B.grotanellii-b
B. madagascariensis B.lamium
B.debilis - B.landfolia-a
B.landfoliab
Fig. 7 Strict consensus of 122 trees produced from analysis of the data matrix in Table 1 . REVISION OF BRILLANTAIS/A 83
TAXONOMIC ACCOUNT tudinal groove down centre of each valve, green with red apex when immature, brown to black when dry, glabrous or glandular and/or eglandular pubescent, elastically dehiscent. Seeds rounded to
Brillantaisia P. t. Beauv., Flore d'Oware 2: 67. 100, fig. 2 (1818); slightly kidney shaped, flattened, each held on a hardened, hook- in Burkill Fl trap. afr. 5: 37 (1899); Durand & Durand, Syll.fl. shaped retinacula, covered with adpressed trichomes which are congol: 416-418 (1909); Chevalier, Explor. hot. Afrique occ. hygroscopic and expand rapidly, producing mucous when wet. franc. 1: 493 (1920); Robyns, Fl. pare nat. Albert (1947); Heine DISTRIBUTION. Throughout tropical Africa with two species ex- in Fl. W. trap. Afr. 2nd ed., 2: 405 (1963); Heine in Fl. Gabon 13: tending to Madagascar. 28 (1966); Benoist, Fl. Madag.fam. 182: 28 (1967);Agnew, Upl. Kenya wild fls: 583 (1974); Troupin, Fl. pi. lign. Rwanda: 84 HABITAT. The distribution of Brillantaisia species is centred in the (1982); Champluvier in Fl. Rwanda 3: 444 (1985). Type species: Guineo-Congolian rain forests with species also occuring in montane Brillantaisia owariensis P. Beauv., Flore d'Oware 2: 68, t. 100: rain forests and woodland throughout Africa. Plants of Brillantaisia
fig. 2(1818). are often found in mesic sites or rooted in water and when in drier savannah type vegetation only occur in shady, wetter places. In West Belantheria Nees in DC., Prodr. 11: 96 (1847). Type species: Africa Brillantaisia is commonly found in disturbed areas (e.g. Belantheria belvisiana Nees in DC., Prodr. 11: 96 (1847). roadsides, plantations, farmland). Leucoraphis Nees in DC., Prodr. 11: 97 (1847). Type species: Leucoraphis vogeliana Nees in DC., Prodr. 11: 97 (1847). Brillantaisia is a genus of 12 species several of which are wide- Ruelliola Baillon in Bull. Mens. Soc. Linn. Paris 2: 852 (1890); spread and tend to be very variable morphologically, and is most Baillon, Hist. pi. 10: 427 (1891); Lindau in Nat. Pflanzenfam. diverse in the Guineo-congolian forests. Many species have previ- 4(3b): 307 (1895). Type species: Ruelliola grevei Baillon in Bull. ously been described in the genus based on local variation which has Mens. Soc. Linn. Paris 2: 852 (1890). been shown to be continuous by extensive study of specimens. The variation within each species is discussed in detail after each de- Erect to prostrate herbs to erect, sturdy shrubs. Stems square, usually scription. A full list of specimens studied is provided in Sidwell swollen at the nodes, glabrous to densely pubescent; cystoliths (1997) and a constantly updated list is available from the author. longitudinal, narrowing towards one end. Leaves opposite, decussate, petiolate, broadly ovate to ovate, occasionally elliptic or linear- Key to fertile specimens of Brillantaisia lanceolate; leaf base cordate or truncate to cuneate; apex acute or acuminate; entire to irregularly toothed; glabrous to pubes- margin 1 . Inflorescence a contracted thyrse with short peduncles, forming a spike; cent above and below; indumentum of multicellular, eglandular flowers/fruit in verticillasters 2 trichomes, evenly pilose to dense and villose above, mainly on midrib Inflorescence a lax thyrse with elongated, clearly visible peduncles and lateral veins below; cystoliths usually visible, with a primary 3 hand lens, above and below over entire leaf surface. Petiole usually 2. Bracts with and winged, lamina decurrent in top half of petiole, though occasionally glandular eglandular trichomes, linear-ovate; style often towards 1 1. B. grottanellii broadly winged to base; petiole wing margin entire, occasionally pubescent apex toothed towards a modified dichasium apex. Inflorescence variously Bracts lacking glandular trichomes, ovate to broadly ovate; style pubes- lateral with forming either a terminal thyrse and/or axillary thyrse cent only at base 12. B. madagascariensis vegetative growth occasionally continuing above flowering axes; 3. Bractlets obovate-rounded, shortly petiolate; corolla lips less than twice dichasial branching usually lax, becoming monochasial towards base the length of the corolla tube; sepals spathulate; flowers with stiff, of contracted with flowers inflorescence, occasionally forming spike unicellular trichomes on inner surface of lower lobe 4 appearing in verticillasters. Bracts on main inflorescence axis ovate bractlets linear; corolla more than twice the length of the corolla to broadly ovate, foliaceous, equal in size to the uppermost leaves at lips tube; sepals linear or spathulate; flowers lacking unicellular trichomes base of inflorescence, becoming smaller towards the apex of the on inner surface of lower lobe 5 inflorescence; bractlets on lateral inflorescence branches linear to obovate. Calyx with five equal or unequal, linear to spathulate sepals, 4. Flowers small, 12-25 mm long, inflorescence highly branched; leaves 1.25-1.8 times than wide 1. B. dorsal sepal longer and usually broader than the four lateral sepals; longer pubescens hirsute, with and eglandular trichomes. Corolla usually glandular Flowers large, 30-40 mm long; inflorescence branched 3-4 times; two- blue or violet, white; 2. lipped, purple, magenta, occasionally leaves 1 9-3.0 times longer than wide B. riparia corolla tube cylindrical, often paler than corolla lobes, with two entire or cre- com- 5. Leaves lanceolate to elliptic, occasionally ovate, barely brown-yellow markings in throat; upper lip hooded, laterally nate; inflorescence narrow terminal thyrse branching 1-2 times two-lobed at often on outer pressed, apex, glandular pubescent 9. B. lancifolia reflexed at surface; lower lip broad, ridged and bullate above, edges, Leaves ovate to ovate, entire or clearly toothed; inflorescence three-lobed at apex, occasionally with stiff trichomes on adaxial broadly lax, open, terminal or lateral thyrse branching 2-many times 6 surface, two membranous lateral pouches at base of lower lip form a tube. Androecium with two hinge with the apex of the corolla 6. Fruit glabrous or with a few erect trichomes at the apex; leaves entire or often 7 posterior stamens fertile; filaments white, flattened, sparsely subentire two anterior pubescent towards the base; anthers sagittate, dorsifixed; Fruit pubescent; leaves toothed, or if leaves entire to subentire, inflores- stamens reduced to slender staminodes, often with membranous cences slender, lateral, thyrse with apex usually reverting to vegetative anther at minute or absent. Gynoecium vestigial apex, occasionally growth 8 towards with slender, linear style, often sparsely pubescent base; fruit 22-30 x 2-5 lower 7. Leaves pubescent particularly towards petiole; mm, stigma a single flattened lobe at the end of the style, second, with 14-16 seeds per locule 8. B. lamium lobe reduced to minute tooth; ovary two-locular on annular disc, numerous. Fruit a fruit 18-22 x 1-2 with 18-28 seeds usually pubescent, placentation axial; ovules Leaves usually glabrous; mm, per locule...... 10. B. debilis linear, convex, two-valved capsule, pointed at apex with deep longi- 84 K. SIDWELL
8. Fruit with 8-10 very rarely to 16 seeds per locule; woody shrub or small Straggling annual, highly branched, aromatic herb, (0.3-)0.45-1 m often black 6. B. in tree, leaves drying kirungae tall, smelling minty or balsamic. Stems 2-5(-14) mm diameter, covered in short glandular trichomes and longer fine, white eglandular Fruit always with more than 1 2 seeds per locule; herb to 2 m tall, not trichomes. Leaves broadly ovate to ovate 50-1 10 x 85-160 mm, drying black 9 often caducous; leaf base cuneate to shortly attenuate, occasionally 9. Flowers 15-25 mm to 35 mm); fruit with 20-24 seeds long (rarely per truncate; apex acute; margin entire, occasionally slightly crenate; rather when often toothed at the of locule; leaves thin, papery dry, top lateral indumentum of silky, eglandular trichomes above and below; the petiole wing 7. B. vogeliana veins 7-9(-13) each side of midrib; petiole to 75 mm long, winged decurrent on in Vi-Vi. Flowers 30-45(-50) mm long; fruit with 1 2-20 seeds per locule; leaves towards lamina, wing narrow, petiole upper not thin and papery when dry, petiole wing usually entire 10 Inflorescence highly branched open thyrse with lateral inflores- cences to base of stem, branches slender, rachis indumentum visible trichomes 1 0. Lower corolla lip with clearly multicellular on upper irregularly glandular-pubescent with longer eglandular trichomes. surface 3. B. oligantha Bracts obovate to elliptic, to 35 x 45 mm, petiolate, irregular Lower corolla lip glabrous 1 1 glandular pubescent on both surfaces; bractlets obovate-rounded, to 5x8 mm, persistent. Sepals unequal, spathulate, larger upper sepal 1 1 . Leaf margin subentire or with small serrations; petiole wing tapering 12-14 x 0.5 mm, 1 .5-2 mm wide at broadened apex, smaller lateral gradually to base of petiole: lower corolla lip strongly reflexed, centre of 4. 8-10(-13) x < 0.5 mm, of slender eglandular lower lip bent up to 90 B. stenopteris sepals pubescence trichomes with shorter glandular trichomes, denser towards spathulate Leaf margin usually strongly toothed; petiole wing tapering gradually apex. Corolla purple to pale purple/pink occasionally with white to broad and toothed; lower corolla lip not not strongly reflexed upper lip, tube 5-10 mm long; upper lip 8-13(-15) mm long, .... 5. B. owariensis pubescent externally; lower lip 7-12(-15) mm; trichomes on lower
lip white, purple at apex; apical lobes 1 x 1 mm, central lobe smaller. Androecium with filaments 6-8 mm anthers 1-2.5 mm 1. H rill;i nt aisia pubescens T. Anderson ex Oliv. in Trans. Linn. long; long; staminodes minute < 0.5 mm or absent, Soc. London 29: 1 25 ( 1 875); Burkill in Fl. trop. afr. 5: 38 ( 1 899); long glabrous. Gynoecium with 8-9 mm 1 mm 2-3 mm Durand & Durand, Syll.fl. congol: 417 (1909); De Wild., Contr. style long; stigma long; ovary long, 10-15 x 2-2.5 mm with 8-16 seeds FL Katanga 1: 143 (1913); Benoist in Cat. pi. madag. 13(1939); glandular pubescent. Capsule brown when Vollesen in Opera Bot. 59: 79 (1980); Vollesen & Brummitt in per locule, yellow dried, sparsely glandular pubescent with trichomes towards the Kew Bull. 36: 571 (1981). Type: Tanzania, Khutu, Kirengwe, longer eglandular apex.
Grant s.n. (K!-holotype). DISTRIBUTION. East and Central Africa, Malawi, Zambia, Zimba- 8. in Vollesen & Brummitt Fig. Map (1981: 571). bwe, Mozambique and Tanzania, west to Zaire, Madagascar.
HABITAT. Sandy dry places, river beds, shade on forest floor or in Brillantaisia rutenbergiana Vatke in Abh. Naturwiss, Vereine, savannah woodlands under, for example, Adansonia digitata and Bremen 9: 131 (1885); Palacky, Cat. pi. madag. 3: 57 (1907). Faidherbia albida trees, or with Acacia, Albizia, Parkia, Lepis- Type: Madagascar, Andranovaka, Rutenberg s.n. (P-holotype). anthes and Terminalia (Abdallah & Vollesen 95/176 K). Locally Ruelliola grevei Baillon in Bull. Mens. Soc. Linn. Paris 2: 852 common; 0-1000 m. (1890); Baillon, Hist. pi. 10: 427 (1891); Lindau in Nat.
Pflanzenfam. 4(3b): 307 ( 1 895).Type: Madagascar, Greve 26 (P!- SELECTED COLLECTIONS National holotype; P!-isotype). DEMOCRATIC REPUPUBLIC OF THE CONGO: Upemba Park, 17 June 1948, 700 m, de Witte 03986 (BR, K). TANZANIA: Brillantaisia anomala Lindau in Pflanzenw. Ost-afr. C: 366 (1895); Kaswabilenga, Lindi, 1 1 September 1934, Schlieben 5323 (G, H, LISC, M, MO, PRE, S, US, Lindau in Bot. Jahrb. Syst. 24: 312-313 (1898); Lindau in Nat. Z); 2 km NW of Kingupira, 5 June 1975, Vollesen 2392 (C); 6 km along Pflanzenfam. 4(3b): 296 (1895). Type: Mozambique, Villa Kisiwani-Mnazi road, 800 m, 7 May 1995, Abdallah & Vollesen 95/176 (K). Gouveia, de Carvalho s.n. (COI-holotype). MOZAMBIQUE: Maringua, Dabi river, 25 June 1950, Chase 2547 (BM). B. pubescens var. rutenbergiana (Vatke) Benoist in Cat. pi. madag. MALAWI: Salima, Chipoka near Ilala Quarry, 25 May 1972, Salubeni 1803 13(1939). (MO, SRGH); Chimpakati village, Mlunguzi River, 27 June 1987, Usi & Anderson ex Benoist in Fl. Hygrophila pubescens (T. Oliv.) Madag. Kaunda 606 (M AL). ZAMBIA: Petuake district, 5 September 1 947, Greenway fam. 182 1: 36 (1967) non Nees (1847). & Brenan 8044 (PRE); NW of Mwern-na-ntipa, 7 August 1962, Tyres 339 (SRGH). ZIMBABWE: Binga district, September 1955, Davies 1426 (MO, Icones: in Trans. Linn. 29: 125 Anderson Soc. London pi. pro parte SRGH); Urungwe district, 20 September 1981, Pope 1987 (MO, SRGH).
(1875); Benoist in Fl. Madag. fam. 182. 1: fig. IV 14-18 (1967). MADAGASCAR: Perrier 94 1 6 (P); Madirobe, July 1 9 1 2, Kaudern s.n. (S).
Table 3 Four characters that differ between plants of Brillantaisia pubescens and B. riparia.
Character B. pubescens B. riparia
Inflorescence type very highly branched branched 3-4 times
Corolla length to 25 mm 30-40 mm long
Hinge at apex of corolla tube not well developed well developed
Leaf shape ovate elliptic REVISION OF BRILLANTA1S1A 85
8 Brillantaisia T. Oliv. A. Habit x B. leaf x C. Bract and bractlets x 1 Fig. pubescens Anderson ex 0.7; Lower 0.7; ; D. Gynoecium x 3; E. Corolla dissection showing androecium x 3; F. Capsule x 4. 86 K. SIDWELL
Brillantaisia pubescens is one of the more distinctive species in the 20-25 mm long, glabrous below, with straight erect stiff trichomes genus. The highly branched inflorescence, small flowers and rounded on centre of inner surface, trichomes white with a dark purple-black to blunt or notched persistent bracts are not found elsewhere \nBrillantaisia. Brillantaisia tip; apical lobes 2 x 2-3 mm, triangular, rounded anthers pubescens sensu lato was split into two subspecies by Vollesen & at apex. Androecium with filaments (10-)20-22 mm long; Brummitt (1981) on the basis of flower size. I consider the large 4-5 mm long; staminodes 3-4 mm long, pale, translucent, difficult flowered plants to be a different species (B. riparia) and treat B. to see with the naked eye, broadened slightly at apex sometimes to pubescens s.s. as including only those plants with smaller flowers. curved vestigial anther. Gynoecium with style to 30 mm long, stiff Two specimens of B. pubescens have slightly larger flowers than upward pointing trichomes towards base; stigma 2 mm long; ovary most (Tinley 2639, SRGH; Thera & Kaunda 313, PRE); however, 3 mm long glandular pubescent. Capsule 14-19x2 mm with 8-12 the leaf shape and venation are that of B. pubescens and this seeds per locule, light brown, covered with simple short trichomes variation does not blur the boundary between B. pubescens and B. and longer, glandular trichomes. riparia. The original illustration of the small flowered B. pubescens DISTRIBUTION. Mozambique and Malawi. by Fitch (Anderson 1875: pi. 125) almost certainly contributed to confusion over species delimitation as the type specimen was repre- HABITAT. Damp, shady places; 1000 m. sented as more, flowers than it has. It having many larger actually SELECTED COLLECTIONS appears that large flowered specimens of B. riparia have been MOZAMBIQUE: Vila Cabral, 8 September 1942, Mendonqa 680 (LISC); identified as B. pubescens by reference to the illustration. There are Nampula, 3 October 1942, Mendon^a 1219 (LISC); between Namapa and several characters which of the two taxa can be by plants clearly Chiure, 19August \948.Barbosa 1812 (LISC); Niassa, Vila Cabral, Meponda, distinguished (Table 3). 9 September 1958, Monteiro 46 (LISC). MALAWI: Kasungu, 27 August 1946, Brass 17442 (BM, MO, NY, SRGH, US); Machinga, 19 October 1979, Banda & Salubeni 1571 Liwawazi river, 3 2. Brillantaisia riparia (Vollesen & Brummitt) Sidwell, comb. (MO, SRGH); September 1986, Kaunda & Usi 429 (MAL); Liwonde, 9 1988, Banda & Kaunda nov. Type: Malawi, 13 miles N. of Kasungu, Dwangwa River, September 3452 (MO). Pawek 3908 (K!-holotype).
9. in Vollesen Brummitt 1 98 1 : 57 1 Fig. Map & ( ). Brillantaisia riparia was considered a variety of B. pubescens by Vollesen & Brummitt (1981). Due to the difference in flower size Brillantaisia pubescens var. riparia Vollesen & Brummitt in Kew and shape, leaf shape, leaf venation and inflorescence structure, I Bull 36: 571 (1981); Vollesen in Opera Bot. 59: 79 (1980). consider B. riparia a separate species from B. pubescens. The two Icon: Anderson in Trans. Linn. Soc. London 29: 125 pi. pro parte species are clearly very closely related yet are easily distinguished in (1875). the field. Brummitt (pers. comm.) noted that B. riparia should be compared with B. oligantha. Brillantaisia riparia and B. oligantha Annual viscid-pubescent, aromatic, rather woody herbs to 0.5-1 m are superficially similar, both having lateral inflorescences, trichomes tall. Stems erect or ascending; pilose with eglandular and glandular on the lower lip of the corolla and flowers of roughly equal size. trichomes. Leaves oblong, elliptic to ovate 20-25(-52) x 39-59(- However, there are several differences in gross morphology (Table 95) mm, often fallen below; upper most leaves of main axis oblong, 4) that clearly separate the two. 5-8 x 10-15 mm; base gradually tapering, cuneate; apex acute, rounded at tip; margin entire to slightly crenate; glabrous to pilose; 3. Brillantaisia oligantha Milne-Redhead in Mem. New York Bot. lateral veins 4-9(-l 1) each side of midrib; petiole 14-22 mm long, Card. 9: 20-21 (1954); Binns, Check List Herb. Fl. Malawi: 12 lamina decurrent on petiole forming very narrow wing sometimes to (1968). Type: Malawi, Nchisi Mountain, 30 January 1946, Brass base. Inflorescence leafy-bracteate terminal thyrse, with lower lat- 17042 (K!-holotype; BM!, MO!-isotypes). eral panicles, occasionally with vegetative growth continuing at Fig. 10. Map 1. apex; rachis covered in short simple trichomes with longer fine, non glandular and stalked glandular trichomes. Bracts as leaves; bractlets Viscid herb 0.3-1 .8 m tall. Stems 2-2.5 mm across, slightly winged elliptic-obovate, rounded at apex, margin entire, glandular trichomes on angles of stem, pubescent, with straggling eglandular and shorter and slender white trichomes above and below. Sepals unequal, larger multicellular glandular trichomes. Leaves ovate to broadly ovate, upper sepal linear-spathulate, 14-18 x 1 mm; smaller lateral sepals 36-65(-130) x 87-160(-210) mm; leaf base cuneate to attenuate; spathulate, 9-14 x 0.5-1 mm. Corolla purple to blue-purple; corolla apex acuminate; margin subentire to shallowly crenate or dentate; tube 13-15x2 mm; upper lip 20-25 mm long, glabrous; lower lip subglabrous to roughly pubescent; 7-10(-16) lateral veins each side
Table 4 Four characters that differ between plants of Brillantaisia riparia and B. oligantha.
Character B. riparia B. oligantha
Inflorescence branching stout, straight slender, curving
Corolla tube straight, long, narrow inflated, short, broad,
Lower corolla lip trichomes straight, single celled multicellular
Lower corolla lip shape gibbous, not strongly reflexed strongly reflexed REVISION OF BRILLANTA1SIA 87
Fig. 9 Brillantaisia riparia (Vollesen & Brummitt) Sidwell. A. Habit x 0.7; B. Corolla dissection showing androecium x 2; C. Capsule x 3.2; D.
Gynoecium x 2; E. Bract and bractlets x 1 . 88 K. SIDWELL
Fig. 10 Brillantaisia oligantha Milne-Redhead. A. Habit x 0.7; B. Corolla dissection showing androecium x 2.5; C. Capsule x 1 .8; D. Gynoecium and calyx x 2.5; E. Bract and bractlets x 3. REVISION OF BR1LLANTAISIA 89
Map 1 The distribution of Brillantaisia oligantha Milne-Redhead.
of midrib; petiole (15-)40-70(-120) mm long, lamina decurrent on eral axillary inflorescences only. The lower corolla lip is more petiole forming a slender wing almost to the base of the petiole. strongly reflexed in B. oligantha than in other species of the genus Inflorescence lateral few flowered thyrse to 45-75 mm long, branches and has multicellular trichomes on the adaxial surface. Previously very slender, branching 34 times, rachis pubescent. Bracts as thought to be restricted to the Nchisi mountains of Malawi, collec- leaves; bractlets narrowly obovate (0. 3-) 1.5 4 x (0.8-)2-9 mm, tions from Mughese in Malawi are clearly the same species. glandular pubescent above and below. Sepals unequal; linear to Brillantaisia oligantha can appear morphologically similar to B. spathulate, larger upper sepal 6-10 x 1 mm, smaller lateral sepals 5- riparia, both species having entire leaves and fine white silky 9 x 0.5 mm, pubescent. Corolla deep blue to purple, sometimes trichomes on vegetative parts, however, B. oligantlia has a much
1 a more tinged brown on the upper lip, corolla tube 3 x 7.5 mm; upper lip 3- shorter, broader corolla tube, a more hooded upper lip, and 17 mm long, sparsely pubescent outside; lower lip 14-18 mm long, slender, less branched inflorescence. This species is also closely strongly reflexed with lateral fold down centre and deep hinge at related to B. stenopteris Sidwell, sp. nov. from Tanzania. However, base, multicellular trichomes on inner surface; apical lobes to 2.5 x the new species has larger flowers, lacks any trichomes on the lower
3 mm. Androecium with filaments 10-15 mm long, pubescent at lip of the corolla and is found in a different habitat to B. oligantha. base; anthers 1-4 mm long; staminodes 1 mm long with small 4. Brillantaisia stenopteris Sidwell, sp. nov. Type: Tanzania, vestigial anther. Gynoecium with style 1-1.5 mm long, pubescent Morogoro region, Kombola, 8 July, 1933, Schlieben 4068 (LISC!- along the entire length; stigma 20-25 mm long; ovary 3 x 1 mm, holotype, MO!, PRE!-isotypes) covered in short glandular trichomes. Capsule 20-27 x 1 -2 mm with Fig. 1 1. Map 2. 10-12 seeds per locule, very sparse, short, glandular trichomes. Herba perennis usque ad 2 m alta. Folii lamina ovata vel latiovata DISTRIBUTION. Restricted to the Nchisi and Mughese mountain 25-144 mm longa, 40-220 mm lata, apice acuminata, basi cuneata, ranges in Malawi. margine obscure serrata, lamina 1 1-16 venatus, petiolo breviter semi- HABITAT. Shady places in wet forest, or in dry evergreen or decurrente; petiolo 15-90 mm longo. Inflorescentia ad 20 cm longa, evergreen forest, also along roadsides in these areas. Locally cymosa, terminalis, interdum vegitativus ad apice. Corolla 24-30 common; 1000- 1400m. mm longa, tubus circiter 8-9 mm longus; labium posticum 18-22 mm labium anticum 17-23 mm longum, glabrum. Fructus SELECTED COLLECTIONS longum; incognitus. Species nova aflUnisBrillantaisia oligantha Milne-Redh. MALAWI: Nchisi Mountain, 29 July 1946, Brass 17021 (NY); 30 July 1946, sed floribus majoribus et labium anticum sine pilis longis septatis Brass 17042 (BM, K, MO); 10 July 1960, Chapman 827 (BM); 1 September differt. 1970, Salubeni 1485 (SRGH); Salubeni 1487 (MAL. PRE, SRGH): Chitipa
1 2984 district, 1 2 1 977. 2836 (MO):ftw** (K, Mughese, September Phillips Herbs to 2 m tall. Stems erect, 2 mm in diameter, pubescent, LISC, MAL, MHU, MO, SRGH). trichomes white, eglandular with shorter glandular trichomes. Leaves with lat- ovate to ovate 25-140 x 40-220 mm: leaf base cuneate; Brillantaisia oligantha is an easily distinguished species, broadly apex 90 K. SIDWELL
1 8 1 acuminate; margin serrate, occasionally subentire; adaxial surface 5. Brillantaisia owariensis P. Beauv., Flore d'Oware 2: 68 ( 8); subglabrous to roughly pubescent, abaxial surface with fine non Bentham in Niger Fl.: 477 (1849); Hooker in Bot. Mag.: 79: t. glandular trichomes; 11-16 lateral veins on each side of the midrib; 4717 (1853); Durand & Schinz, Etudes fl. Congo 1:217 (1896); petiole 1 5-90 mm long, lamina decurrent on the petiole forming a Burkill in Fl. trop. afr. 5: 40 (1899); Durand & Durand, Syll.fl. narrow wing, often to the base of the petiole. Inflorescence open congoi: 417 (1909); Benoist in Bull. Soc. Bot. France 60: 335 panicle, sometimes becoming vegetative at the apex, to (4-)7-20 cm (1913); Chevalier, Explor. hot. Afrique occ. franc . 1: 493 (1920); long; lateral branches slender, few flowered. Bracts as leaves or Hutchinson & Dalziel in FL W. trop. Afr. 2: 254 (1931); Heine in ovate, sessile; bractlets ovate-linear 7-12 x 1-3 mm. Calyx unequal, Fl. W. trop. Afr. 2nd ed., 2: 406 (1963). Type: Nigeria, Benin, sepals slightly spathulate, larger upper sepal 10-12 x 1-2 mm, Agathon, Palisot de Beauvois s.n. (Gl-lectotype; G!- smaller lateral sepals 8-10 x 1 mm; covered in long non glandular isolectotypes). and short glandular trichomes. Corolla purple, corolla tube 8-10x3 Belantheria belvisiana Nees in DC. Prodr. 11: 96 (1847). Type: as mm, to 18-22 mm outside; upper lip long, glandular pubescent for B. owariensis. lower lip 17-23 mm long, glabrous on inner surface. Androecium Brillantaisia patula T. Anderson in J. Proc. Linn. Soc. Bot. 7: 21 with filaments 25 mm long; anthers 4 mm long; staminodes 1 mm (1864); Burkill in Fl. trop. afr. 5: 41 (1899); Hiern in Cat. afr. pi. long, minute, lacking vestigial anther. Gynoecium with style 30 mm 1: 807 (1900); De Wild., Etudes fl. Bas- Moyen- Congo 1: 314 long; stigma 4 mm long; ovary 6 mm long, densely pubescent. (1903-1906); Durand & Durand, Syll.fl. congoi.: 417 (1909); not known. Capsule Benoist in Bull. Soc. Bot. France 60: 335 (1913); DeWild., Contr. Fl. Katanga 1: 143 & 2: 144 (1913); Mildbraed, Wiss. Erg. zweit. DISTRIBUTION. Restricted to the Morogoro region of Tanzania. deut. Zentr.-Afr. Exped., Bot. (1922); Hutchinson & Dalziel in FL
HABITAT. Rain forest; 1000m. W. trop. Afr. 2: 254 (1931); Exell, Cat. pi. S. Tome: 260 (1944); Robyns, Fl. pare. nat. Albert. 2: 269-270 (1947); Heine in Fl. W. COLLECTIONS EXAMINED trop. Afr. 2nd ed., 2: 406 ( 1 963); Heine in FL Gabon 13: 94, pi. 1 9, TANZANIA. Morogoro region, Kombola, 8 July 1933 Schlieben 4068 figs 1-6 (1966); Champluvier in Fl. Rwanda 3: 444-448 (1985). (LISC. MO, PRE); Kilosa district, 13 November 1987, Po$s et al. 8722 Type: Congo, Smith s.n. (K!-holotype; BM!, P!-isotypes). (UPS). B. alata T. Anderson ex Oliv. in Trans. Linn. Soc. London 29: 125
Brillantaisia stenopteris is very closely related to B. oligantha from (1875); S. Moore in J. Bot. 18: 197 (1880); Durand & Schinz, the Nchisi and Mughese mountains in Malawi. The two species have Etudes fl. Congo 1: 216-217 (1896); Durand & DeWild., in Bull. similar ovate, cuneate leaves with a long slender petiole wing. Soc. Roy. Bot. Belgique 36: 83 (1897); De Wild., Miss. Em.
However, they can be easily distinguished due to the larger flowers Laurent 1: 1 82 (1905); DeWild., Etudesfl. Bas- Moyen- Congo 1: and absence of trichomes on the inner surface of the lower corolla lip 314 (1903-1906); De Wild., Contr. FL Katanga 1: 143 (1913). of B. stenopteris. The corolla tube of B. stenopteris is longer and Type: Uganda, Unyoro, Speke & Grant 583 (K!-holotype). thinner than that of B. oligantha, resembling the corolla tube of B. B. salviiflora Lindau in Bot. Jahrb. Syst. 17: 101 (1893); Burkill in riparia, which is also closely related to B. stenopteris. FL trop. afr. 5: 41 (1899); Benoist in Bull. Soc. Bot. France 60:
Map 2 The distribution of Brillantaisia stenopteris Sidwell. REVISION OF BRILLANTAISIA 91
Fig. 11 Brillantaisia stenopteris Sidwell. Habit x 0.7; B. Corolla dissection showing androecium x 2.5; C. Gynoecium and calyx x 2.5; D. Bract and bractlets x 3. 92 K. SI DWELL
with in 336 (1913). Type: Togo, Bismarksburg, Buttner 341 (B|- pale purple to deep blue-purple often yellow markings x 2-6 18- holotype). throat; tube (5-)7-10(-14) mm long; upper lip (13-) B. nitens Lindau in Bot. Jahrb. Syst. 17: 102 (1893); Burkill in Fl. 30(-60) mm long, outer surface of upper lip glandular pubescent; lobes trap. afr. 5: 41 (1899); Lindau in Wiss Erg. deut. Zentr.-Afr. lower lip (14-) 18-30(55) mm long, glabrous; apical (l-)2- with filaments Exped., Bot. 2: 292 (191 1); Benoist in Bull. Soc. Bot. Fr. 60: 336 5(-10) mm long. Androecium (20-)25-35(-40) staminodes 15- (1913); Mildbraed, Wiss. Erg. zweit. deut. Zentr.-Afr. Exped., Bot. mm long; anthers (2-)5-7(-10) mm long; (10-) anther at the (1922); Hutchinson & Dalziel in Fl. W. trap. Afr. 2: 254 (1931); 25(-30) mm long with small vestigial apex. Robyns, Fl. pare. nat. Albert. 2: 270-272 (1947); Heine in Fl. W. Gynoecium with style (15-)20-35(-45) mm long; stigma 2-4(-6) 3-7 trap. Afr. 2nd ed., 2: 406 (1963); Blundell, Wildfl. Kenya: 104 mm long; ovary mm long, pubescent. Capsule (15-)18-26(-
in Fl. 3: 448 1 Wild x 2-3 with seeds locule; covered ( 1 982); Champluvier Rwanda ( 985); Blundell, 33) mm (10-)12-18(-24) per of Preuss 847 with and trichomes. fl. E. Afr.: 389 (1987). Type: Cameroon, W. Buea, eglandular glandular (Bt-holotype; K!-isotype). DISTRIBUTION. tropical Africa. B. dewevrei De Wild. & Th. Dur. in Durand & De Wild., in Bull Soc. Throughout
Bot. 38: 44-45 ( 1 899); Burkill in Fl. 5: 5 1 Roy. Belgique trop. afr. HABITAT. Moist montane forest particularly in open shady places 1 De Wild. & Durand, Ann. Mus. Ser. 3 Bot. 1 : 1 74- ( 899); Congo where competition is reduced in recently disturbed areas; 600-1600 175 (1901); De Wild. & Durand in Bull. Herb. Boissier 1: 833
(1901); Durand & Durand, Syll. fl. congol.: 416-417 (1909). SELECTED COLLECTIONS Type: Congo, between Lukolela an Gombi, Dewevre s.n. (BR!- SIERRA LEONE: 18 November 1965, Adam 22109 (MO, P); 2 holotype). January 1966, Adam 22916 P); 27 November 1965, Adam 27084 (MO); Mt. 1 Fl. (MO, B. nyanzarum Burkill in Fl. trop. afr. 5: 39 ( 899); Robyns, pare. Nimba, 14 December 1 966, Bos 2397 (WAG); 825'W732'N, 18 December nat. Albert. 2: 272 (1947); Andrews, Fl. pi. Sudan 3: 171 (1956); 1967, Geerling & Bockdam 1859 (C, MO, WAG); LIBERIA: 10 January 1 in Agnew, Upl. Kenya wildfls: 583 & 584 ( 974); Synnott Comm. 1965, Adam 20549 (BR, K, MO, P, UPS); Grand Cape Mount Co., 21 Inst. Occ. Papers 27: 68 ( 1985). Type: Kenya, Kavirondo, Forestry December 1947, Baldwin 10778 (K, MO); 6 November 1947, Baldwin 101 82 Scott-Elliot 6999 (K!-holotype). (K, MO); Loffa country, 21 December 1966, Bos 2552 (K, WAG). IVORY Burkill in Fl. 5: 41 Benoist in Bull. B. leonensis trop. afr. (1899); COAST: 24 November 1909, Chevalier 22421 (P); 20 km de Man sur route Soc. Bot. France 60: 335 (1913); Hutchinson & Dalziel in Fl. W. de Danane, 23 November 1956, de Wilde 859 (WAG); MontTonkoui, pres de Trop. Afr. 2: 254 (1931). Syntypes: Sierra Leone, Scott-Elliot Man, 3 December 1985, Ake Assi 17170 (G). GHANA: Bauchi Plateau, 17 148 3990a (K!), Windwood Reade s.n. (K!), Don s.n. (BM!). April 1955, Morton K416 (K, WAG); Kofordua, Dalziel (C, E, K, M, MO, PRE, WAG). NIGERIA: Mandaga, Nambila Plateau, 4 January 1955, B. patula van welwitschii Burkill in Fl. trop. afr. 5: 41 (1899); Hiern Latilo & Daramola FHI 28992 (FHO); Shasha Forest Reserve, Richards in Cat. afr. pi 1: 807 (1900). Syntypes: Angola, near Sange, 3108 (BM, MO, NY); 1 1 December 1976, Ariwaodo 33. CAMEROON: 13 Quiapoza and Cuango Rivers, Welwitsch 5149 (BM!), Quibolo, February 1927, Dalziel 8232 (US); Buea, Preuss 1029 (BM, M); Dschang, Welwitsch 5182 (BM!, K!, P!). December 1938, Jacques-Felix 2600 (P); 31 km on Ebolowa-Ambam Road, B. mahonii C.B. Clarke in Bull. Misc. Inform. 1906: 251 (1906); S. 1 September 1974, de Wilde 7544 (WAG); Lake Oku, 613'N 1028'E, Bot. 45: 89 Moore in/ (1907); Lind, Comm.fl. pi Uganda: 140, Thomas 4381 (MO, YA); GABON: Woleu-Ntem Province, 14 November s.n. fig. 82 ( 1962).Type: Uganda, Entebbe, Mahon (K!-holotype). 1933, Le Testu 9395 (BR); CENTRAL AFRICAN REPUBLIC: 10 Decem- B. bauchiensis Hutchinson & Dalziel in Fl. W. trop. Afr. 2: 253 ber 1936, Eckendorfll (P); CONGO: Sangha, 2N 1355'E, 27 November
(1931); Hutchinson & Dalziel Fl. W. trop. Afr. 2nd ed., 2: 407. 1991, Thomas 9005 (MO); DEMOCRATIC REPUPUBLIC OF THE CONGO: 1905 Kivu, Lac Mokoto, 27 Syntypes: Nigeria, Bauchi Plateau, Lely PI 29 (K!), Dent Young 12January 1914, Bequaert (BR); July 1953, van cler Ben 648 (BR); Yabahondo October 195 (K!). Yangambi region, village, 1952, Germain 8141 (BR, W, Z). RWANDA: Rwaza, SE de Ruhengeri, 23 Icones: P. Beauv., Flored'Oware 2: t. 100 fig. 2 (1818); Anderson in February 1972, Auquier 2684 (BR). BURUNDI: 2 June 1969, Lewalle 3795 3 1131 Trans. Linn. Soc. London29: pi. 124 ( 1 875); Hooker inBot. Mag.: (BR). UGANDA: Bigera River, August 1906, Bagshawe (BM). KENYA: December Dale 3208 1932, Brodhurst- 79: t. 4717 (1853) excluding glabrous fruit; Heine in Fl Gabon, 1933, (BR, FHO); January
HU1693 (Z).TANZANIA: 26January 1 99 1 , Ka \ombo 1072(MO).ANGOLA: Acanthacees: 13: pi. 19 figs 1-6 (1966); Agnew, Upl. Kenya wild 24 June 1958, Montiem, Santos & Murta 205 (PRE). fls: 584 (1974); Blundell, Wildfl. Kenya: pi. 264 (1982);
Champluvier inf/. Rwanda 3: fig. 1 39: 2, 3 A-3D ( 1 985); Blundell, Brillantaisia owariensis is the most widespread and variable of all Wild./?. E. Afr.: pi. 849 (1987). species in the genus and has caused many problems of species delimitation for botanists over the After extensive herbarium Herbs or woody herbs to l-3(-4) m tall, viscid, often aromatic. years. Stems l-8(-15) mm in diameter, glabrous or pubescent towards studies, supplemented by field work in Cameroon and Tanzania, I consider it to this into discrete taxa and apex. Leaves ovate to broadly ovate (35-)50-140(-210) x (25- impossible split group )40-120(-185) mm; leaf base cordate to cuneate; apex acuminate recognize it as a very diverse 'oc/z/o-species' (sensu White, 1962: this is the viable sometimes markedly so; margin toothed, usually irregularly ser- 79) and accept that, although unsatisfactory, only rate, crenate towards apex with or without secondary toothing; option without further extensive studies in the field. Brillantaisia eglandular pubescent above and below, sometimes densely pubes- owariensis sensu meo encompasses all fairly robust, woody cent; lateral veins 6-19 each side of the midrib; petiole Brillantaisia plants with a paniculate inflorescence, toothed leaves fruit. variation (10-)30-90(-130) mm long, lamina decurrent in upper !/2-% with a winged petiole and pubescent Morphological within the in certain ofAfrica. length of petiole, occasionally entire length of petiole. Inflores- group appears more pronounced parts In East for clear differences can be cence a many flowered terminal panicle (9-) 1 5-40(-60) cm long Africa, example, morphological with 4-18 nodes; rachis glandular pubescent. Bracts ovate, (3- observed between different isolated groups and two representative like of )20-50(-75) x (5-)15-25(-40) mm, usually falling early; bractlets specimens may look nothing one another when the extremes linear oblong, to 20 x 5 mm, eglandular pubescent. Sepals morphological variation are observed. However, when morphologi- cal variation is studied the character subequal to unequal, linear, (5-)9-15(-22) x 1 mm long, rounded accross whole of Africa, to slightly spathulate at the apex, glandular pubescent. Corolla differences break down and distinct groups cannot be distinguished. REVISION OF BRILLANTAISIA 93
Variable characters include leaf shape, leaf base shape, leaf margin, truncate to attenuate or cuneate; apex acute to acuminate; margin number of lateral veins, calyx shape, inflorescence density, flower irregularly toothed, teeth single or double, smaller becoming crenate size, fruit length/width and fruit pubescence. The morphological towards the apex, rarely entire; puberulous; lateral veins 7-13 each variation within Brillantaisia owariensis is informally described side of the margin, lower laterals closer together; petiole (0-)10-140 below in terms of two main groups: the nitens group and the patula mm long, lamina gradually decurrent on top '/2-% of petiole, upper group. pair of leaves at base of inflorescence sessile. Inflorescence a Plants which tend to differ from typical B. owariensis in possess- terminal thyrse 15-25(^0) cm long, lateral branches erect, at 30-50 ing rather finely toothed leaves, a somewhat narrow petiole wing, degrees to the main rachis, flowers crowded at end of lateral larger number of lateral nerves and a relatively dense inflorescence branches; rachis shortly pubescent; trichomes, red-brown, non glan- could be assigned to the nitens part of Brillantaisia owariensis. dular, sometimes straggly purple trichomes. Bracts broadly ovate to
Brillantaisia nitens was described by Lindau (1893) as being dis- ovate, occasionally elliptic, base slightly amplexicaul, apex rounded tinct from all other Brillantasias due to the leaf pubescence, which is to acute; bractlets ovate(-linear), falling late or remaining in fruit. comparatively dense for the genus. Plants with characters of the Sepals unequal, larger upper sepal linear, rounded at tip or slightly nitens group tend to occur in the forests of Cameroon and further spathulate, (8-)10-14(-19) x l-3(-5) mm, often twice as wide as smaller west in tropical Africa. Plants that are slightly more robust than lateral sepals (clearly seen in bud), occasionally very large; typical Brillantaisia owariensis, with velvety-tomentose leaves, lateral sepals linear 8-15 x 1 mm; densely pubescent dorsally. have previously been distinguished as Brillantaisia bauchiensis in Corolla pale blue to bright blue-purple or pink-purple, with darker x Nigeria and B. leonensis in Sierra Leone. Specimens with longer markings on lower lip; corolla tube 6-10 3-6 mm, occasionally covered in calyx lobes and sub-persistent bracts were previously assigned tofi. inflated; upper lip 19-35 x 6-10 mm, evenly dense, x mahonii. Specimens from East Africa that generally have the same eglandular and glandular trichomes; lower lip 19^40 10-18(-25) x morphological characters as those of the nitens group, but differ in mm, ridged on upper surface; apical lobes triangular, 3-10 2-8 anthers the presence of more persistent bracts, have often been identified as mm. Androecium with filaments 20-25(-30) mm long; (3- B. nyanzarum. )5-8 mm long; staminodes 8-12 mm, very slender often with well The patula group: The most distinct node of variation within developed vestigial anther. Gynoecium with style 25^0 mm long; Brillantaisia owariensis is seen on specimens that can be informally stigma 2 mm long; ovary 5-10 mm, covered in tangled glandular trichomes. described as the patula group. These plants have a very open, lax, trichomes, sometimes also with non glandular Capsule seeds 'zigzag' inflorescence architecture and often have large flowers and 30-40 x 2-5 mm with 8-10(-16) per locule; black, glandular often black. a linear calyx. Specimens with these characteristics do not seem to puberulous. Specimens drying be restricted to one area of Africa and have been collected from DISTRIBUTION. East and central Africa, from Kenya south to Zim- Guinea to Zaire and on Sao Tome. Mostpatula specimens have been babwe and Mozambique, west to Uganda and Eastern Zaire. collected from the Congo basin. Leaves of the patula 'node' often, in altitude and montane but not always, have a distinct wing to the base of the petiole HABITAT. In gaps mid forest, forming 1000-3500 m. (Mocquery's 115, Z; Richards 3018, BM) which is occasionally thickets, often along streams. Very local; amplexicaul. SELECTED COLLECTIONS DEMOCRATIC REPUPUBLIC OF THE CONGO: Kivu, 27 July 1959, 6. Brillantaisia kirungae Lindau in Gotzen, Durch Afrika von Ost Cambridge Congo Expedition 146 (BM, LISC, US). RWANDA: Pare des der Veken 10350 BURUNDI: 29 nach West, Sonderabdr.: 9 (1896); Burkill in Fl. trap. afr. 5: 42 Volcans, 27 July 1974, Van (BR). Rwegura, Lewalle 3640 22 June 1980, Reekmans 9408 : 1969, (BR, G); Bubanza, (1899); Durand & Durand, Syll. fl congol. 417 (1909); Robyns, May (K, MO, UPS). SUDAN: Imatong Mts, Gilo Village, 8 November 1980, Friis Fl. pare nat. Albert 2: 269 (1947). Type: Zaire, Mount Kirunga, & Vollesen 58 (C). UGANDA: Ruwenzori Mts, Namwamba Valley, 17 von Gotzen 48 (Bf-holotype). January 1935, Taylor 3156 (BM, MO, NY, S). KENYA: 18 March 1977, 12. Fig. Faden & Faden 77/919 (BR, US, WAG). TANZANIA: Rungwe, Kiwara
Burkill in River, 9 1 949, Greenway & Eggeling 8394 (PRE); Morogoro, Bunduki, B. ulugurica Lindau in Bot. Jahrb. Syst. 22: 112 (1897); August 1 1 95 1 8594 River, 1 Checklist brit. 7 August , Greenway & Eggeling (FHO); Nugwi 9August Fl. trop. afr. 5: 43 (1899); Brenan & Greenway, 1952, Carmichael 102 (FHO); Mbisi Mts, 17 June 1960, Leach & Brunton 5 terr. 2: 6 Tanzania, Uluguru empire Tangan. (1949). Syntypes: Kilosa 10069 (SRGH); Ufipa, 6 August 1960, Richards 12983 (SRGH); Mts, Stuhlmann 8850 (B|) & Stuhlmann 9224 (B|) District, Ukaguru Mts, 7 August 1972, Mabberley 1403 (K), Mufindi Tea Check- B. Burkill in Fl. trop. afr. 5: 42 (1899); Binns, subulugurica Estate, 17 August 1984, Bridson & Lovett 544 (MO). MOZAMBIQUE: list herb. Malawi: 12 Makua, fl. (1968). Type: Mozambique, Namuli Peaks, 26 July 1962, Leach & Schelpe 11470 (LISC, SRGH); Namuli Hills, Last s.n. (K!-holotype). MALAWI: Nchisi Mts, 4 September 1929, Bunt Davy 1228 (FHO); Kafwimba ZIMBABWE: B. grandidentata S. Moore in/ Bot. 45: 331 (1907). Type: Uganda, Forest, 4 July 1973, Pawek 6978 (C, PRE, SRGH, UPS). Plowes 2261 Vumba, 20 Toro, Fort Portal, Bagshawe 1270 (BM!-holotype). Ngoruma reserve, 14 August 1962, (LISC, SRGH); Mutter 802 B. cicatricosa var. kivuensis Mildbr. in Bull. Jard. Bot. Etat 17: 86 September 1968, (SRGH). Albert 2: 270 Zaire, (1943); Robyns, Fl. pare nat. (1947). Type: Brillantaisia kirungae is the largest and most woody of the species Lake Magera, de Witte 1434 (BR!-holotype; BR!-isotype). of Brillantaisia and is easily recognized by the woody or shrubby 583 non Lindau in a B. nitens sensu Agnew, Upl. Kenya wildfls: (1974), habit; highly irregularly toothed leaves; large flowers crowded (1893). erect inflorescence; broad upper calyx lobe and large, few-seeded is variable in leaf and 1A-1D fruit. This species rather shape toothing (most Icones: Champluvier in Fl. Rwanda 3: fig. 139, (1985); collections lack the larger lower leaves, so measurements given Fischer & Hinkel, Nat. env. Rwanda: fig. 68 (1992). above need refining after more field studies); the degree of contrac- the trees m tall tion of the inflorescence; density of the indumentum on both Stout shrubby herbs 1-2 m tall to shrubs or small 2-5(-7) sometimes inflorescence and fruit; and flower size, particularly the size of the with bole 1.4 m high and 12 cm diameter, viscid and lobes at the of the lower Previous treatments ofBrillantaisia aromatic. Stems erect, 5-20(-120) mm across, shortly puberulent. apex lip. base cordate or have all divided it into more than one species. For example, Leaves, broadly ovate 60-320 x 50-240 mm; leaf kirungae 94 K. SIDWELL
Fig. 12 Brillantaisia kirungae Lindau. A. Habit x 0.7; B. Lower leaf x 0.7. C. Corolla dissection showing androecium x 1 .4; D. Gynoecium x 1 .4.
E. Capsule x 1 .4; F. Bract and bractlets x 1 . REVISION OF BRILLANTAISIA 95
Burkill (1899) recognized four species: Brillantaisia cicatricosa checked. It may well be the case that Lindau based his observation with subglabrous sepals and numerous seeds; B. kirungae with on the illustration of Fitch in Anderson (1875) which has caused lanceolate bracts; B. ulugurica with large flowers and large lobes at considerable confusion in Brillantaisia (see p. 86). Brillantaisia the apex of the lower corolla lip and a new species, B. subulugurica cicatricosa sensu Burkill ( 1 899) adds to the ambiguity of this name which was considered distinct from B. ulugurica due to the shorter as he describes the species as having entire, glabrous leaves, a loose corolla tube. More recently, Champluvier (1985) recognized two inflorescence, nearly glabrous sepals and 1 6-20 seeds, whereas the species, B. kirungae and B. cicatricosa, separated on the size of the species recognized here has irregularly toothed pubescent leaves and upper sepal. All of the characters used by previous authors to divide a crowded inflorescence. A variety of B. cicatricosa Lindau was this species are continuously variable when the entire geographical formally recognized by Mildbraed ( 1 943) on the presence of glandu- variation of the group is accounted for. In his treatment of the genus lar trichomes on the calyx. This taxon is illustrated in Flore du for the Flora ofTropical EastAfrica and Flora Zambesiaca, Vollesen Rwanda (Champluvier, 1985) and clearly falls within B. kirungae as (in prep a & b) recognizes two species within this group, B. delimited here. However, I am not certain that Mildbraed saw the cicatricosa and B. ulugurica. He considers plants of B. cicatricosa type specimens of B. cicatricosa or that he understood Lindau's to have irregularly large-toothed leaves; smaller lower corolla lip concept of that species. Therefore, as the protologue offl. cicatricosa lobes; smaller seeds, to lack glandular trichomes on the fruit and to Lindau is ambiguous and the description of B. cicatricosa by Burkill have a more northerly distribution from Kenya, Uganda, Burundi, does not fit this species, I treat this name as dubious, and use the Rwanda and Zaire. Brillantaisia ulugurica sensu Vollesen is recog- second earliest name, B. kirungae, for this species. nized as distinct on the basis of smaller irregular teeth on the leaves; leaf margin becoming entire towards the apex; larger lobes on the 7. Brillantaisia vogeliana (Nees) Benth. in Hooker, Niger FL: 477 lower corolla lip; larger seeds; both glandular and non glandular (1849); Anderson inJourn. Linn. Soc. Bot. 7: 21 (1864); Burkill trichomes on the and a more in fruit; widespread geographical range in Fl. trap. afr. 5: 40 ( 1 899); Benoist in Bull. Soc. Bot. France 60: Zimba- Uganda, Burundi, Zaire, Tanzania, Malawi, Mozambique, 335 (1913); Hutchinson & Dalziel in Fl. W. trop. Afr. 2: 254 B. bwe and Zambia. Two characters used by Vollesen to distinguish (1931); Exell, Cat.pl. S. Tome: 260(1944); Mildbraed, Wiss. Erg. in cicatricosa from B. ulugurica are not overlapping or continuous zweit. deut. Zentr.-Afr. Exped., Bot.: 192 (1922); Robyns, Fl. size fruit hair I have studied his species descriptions: seed and type. pare. nat. Albert 2: 212 (1947); Heine in Fl. W. trop. Afr. 2nd ed., the two species across their entire range which includes specimens 2: 406 (1963); Heine inFl. Gabon 13: 92 (1966). Type: Fernando from Sudan and Angola. It appears that although some 'cicatricosa- Po, Vogel 179 (K!-holotype; K!-isotype). smaller toothed leaves with like' specimens do have flowers, deeply Fig. 13. a more cordate base, smaller seeds and no glandular tricomes on the Leucoraphis vogeliana Nees, in DC., Prodr. 11: 97 (1847). capsule, these characters are continuously variable and there is no Brillantaisia molleri Lindau \nBot. Jahrb. Syst. 17: 99 ( 1 893). Type: morphological data to suppport separatation of the two species. Sao Tome, Moller 33 (87) (Bt-holotype; COI!, K!-isotypes). A new species from Tanzania was mentioned by Vollesen (in prep. B. preussii Lindau in Bot. Jahrb. Syst. 17: 100 (1893). Syntypes: a) as closely related to Brillantaisia ulugurica. Plants belonging to Cameroon, Barombi, Preuss 320 (K!-isosyntype) Buea, Preuss the new putative new species were recognized primarily by the 998 (BM!, K!-isosyntypes) presence of long, rather synthetic looking, purple, eglandular B. soyauxii Lindau in Bot. Jahrb. Syst. 17: 101 (1893); Burkill in Fl. trichomes on the panicle and calyx, and by the larger capsule (no 60: 335 trop. Afr. 5: 39 (1899); Benoist in Bull. Soc. Bot. France measurements given). Several intermediates between B. ulugurica (1913); non Heine in Fl. Gabon 13: 84 (1966). Type: Gabon, sensu Vollesen, with short puberulent trichomes on the panicle and Munda, Sibange Farm, Soyaux 454 (Bt-holotype; E!, H!, short glandular and eglandular trichomes on the calyx, and the studied US!,WAG!-isotypes). purple haired sp. nov. cf. B. ulugurica have been (e.g. B. schumanniana Lindau in Bot. Jahrb. Syst. 17: 102-103 (1893). MOZAMBIQUE: Leach & Schelpe 1 1470; MALAWI: Chapman & Cameroon, Braun 47 (Bf-holotype). and I consider the Type: Chapman 921 \ Mt Mulanje; Burn Davy 1228) variation to one Within the species recog- in Fl. represent species. single Icones: Heine in Fl. Gabon 13: pi. 18, figs 1-6 (1966); Heine, nized in this treatment, with a more lax inflorescence, more plants W. trop. Afr. 2nd ed., 2: 406, fig. 300 (1963). slender sepals and smaller flowers tend to occur further north-north Pare Erect herbs to 0.2-1 m tall. Stems erect, 2-4 mm in diameter, west (SUDAN, Gilo Village, Friis & Vollesen 58 C; RWANDA, near leaf fused across each node, des Volcans, Van der Veken 10350, BR; UGANDA, Kanaba Pass, branching base, petioles gla- brous or with a of trichomes at the nodes. Makerere College 29) whereas larger more robust plants are more shortly pubescent fringe ovate to ovate x mm, commonly from Malawi and Mozambique. Leaves broadly 35-100(-l 10) 45-120(-150) as men- below, evenly pilose; leaf base truncate Typification of this species is somewhat problematic papery, bright green, paler acumi- I with her decision to retain or cordate, occasionally shortly attenuate; apex slightly tioned by Champluvier ( 1 985). disagree sensu meo have nate, rounded; irregularly serrated, dentate at base of the epithet cicatricosa. Specimens of B. kirungae margin the orfl. lamina around the widest part of the leaf, crenate towards apex historically been named asfi. cicatricosa, B. kirungae, ulugurica, were of the leaf; lateral veins 7-13 each side of midrib, prominent; however, the type specimens of all three of these names B. cicatricosa (30-)40-120 mm, lamina decurrent on upper third of peti- destroyed in Berlin. The protologue of the earliest name, petiole ole, often toothed at Inflorescence a terminal lax thyrse Lindau, is somewhat ambiguous and does not fit the species descrip- wing top. in 1 cm with smaller lateral inflorescences tion above. Although the Latin diagnosis of B. cicatricosa mentions (8-) 2-20(-30) long, with characters do not fit the axils of lower leaves; rachis shortly glandular pubescent the large upper sepal of this species, several often seeds in the slender non trichomes. Bracts to 7 x 15 mm, group well, notably subglabrous sepals and 16-20 glandular with Lindau men- early; bractlets linear elliptic to 2 mm long, glabrous capsule. The original description of B. cicatricosa falling however of irregular glandular trichomes. Sepals unequal, slightly tioned that the species was close to 5. pubescens T. Anders., fringe observed larger upper sepal 6-15 mm long, smaller lateral sepals without a type specimen of B. cicatricosa the variation by spathulate, be mm viscid glandular pubescent. Corolla clear Lindau when he made this seemingly misplaced statement cannot 6-10(-12) long, 96 K. SIDWELL
Fig. 13 Brillantaisia vogeliana (Nees) Benth. A. Habit x 0.7; B. Corolla dissection showing androecium x 3; C. Capsule x 2; D. Gynoecium x 3; E. Bract
and bractlets x 1 . REVISION OF BRILLANTAISIA 97 mauve-violet, white in throat with lateral yellow spots; corolla tube forms of this species are fairly common. Exell (1944: 261) noted paler, tube 2^4 x 5-10 mm, slightly shorter than the calyx lobes; the two colour forms and stated They seem to be identical except upper lip 1 l-15(-25) mm covered with glandular and non glandular for the flower-colour and they always grow together, the purple- trichomes on outer surface, more so towards the apex; lower lip 10- flowered form being rather more common'. 15(-25) mm long; apical lobes to 1 x 1 mm, central tooth broader than the laterals, glabrous. Androecium with filaments 12-17(-24) 8. Brillantaisia lamium (Nees) Benth. in Hooker, Niger FL: 477 anthers mm staminodes 6-10 mm mm 1 1 in Cat. long, glabrous; 2-4(-6) long; ( 849); Burkill in FL trop. Afr. 5: 38 ( 899); Hiern afr. pi. flattened at to with long, tip vestigial anther, glabrous. Gynoecium 1: 807 (1900); Durand & Durand, Syll.fl. congol.: 417 (1909); style 15-20 mm long; stigma 1.5-3 mm long; ovary 3-4 mm long. Benoist in Bull Soc. Bot. France 60: 334 (1913); Chevalier, 17-21 x 1-2 with 20-24 seeds in each Capsule (10-) (-24) mm Explor. hot. Afrique occ. franc,. 1: 493 (1920); Mildbraed, Wiss. indumentum of slender white non trichomes and locule, glandular Erg. zweit. deut. Zentr.-Afr. Exped., Bot. 2: 90(1922); Hutchinson short trichomes. glandular & Dalziel in FL W. trop. Afr. 2: 254 (1931); Exell in Cat. pi. S.
Tome: 260 (1944); Heine, in FL W. trop. Afr. 2nd ed., 2: 406 DISTRIBUTION. Throughout West Africa and on the islands of the (1963); Heine inFl. Gabon 13: 88(1966).Type:A/we// 1842 (K!- Gulf of Guinea across Central African Repulic and Congo to Sudan, lectotype) (P!-isolectotype). Uganda and Kenya. Fig. 14.
HABITAT. A weedy species common in wasteland, clearings, plan- 1 for Leucoraphis lamium Nees in DC., Prodr. 11: 97 ( 847). Type: As tations and along roadsides; 60-1200 m. Brillantaisia lamium.
SELECTED COLLECTIONS Brillantaisia palisotii Lindau in Bot. Jahrb. Syst. 17: 99 (1893); in Brot. 10: 146 Lindau in Nat. IVORY COAST: Mont Momi, 29 October 1 966, AkeAssi 9 1 36 (G). GHANA: Henriques Bol. Soc. (1893).
1 1 1 Sierra Atewa Range N. of Kibi, 3 November 1990, Manktelow, Steiner &Amponsah Pflanzenfam. 4(3b): 296, fig. 9 ( 895). Syntypes: Leone, 89 (UPS). CAMEROON: N'Kolbisson, 3 November 1963. de Wilde & de Freetown, Preuss 16 (B); Sierra Leone, Freetown, Afzelius s.n.
1 130 8 km W. of Yaounde, Wilde-Duyfies (K, MO, WAG, YA); Nkolbison, (UPS!); Togo, Bismarksburg, Buttner 231 (B), Burner 320 (B); Mount Minloua, 19 November 1964, 1 1964 (P); 17 km Kribi- Raynal along Cameroon, Mungo, Bucholz s.n. (BM!); Sao Tome, Quintas Lolodorf road, 22 September 1969, Bos 5387 (C, K, MO, PRE, UPS, WAG, 1235(88) (BM!, BR!, COI, Z!), Quintas 1 132(89) (COI); Angola, YA); Chutes de Ntem, 40 km ESE of Campo, 10 December 1979, Letouzey Luanda, Pogge 1123(7). 15340 (P); Sao Nicolau, 9 January 1980, de Wilde, Arends & Groenendijk 3 B. eminii Lindau in Bot. Jahrb. Syst. 17: 103 (1893); Lindau in (WAG); crossing of road from S. Joao dos Angolares to Ribeira Peize, 12 Pflanzenw. Ost-Afr. C: 366 (1895 d); Burkill in FL trop. Afr. 5: 38 January 1980, de Wilde, Arends & Groenendijk 95 (WAG); Kumba area, 13 84 Brenan & October 1984, Thomas & Nemba 4048 (MO). BIOCCO: October 1911, (1899); Jex-Blake, Wildfl. Kenya: 103, fig. (1948); Terr. 2: 5 Mildbraedl028 (H). CENTRAL AFRICAN REPUBLIC: 27 km S. of Nola, Greenway, Checklist brit. empire 5 Tangan. (1949);
29 November 1965, LeeuwenbergllQQ (BM, C, K, LISC, WAG). CONGO: Andrews, FL pi. Sudan 3: 172 (1956); Synnott, Comm. forest. - 28 Sulongo, 22 March 1959, Everard 6002 (PRE); Piste Meya Mpassa, inst. occ. papers 27: 68 (1985). Syntypes: Tanzania, Bukoba, Farron 4085 SAOTOME: Famosa, 5 1949, April 1965, (P). January Espirito Stuhlmann 3664 (K!); Stuhlmann 3995 (Bt). Santo 68 (BM); Monte Cape, 5 March 1968, Espirito Santo 4298 (LISC); T. B. owariensis sensu Hook, in Bot. Mag.: t. 4717, fig. 3 (1853); road from Nora Moka to Manuel Jorge River, 1 5 October 1 993, Figueiredo & Anderson in Journ. Linn. Soc. Bot. 7: 339 (1863); Engler in Bot. Arriegas 33 (FHO, K, LISC). GABON: Belinga Mines de Per, 25 June 1966, Jahrb. Syst. 7: 339 (1886), non P. Beauv. (1818). Halle 4058 (P); Boka-Boka, Mount Bengoue, 4 March 1979, Florence 1706 Wild. T. Durand in: T. Durand De Wild. Bull 5 B. subcordata De & & (P); Woleu-Ntem, 2 May 1986, Louis 2077 (WAG); 10 km from Belinga, Soc. Bot. 38: 44-^5 (1899); De Wild. & Durand December 1986, Bos, van derLaan & Nzabi 10755 (K, US, WAG). DEMO- Roy. Belgique 1: DeWild. Miss. Em. Laurent CRATIC REPUPUBLIC OF THE CONGO: Dyuma, Pare National Albert, Ann. Mus. Congo 175-176 (1901); 418 11 October 1955, de Witte 12827 (MO). SUDAN: Imatong Mountains, 1: 182, t. 45 (1905); Durand & Durand, Syll. fl. congol.: Talanga, 1 December 1980, Friis & Volleson 612 (C). UGANDA: Budongo (1909). Type: Zaire, Bokakata, Dewevre 802 (BR!-syntypes). 8 November forest, December 1935, Eggeling 3323 (K); Mabira forest, 1938, B. subcordata var. macrophylla DeWild. &T. Durand. in Contrib.fl. KENYA: forest, 1 1 December 1956, Verdcourt on the Loveridge25 (K). Kakamega Congo 1, 2: 47 (1900). Type: Locality uncertain, village 1683 (K). Congo river, Duchesne 13 (BR!-holotype).
Brillantaisia is often locally abundant as a weed along vogeliana Icones: Lindau in Nat. Pflanzenfam. 4(3b): fig. 119A-G (1895); road and sides and in clearings in secondary vegetation, small Fl path Heine, in FL W. trop. Afr. 2nd ed., 2: fig. 300 (1963); Heine, leaves that are farms, plantations, etc. It is readily identified by Gabon: 13: pi. 17 figs 4-8 (1966). often often cordiform, characteristically irregularly toothed and rela- Perennial herbs (7-)20-100(-200) cm tall, creeping to erect, occa- with a toothed petiole; the highly branched inflorescence; 4.5 a clear mauve-violet and with adventitious roots at nodes. Stems mm tively small flowers which are usually sionally climbing which have numerous in diameter, occasionally with longitudinal wings on angles, gla- paler in the throat, and the pubescent fruit with B. brous to Leaves ovate to broadly ovate seeds per locule. This species might be confused velvety-tomentose. x mm or rarely oblong and owariensis, on the basis of the lax inflorescence and toothed (12-)25-80(-l 10) (20-)45-100(-180) with 55-90 x 20-30 mm, to tomentose with white silky leaves, and Vollesen tends to place East African specimens sparsely pilose trichomes to below; leaf base rounded teeth and a wider inflorescence into that species. Plants of above, pilose velvety-tomentose in cordate, subcordate or truncate occasionally shortly attenuate; apex B. vogeliana differ, from those of B. owariensis, possessing contracted to leaf acute with short acuminate tip or obtuse and abruptly thinner, papery leaves with a more dentate, irregular margin a acuminate entire, rarely subentire to slightly crenate; and pronounced dentate margin at the apex of the petiole wing; tip; margin of leaves at smaller flowers and lateral veins 6-10(-13) each side of midrib; upper pair more highly branched, slender inflorescence; from the base of terminal inflorescence sessile, petiole (0-)8-80(-l 10) mm fruit with more seeds. A few specimens of B. vogeliana of than B. unwinged or lamina decurrent close to apex petiole. Inflo- Congo basin have small flowers but larger leaves typical long, White-flowered rescence a terminal, open, branching thyrse, 12.5-35 cm long. vogeliana and seem to approach B. owariensis. 98 K. SIDWELL
x C. x D. x Fig. 14 Brillantaisia lamium (Nees) Benth. A. Habit x 0.7; B. Corolla dissection showing androecium 2; Gynoecium 2; Calyx 2.5;
E. Capsule x 1 .5; F. Bract and bractlets x 1 . REVISION OF BRILLANTAISIA 99 often with smaller lateral panicles below, primary branches of weedy. 'In moist exposed woody situations almost everywhere' (on inflorescence spreading almost at right angles to main axis, rachis Princes Island, Hiern, 1900: 807). Common; 200-1600 m. usually sparsely covered in capitate glandular trichomes. Bracts SELECTED COLLECTIONS ovate, adaxial surface pilose, often with slender multicellular SIERRA LEONE: Njala, 30 October 1928, Deighton 1419 (BM, K, MO). trichomes, abaxial surface glabrous; bractlets rhomboid to linear, LIBERIA: Louisiana, 1 1 November 1966, Bos 2310 (WAG); 10 miles N. of sessile, margin entire, adaxial surface covered with multicellular Monrovia, September 1 970, Jansen 2207 (WAG). IVORY COAST: Sassandra, non-glandular trichomes, abaxial surface smaller glabrous; becoming 1 8 June 1 963, de Wilde 280 (K, WAG, Z); Foret de YAPO, 8 km S. of Becedi- and linear towards ends of lateral branches. Sepals sub-equal, linear, Bugnan, 6 August 1963, de Wilde 641 (WAG); 27 October 1976, Fabrigues apex rounded, 4-7 mm long, green often tinged purplish; glabrous or 3306 (P). GHANA: Banyimade, 10 September 1956, Cudjoe, 161 (WAG). with a few long glandular trichomes. Corolla deep bright blue- TOGO: Kpandu, 1924, Robertson 62 (BM). NIGERIA: Oban, 191 1, Talbot or violet 981 (BM, K); Bauchi Plateau, 1930, Lely 652 (MO); Cross River, 27 purple occasionally pale lilac, one lip, often the upper; may August September 1985, Chile 27 (MO). CAMEROON: Bibundi, October 1891, be paler than the other; tube 5-8 x 2-4 mm, pale than lips, flushed Junger 248 (UPS); Bipindi, 1896, Zenker 1104 (BM); NW of Tibati, 9 inside with yellow, rarely completely white; upper lip (15-)22-30(- September 1963, Letouzey 5666 (P); Bakaka forest, 27 August 1971, 35) mm long, outer surface finely covered in glandular multicellular Leeuwenberg 8206 (H, LISC, MO, UPS, WAG); Moundjo, 27 October 1975, trichomes; lower lip (13-)19-27(-30) mm inner surface long, de Wilde 8522 (WAG). BIOCCO: Malabo, near Bae Basula, 13 September occasionally sparsely pubescent; apical lobe to 2 x 3 mm.Androecium 1986, Carvalho 2445, (BM); Sooye", S. of Mamon, 16 February 1945, with filaments 12-35 anthers 4.5-6 mm long, usually white; mm Roberty 6590 (G, Z). CENTRAL AFRICAN REPUBLIC: Koumbala, 25 long; staminodes exserted for 4-5 mm from corolla tube, small October 1983, Fay 6055 (MO); Bambari, November 1923, Tisserant 1352 vestigial anther. Gynoecium with style to 40 mm long, sparsely (P); Tisserant 3167 (BM, P). SAO TOME: 18 March 1968, Espirito Santo
4312 Monte 14 Monod 1 1681 DEMO- pubescent at base, stigma 3-5 mm long; ovary 3-4 mm long. (LISC); Cape, August 1956, (BM). CRATIC REPUPUBLIC OF THE CONGO: 26 June 1914, 4836 Capsule 22-30 x (2-)3-5 mm with 14-16 seeds per locule, blunt Bequaert (BR). BURUNDI: Ruyigi, 3 May 1980, Reekmans 9060 (MO, SRGH). tipped, green, tinged red at apex when immature, brown-black when SUDAN: Imatong mountains, Talanga, 2 December 1980, Friis & Vollesen mature and/or dried; glabrous or with a few erect, non-glandular 1 1 1 1 6 9 (C); Imatong Mountains, Acholi Hills, November 98 , Howard UTT 6 trichomes at the apex, covered in black gland dots. (C). ETHIOPIA: Bidgood s.n. (K). UGANDA: shores of Lake Nabugabo, 1 3 November 1934, Taylor 1660 (BM); Malabigambo forest, 15 August 1950, DISTRIBUTION. Throughout wet forest regions of west and central Dawkins D6 1 6 (BM). KENYA: Itare river, October 1940, Copley B 1 1 88 (G); Africa, east to southern Sudan, Zaire to northern and through Angola Nairobi, 12 October 1972, Hansen 713 (C). ANGOLA: Loanda, 1903, through Uganda to northwest Tanzania. Gossweiler 131 (BM).
HABITAT. Forest clearings, shade beside rivers or streams, on lake Brillantaisia lamium is a widespread species that is readily recog- shores or in swamps, often waterlogged/with roots in water, dis- nized by the cordate leaves with an entire margin and unwinged turbed areas, roads, paths and plantations. In west Africa often very petiole. The inflorescence is open and rather sparsely flowered with
Map 3 The distribution of Brillantaisia lancifolia Lindau. 100 K. SIDWELL
Fig. 15 Brillantaisia land/alia Lindau. A. Habit x 0.7; B. Corolla dissection showing androecium x 2; C. Gynoecium x 2; D. Calyx x 2; E. Bract and bractlets x 2; F. Capsule x 2.2. REVISION OF BRILLANTAISIA 101 few lateral branches that tend to be at right angles to the central axis. Thomas 2775 (MO). GABON: Cristal Mts, 21 January 1968, Halle & Villiers 4626 7 June Le Testu 5943 Fruit are characteristically glabrous but occasionally with a tuft of (P); Haute-Ngounye, 1926, (BM, BR, P). trichomes at the apex. Very few specimens have been collected of Brillantaisia lancifolia. East African specimens ofBrillantaisia lamium are often far more However, from available material and without further opportunity robust and pubescent than West African representatives, which tend for field study, I maintain this species as clearly distinct from all to be slender herbs; however, the variation appears to be continuous other species of Brillantaisia. Brillantaisia lancifolia has lanceolate and there are no morphological and/or geographical data to support leaves unlike any other species in the genus, the plants tend to be subdivision of the species. Vollesen (in prep, a) notes 'A similar delicate and leafy, with numerous axillary vegetative shoots. The where eastern plants from higher altitudes have larger flow- pattern nodes are regularly spaced and the internodes rarely exceed 40 mm. ers than western lowland is known from other Acanthaceae, plants The inflorescence is also very delicate with few flowers. Few of the alboviolaceum '. Brillantaisia lamium can form e.g. Anisosepalum specimens studied have fruit and additional counts of seeds per in areas of cleared or large weedy populations recently vegetation locule are needed. Two collections from Cameroon (Leeuwenberg plantations of oil palm, cocoa or bananas, and in open grassy areas 8994, WAG;Hepper 1 332, S) are intermediate in leaf shape between at altitude. In Cameroon slender low growing forms with very higher B. lancifolia and the less woody forms of B. owariensis; these purple flowers were cultivated as an ornamental in small deep specimens have ovate-elliptic leaves with a cuneate leaf base. How- on Mount Cameroon. Collections from village compounds Kenya ever, the flowers on these specimens are far larger than those of are of naturalized plants from Uganda and in swamp areas around typical B. lancifolia and have been provisionally determined as part Nairobi the is now quite common. species of the large and variable B. owariensis group.
9. Brillantaisia lancifolia Lindau in Bot. Jahrb. Syst. 17: 98 Engl. 10. Brillantaisia debilis Burkill in Fl. trap. Afr. 5: 39 (1899); mFl. 5: Benoist in Bull. Soc. (1893);Burkill trop.Afr. 40(1899); Benoist in Bull. Soc. Bot. France 60: 335 (1913). Type: Bot. France 60: 335 Heine in Fl. W. 2nd ed., 2: (1913); trap. Afr. Cameroon, Efulen, Batanga, Bates 350 (Ki-lectotype, BM!, 406 Heine in Fl. Gabon 13: 86 (1966). Gabon, (1963); Type: MO!, Z!-isotypes). Sierra del Mann 1688 (Bt-holotype, Kl-lectotype, P!- Crystal, Fig. 16. Map 4. isotype). Icon: Heine, Fl Gabon, Acanthacees: 13: pi. 16 figs 1-3 (1966). Fig. 15. Map 3.
B. sensu Heine in Fl. Gabon 13: 84, pi. 17, figs 1-3 (1966) B. talbotiiS. Moore Cat. pi. TalbotOban distr.: 75 (1913); Hutchin- soyauxii non Lindau (1893); non Burkill (1899). son & Dalziel in Fl. W. trap. Afr. 2: 254 (1931). Type: S. Nigeria, Oban Talbot 2000 (BM!, K!-syntypes). Leafy herbs 0.2-1 m tall. Stems 2-5 mm in diameter, often highly branched, rooting at nodes. Leaves elliptic-ovate to rhomboid, 45- Icon: Heine, Fl Gabon, Acanthacees: 13: pi. 19 figs 1-3 (1966). 1 10 x 25-58 mm, dull green, veins pale green to white; glabrous; tall. 2-3 in Perennial herb to 0.3-0.5 m Stems mm diameter, gla- lateral branches with smaller leaves 18^0 x 11-16 mm; leaf base brous to bases fused across densely pubescent, petiole forming ridge cuneate; apex acuminate; margin entire to shallowly crenate; lateral nodes. Leaves to lanceolate x 8-30(- ovate-elliptic 25-130(-190) veins 4-8 on each side of midrib; petiole 22^0(-80) mm, lamina smaller on or 50) mm, axillary shoots, glabrous very sparsely decurrent towards base of petiole. Inflorescence a few flowered base acute; entire to shallowly to pubescent; cuneate; apex margin very terminal panicle (50-)80-140(-180) mm long; rachis glabrous
serrated or subentire towards entire towards base; x 1 rounded crenate, apex, glandular pubescent. Bracts ovate-obovate, 4-22 3- mm, main lateral veins 5-8 on each 5-22 mm side; petiole long, unwinged, at apex; bractlets 1-5 mm long, narrowly oblong-obovate, glabrous lamina decurrent at a delicate, few-flow- apex. Inflorescence open, to glandular pubescent, pedicel 1-2 mm long. Sepals subequal, rachis Bracts with ered panicle 50-150 mm long, glandular pubescent. linear, rounded at apex, 5-10 x 0.5-1 mm, pale green, fringed often bractlets if linear lanceo- violet to caducous, linear-lanceolate, present unequal, purple, glandular trichomes. Corolla deep bright late with rounded linear, tube 2-3 x 5-12 apex, sparsely pubescent. Sepals unequal, purple with pale green or yellow spots in throat; towards the sepal 6-10 mm outer surface of tapering gradually apex, larger upper mm; upper lip 20-28 x 7-10 mm long, upper lip to 5-8 with long, smaller lateral sepals mm long, pubescent, capitate covered in short entangled, glandular trichomes, more so at apex; trichomes. Corolla blue to violet, often with brown glandular deep lower lip 19-25 mm long, glabrous above, sparsely glandular pubes- corolla tube sometimes with marking in throat; 5-10x2 mm, pale purple cent below; apical lobes triangular to 2 x 2 mm. Androecium x 2-3 tinged green; upper lip 10-15(-20) mm, glandular pubescent, filaments whitish to 20 mm long; anthers 3-4 mm long; staminodes of lower 10-18 x 6-8 mm, anther at particularly of edge upper lip; lip apical 5 mm long, densely glandular with a small vestigial apex. 1-2 x 1 mm. Androecium with filaments to 2 to 4 teeth rounded-triangular Gynoecium with style 25 mm long; stigma mm long; ovary staminodes x with 18-28 seeds 20 mm long, anthers 3-4 mm long; usually prominent, mm long, glabrous. Capsule 18-22 1-2 mm per with 15-28 around half length of the corolla lips. Gynoecium style locule, brown, glabrous. mm, stigma 1 mm, ovary 3 mm long, shortly glandular pubescent. when DISTRIBUTION. Cameroon and Gabon. Capsule 15 x 1-1.5 mm with c. 9-10 seeds per locule, brown dry, evenly glandular pubescent. HABITAT. Along rivers in primary forest in rocky, mossy places. 400-500(-900) m. DISTRIBUTION. Restricted to a few montane regions in Nigeria, Cameroon and Gabon. SELECTED COLLECTIONS CAMEROON: Yaounde, Bitye, Bates 1004 (BM, MO); Lolodorf, 1896, HABITAT. On rocks, in dark ravines or at edges of waterfalls; damp Staudt 240 (G, K, NY, S, WAG); Akoakas, 15 February 1963, Raynal & 300-700 m. 50 km S. of Batouri, 24 1 963, Letouzey Raynal 97 1 9 (P); near Molobo, July 4 March 1964, de Wilde & de 2072 SELECTED COLLECTIONS 5472 (P); Kwoeinvom, Wilde-Duyfies (BR. MO, SRGH, WAG, Z): 3 km N. of Lomie, 8 September 1965, NIGERIA: Oban, 1912, Talbot2WO (BM, K). CAMEROON: Ebone-Yabassi, Leeuwenberg 6560 (LISC, MO, WAG); Nvem, near Bitye. 20 October 1967, 27 December 1967, Bamps 1632 (BR); Mount Cameroon, above Batoke, 102 K. SIDWELL
x Fig. 16 Brillantaisia debilis Burkill. A. Habit 0.7; B. Corolla dissection showing androecium x 1.8; C. Capsule x 2; D. Gynoecium x 1.8; E. Bract and
bractlets x 1 . REVISION OF BR1LLANTAISIA 103
Map 4 The distribution of Brillantisia debilis Burkill.
Letouzey 8159 (P); 5 km E. of Meyo Centre, 24 March 1970, Letouzey 10229 Erect aromatic herb to l-2(-2.5) m tall. Stems 2-4 mm in diameter, 11 October de Wilde 7635 Rocher de (P); N'Koemvon, 1974, (WAG); rooting at nodes, covered in rather dense short, weak trichomes. Akoakas, 27 March 1981, Meijer 15307 (MO); Cult. WAG, 20 December Leaves ovate to broadly-ovate (35-)90-225(-250) x ( 10-)40-1 10(- 1985, van derLaan 986 (WAG). GABON: near lake Moudiboubacoudou. 6 1 50) mm, olive green to dull dark green above, paler below, glabrous April 1927, Le Testu 6478 (BM); Le Testu 9135 (BM); Monts de Cristal, 3 to pubescent above and below; base cuneate; apex acuminate; February 1968, Halle & Villiers4891 (P); Akoga, 8 February 1968, Halle & margin crenate to serrate, occasionally serrulate; 8-13 veins each Villiers 5047 (P); Cristal Mountains, 15 km NE of Asok, 21 August 1978, side of the midrib; petiole (10-)20-70(-1 10) mm, lamina decurrent Breteler & de Wilde 1 87 (MO, WAG); Cristal Mountains, 2 1 January 1 983, de !/2-% of a terminal Wilde et al. 133 (WAG); W of Belinga, 18 July 1985, Bos, van der Laan & along length petiole. Inflorescence spike (60- with smaller lateral flowers Nzabi 1 0696 (BR, WAG); 24 km SE of Medouneu, 5 February 1 986, Reitsma, )80-200(-340) mm long spikes below, Reitsma & Louis 1880 (NY, WAG); Wolou-Ntem province, Chantier Oveng in distinct verticillate whorls, rachis often visible between near Mitzic, 6 May 1986, Louis 2161 (WAG). verticillasters. Bracts ovate to linear-lanceolate, entire or crenate, rounded at apex, densely pubescent with glandular and eglandular Plants of Brillantisia debilis have few flowers and are often very trichomes, bractlets linear, trichomes as for bracts. Sepals subequal, leafy, the vegetative parts of plants of this species are entirely linear rounded at apex, 0.5-1 x 12-20 mm, densely covered in long glabrous and the leaves are often purple underneath. The inflores- glandular and eglandular trichomes. Corolla dark blue-purple to cence may be glandular-pubescent on the rachis, peduncles, calyx violet or bright blue-lilac; tube 8-1 3 x (2-)3-5 mm; upper lip 20-30 and corolla. Many collections of this species have been mistakenly x 4-6 mm covered in short glandular trichomes, which are slightly determined asfl. soyauxii. In Flare du Gabon, Heine ( 1 966) used the longer at the apex of the petals; lower lip 22-30 x 14-20 mm; apical name B. soyauxii to describe a species which is glabrous except for lobes to 7 mm long, acute. Androecium with filaments 35^40 mm the pedicels, calyx and, occasionally, the peduncles. The illustration long; anthers 6-7 mm long; staminodes 5-10 mm long Gynoecium of B. soyauxii in that work (Heine 1966: 87, 1-3) clearly fits the with style 35 mm long pubescent to stigma, stigma 4 mm long. description of B. debilis given here. The type specimen of B. Capsule 20-22 x 2-3 mm with 8 seeds per locule, brown-black, soyauxii has pubescent fruit, and a highly branched, many flowered covered with sparsely glandular trichomes. inflorescence unlike B. soyauxii described in the Flore du Gabon flowers fruit. Other (Heine 1966: 84-85) with few and glabrous DISTRIBUTION. Ethiopia and possibly Sudan. characters on the type specimen of B. soyauxii are inconsistent with HABITAT. Montane forest, beside streams in shade, in disturbed the description of Heine (1966), including the larger, broader, collected from plantations; 1500-2000 m. spathulate upper calyx lobe and lower number of seeds in the fruit. ground, occasionally Soyaux 454 is certainly a plant belonging to B. vogeliana. SELECTED COLLECTIONS ETHIOPIA: Kaffa Prov., Beletta forest. 13 February 1957, Mooney 6725 11. Brillantaisia Pichi-Sermoli in Miss. Stud. Lago grottanellii (ETH): Wellega region near Dembidollo, Anfilo, 6 March 1957, Mooney Tana 7. Recherche Botaniche 1: 254-255 Cufodontis, (1951); 6890 (S): 7 km NW of Tippi airstrip, 14 January 1962, Meyer 7995 (US); 35
Tucur 1 Enum. pi. Aeth.: 930 (1964). Type: Ethiopia, Dinghia km W. of Lekemeti. 1 3 April 1 966. de Wilde & de Wilde-Duyfjes 0768 (MO, forest, Pichi-Sermoli 2099 (FT-holotype, BM!-photo). WAG); Kaffa Prov., Bonga, 9 January 1972. Ash 1454 (ETH, MO, UPS, Kaffa 57 km from Jimma on river track, 8 Map 5. WAG);. Prov., Sheki-Gogeb 104 K. SIDWELL
Table 5 Six characters which differ between plants of Brillantaisia grottanellii and B. madagascariensis.
Character B. groUanellii B. madagascariensis
Bract shape lanceolate-ovate broadly ovate-ovate
Glandular trichomes on present absent
bract
Trichomes on style along more than half length at base of style only
of style
Leaf margin subentire-serrate entire or crenate
Upper sepal shape linear-spathulate linear
Pollen ornamentation smooth irregularly spiny
December 1972, Friis et al. 1677 (C); Mt. Karkarha, 18 February 1976, Ash Usambara Mountains, Gonja, Handei, Hoist 4216 (COI, G!, K!, 3402 (MO, UPS); Gofa region, 10 km from Sawla, 27 December 1983 M!, P!,W!, WAG!, US!, Z!). Ensennu Zerihun 649 43 km from MizanTeferi on road & (C. ETH); toTepi, B. verruculosa Lindau in Bot. Jahrb. Syst. 22: 1 13 (1897); Burkill in 27 January 1984, Ensennu et al. 745 (ETH); 37-41 km on the track from FL trap. Afr. 5: 43 (1899); Mildbraed, Wiss. Erg. zweit. deut. Gore to Masha, 1 February 1984, Ensermu et al. 821 (ETH); 27 km from Zentr.-Afr. Exped., Bot.: 192 (1922); Robyns Fl. pare. nat. Albert Bebeka coffee plantation on road to Guraferda, 7 November 1985 Puff & 2: 269(1947); Heine inF/. Gabonll: 89 (1966). Type: Cameroon, Ensermu 861107-4/6 (ETH); Wellega Region, Bedelle/Arjo road-Didessa Yaounde, Zenker & Staudt 166 (Bf-holotype). road crossing, 16 January 1990, Friis et al. 6023 (ETH); Kaffa Prov., B. S. Moore in J. Bot. 46: 312-313 (1908). Type: Beletta forest, 18 January 1990, Friis et al. 6041 (ETH); SUDAN: Torit bagshawei 1387 Distr., near Katire, 9 February 1950, Jackson 1144 (BM, MO), identity Uganda, Bugoma forest, Bagshawe (BM!-holotype). uncertain. B. majestica Wernham in J. Bot. 54: 229 (1969). Type: Cameroon, Buea, Mount Cameroon, Bates 817 (K!-holotype). This species is clearly very closely related to Brillantaisia mada- gascariensis and, early in this study, was thought to be the same Icones: Heine, Fl Gabon, Acanthacees: 13: pi. 18 figs 7-15 (1966); species. However, after detailed study of gross morphology and Benoist inF/. Madag. fam. 182, Acanthacees: fig. IV 8-13 (1967). pollen characters, I recognize this species as distinct. Discussion Erect or scrambling much branched herb 0.2-2 m tall. Stems 2-5 (- with Dr Ensermu Kelbessa from Addis Ababa University who has 8) mm in diameter, shortly pubescent, more densely so at nodes, studied these plants in the field convinces me that the madagas- glabrous below. Leaves ovate to broadly ovate, 40-150(-180) x 30- cariensis-grottanellii group is best divided. Characters differentiating 90(-130) mm, lower leaves becoming much larger, to 400 x 500 B. grottanellii from B. madagascariensis are shown in Table 5 and mm, usually sparsely pubescent, with short white or translucent Figure 5. trichomes on midrib and main veins, occasionally densely pilose; leaf base attenuate, occasionally truncate or cordate; apex acute- 12. Brillantaisia madagascariensis T. Anderson ex Lindau in Bot. acuminate; margin entire, occasionally slightly crenate; 8-12(-14) Jahrb. Syst. 17: 103 (1893); Burkill inF/. trop.Afr. 5: 43 (1899); main lateral veins each side of midrib; petiole 6-90 mm long, 3: hot. Palacky, Cat. pi. Madag. 57 (1907); Chevalier, Explor. 2 winged for /3 of length. Inflorescence a terminal spike with smaller Afrique occ.franq. 1: 493 (1920); Hutchinson & Dalziel in Fl. lateral spikes below (20-)40-220(-350) mm long, flowers crowded W. trap. Afr. 2: 254 (193 1); Brenan & Greenway, Checklist brit. into dense verticillate whorls, rachis densely pubescent, not usually empire 5 Tangan. Terr. 2: 5 ( 1 949); Heine in Fl. W. trap. Afr. 2nd visible between verticillasters. Bracts ovate to elliptic, apex rounded ed., 2: 406 (1963); Cufodontis, Enum. pi. Aeth: 930 (1964); to acuminate, green with purple tinge, persistent. Sepals subequal, Benoist in Fl. Madag. fam. 182 1: 29 (1967); Agnew in Upl. slender tapering to apex or linear and rounded at the apex, occasion- Kenya wild/Is: 583 (1974); Champluvier in Fl. Rwanda 3: 446 ally spathulate, larger upper sepal 10-30 x 1-2 mm, very slightly (1985). Type: Madagascar, Betsileo, Nandihizana, Hildebrandt longer than the other four; lateral sepals 8-25 x 1-2 mm; green, red 3901 (Bt-holotype; BMI-lectotype; G!, M!, P!, W!, Z!- to purple at apex, densely pubescent on abaxial surface, trichomes isolectotypes). white, less dense shorter trichomes on adaxial surface. Corolla
Brillantaisia spicata Lindau in Bot. Jahrb. Syst. 20: 4 (1894); in variable shades of blue-purple, violet, magenta or white, with lateral
Abh.. Preuss. Akad. Wiss. 50 & 54 (1894); in Wiss. Erg. deut. yellow spots in the throat; corolla tube to 15 mm long; upper lip 15- Zentr.-Afr. Exped., Bot. 2: 293 (1911). Syntypes: Tanzania, 30 x 5-8 mm, glandular pubescent outside occasionally with long Usambara Mountains, Bangarra Lutindi, Hoist 3316 (COI, K!); trichomes; lower lip 1 5-25 x 1 0- 1 2 mm, apical lobes 2-5 x 2-3 mm, REVISION OF BRILLANTA1SIA 105 often short, rounded-blunt but can be longer, narrowly triangular Lambinon 78/321 (BR). KENYA: Kakamega forest, 17 November 1975, s.n. TANZANIA: Mt Surra 10 Novem- with acute apex. Androecium with filaments 20-22 mm long, white; Cunningham (LISC). Meru, Forest, ber Richards 24663 (C, above tea estate N. of Derema, anthers 3-6(-8) mm long, creamy yellow or brown, pubescent along 1969, M); Mshituni, 5 October 1979, Kibuwa 5191 (UPS). MADAGASCAR: August 1902, one side; staminodes 5-10 mm long. Gynoecium with style 25 mm Perrier 9227 (P); near Doanyanala, 25 January-25 February 1949, long, often persistent when corolla has fallen, stigma 2.5 mm long; Humbert 23090 (P); 6 December 1963, Rakotozafy 287 (P). ovary 3-5 mm long. Capsule 1 8-30 x 3-5 mm with 6-8 seeds per locule, black-brown, glandular puberulous with longer scattered Plants of this species are very distinctive in flower and fruit, with a eglandular trichomes. narrowly spicate inflorescence, flowers in verticillasters, persistent bracts and linear, pubescent calyx lobes. Brillantaisia verruculosa DISTRIBUTION. West Africa the basin Throughout through Congo has been upheld as a species by several recent workers, notably to and south into also in Ethiopia Tanzania, Madagascar. Heine (1966), solely on the basis of the floral colour being white rather than purple. White flowers are found in three other species of HABITAT. Montane forest, often weedy in plantations, recently Brillantaisia and as this is the only character on which Heine split B. logged areas, clearings or pathsides; 500-1200 m. madagascariensis, I maintain this variation under the earliest name. SELECTED COLLECTIONS
LIBERIA: Yekepa, Yiti village, 24 January 1965, Adam 20734 (K, MO). BIOCCO: November 1911, Mildbraed 7126 (H). CAMEROON: 200'N 1215'E, 2 January 191 1, Mildbraed 4\41 (H). GABON: Haute-Ngounye, NOMEN DUBIUM 1924-1927, Le Testu 6431 (BM, MO); Lastoursville region, 1929-1931, Le Testu 8853 RWANDA: near 4 (BM, P). Nyungwe forest, Rukuzi, May Brillantaisia cicatricosa Lindau in Bot. Jahrb. Syst. 20: 4 (1895) 1972, Bouxin 1577 (BR). BURUNDI: on Rwandan border, Mabaya-Hua nomen dubium; Lindau in Pflanzenw. Ost-Afr. C: 366 (1895); 22 June 1969 Lewalle 3783 (BR, G, SRGH); Mabayi valley, Bubanza, 16 Burkill in Fl. trap. Afr. 5:39 ( 1 899); Mildbraed, Deutch Zent. Afr. October 1971, Reekmans 1 136 (BR). SUDAN: Torit District, Lotti forest, Exped. (1922); Robyns, Fl. pare, nat Albert 2: 270 (1947); 2 January 1950, Jackson 1023 (BM); Talanga, 1 December 1980, Friis & Troupin, Fl. pi. lign. Rwanda: 84 (1982); Champluvier in Fl. Vollesen 613 (C). ETHIOPIA: Kaffa Prov., Abetu valley, near Belita, 18 Rwanda 3: 446 (1985); Fischer & Hinkel, Nat. Env. Rwanda: 81, October 1954, Mooney 6071 (ETH, K, S); 10 km E. of Bonga along Jimma 68 W. of Ruwenzori Mountains, road, 22 December 1965, de Wilde 9415 (C, DSM, ETH, MO, UPS, WAG); fig. (1992). Type: Zaire, slope 2301 Problems with Kaffa Prov., Bonga, 20 November 1970, Friis, Hounde & Jacobsen 369 Buhtahu Valley, Stuhlmann (Bt-holotype). (C); IllubaborProv., 22 km S. of Gore, 19 December 1972, Friis et al. 1882 the application of this name are described in detail above (p. 95). (C); UGANDA: Budongo Forest, 7 February 1935, Taylor 3327 (BM, Brillantaisia hirsuta T. Anderson in Journ Agric.-Hort. Soc. NS 1: MO); Mabira Forest, 8 November 1938, Love ridge 26 (MO). DEMO- 267 This name refers to a introduced to the CRATIC REPUPUBLIC OF THE CONGO: Kasongo-Kindu road, km 60, (1868). plant Royal Botanic Calcutta from Kew and which cannot now be 29 June 1952, Germain 7826 (PRE); Valley de la Mbizi, near Burora, 10 Gardens, that 'It is on the May 1957, Gutzwiller 841 (BR); Kivu, W. of Kaluzi Mts, 26 April 1978, traced. Anderson wrote of this species common
Map 5 The distribution of Brillantaisia grottanellii Pichi-Sermoli. 106 K. SIDWELL
Euramerica: an on Annals the Missouri banks of streams in Western equatorial Africa. It differs from B. essay palaeobotanical phylogenetics. of Botanical Garden 79: 500-559. owariensis in its very prostrate habit, much smaller non decurrent Benoist, R. 1967. Acanthacees. In H. Humbert (Ed.), Flore de Madagascar et des in the smaller size of its flowers which are not e leaves and Comores: 182 famille. Paris.
to Brillantaisia J. Flora: 477^478. glandular'. This description may refer vogeliana Bentham, G. 1849. Brillantaisia. In W. Hooker. (Ed.) Niger London. which has sparsely pubescent flowers, however, as this is the only 1 876. Acanthaceae. In G. Bentham, & W. J. Hooker (Eds), Genera plantarum: information regarding the the identity of the plant in Calcutta, 2: 1060-1122. London. of this name remains uncertain. application Blackmore, S. & Tootill, E. (Eds) 1984. The Penguin dictionary of botany. Middlesex. Bremekamp, C.E.B. 1944. Materials for a monograph of the Stobilanthinae der Nederkmdse Akademie van Wetenschappen transferred to (Acanthaceae). Verhandelingen Species Hygrophila Afdeeling Natuurkunde 41: 1-305. 1953. The delimitation of the Acanthaceae. Proceedings Koninkijke Nederlandse Akademie van 56: 533-546. Brillantaisia borellii Lindau in Bot. Jahrb. Syst. 33: 1 86 ( 1 904). This Wetenschappen 1965. Delimitation and subdivision of the Acanthaceae. Bulletin the all other of species was considered to be very different from Botanical Survey of India 1: 21-30. Brillantaisia due to the presence of four fertile stamens, species Bin-kill, L.H. 1899. Brillantaisia. In W.T. Thistleton-Dyer, (Ed.). Flora of Tropical leaves and the lack of an corolla very narrow upper lip. Africa: 5: 37. London. Carlquist, S.& Zona, S. 1988. Wood anatomy ofAcanthaceae: a survey. Aliso 12: 201- in Bot. Jahrb. 24: 3 1 3 1 Brillantaisia didynama Lindau Syst. ( 898). 227. Considered by Lindau to be clearly distinct from all other species Champluvier, D. 1985. Brillantaisia. In G. Troupin, (Ed.) Flore du Rwanda: of Brillantaisia by virtue of having four fertile stamens and very Spermatophytes 3: 444-448. Tervuren. Chappill, J.A. 1989. Quantitative characters in phylogenetic analysis. Cladistics 5: distinctive Lindau maintained this species within the pollen, 217-234. and it close to B. lancifolia due to the few-flowered genus placed Cramer, L.H. 1991. Brillantaisia P. Beauv. (Acanthaceae), a new generic record for inflorescence and lanceolate leaves. Asia. Kew Bulletin 46: 335-338. Cranston, P.S. & Humphries, C.J. 1988. Cladistics and computers: a chironomid in Bull. 46: 335-338 Brillantaisia thwaitesii Cramer Kew (1991). conundrum? Cladistics 4: 72-92. Cramer transferred Plaesianthera thwaitesii (C.B. Clarke) Livera Daniel T.F. 1984. Revision of Stenandrium (Acanthaceae) in Mexico and adjacent Annals the Missouri Botanical Garden 71: 1028-1043. to Brillantaisia as he considered it close to Brillantaisia pube- regions. of De, A. I960. Cytological, anatomical and palynological studies as an aid in tracing scens. Recent studies (Sidwell, in press a) have shown that the affinity and phylogeny in the family Acanthaceae. IV. Transactions of the Base is best in until a needed mono- species placed Hygrophila greatly Research Institute Calcutta 31: 17-29. graph of that genus is undertaken. De Pinna, M.C.C. 1996. Comparative biology and systematics, some controversies in retrospective. Journal of Comparative Biology 1: 3-16.
Doyle, J.A. & Donoghue, M.J. 1 986. Seed plant phylogeny and the origin ofangiosperms: Species transferred to Eremomastax an experimental cladistic approach. The Botanical Review 52: 32 1^-3 1 . Ensermu, K. 1 990. Justicia sectAnsellia (Acanthaceae). Acta Universitatis Upsaliensis Symbolae Botanicae Upsaliensis 29: 1-96. Brillantaisia Lindau in Bot. Jahrb. 24: 313-314 (1898). fulva Syst. Erdtman, G. 1960. The acetolysis method. A revised description. Svensk Botanisk
Tidskrift 54: 561-564. Exell, A.W. 1944. Catalogue of the vascular plants of S. Tome (with Principe and Annobon). London. ACKNOWLEDGEMENTS. I would like to thank Robert Scotland and Kaj Ezcurra, C. 1993. Systematics of Ruellia (Acanthaceae) in southern south America. Vollesen for the course of PhD Thanks also supervision throughout my study. Annals of the Missouri Botanical Garden 80: 787-845. to John Baker, Ian Gourlay, Julie Hawkins, David Mabberley, Alison Strugnell Farris, J.S. 1988. Hennig86: version 1.5. Published by the author, Jamaica Plains. and Rosemary Wise in Oxford; Alex Ball and Louisa Jones in the SEM unit Furness, C.A. 1994. The pollen morphology of Hygrophila and Brillantaisia Acta Botanica Gallica 141: 267-278. at The Natural History Museum; everyone at Limbe Botanic Gardens and the (Acanthaceae: Ruellieae). Gift, N. & Stevens, P.F. 1997. Vagaries in the delimitation of character states in National herbarium in Martin Cheek and Alan Paton at Kew and Cameroon; - quantitative variation an experimental study. Systematic Botany 46: 1 12-125. Axel Poulsen in Copenhagen. This research was funded by BBSRC research Graham, V.A.W. 1988. Delimitation and infra-generic classification of Justicia with additional financial assistance from Wolfson grant 93309762, College (Acanthaceae). Kew Bulletin 43: 551-624. to cover seeds and fruits Oxford and the University of Oxford Hardship Fund. Funding Grubert, M. 1 974. Studies on the distribution of myxospermy among 8: 315- materials and tuition for all illustrations was received, with thanks, from the of angiospermae and its ecological importance. Acta Biologica Venezuela 551. Linnean Society of London's A.G. Side fund for research in systematic Hauser, D.L. & Presch, W. 1991. The effect of ordered characters on phylogenetic biology. reconstruction. Cladistics 7: 243-265.
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Palisot de Beauvois, A.M.EJ. 1818. Flore d'Oware el de Benin en Afriaue. 67-68, t.100. Paris. Pimentel, R.A. & Riggins, R. 1987. The nature of cladistic data. Cladistics 3: 201- EXSICCATAE 209.
Platnick, N.I., Griswold, C.E. & Coddington, J.A. 1991. On missing entries in Abdullah & Vollesen 95/176, pubescens (K). cladistic analysis. Cladistics 7: 337-343. Achien 524, lamium (BR). Pleijel, F. 1995. On character coding for phylogeny reconstruction. Cladistics 11: 309- 315. Adam 2968, owariensis (K, MO); 4782, lamium (BR, K); 4830, lamium lamium lamium 7351, lamium Punt, W., Blackmore, S., Nilsson, S. & Le Thomas, A. 1994. Glossary of pollen and (MO); 5286, (MO); 6910, (MO); (MO); spore terminology. LPP Contributions series No. I. Utrecht. 16325, lamium (K. P); 16453, lamium (K); 20464, lamium (K); 20549, Quicke, D.L.J. 1993. Principles and techniques of contemporary taxonomy. London. owariensis (BR, K, MO, P. UPS); 20734, madagascariensis (K. MO); L. 1883. Werth der Pollenbeschaffenheit bei den Radlkofer, Ueberdie systematischen 22019, owariensis (MO, P); 22262, lamium (MO); 22916, owariensis Acanthaceen. 5. B. Akademie der math-phys. Cl. 13: 256- buyer. Wissenschaften., (MO, P); 23291, lamium (MO, PRE); 241 16, lamium (MO); 24302. 314. lamium (MO); 24663, lamium (MO); 24795, lamium (MO, PRE); Raj, B. 1 96 1 . Pollen morphological studies in the Acanthaceae. Grana Palynologica 3: 25258, owariensis (MO); 26520, lamium (MO); 26558, lamium (MO, 1-108. PRE); 27084, owariensis (MO); 30140, lamium (MO). Ramos, T. 1997. Tree Gardener: version 2.2. Sao Paolo. Adames 73, lamium 694, lamium 791, owariensis (K, 1 . (K); (K); UPS). Rickett, H.W. 1 944. The classification of inflorescences. Botaw/a;/ Review 10: 1 87-23 FHI lamium 1 955. Materials for a dictionary of botanical terms-Ill, Inflorescences. Bulletin of Adebusuyi 43972, (K). the Torrev Botanical Club 82: 419^45. Afzelius s.n., lamium (UPS). 108 K. SIDWELL
Ake Assi 3244, lamium (G); 3333, owariensis (G); 5502, owariensis (G); 187, 7186, lamium (G); 9136, vogeliana (G); 9243, lamium (G); 1221 1, Breteler si al. 283, owariensis (K). owariensis (G, Z); 15745, lamium (G); 17170, owariensis (G). Bridson 610, madagascariensis (K, MO). Alluard 309. owariensis (P); 390, owariensis (P). Bridson & Lovett 544, kirungae (MO). Anderson 1383. lamium (K). Brodhurst-Hill 693, owariensis (Z). Andrada 1811, kirungae (LISC). Brown 2107, owariensis (MU). Amr// 1842, lamium (K). Brown & Brown 33a, lamium (US); 316/b lamium (US). Archibald 2589. kirungae (C). Brunt 167, owariensis (K). Arena's & Groenemlijk 3, vogeliana (WAG); 95, vogeliana (WAG). Brynaert 406, pubescens (BR). Ariwaodo 33. owariensis (MO): 77, owariensis (MO). Bunting s.n., lamium (BM, MO). As/? 1454. grottanellii (ETH, MO, UPS, WAG); 3402, grottanellii (MO, Burbidge 473, owariensis (K). UPS). Burrows 769, kirungae (SRGH). Auauier 2684, owariensis (BR). Bwm 9, lamium (K); B16, owariensis (K). Bahault s.n. owariensis (P); s.n. kirungae (P). Burn Davy 1228, kirungae (FHO). Bagshawe 1131, owariensis (BM); 1270, kirungae (US); 1342, owariensis Butayo s.n., owariensis (BM). (BM); 1381, madagascariensis (BM); 1506, owariensis (BM). Buttner 353, owariensis (W). BrtWuw 10182, owariensis (K, MO); 10778, owariensis (K, MO); 5993, Cambridge Congo Expedition 146, kirungae (BM, LISC, US). /wi/M/n (K, MO); 6175, /a/w/Mm (K); 6339, /am/w/n (K); 7070, /am/Mm Cardosode Matos 1 52, owariensis (LISC). (K, MO). Carmichael 102, kirungae (FHO). ft/// 62. kirungae (MAL, SRGH). Carpenter 312, owariensis (K). Carvalho /am/nm fiwwp.v 1632, lancifolia (BR); 2592, /a/n/wm (K). 2359, vogeliana (K); 2445, (BM, K, MO, S); 3766, owariensis <& Kaunda 3452, riparia (MO). (K, MO). 1 owariensis 1 owariensis 1 c$ Salubeni 1571, r/paria (MO, SRGH). Caw 1471, (K); 1723, (K, MO); 1790, owariensis Barbosa 1812. /-/panY/ (LISC); 2042, pubescens (LISC); 2395, pubescens (K). (LISC). Chaloner 342c owariensis (K). HNC191, owariensis (K); 827, (BM); 5053, fiarfcr 1 131. /tfw/M/n (K); 1418, /a/nw/H (K). Chapman oligantha owariensis owariensis (K); 5258, owariensis (K). Barton (?) 103. /;;/// (K). (K); 5141, & 8862, (E, FHO, MO, PRE); 9271, #/ 216. vogeliana (BM, E, K, G, Z); 350, de Wilde, A rends & 3, liana Gmenendijk voge (WAG); 95, liana 1 voge Fischer 8 , lamium (K). (WAG). Fishlock 16, lamium (K); 55, owariensis (BM, K). de Wilde & de 1 130, Wilde-Dufjes vogeliana (K, MO, YA); 2072, debilis Flanigan 193, owariensis (NY). (BR, K, MO, SRGH, WAG, YA, Z); 9415, madagascariensis (UPS, Florence 1706, vogeliana (P). WAG); 10768, gmttanellii (MO, WAG). For/Ms 2485, owariensis (P, YA). de Wilde & Voorhoeve 3772, lamium (K). Frederick 1 8439, madagascariensis (BR); 949 , owariensis (BR). de Wilde, de Wilde & de 1 owariensis Wilde-Duyfjes 152, (MO, PRE, WAG, Friedberg 1, vogeliana (K). YA); 3912, owariensis (H, MO, 4146, owariensis Fr/7s <& WAG); (K, WAG); Vollesen 58, kirungae (C); 483, owariensis (C); 612, vogeliana 4198, owariensis (WAG); 4513, owariensis (WAG); 4557, owariensis (C); 613, madagascariensis (C); 619, lamium (C); 1299, owariensis (C). (WAG). Friwetal. 1677, gmttanellii (C); 1 882, madagascariensis (C, ETH); 1908, de Wilde et al. 133, debilis (WAG). madagascariensis (C, ETH); 6023, gmttanellii (ETH); 604 1 , gmttanellii de Wine 1434, kirungae (BR); 1508, kirungae (BR); 1539, kirungae (BR): (ETH). 3986, pubescens (BR, K); 9697, owariensis (BR); 10747, kirungae Fr//s, Gilbert & Vollesen 4109, madagascariensis (ETH, UPS). 10872, owariensis 1 (MO); (BR); 1065, kirungae (MO); 1 1640, Fri/X Hounde & Jacobsen 369, madagascariensis (ETH). kirungae (BR); 1 1862, owariensis (BR); 12148, kirungae (C); 12437, Froment 385, owariensis (BR). owariensis 1 268 1 (BR); , madagascariensis (BR); 12827, c/ vogeliana Garley 464, kirungae (LISC). (BR, MO); 12907, owariensis (BR, MO); 12919, owariensis (BR). Garrett s.n., owariensis (K). Decary 2047, madagascariensis (P); 7782, pubescens (P); 8208, Gassy Leon 2002, lamium (K). pubescens (MO, NY); 14.278, madagascariensis (P); s.n., GWte 27, /a/n/MOT (MO). madagascariensis (US). Geerling & Bockdam 1533, /am/H/n (C, MO, WAG); 1739, /ammw (K, Deighton 356, owariensis (K); 1419, lamium (BM, K, MO); 3571, MO, WAG); 1859, owariensis (C, MO, WAG); \95],lamium (C, MO, owariensis (K); 5169, lamium (K); 5897, owariensis (K); 6141, lamium WAG). (K); 6152, fam/M/n (K); 1419, /am/Mm (K). Gelinger 2869, kirungae (Z). Demeuse 23, owariensis (BR). Gentry 1 1720, madagascariensis (MO). Demoulin 61, lamium (K). Geo Watt 7049, /a/n/M/n (BM, MO Dennett 34, owariensis (K). Gerard 1954, lamium (K); 3626, lamium (K); 5341, lamium (BR, K). et al. 1 94, lamium (K). Gereau 5561, lamium (K); 5629, owariensis (K); 5657, owariensis , owariensis (BM, BR); 3429, c/ vogeliana (BR). (K). Dinklage 2624, lamium (H). Germain 1952, madagascariensis (PRE); 3915, kirungae (BR); 4934, Don s.n., vogeliana (BM); s.n., owariensis. madagascariensis (BR); 7826, madagascariensis (BR); 8141, Dowsette-Lemaire 999, kirungae (MAL). owariensis (BR). G/7be/7 cfc Drummond & Hemsley 4556, madagascariensis (S); 4584, lamium (S); Afev//n 6612, kirungae (PRE). G/V// 5 1 s.n., kirungae (S). 9, kirungae. Dschang College 1712, owariensis (K). Cocker 2, vogeliana (MO, US). Duchesne 13, lamium (BR). Goldsmith 54161, kirungae (MO). Dujandin 409, owariensis (BR). Gomes & Sousa 4137, riparia (PRE). Goosens 1313, owariensis owariensis Diimmer241, owariensis (MO, US, Z); 587, c/ owariensis (P, US); 1048, (BR); 2369, (BR); 6030, lamium lamium (US, Z); 3975, madagascariensis (US). (K). DHAU/OS- FHI 15323, /a/n/wm (K). Gossweiler 131, lamium (BM); 500B owariensis (BM); 1048, owariensis Dunlap 25, owariensis (K). (BM). Dusen s.n., lamium (S). Goudot s.n., madagascariensis (G). Eckendorfl3, owariensis (P). Greenway & Brenan 8044, pubescens (PRE). Eggeling 402, /am/w/n (FHO); 1380, /am/ww (FHO); 1487, owariensis Greenway & Eggeling 8394, kirungae (PRE); 8594, kirungae (FHO). (FHO); 3323, vogeliana (K). Greenway & Farquhar 8634, madagascariensis (NY, PRE). Eijnatten 2130, lamium (WAG). Greenway & Kanuri 12107, madagascariensis (PRE). Ekwuno, Osanyinlusi & Okoro 124, owariensis (MO). Grew 26, pubescens (P). Elskens 7, owariensis (BR); 76, owariensis (BR); s.n., owariensis (BR). Gmenendijk & Dunge 466, riparia (SRGH). Emwiogbon FHI 87191, /a/n/wm (MO); 66079, owariensis (K). Guile 1733, owariensis (MO). Emwiogbon & Osanyinlusi FHI 87313, lamium (MO). Guinea 482, vogeliana (K). Ensermu et al. 745, gmttanellii (ETH); 821, gmttanellii (ETH). Gutzwiller 841, madagascariensis (BR); 1612, madagascariensis (BR). Ensermu & Zerihun 649, gmttanellii (C, ETH). //.V/, P652, /am/Mm (K). Espirito-Santo 67, lamium (PRE); 68, vogeliana (BM); 3870, lamium //aaren 2483, /am/urn (G). (LISC); 4312, /a/n/jim (LISC); 4727, fa/mum (LISC); 4298, vogeliana Hall-Martin 8 1 9, pubescens (PRE). (LISC); 5118, /am/wm (LISC). //a//affi 128, /am/M/n (K). //a//e owariensis Efwge (ft Thomas 442, owariensis (K, MO); 45 1, owariensis (MO). 2382, (P); 3130, owariensis (P); 3337, vogeliana (P); Evrarc/2681, lamium; 6002, vogeliana (BR, K); 6004, vogeliana (PRE). 3282, owariensis (P); 4058, vogeliana (K, P). Ere// 88, vogeliana (BM, K); 100, vogeliana (BM, K); 533, /amwm (K). //a// 1, kirungae (SRGH). Hanke 415, lamium (G, Z). Fabrigues 3306, lamium (P). Hansen 713, lamium (C); 937, kirungae (C). Fatten e FaoVn 77/919, kirungae (BR, US, WAG). Harley 678, lamium (K); 1631, owariensis (K). Fairbairn 1934, owariensis (MO). Harris 1 16, kirungae (DSM). Fanshawe Fl 1642, kirungae (FHO). Harris & Mwasumbi 2462, madagascariensis (DSM). Farquhar42, lamium (K). //arm eft Pofj 3227, pubescens (DSM); 3274, kirungae (DSM, US). 1 Farron 4085, vogeliana (P); 4582, vogeliana, //eaa 16, owariensis (K); 60, lamium (K). Fay 1844, /am/Mm (K); 6055, /am/Mm (K, MO). Hedin s.n., owariensis (P). Federicq 8116, kirungae (BR). Hedren, Iversen, Mziray & Pocs 841 18, madagascariensis (UPS). Figueiredo & Arriegasll, lamium (FHO, K, LISC); 32, vogeliana (K); 33, Hendrickx 77, owariensis (BR); 4067, owariensis (PRE); 7226, owariensis vogeliana (FHO, K, LISC); 66, vogeliana (K). (BR). 110 K. SIDWELL Hepburn 71, lamium (K). Leac/z /Yv/;z# s.n.. lamium (K). LISC, MO, PRE, UPS, WAG, YA); 10628, vogeliana (K, MO, WAG, Iversen & Steiner 86667, madagascariensis (UPS). YA). Jackson 1023, madagascariensis (BM); 1 144, c/ grottanellii (BM, MO). Lejoly 2281, kirungae (BR); 84/409, kirungae (BR). Jacques-Felix 1246, owariensis (K, P); 1 188, /am/w/n (P); 2600, Lely 129, owariensis (FHO, K); 469, lamium (K); 652, lamium (MO). owariensis (P); 8087, /amiM/rz (K, YA); 9135, vogeliana (K, YA); 9136, Leonard 771, lamium (MO); 1678, kirungae (BR). /am/Mm (P, YA). Letouzey 1 178, madagascariensis (H); 3561, owariensis (P, YA); 5472, Jaeger 7770, lamium (K). ye/frey 109, vogeliana (K). (P). Johnson 965. lamium (K). Lewalle 795, kirungae (BR); 3575, kirungae (BR, NY); 3640, kirungae Johnston s.n., lamium (K). (BR, G); 3778, kirungae (BR, NY); 3783, madagascariensis (BR, G, yo/i?s 127, owariensis (K). SRGH); 3795, owariensis (BR); 5613, /am/w/w (K); 5740, kirungae Jongkind, Schmitt & Abbiw 1799, owariensis (K). (BR). Jordan 375, lamium (K). 2989, /am/urn (PRE). /am/Mm /am/w/n (S, 258, 1 owariensis Jungner 181, (S, UPS); 229, (UPS); 248, UPS); , /a/n/w/n (K, MO, WAG); 344 , (K, PRE). vogeliana (UPS). Longfield 21, owariensis (BM); 66, kirungae (BM, US). Juniper, Jefford & Mgaza 256, owariensis (MO). LOMJS 118, vogeliana (WAG); 1307, c/ vogeliana (BR, MO); 1672, Kahurananga, Kibuwa & Mungai 2631, owariensis (M, UPS). owariensis (BR); 2077, vogeliana (WAG); 2161, 1 3 1 1 Kassner ( )2908, vogeliana (E, H, Z); 3, kirungae (H). owariensis (BR, PRE); 10839, madagascariensis (C); 10841, c/ Katende 831, vogeliana (BM); K522, kirungae; 969, lamium (MO); vogeliana (BR); 12194, madagascariensis (BR, C, MO); 12350, K2503, kirungae (MO); 2824, vogeliana (MO). owariensis (BR); Kaudern s.n., pubescens (S). Loveridge 25, vogeliana (K); 26, madagascariensis (MO). Kaunda & Usi 429, r/par/a (MAL). Lovef? c Love/7 2062, kirungae (MO). Kavombo 1072, owariensis (MO). Lowe 1804, /aw/Mm (K); 1809, /am/w/n (K); 2955, owariensis (K). Keay FHI 28233, owariensis (K); FHI 28264, owariensis (K). Ludanga 1337, pubescens (C). <6 Keay FHI 20068, /am/M/n (FHO). Luxen 373, owariensis). (S Onochie FHI 20245, /am/M/rz (K). Lye 1657, owariensis (MHU); 2768, /am/w/n (UPS); 5555, kirungae Kennedy \57\, lamium (FHO, K, US); 2672, /am/m (K). (SRGH). Ke//oo? 3735, owariensis (PRE); 4374, kirungae (SRGH). M. /. B. 629, owariensis (BR). Kersting A 108, owariensis (BM). Mabberley 1403, kirungae (DSM, K). Kibuwa 5191, madagascariensis (UPS), MacGregor 265, lamium (K); 333, lamium (K); 402, owariensis (K). /ftng 39B /a/njMW (K); 1 14B owariensis (K). Magnus s.n., owariensis (H). owariensis Kitson s.n., owariensis (BM, MO, NY). Maitland \ 10, vogeliana (K); 199, owariensis (K); 207, (K); 1 /am/Km Koechlin 289 , owariensis (P). 757, vogeliana (K); 1619, (K). A>M//850, owariensis (WAG). Malaisse & Goetghebeur 242, owariensis (BR). La Crak 3122, pubescens (MO); 31 14, nparia (MO); 4083, kirungae Manktelow, Steiner & Amponsah 89, vogeliana (UPS). (MO); 4600, kirungae (MO). Mann 36, vogeliana (K); 1688, lancifolia (K); 1861, /amium (K); 1959, Lamb 66, owariensis (K); s.n., lamium (UPS). owariensis (K). Lambinon 78/321, madagascariensis (BR). Manning 67, lamium (K, MO); 398, lamium (K, MO); 441, vogeliana Lane Poo/e 392, lamium (K). (MO); 1045, owariensis (MO); 1046, /am/Mm (K, MO); 1049, Lang Brown 9, kirungae (FHO); 81, owariensis (FHO). owariensis (K, MO). La/z7o 97, owariensis (K); FHI 43460, /amzwnz (K); FHI 67539, /aw/Mm Marmo 1 14, owariensis (K); 251, lamium (K). (K). Maf/zey < Scno/z 312, /a/n/w/n (K). Lan/o < Daramola FHI 28926, owariensis (FHO, K); FHI 28969, Mbago 1030, owariensis (DSM). owariensis (K); FHI 28992, owariensis (FHO). McClintock 222, owariensis (K). La?//o <* Fagbemi FHI 64782, /am/urn (K). McEdery 7627, owariensis (BM). Law/or < //a// 659, /am/Mm (K). Mearns 2465, owariensis (US). Le TesfM 9, owariensis (BM); 1010, owariensis (FHO); 3541, owariensis Meyer 15307, deW//s (MO). (BM, P); 3557, /am/Mm (K); 5943, lancifolia (BM, BR, P); 6431, Me/Ar/e 590, /am/urn (K); 953, /amiwm (K). madagascariensis (BM, MO); 6478, ae/Wis (BM, P); 8853, Melville & Hooker 85, /am/Mm (K); 357, /am/wm (K). madagascariensis (BM, P); 9135, debilis (BM); s.n., lamium (BM); Mendonqa 680, ripana (LISC); 1219, r/paria (LISC); 2110, kirungae 9395, owariensis (BR). (LISC). REVISION OF BR/LLANTAISIA 111 Methuen 80, kirungae (LISC). Pope 1987, pubescens (MO, SRGH). Meurillon 402, lamium (K, P); 521, owariensis (P); 859, owariensis (BR); Pou/sen et al. 523, kirungae (C, FHO, K, MU). 886, owariensis (BR); 1998, vogeliana (K). Preuss s.n., owariensis (BM); 998, vogeliana (BM); 1029, owariensis Meurillon et al., 1970, /a/mwm (K); 203\, lamium (K). (BM, M). Meyer 7995, grottanellii (US); 8131, madagascariensis (US). Price <& van.v 69, deb<7/.s (K); 1 \ , debilis (K). M/c/ze/ 2370, owariensis (MO); 5522, kirungae (BR). Procter 428, owariensis (PRE). owariensis owariensis Migeod 27, (K); 359, (BM); Migeod s.n., Pwjff < Ensermu 861 107-4/6, grottanellii (ETH). owariensis (BM). P>'/ze \\,lamium (K); 109, owariensis (K); 138, /am/Mm (K, P, S). Mildbraed 3866, vogeliana (H); 4147, madagascariensis (H); 5487, Quintas 1235(88) /awiwrn (BM, Z). owariensis (BR); 5782, owariensis (H); 7028, vogeliana (H); 7126, Raimano, Matos & Figueira 254, owariensis (LISC). owariensis madagascariensis (H); 7353, (K). Rakotozafy 287, madagascariensis (P). M//ne 140, /am/Hm (W). Ratabu 17, madagascariensis (MO). lamium 1 owariensis Mocquery 9, (G, Z); 96, vogeliana (G, Z); 15, (G, Z). Rattray s.n., lamium (K). 1 1 /am/wm 1 Monoa 1681, (BM); 11718, vogeliana (BM); 1857, vogeliana flayna/ 1 1964, vogeliana (P); 20477, kirungae (K). (BM); 1 1881, vogeliana (BM). flayna/ cfc flayna/ 9719, aeW/w (P, YA); 10009, vogeliana (P, YA); 12167, Montiero 46, riparia (LISC). owariensis (P, YA); 12966, owariensis (P, YA). Montiero & Murta owariensis owariensis 198, (PRE); 205, (PRE). Reekmans 1 136, madagascariensis (BR); 1986, kirungae (MO); 5237, Montiero, Santos & Murta 205, owariensis (PRE). kirungae (C, MO); 5918, /am/Mm (K, MO, PRE); 7028, /a/mum (K, Mooney 6071, madagascariensis (ETH, K, S); 6725, grottanellii (ETH); MO); 8288, owariensis (MO, PRE, SRGH); 9060, /a/n/w/n (K, MO); 6890, grottanellii (S). 9408, kirungae (K, MO, UPS); 10009, kirungae (K, MO); 10017, Moriarty 18, kirungae (MAL). kirungae (K, MO, PRE, UPS); 10073, kirungae (K, MO, UPS); 10604, lamium lamium FHI Morton A36, vogeliana (K); Al 14, (K); 327, (BM, , kirungae (K, MO, PRE, S, UPS). GC, K, SL, WAG); K416, owariensis (K, WAG); A1032, /aw/Mm (K); Reitsma & Reitsma 1048, vogeliana (WAG). A 1643, /am/ww (K); Morton GC8213, vogeliana (K); GC9378, /amwm Reitsma, Reitsma & Louis 1880, deWfo (NY, WAG). (K). Reynders 411, kirungae (PRE). Morton cfe Gledhill SL242, owariensis (K). Richards 3108, owariensis (BM, MO, NY); 12983, kirungae (SRGH); Morton < Jarr SL2394, /am/Mm (K). 24663, madagascariensis (C, M). Mucke 257, madagascariensis (PRE). Robertson 62, lamium (BM). Mullenders 2087, /am/Mm (BR). Roberty 6590, /am/urn (G, Z); 6652, owariensis (G); 6946, /aw/urn (G, Z); Mw7/er 802, kirungae (SRGH); 2353, kirungae (LISC). 10695, owariensis (G, Z); 12613, /a/w/w/n (G, MO, Z); 13020, vogeliana Mwasumbi 1 1874, kirungae (DSM). (G); 15849, lamium (G); 15991, owariensis (G); 16148, owariensis (G); Mwasumbi & Munyenyembe 13883, pubescens (DSM). 16202, /am/Mm (G); 16210, owariensis (G, Z); 16412, owariensis (G). Nannan 33, owariensis (BM); 164, lamium (BM, BR). Robevazalia 1 1530, madagascariensis (P). Napper 1515, lamium (PRE). Robson & Fanshawe 537, kirungae (LISC, SRGH). Nemba & Mambo 622, vogeliana (MO). Robyns 151, owariensis (BR); 2098, owariensis (BR). Newberry s.n., lamium (K). Rodgers-Hall 1484, kirungae (DSM). Newbold & Harley 4289, owariensis (MO). Rosevear 41-29, /am/Mm (K); 68-31, owariensis (FHO); 82-29, /am/Mm Newman & Whitmore 574, kirungae (NY). (K). 201 owariensis /am/Mm Nkongmeneck , (P, YA). Saeger 1431, (C, MO). Nolde 221 , owariensis (BM). Sa/y 6312, madagascariensis (P). Norman 103, owariensis (S); s.n., madagascariensis (MHU Salubeni 1485, oligantha (SRGH); 1487, oligantha (MAL, PRE, SRGH); Offerman et al. 648, /am/wm (BR, K). 1489, kirungae; \8Q3, pubescens (MO, SRGH); 1 842, pubescens (MO). O&z/or FHI 35870, /omium (K). Salubeni & Kaunda 4523, pubescens (MAL). Oldeman 108, /am/wm (K, WAG). Salubeni & Tawakali 2334, riparia (MO). Onochie FHI 34064, /am/Mm (K); FHI 40410, owariensis (K); FHI 40416, Samai 579, lamium (K). lamium (K). Sanford46\l, lamium (MO); 5812, lancifolia (K). Osmaston 2491, owariensis (BM). Soft/We 953, /am/wm (G). Overlaet 544, madagascariensis (BR); 1052, madagascariensis (BR). Savory FHI 36599, owariensis (K). FHI owarten.s/.v Paget-Wilkes 594, kirungae (MO). Savory <6 feay FHI 25015, owariensis (K); 25056, (K); Pate/ < Kwatha 2856, pubescens (MAL). FH25017, owartenm (K). Paw/o 646, kirungae (BR). Saver 248, /am/ww (G, K, Z). /aw/w/n Pawefc 6978, kirungae (MO, PRE, SRGH, UPS); 10135, kirungae (MO); Scaer/a 31 12, /am/wm (P); 3284, (P). Schlieben M, LISC, S, Z); 2767, kirungae (G, S, Z); 1 17'17, pubescens (LISC, MO, SRGH); 12984, oligantha (K, LISC, 1358, kirungae (G, MAL, MHU, MO, SRGH). 4068, stenopteris (LISC, MO, PRE); 4069, madagascariensis (G, H, Pearson 2273, owariensis (BM). LISC, M, P, S, Z); 4108, kirungae (G, LISC, M, S, Z); 5323, pubescens PRE, S, Z). PeraW < /fitowa 9242, kirungae (LISC); 9435, kirungae (LISC). (G, H, LISC, M, MO, et al. /am/M/ Perrier 1066, pubescens (P); 9227, madagascariensis (P); 9261, Schmidt 2090, (K). lamium madagascariensis (P); 9416, pubescens (P); 9345, madagascariensis Scnne//6513, (K). (P). Schweich 2423, kirungae (LISC, PRE). Scott E///O? owariensis 3990a owariensis (K). Peter < TM/ey 58, owariensis (K). 3990, (BM); lamium Peyre de Fabrigues 3306, lamium (P). Scurlock 17, (K). 346, 716, owariensis (BR). PM/i/w 2836, oligantha (MO); 2846, kirungae (MO, Z); 3507, kirungae Serer 97, c/ vogeliana (BR): vogeliana; owariensis (MAL, MO). Sharland 362, owariensis (K); 1 155, owariensis (K); 1202, /aw/Mm 1736, owariensis 1802, Pierlot 146, owariensis (PRE); 1649, madagascariensis (BR). (K); 1411, (K); (K); P/fe 2069, lamium. (K); 2190, /am/Mm (K); 2658, /am/Mm (K). Sarpe B8833, kirungae (G). S/zavv 26, owariensis (K). Pimenta s.n., kirungae (LISC). S/ra lamium P/owes 2261, kirungae (LISC, SRGH); 2456, kirungae (LISC, PRE). 670, (P). Sma// 212, /am/wm (K); 243, /am/wm (K, P). Pocs et al. 8722, stenopteris (UPS). 32, (BR); 54, lamium). Po/e Evans c& Erens 1796, kirungae (BR, E). Smeyers kirungae 12 K. SIDWELL Smith s.n., owariensis (K). P); 3167, /am/urn (BM, P); 3245, owariensis (P); 3556, /amw/n (LISC, Smythe 240, owariensis (K). P). Snowden 954, kirungae (G, MO); 1847, owariensis (G). Tom? 486, r<>ar/a (LISC); 4586, kirungae; 35354, kirungae (LISC). SovaM.v 1 18, owariensis (M. W): 454, vogeliana (E, H, US). Torre 1 1 Stanton 38, owarienxis (EM, SRGH). Toussaint 23 , c/ vogeliana (BR). Stauble 952, lamium (G). Troupin 392, owariensis (BR); 395, owariensis (BM); 1442, lamium (K); S/aurfr 240, dVW//.v (G, K. NY, S, WAG). 2468, vogeliana (BR); 2986, kirungae (BR); 4580, owariensis (BR, K); 471 owariensis owariensis owariensis Stauffer4f>, kirungae (PRE, UPS, Z). 1, (BR, K); 7523, (BR); 9332, S/o/z 152. kirungae (G, M, MO, NY, S, W, Z). (BR); 9393, kirungae (BR); 10371, kirungae (BR); 1 1058, kirungae S/one 63, lamium (K); 86, lamium (K). (BR); 1 1723, kirungae (MO, US). owariensis Strid 2916, kirungae (S). Tufnell s.n., (BM). owariensis owariensis Stubbings 200, lamium (K). Tw/ey 1001, /a/n/M/n (K); 1003, (K); 1934, (K). Swarbrick 293, vogeliana (E, YA): TS2822, /am/Mm (K). Tyres 339, pubescens (SRGH). /a/n/Mm .Svnse 31395. kirungae (BM, MO, S, US). (//or FHI 30224, (K). Ta/fcof s.n., /awi/H/N (K, Z); 981, /am/ww (BM, K, Z); 993, owariensis (K); (7s/ c Kaunda 606, pubescens (MAL). 1393, owariensis (MO); 1912, owariensis (MO); 2000, lancifolia (BM. van der Ben 648, owariensis (BR); 1 120, owariensis (BR, SRGH). K). van Jer Laa/i 986, debilis (WAG), Talbot & Talbot s.n., owariensis (K); 1270, owariensis (K). van der Ve&en 10350, kirungae (BR). /a/m'H/n 7rt/on 1 172, lamium (BR). van //arfen 192, (H, K, WAG), Tav/or s.n., madagascariensis (MO); 1660, lamium (BM, MO, S); 2477, van Meer 303, owariensis (BM, MO, WAG). owariensis (BM, MO); 2516, owariensis (BM); 3156, kirungae (BM, Vanden Hondt 58, kirungae (BR). MO, S): 3203, owariensis (BM); 3284, owariensis (S); 3327, Vanderijst s.n., lamium (BR). madagascariensis (BM, MO); 3384, owariensis (BM). Verdcourt 1674, kirungae (PRE); 1683, vogeliana (K). TV/era c6 Kaunda 313. pubescens (PRE). Vermoesen 1 852, c/ vogeliana (BR); 1 854, owariensis (BM). Thijsen 276, lamium (WAG). Voge/ 17, lamium (K). TTioen 4829, pubescens (BR). Vollesen 2392, pubescens (C). Thomas, A.S. 1595. owariensis (BM); 1768, owariensis (BM); 2845, Vbornoeve 726, owariensis (WAG). owariensis (MO); 2952, owariensis (MO, PRE, YA). Vw;y 330, madagascariensis (WAG). rnoma.?, D.W 1722, /am/wwi (K); 2091, /am/H/n (K); 2096, /am/urn (S); Watmough 707, pubescens (SRGH). 2320, /am/M/n (K); 2502, /a/w/wm (BM, MO, NY); 2609, vogeliana (MO, Watt 7384, vogeliana (BM). YA); 2762? /am/urn (W); 2775, lancifolia (MO); 2845, owariensis We/w/tec/2 5149, owariensis (BM); 5150, owariensis (BM); 5182, (MO); 2935. /am/Mm (K); 2952, owariensis (K, MO, PRE); D.W 3790, owariensis (BM, G); 5205, lamium (BM). kirungae (MO); 4381. owariensis (MO, YA); 7149, owariensis (MO); Wn/re 8520, owariensis (FHO); 13908, kirungae (FHO). 9005, owariensis (MO). Whyte s.n., lamium (K). Thomas, D.W. & Madeod 5 1 06, owariensis (MO); 5145, owariensis WW 3566, kirungae (NY). (MO). Williams s.n., owariensis (US); 33, kirungae (MO); 216, kirungae (LISC, Thomas, D.W cfc Mamto 4235, /a/n/wm (K, MO, PRE). SRGH). Thomas, D.W., Mambo & Nemba 4987, owariensis (MO, NY). Winkler 505, vogeliana (Z). Thomas, D.W. & Nemba 4047, /am/H/n (MO); 4048, vogeliana (K, MO, Wollaston s.n., owariensis (BM). YA). Wr/g/tf 2463, /aw/wm (FHO). 1 TTzomas, N.W. 102, owariensis (K); 2034, owariensis (K); 2285, Wy/a 679, /am/Mm (BM). owariensis (K). Yamada 261, lamium (K). Thompson & Rawlins 1460, lamium (MO). Kzfes 35, owariensis (K). 77zw/in < Mzoro 2904, kirungae (UPS); 3181, kirungae (MO, UPS). Young 651, kirungae (BM). Tindall 29, owariensis (K). Zenjfeer 286, owariensis (C, MO, US, Z); 396, /anu'wrn (C, G); 513, 7In/ey 2639, pubescens (LISC, MO, SRGH). vogeliana (K); 1 104, /am/wm (BM, E, K). Tisserant 1352, /am/ww (P); 2346, /am/Mm (BM); 2743, c/ vogeliana (BM, Zenker & Staudt 520, owariensis (S, Z). REVISION OF BRILLANTAISIA 113 SYSTEMATIC INDEX Accepted names are in roman, and synonyms in italic; new names and principle references are in bold. Belantheria Nees 83 madagascariensis T. Anderson ex Lindau 83, 104 .ioyauxii.ien.iu Heine 101 belvisiana Nees 83, 90 mahonii C.B. Clarke 92 spicata Lindau 104 P. Beauv. Brillantaisia 83 majestica Wernham 1 04 stenopteris Sidwell 84, 89, 90, 91 alata T. Anderson ex Oliv. 90 molleri Lindau 95 subcordata De Wild. & T. Durand 97 anomala Lindau 84 nitens Lindau 92 subcordata var. macmphylla de Wild. & Th. Dur. bagshawei S. Moore 104 sensu Agnew 93 97 bauchiensis Hutchinson & Dalziel 92 nyanzarum Burkill 92 subulugurica Burkill 93 borellii Lindau 106 oligantha Milne-Redhead 84, 86, 88, 89 talbotii S. Moore 101 cicatricosa Lindau 95, 105 owariensis P. Beauv. 83, 84, 90 thwaitesii Cramer 106 cicatricosa var. kivuensis Mildbr. 93 sensu Hook. 97 ulugurica Lindau 93 debilis Burkill 83, 101, 102, 103 palisotii Lindau 97 verruculosa Lindau 104 dewevrei De Wild. & Th. Dur. 92 panda T. Anderson 90 vogeliana (Nees) Benth. 84, 95, 96 didynama Lindau 106 patula var. welwitschii Burkill 92 Hygrophila 68 eminii Lindau 97 preussii Lindau 95 pubescens (T. Anderson ex Oliv.) Benoist 84 fulva Lindau 106 pubescens T. Anderson ex Oliv. 83, 84, 85 thwaitesii (Benth.) Heine 106 grandidentata S. Moore 93 pubescens var. riparia Vollesen & Brummitt 86 Leucoraphis Nees 83 grottanellii Pichi-Sermoli, 83, 103. 105 pubescens var. rutenbergiana (Vatke) Benoist 84 lamium Nees 97 hirsuta T. Anderson 105 riparia (Vollesen & Brummitt) Sidwell 83, 86, 87 vogeliana Nees 83, 95 kirungae Lindau 84, 93, 94 rutenbergiana Vatke 84 Plaesianthera (C.B.Clarke) Livera 68, 106 lamium (Nees) Benth. 83, 97, 98 salviiflora Lindau 90 thwaitesii (C.B.Clarke) Livera 106 lancifolia Lindau 83, 99, 100, 101 schumanniana Lindau 95 Ruelliola Baillon 83 leonensis Burkill 92 soyauxii Lindau 95 grevei Baillon 83, 84 Bull. mil. Hist. Miis. Loml. (Bot.) 28(2): 1 15-150 Issued 26 November 1998 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 6. Genera [Q] R-Z, and an update of current names for non-geniculate Corallinales WILLIAM J. WOELKERLING Department of Botany, La Trobe University, Bundoora, Victoria 3083, Australia GEORGE W. LAWSON, JAMES H. PRICE AND DAVID M. JOHN Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD WILLEM F. PRUD'HOMME VAN REINE Research Institute Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden, The Netherlands CONTENTS Introduction 1 15 Species list 1 17 An update of current names for non-geniculate Corallinales reported from West Africa 129 Numerical list of references 141 References ... 143 SYNOPSIS. This paper assembles and, so far as is possible without extended field and herbarium studies, examines critically Africa. whole the validity of records of marine and brackish-water Rhodophyta (Florideae) for the western coast of tropical The 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, 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. All names used for non-geniculate Corallines in earlier papers in the series are updated. More statements of distri- INTRODUCTION arranged alphabetically. generalized bution follow the specific country list. Complete distribution patterns require a scan of records under all names by which a The area dealt with in this final of the series is identical with that part species is known for this or adjacent areas. Hence, generalized covered in earlier Price et al., 1978, parts (Lawson & Price, 1969; distribution statements are included verbatim since it is not 1986, 1988, 1992; John et al., 1979, 1994; Lawson et al., 1995). always clear for precisely which countries within the area they Relevant names and their earlier and country employed equivalents, establish records. In all cases, numbers within parentheses the names of island are either listed in the or both groups, legend after the names refer to corresponding numbers in the refer- listed and shown on the in 1 . Genera with the initial letters map Fig. ences. In the present reference list, for ease of readjustment and constituent are listed alphabetically. [Q-]R-Z species from other parts, references have not been renumbered but Each main in the first of this consists of: entry part paper simply omitted or added and additionally numbered appro- lists references are the priate. However, of only partially (i) The major bold heading, representing currently-accepted between different of the overall list since name and authorities. interchangeable parts had a different Presenta- are in brackets some earlier parts numbering system. (ii) Subsidiary italicized headings. These square the in the tion of the references follows that from previous part and essentially subdivide the overall entry. They represent authors and in having first a numerical sequence presenting only different ways in which the species has been referred to past dates, followed by separate listing of the full references in publications for the area. Incorrect citations from past litera- order. and expeditionary sources, as ture have been maintained in these subsidiary heads so that alphabetical Manuscript well as works in are also included in the there shall be no doubt as to which record we attribute to which currently press was 'References'. species or lower taxon level; only when clarification notes were in cases and in which (iv) Additional qualifying required many required have changes been made in subhead citation, entries or individual of entries notes. may be found below whole parts case an explanation is given in intermediary or terminal refer. References in the notes are island to which they specifically (iii) The distributional data, with countries and groups The Natural History Museum, 1998 116 W.J. WOELKERLING ET AL. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 117 cited reference number '= by when they contain species records Note. Noted by Prud'homme van Reine (598) as tetrasporophyte of and by author name, date of publication, and sometimes page Atractophora hypnoides" on basis on an abstract by Maggs et al. (1983) and a numbers (after colon) when they do not. note by Irvine (273:35). Species nomenclature has been revised as far as and possible Rhodomela episcopalis Montagne the complete author citation derived from Brummitt & Powell See Halopirys incurvus (Hudson) Batters. (1992) is given for each currently accepted combination. The Rhodomela subsidiary italicized headings and any other discarded combi- lycopodioides (L.) C. Agardh nations that See Fucus Flor. Dan. require reference are included as cross-referencing lycopodioides Note. On nomenclatural the (90) record for the entries to the currently accepted names in the overall list. The purely grounds Bory Canaries would probably be attributable to Rhodomela lycopodioides. necessarily preliminary nature of all the treatments presented However, this is biogeographically extremely unlikely to be correct and the has been emphasized for each previously published part of the record may well represent a report of Halopitys incurvus (Hudson) Batters list and applies no less to this final contribution to the series. As (q.v.). indicated in previous parts, critical updating is kept firmly in Rhodomela C. mind for the whole work, although feasibility of that process pinastroides (Gmelin) Agardh See incurvus Batters. will remain the final determinant at the appropriate time. We Halopitys (Hudson) would appreciate notification of any detected errors and omis- Rhodomela pinastroides (Gmelin) C. Agardh van episcopalis sions in of the any parts. Montagne See Halopitys incun'us (Hudson) Batters. billda & Woodward) SPECIES LIST Rhodophyllis (Goodenough Kiitzing See Rhodophyllis divaricata (Stackhouse) Papenfuss. Reinboldiella J. poeppigiana (Grunow) Feldmann & Feldmann- Rhodophyllis capensis Kiitzing Mazoyer See Rhodophyllis reptans (Suhr) Papenfuss. See Ceramium poeppigianum Grunow. Rhodophyllis divaricata (Stackhouse) Papenfuss Additional record: Cape Verde Islands (639). Canaries (18;380;633;745;747). Rhabdonia decumbens (Grunow) Grunow Mauritanie (624). 1 1 1 Canaries (37;70;7 ; 3 ;375 ;493 ;598). [As Rhodophyllis bifida (Goodenough & Woodward) Kiitzing] Cape Verde Islands (37;70;131;375;598). Canaries (598). [As Meristotheca? decumbens Grunow] 'Nordwestafrika' (499). Canaries (439). Rhodophyllis gracilarioides M. Howe & W.R. Taylor [As Rhabdonia decunbens Grunow] Ghana (290;292;299;350;376;377;586). Canaries (191;227;493). Senegal (59;722). Note. For discussion on names, see Meristiella echinocarpa ( Areschoug) 'in tropical parts of the Atlantic Ocean/ (350;586). Cheney & Gabrielson for Cape Verde Island specimens. Prud'homme van - 'Tropical Africa (N. Gambia Congo River)' (598). Reine et al. (663) have re-investigated the Macaronesian algae studied by Note. Bodard & Mollion (59) indicated that this alga is known otherwise Piccone (see 439, 450, 451) and Askenasy (see 37, 38) and concluded that only from Brazil, a comment clearly now outdated. Askenasy had erroneously identified both Meristiella echinocarpa (from the Cape Verde Islands) and Meristotheca! decumbens (from Madeira, the Rhodophyllis reptans (Suhr) Papenfuss Canaries and also the Cape Verde Islands) as Mychodea schrammii. Namibia (36B;707). [As Rhodophyllis capensis Kiitzing] Rhabdonia sp. Namibia (500). See Gigartina flagelliformis (Sonder) Sender. Rhodophysema africana D.M. John & G.W. Lawson Rhodochorton sp. (217;294;352;532;586). See Audouinella. Angola Gabon (217;294;350;532;586). Additional record: Cape Verde Islands (652). - Tropical Africa (N. Gambia Congo river)' (598). Rhododiscus pulcherrimus P. & H. Crouan [As Rhodophysema sp.] Canaries (232B;598). Angola (352). Fig. 1 The coastline of tropical West Africa and the offshore islands. West the often Rio de Oro, the 1, Salvage Islands; 2, Canary Islands; 3, Western Sahara [=former Spanish Sahara, Spanish Africa](includes quoted Guinea-Bissau 8, 9, Sierra southern region of the country, but excludes Ifni); 4, Mauritanie; 5, Sene'gal; 6, Gambia; 7, [^Portuguese Guinea]; Guinee; 15, 16, Cameroun; 17,* Bioko Leone; 10, Liberia; 1 1, Cote d'lvoire; 12, Ghana; 13. Togo; 14, Benin [=Dahomey]: Nigeria; [=Macias Nguema 21, Gabon; 22,** of the 23, Biyogo, Fernando Poo]; 18, Principe; 19, Sao Tome; 20,* Equatorial Guinea [=Spanish Guinea]: Republic Congo; Ascension Island; 28. St Helena; 29, Annobon Cabinda; 24, Zaire [=Congo Republic]; 25, Angola; 26, Namibia [=South West Africa]; 27, [Pagalu]. omitted from this but are included in the list that The Cape Verde Islands, which lie immediately to the west of Dakar (Senegal), have been map species follows. = administered as Guinea. Bioko is entered where *Nos 17 (Bioko) and 20 (Spanish Guinea, Rio Muni) are now jointly Equatorial separately, appropriate, in the species list. a coastal of West Africa. Its to have **Loango, a name much used by earlier collectors such as Welwitsch, was formerly region application appears and Zaire Because far the and rockiest included much of the coastline of the Republic of the Congo (22), as well as of Cabinda (23) (24). by longest part all marine records from to the of the Loango coast lies now within the Republic of the Congo we have attributed algal Loango Congo. AL. 118 W.J. WOELKERLING ET H. Note. This species closely resembles Rhodophysema elegans (P. & Canaries (44). For further informa- Crouan ex J. Agardh) Dixon: see John & Lawson (294). [As IHalymenia clavaeformis Suhr] Masuda Ohta tion on the genus see South & Whittick (532) and & (1981). Canaries (401). [As Rhodymenia palmata (Esper) Greville] Rhodothamniella codii (Crouan) J. Feldmann Canaries (375). See Audouinella codii (P. & H. Crouan) Garbary. 1 that the Canarian material was a Note. Montagne (40 ) commented single to Webb under the name Delesseria Rhodymenia ardissonei J. Feldmann juvenile individual sent by Despreaux record was a case of mistaken Canaries (634:635:648). lactuca. B0rgesen (70) believed Montagne's identification for Rhoodymenia palmata. Possibly all these records relate to P.A. Rhodymenia caespitosa Dangeard Palmaria palmata (L.) Kuntze but there are reservations. See the earlier note Canaries (635). to P. palmata. Rhodymenia capensis J. Agardh Rhodymenia palmetta (Esper) auct. See Epymenia capensis (J. Agardh) Papenfuss. See Rhodymenia pseudopalmata (J.V. Lamouroux) P.C. Silva. Rhodymenia corallicola Ardissone Rhodymenia pseudopalmata (J.V. Lamouroux) P.C. Silva See Rhodymenia ligulata Zanardini. Angola (273:352). Canaries (13;38B;38D;108;225;226;227;229;232B;273;306B; Rhodymenia holmesii Ardissone 379;490;546;633;634;635;662;663;684;685;747). Angola (unpublished). Cape Verde Islands (38B;38D). Canaries (663). to Gabon (294?;350?;586?). Note. Many plants under this name from Angola appear correspond R. Ghana (299;350;376;377;586;695). rather closely to Rhodymenia pseudopalmetta var. pseudopalmetta [= towards the Mauritanie pseudopalmata}, but a few with narrow fronds often tapering (38B;38D;349;624). 'stolons' as rounded apices have sufficient development of apparent proc- Salvage Islands (38B;38D). known esses from the frond to show strong resemblance to plants previously Senegal (38B;38D;55;56;57;59;399;654;722). latter is now as R. pseudopalmetta var. ellisiae (Duby) Guiry. This taxon Sierra Leone (295;350;586). Ardissone. Prud'homme van Reine et placed in synonymy with R. holmesii Western Sahara (38B;38D;349). al. (663) re-examined Piccone's specimens of R. palmetto (Esper) Greville '. . . Afrique noire . . .' (59). and renamed some of them as R. holmesii, pointing out that this species was . . . '. . . Atlantique (de 1'Angleterre aux Canaries) (33). a new record for Macaronesia. In all probability, records for Namibia and '. . . in and seas.' ex warm temperate tropical (350;586). Ascension given under Schottera nicaeensis (J.V. Lamouroux Duby) 'SubtropicalAfrica [Senegal (N. of Gambia); Mauritania; former W. Guiry & Hollenberg should also be attributed here. Sahara]' (598). Zanardini - Rhodymenia ligulata 'Tropical Africa (N. Gambia Congo river)' (598). of Mauritania; former W. [Senegal (N. Gambia); I. Britanicas hasta . . .' 'SubtropicalAfrica '. . . O. Atlantico (desde Canarias) (747). Sahara]" (598). [As Rhodymenia pseudopalmetta (J.V. Lamouroux) P.C. Silva var.] corallicola Ardissone] [As Rhodymenia Senegal (59). Mauritanie (269;349;516). [As Rhodymenia palmetta Esper] Verde Islands (145). Rhodymenia linearis J. Agardh Cape Greville] Namibia (36B;707). [As Rhodymenia palmetta (Esper) was Canaries pro parte;499;547;663). Note. Wynne (36B) suggested that the general habit of this alga (70;191;252;375;390;439 that the Verde Islands 1;499). similar to Epymenia capensis (J. Agardh) Papenfuss, save tetraspor- Cape (38;252;41 sori near the angia did not occur in special proliferations but rather in rounded Ghana (153;338;537). the apices of frond segments. Stegenga et al. (707: 363-364), however, regard Mauritanie (252). between this natalensis as and differences alga andRhodymenia insignificant Senegal (123). have included all material in the latter species. 'From the English Coast to the Canary Islands . . .' (70). 'Nordwestafrika' Rhodymenia multipartita (Clemente) Montagne (499). 'Westafrika' See Gracilaria multipartita (Clemente) Harvey. (499). Note. Some specimens recorded in 439 are Rhodymenia holmesii (see Rhodymenia natalensis Kylin 663). Namibia (36B). [As Rhodymenia palmetta Greville] 'Southern Africa' (707). Angola (42). some reservations in material Note. Wynne (36B) expressed assigning Guinea-Bissau (529). to this due to the colder water conditions in Namibia to species, compared [As Rhodymenia palmetta J. Agardh] those of the eastern African coastline. Thalli are broader, with a more fan- Angola (41). stoloniferous shaped aspect, with larger tetrasporangia, and without the basal [As Rhodymenia palmetta Greville var. fusco-purpurea P.A. growths found in Rhodymenia pseudopalmetta, which R. natalensis other- wise resembles. Dangeard] Senegal (49;59;122;182). Greville Rhodymenia palmata (L.) [As Rhodymenia palmetta Greville var.] Canaries (401). Canaries (439). . . '. . . bis zu dem Kanaren Atlantischen Ocean (Skandinavien .)' [As Rhodymenia palmetta} (37). Senegal (411). 'Coast of the Gulf of Guinea'. (269). Note. Irvine (273) commented: 'British Isles to at least Morocco; Azores; the coasts of and South African records doubtful. Records '. . . from the Canary Islands and Mediterranean Sea to Canary Isles; Mediterranean to of Lawson & John Norway and Ireland.' (268). from elsewhere apply other species Rhodymenia'. noted: from the to [As Rhodymenia palmata Greville] (350,586) 'Many plants region appear correspond closely RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 119 with variety pseudopalmata, but a few of those with narrow fronds often [As Rythiphloea tinctoria (Clemente) C. Agardh] towards the rounded show some resemblance to R. holmesii tapering apices Canaries (229). Ardissone (= R. pseudopalmata variety elisiae in & (Duby) Guiry Guiry [As Rytiphloea tinctoria (Clemente) C. Agardh] Hollenberg)'. Canaries (13; 177). The record from Gabon (294,350,586) is doubtful because the specimens Salvage Islands (231). were sterile. '. . . .Atlantic Ocean (. . . African coasts . . .)' (177). Rhodymenia pseudopalmetta (all varieties) [As Rytiphloea tinctoria C. Agardh] See Rhodymenia pseudopalmata (J.V. Lamouroux) P.C. Silva and Canaries (44). Rhodymenia holmesii Ardissone. Sarcodia ceylanica Harvey ex Kiitzing Rhodymenia sp. [As Sarcodia ceylanica Harvey] Canaries (633). Senegal (122;4l'l). Guinee (529). Note. Various authors, including Silva et al. ( 1987) and Silva etal. (746), Namibia (3 12A;348). consider this variable alga to be equivalent to Sarcodia montagneana (J. Hooker & Harvey) J. Agardh. Ricardia montagnei Derbes & Solier Sarcomenia intermedia Grunow See Erythrocystis montagnei (Derbes & Solier) P.C. Silva. Cape Verde Islands (38; 191). J. Rissoella verruculosa (Bertolini) Agardh '. . . assez communes aux ties du Cap Vert'(38). Canaries (646). Note. Correct name is Platysiphonia intermedia (Grunow) Wynne. For nomenclature, see Silva et al. (746). Rytiphlaea episcopalia (Montagne) Endlicher Sarcomenia miniata J. C. See Halopitys incurvus (Hudson) Batters. (C. Agardh) Agardh [or Agardh] See Platysiphonia delicatula (Clemente) Cremades. Rytiphlaea fruticulosa Harvey Sarcodiotheca divaricata W.R. Taylor See Polysiphonia fruticulosa (Wulfen) Sprengel. Canaries (664). [Rytiphloea] pinastroides auct. Rytiphlaea Schimmelmannia bollei Montagne See Halopitys incurvus (Hudson) Batters. Canaries (227;410;502). Verde Islands Rytiphlaea tinctoria (Clemente) J. Agardh Cape (27;38;70;134;191;390;408;500;502;598;625). Note. in the citation of Verde' instead of [As Rytiphlaea tinctoria (Clemente) C. Agardh] B0rgesen (70), explaining 'Cape 'Canary' Islands stated: 'Schimmelmannia Bollei Mont, is in Engler and Canaries ( 16;26;38B;38D;7 1 ;89; 1 28A; 133; 1 9 1 ;226;227;232B;302; Prantl, Natiirl. Pflanzenfam. I, 2, 1897, p. 507, said to occur "in den 304;306B;375;392;401;439;489;517;556;584;598;634;635;648;662; Gewassern der canarischen Inseln". Dr. O.C. Schmidt, Bot... Mus..., Dahlem 663;684;717;747). bei Berlin, ... informed me that all the specimens found in the Museum Mauritanie (38B;38D;349;5 16;556;624). originate from the Cape Verde Islands.'. Islands that 1 Salvage (38B;38D;375;556;598). Gil-Rodriguez & Afonso-Carrillo (227) indicated B0rgesen (7 ) had Western Sahara (38B;38D). stated the Canaries record to be erroneous, so they listed the record but . . .' their as believed it to be absent. '. . . African coasts, Canary Islands (177). excluded the species from catalogue they et de la Loire a la De Toni (134: 1527), on the Bolle collections from St. '. . . Atlantique meridional (de Brest du sud commenting Nicolas (C.V.I.), stated: 'Species tantum sterilis hucusque reperta. ita ut de Mauritanie) . . .' (222). affmitate vix dijudicare liceat; quoad structuram haec potius ad 5. Frauen- . . .' '. . . Atlantique (de Brest aux Canaries) (190). feldii quam ad S. ornatum adpropinquare videtur'. . '. . . Atlantique (depuis Brest aux Mauritanie)' . . (33). '. . . Atlantico desde Brest a Canarias' (517). Schimmelmannia ornata Schousboe '. . . de Brest aux Canaries . . .' (89). Canaries (389). referred to for the area . record the 'From Brest to the Canary Islands . .' (71). Note. This probably represents species 'Atlantic between the Canaries and the British Isles' (668). as 5. bollei Montagne. .' '...... de Brest aux Canaries . . 1'ocean Atlantique (517). Schizymenia dubyi (Chauvin ex Duby) J. Agardh Africa (N. of Gambia); Mauritania; former W. bis an die 'Subtropical [Senegal '. . . Atlantischer Ozean, von den englischen Sahara]' (598). nordwestafrikanischen Kiisten . . .' (498;499). [As Rythifloea tinctorea] Note. The southern limit of this species appears to be Morocco (34; 118), Canaries (237). but since Schmidt did not define what he meant by North West Africa it is he was Western Sahara and/or Mauritanie. [As Rythifloea tinctorea] just possible that including is Haematocelis rubens J. (33;34). Canaries (237). Tetrasporic phase Agardh [As Rythifloces tinctores] Schizymenia obovata (J. Agardh) J. Agardh Canaries (237). Namibia (36B;348). C. J. [As Ryptiphloea tinctorea (Clemente) Agardh] Note. Seagrief (570) gave Platymenia undulata van obovata Agardh Canaries (229). in synonymy, but Silva et al. (746) suggest Schizymenia apoda (J. Agardh) name. [As Rytiphlaea tinctoria C. Agardh] J. Agardh to be the correct Canaries (547). Schmitziella endophloea Bornet & Batters tinctorea J. [As Rytiphlaea (Clemente) Agardh] Canaries (9;227;582;584). Canaries (663). Note. After studies of relevant type material. Woelkerling & Irvine ( 1 982) in the [As Rytiphlafa tinctoria (Clemente) C. Agardh] concluded that Schmitziella, typified by 5. endophloea, did not belong as a incertae Canaries (21). Corallinaceae and placed it next to the Acrochaetiaceae genus from studies of structure [As Rythyphloea tinctorea (Clemente) C. Agardh] sedis. Subsequently Pueschel (1989: 634), pit-plug and mode of cleavage, considered that Schmitziella may Canaries (229). tetrasporangial 120 W.J. WOELKERLING ET AL. belong in the Gigartinales rather than the Corallinales orAcrochaetiales.The Canaries (2;38B;38D;39;68;117;1 18;128A;191;375;392;489;499;517; taxonomic affinities of Schmitziella hence require further investigation. 547;584). information on material of this in the Woelkerling (730) provides species Congo (249;250;350;586). Thuret-Bornet herbarium in PC. Salvage Islands (38B;38D). Schmitziella sp. Senegal (38B;38D;48;55;59). 'Atlantico de Gran Bretana a Canarias' Cape Verde Islands (366). (517). '. . . la aux . . .' Note. The specimens upon which this record is based need examination in Atlantique (de Novege Canaries) (33). detail to determine their taxonomic disposition. 'From Great Britain down to the Canary Islands' (68). '. . . largement repandue dans tout 1' Atlantique boreal et dans les Schottera nicaeensis (J.V. Lamouroux ex Duby) Guiry & Hollenberg eaux tropicales' (59). Namibia (36B). 'Nordwestafrika', 'Westafrika' (499). [As Phyllophora palmettoides var. Nicaeensis J. Agardh] = [As Ginnania furcellata (C. Agardh) Montagne Halymenia Ascension (37). furcellata Agardh] et this should included Note . According to Irvine (273: 94 seq.) be under Canaries (402). Rhodymenia holmesii. [As Scinaia furcellata (Turner) Bivona var. constricta Pilger] Scinaia Annobon (Pagalii) (139;456;457;496). For a detailed treatment of the see revisionary 5c/a/a-assemblage, [As Ginnania furcellata Montagne] 271 A. Canaries (401). '. . . in mari atlantico ... ad oras . . . Africae meridionalis . . .' (318). Scinaia australis (Setchell) Huisman [As Ginnania furcellata (Turner)] Canaries (724). Senegal (408). Scinaia canaliculata J. Feldmann [As Scinaia pseudocrispa (Clemente) Wynne] Senegal (48;55;59;722). Canaries (724) la . . .' '. . . 1'aire se limite au golfe du Benin et a Mauritanie (59). Note. Nomenclature of this species has often been discussed. Dixon & former Irvine 1 of the that the of Scinaia 'SubtropicalAfrica [Senegal (N. of Gambia); Mauritania; W. ( 970) were opinion epithets forcellata Bivona-Bernardi 1 822: andS. J. 1 85 1 : Sahara]' (598). ( 232) furcellata (Turner) Agardh ( 422) are variants and S. as the correct name. How- Note. Bodard & Mollion (59: 214) referred to this species as 'Feldmann orthographic adopted forcellata ever, to Silva et al. the two have to be considered as mscr', stating (p. 198) '...n'est pas decrit...'. according (746) epithets distinct, with the result that the correct name isS. furcellata. The combination Scinaia (Pseudogloiophlea) capensis Scinaia pseudocrispa (Clemente) Wynne is therefore superfluous. Note. See earlier entry for Ginnania furcellata Montagne. Scinaia halliae (Setchell) Huisman Scinaia caribaea (W.R. Taylor) Huisman See Scinaia caribaea (W.R. Taylor) Huisman. Canaries (633;634;724). Scinaia hormoides Setchell Note. See Reyes et al. (724: 57) regarding the possibility that this record Ghana (350;586). Scinaia halliae Huisman. may represent (Setchell) - Tropical Africa (N. Gambia Congo river)' (598). Scinaia complanata (Collins) Cotton '. . . widespread in tropical seas . . .' (350;586). Canaries (17;128A;598;635;648;662;724). Scinaia (?)johnstoniae Setchell Salvage Islands (38B;38D;598). Gabon (294;350;586). Senegal (48). Ghana (350). [As Scinaia complanata (f. typus)]. Senegal (722). Senegal (182). '. . . in warm temperate and tropical parts of the Atlantic and Pacific Scinaia cottonii Setchell Oceans' (350;586). Africa - Ghana (350;586). Tropical (N. Gambia Congo river)' (598). Note. It is that the Ghanaian more than Senegal (722). possible plants may represent one entity (see 586). '. . . in warm temperate and tropical parts of the Atlantic and Pacific Oceans.' (350;586). Scinaia pseudocrispa (Clemente) Huisman or Wynne Africa Gambia - 'Tropical (N. Congo river)' (598). See Scinaia furcellata (Turner) J. Agardh. forcellata Scinaia Bivona-Bernardi Scinaia verae (C.I. Dickinson) Huisman See Scinaia furcellata (Turner) J. Agardh. Ghana (586). '. . . in tropical parts of the eastern Atlantic Ocean' (586). Scinaia furcellata (Turner) J. Agardh [As Pseudogloiophloea verae (C.I. Dickinson) Papenfuss] [As Scinaia forcellata Bivona-Bernardi] Ghana (350;434;7 18). Canaries (13;89;172;227;232B;271A;547;598). '. . . in tropical parts of the eastern Atlantic Ocean' (350). Cape Verde Islands (38). - Tropical Africa (N. Gambia Congo river)' (598). Salvage Islands (598). Note. 598 recorded this taxon with '?'. Senegal (38). [As Gloiophloea verae C.I. Dickinson] [As Scinaia furcellata (Turner) Bivona-Bernardi] Ghana (154;338;434) Senegal (722). 'SubtropicalAfrica [Senegal (N. of Gambia); Mauritania; former W. Scinaia spp. Sahara]' (598). Canaries (721). [As Scinaia furcellata (Turner) Bivona] Ghana (299;300;376;491). Annobon (Pagalii) (350;456;586). Senegal (59;182;399). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 121 Sebdenia canariensis Soler-Onfs, Haroun, Viera-Rodrfguez & Spermothamnion capitatum (Schousboe) Bornet Prud'homme van Reine See Tiffaniella capitata (Bornet) Doty & Mefiez. See Sebdenia macaronesica Soler-Onfs. Spermothamnion flabellatum Bornet Sebdenia dichotoma Berthold (J. Agardh) Canaries (646). Canaries (105;598;714;748). [As Sebdenia feldmannii] Spermothamnion gorgoneum (Montagne) Bornet Canaries (584). See Tiffaniella gorgonea (Montagne) Doty & Mefiez. Note. Boudouresque et al. (1984) list this taxon as Sebdenia feldmannii Spermothamnion investiens (P. & H. Crouan ex Schramm & Codomier (-Hulymenia dichotoma (J. Agardh) J. Agardh, = Sebdenia Vickers dichotoma Berthold). For an explanation of this, see Codomier (714). See Maze) Halymenia [Chrysymenia] dichotoma. Cameroun (350?;586?). Gabon (294;350;586). Sebdenia feldmannii Codomier Ghana (350?;377;586?). See Sebdenia dichotoma (J. Berthold. Agardh) Mauritanie (624?). Sebdenia macaronesica Soler-Onfs Senegal (55;59;722). Africa of former W. Canaries (748). 'Subtropical [Senegal (N. Gambia); Mauritania; Cape Verde Islands (748). Sahara]' (598). Africa - [As Sebdenia canariensis Soler-Onfs] 'Tropical (N.Gambia Congo river)' (598). Canaries (749). [As Spermothamnion sp.] Ghana (376). Sebdenia rodrigueziana (J. Feldmann) Codomier Note. The plants from Cameroun and Ghana were sterile and determina- Canaries (748). tion is doubtful. Cape Verde Islands (748). Spermothamnion macromeres Collins & Hervey Schmitz Seirospora interupta (J.E. Smith) Mauritanie (624). Canaries (699). Spermothamnion repens (Dillwyn) Rosenvinge Solieria chordalis (C. Agardh) J. Agardh Canaries (5;16;71;191;227;235;306B;490;598). See Solieria filiformis (Kutzing) Gabrielson. Mauritanie (624?). Solieria filiformis (Kutzing) Gabrielson '. . . Atlantique (de la Norvege aux Canaries ...)...' (33). Cape Verde Islands (713). '. . . Atlantique nord, de la Scandinavie aux Canaries . . .' (190). Gabon (586). '. . . coast of Europe southwards to the Canary Islands' (71). Ghana (586;654). '. . . N. Atlantic from Scandinavia to Canaries' (97). Senegal (722). 'W. Norway to Canaries' (711). '. . . in warm temperate and tropical parts of the Atlantic Ocean.' [As Spermothamnion repens var. flagelliferum De Notaris] (586). '. . . Atlantique de la Scandinavie aux Canaries . . .' (189). [As Solieria chordalis (C. Agardh) J. Agardh] [As Spermothamnion repens (Dillwyn) Rosenvinge var. turneri Mauritanie(349?;516?). (Mertens)] Senegal (408). Canaries (71). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. [As Spermothamnion repens var. turneri (Mertens) Rosenvinge] Sahara]' (598). '. . . Atlantique de la Scandinavie aux Canaries . . .' (189). [As Solieria chordalis J. Agardh] [As Spermothamnion repens var. variable] Senegal (38). Canaries (584). '. . . de la Scandinavie aux Canaries . . .' [As Solieria tenera (J. Agardh) Wynne & W.R. Taylor] Atlantique (189). Gabon (294;350). [As Spermothamnion repens var. variabile (C. Agardh) J. Feldmann] '. . . N. Atlantic from Scandinavia to Canaries . . .' Ghana ( 1 87;290;299;300;350;366;376;377;49 1 ;732). (97). Mauritanie (187;349;366;624;732). [As Callithamnion repens (Dillwyn) Lyngbye] Senegal (187;366;732). Canaries (401). turneri '. . . in warm temperate and tropical parts of the Atlantic Ocean' [As Spermothamnion Areschoug] (350). Canaries (71;547). 'North west Africa' (565A). [As Spermothamnion turneri (Mertens) Areschoug] Canaries 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. (133;239;499). '. . . Atlantico ab ad Gades Sahara]' (598). in oceano Helgolandia usque Hispaniae - et ad Canarias insulas . . .' Tropical Africa (N. Gambia Congo river)' (598). (133). . . Atlantischer Ozean bis zu den Kanaren' 'West Africa' (562). '. nordlicher (499). 'Nordwestafrika' . . . . (499). '. tropical West Africa .' (712). [As Agardhiella tenera (J. Agardh) Schmitz] Spermothamnion speluncarum (Collins & Hervey) M. Howe Cape Verde Islands (477). 1 1 1 Canaries ( 1 3;7 ; 9 ;227;598). Ghana (153). Ghana (350;586). Mauritanie (192;252). '. . . element Atlantique-tropical' (191). Senegal (49;59;182). Atlantic . . .' . of the Ocean '. . in warm temperate and tropical parts Note. All records of Solieria chordalis from tropical West Africa are considered doubtful. On the of nomenclatural (350:586). complexities interpretation, - Africa (N. Gambia river)' (598). see Silva et al. (746). Tropical Congo 122 W.J. WOELKERLING ET AL. Spermothamnion turner! Areschoug Spondylothamnion multifidum (Hudson) Nageli 1 1 1 1 1 9 1 See Spermothamnion repens (Dillwyn) Rosenvinge. Canaries ( 7;7 ; 28A; 33; ;227;232B;240;306B;390;584; 598;635). Spermothamnion sp. '. . . Atlantique (de 1'Angleterre aux Canaries) . . .' (33). Angola (352). '. . . Atlantique nord, de 1'Ecosse aux Canaries . . .' (190; 196). Canaries (235;489). 'British Isles to Canaries.' (711). Ghana (299;350;376;377;49 1 ;586). '. . . nell'Atlantico si estende della Gran Bretagna al Marocco, Mauritanie (624). toccando le Canarie . . .' (390). St Helena (644). '. . . English coast southwards to the Canary Islands . . .' (71). Note. There may be other representatives of the genus in Sierra Leone '...... ' NE. Atlantic , the Canaries (668). (295;350;586) different from those reported by Lieberman et al. (376) from including Ghana (cf. Spermothamnion investiens). All specimens found have been [As Sphondylothamnion multifidum Nageli] sterile so that positive identification, even to genus, has not been possible. Canaries (547). Sphaerococcus acicularis (Wulfen) C. Agardh Spongites See Gigartina acicularis (Roth) J.V. Lamouroux. The concept of Spongites adopted in this paper is that of Penrose Sphaerococcus cartilagineus (L.) C. Agardh (1996/7). the of to other See Gelidium cartilagineum (L.) Gaillon. Keys showing relationships Spongites genera of the subfamily Mastophoroideae (to which Spongites confervoides C. Sphaerococcus (L.) Agardh belongs) are provided by Penrose & Chamberlain (1993) and by and var. 6 Turner procerrimus (Esper) Woelkerling (\996b), while Woelkerling (1985) gives an account of See Gracilaria verrucosa (Hudson) Papenfuss (now Gracilariopsis the original collections upon which KUtzing (1841) based the genus Steentoft, L.M. Irvine & Farnham). longissima (Gmelin) and the six species he assigned to it. Paragoniolithon Adey et al. (1982: 12), based on f! solubile (Foslie & Howe) Townsend& Sphaerococcus corneus (Hudson) C. Agardh Adey, Goniolithon solubile Foslie & Howe) is a See Gelidium corneum sensu B0rgesen. Boykins (Basionym: heterotypic synonym of Neogoniolithon (see Woelkerling, 1988: C. Sphaerococcus coronopifolius (Goodenough & Woodward) 141). Agardh absimile & M. Afonso-Carrillo Canaries (70;71;140;184;191;207;439;499;517;584;598;635;663; Spongites (Foslie Howe) Canaries (733). 698). [Lithophyllum absimile Foslie] '. . . Atlantico (de Gran Bretana a Canarias) . . .' (517). Cape Verde Islands (7365). 'Du nord de la Grande-Bretagne aux Canaries . . .' (89). 'Golfe de Guinee' (7366). 'From Great Britain down to the Canary Islands . . .' (70). Note. Uncertainties surround the generic disposition of the type and of [As Sphaerococcus coronopifolius Agardh] certain records of this taxon from tropical West Africa and adjacent islands. Canaries (439). The species was originally described as Lithophyllum absimile Foslie & [As Sphaerococcus coronopifolius C. Agardh f. gracilior] M. Howe in Foslie (207: 7) based on material from Jamaica and was Canaries (547). subsequently transferred to Pseudolithophyllum by Adey ( 1 970: 1 2) and then [As Sphaerococcus coronopifolius (Goodenough & Woodward) to Spongites by Afonso-Carrillo (733: 98). As noted by Afonso-Carrillo (733: Greville] 97), the combination Neogoniolithon absimile (e.g., see Notoya, 1976a: 1 37, '. . . in den warmeren Teilen des atlantischen Oceans . . .' (502). 1976ft: 314; Cabioch, 1972: 272; Gil-Rodriguez & Afonso-Carrillo, 227: 36; [As Sphaerococcus coronopifolium (Goodenough & Woodward) South & Tittley, 1986: 44) has not been validated in accordance with ICBN Art. 33.2 1 see under Afonso-Carrillo Stackhouse] (see Greuter, 994); Spongites wildpretii. (733: 9 1 -93, 1 -6) an account of material in TRH (see Canaries (392;407). figs provided holotype also Woelkerling, 678: 14), but noted (p. 97) that placement mSpongites was [As Gelidium coronopifolius (Goodenough & Woodward) attended by some uncertainty since only senescent conceptacles were Lamouroux] observed. Data on the vegetative thallus of the type provided by Afonso- Canaries (44;71;401). Carrillo (733) suggest that the species most likely belongs to the Corallinaceae, [As Stackhouse] Sphaerococcus coronopifolius subfamily Mastophoroideae (see Woelkerling, 1988: 1 15, 1996ft: 237), but Canaries (227;375). the absence of data on tetrasporangial conceptacle anatomy from the type '. . . British Isles to the Canary Isles . . .' (172). precludes certain generic placement within the subfamily. Afonso-Carrillo Note. The tetrasporophyte is Haematocelis [Ethelia] fissurata P. & H. (733) also concluded that Canary Island specimens referred to absimile by Crouan (719). Lemoine (362) as well as plants deposited atTFC (Departamento de Botanica, Universidad de La Laguna, Canary Islands) [upon which some other records Sphaerococcus intricatus C. Agardh (Gil-Rodriguez & Afonso-Carrillo, 227: 36; Afonso-Carrillo, 576: 139; See the entries for intricata (C. Vickers and Gelidiopsis Agardh) Afonso-Carrillo et al., 582: 30) presumably were based] did not belong to variabilis the note to the Gelidiopsis (J. Agardh) Schmitz, especially Spongites absimile but rather to S. wildpretii (see below). latter (Price et al., 1988). Records ofLithophyllum absimilefrom the Cape Verde Islands (Lemoine, 365: 1071) and from the 'Golfe de Guinee' (Lemoine, 366) require verifica- C. Sphaerococcus norvegicus (Gunnerus) Agardh tion from examination of relevant voucher material. See Gymnogongrus crenulatus (Turner) J. Agardh. Spongites africanum (Foslie) Afonso-Carrillo, Chacana & Sanson Sphaerococcus oligacanthus Kutzing Cape Verde Islands (726). See Gracilaria dentata J. Agardh. Senegal (726). Sphaerococcus rangiferinus Kutzing [As Lithophyllum africanum Foslie] See Gracilaria dentata J. Agardh. Annobon (Pagalu) (397;455;457;500;535). Note. This name has been replaced by Gracilaria rangifera (Kutzing) Bioko (500). Piccone, according to Silva et al. (746). Canaries (353;366;537). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 123 Cape Verde Islands (6;100;353;365;366;455;535). Cape Verde Islands (210;366;598). Mauritanie (359;366). Mauritanie (349;356;359;360;366). Sao Tome (6;455;535). Senegal (6;366). Senegal (198;212;535). [As Lithothamnion fruticulosum (Kutzing) Foslie f. crassiuscula] '. . . Golfe de Guinee . . .' (366). Cape Verde Islands (2 10). [As Lithophyllum africanum Foslie f. intermedia Foslie] Note. Status and disposition uncertain; TRH neotype (Woelkerling, 678: from Brionic Adriatic Sea not studied in detail in a modern Senegal (199;211;212;678;730). 66) Islands, context; in this does not Note. Status and disposition uncertain; holotype from Cap Vert, Senegal in placement \inderSpongitesfruticulosus paper imply with the of the TRH (Woelkerling, 730) and isotype material in PC (Woelkerling, 730) not synonymy type species. Foslie f. studied in detail in a modern context; placement under Spongites africanum [As Lithothamnion fruticulosum (Kutzing) clavulata] in this paper does not imply synonymy with the type of the species. Cape Verde Islands (2 10). 678: [As Lithophyllum africanum Foslie f. truncata Foslie] Note. Status and disposition uncertain; TRH neotype (Woelkerling, 5 1 from the Adriatic Sea not studied in detail in a modern SaoTome(211;212;535). ) context; placement under Spongitesfruticulosus in this paper does not imply synonymy with the Senegal (198;2 12). type of the species. Note. Status and disposition uncertain; holotype from Cap Vert, Senegal in [As Lithothamnium Foslie] TRH (Woelkerling, 730) not studied in detail in a modern context; placement fruticulosum Mauritanie under Spongites africanum in this paper does not imply synonymy with the (354). of former W. type of the species. 'SubtropicalAfrica [Senegal (N. Gambia); Mauritania; [As Lithophyllum (cf.) africanum} Sahara]' (598). The of which is also the of the Cape Verde Islands (598). Note. type Spongites fruticulosus, type has been studied (1985: 135-139, 23- 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. genus Spongites, by Woelkerling figs 32), who (Woelkerling, 1988) placed the genus in the subfamily Sahara]' (598). Mastophoroideae. As noted by Woelkerling (1985: 139), Kutzing's epithet [As Lithothamnion ponderosum Foslie] fruticulosum has been widely misapplied to a species with multiporate Sao Tome (134?;197;265). tetrasporangial conceptacles belonging to the Corallinaceae, subfamily Lithothamnion Foslie] [As proboscideum Melobesioideae and not to the subfamily Mastophoroideae. More recently, Sao Tome (197). Woelkerling (729) concluded that at least some of the collections to which the Porolithon [As africanum (Foslie) Foslie] epithet fruticulosus had been misapplied were conspecific with the type of Annobon (Pagalu) (139). Lithothamnion fasciculatum (Lamarck) Areschoug in J. Agardh. All speci- and its Bioko (350;586). mens upon which published records of Spongites fruticulosus from the West African are based now need to be Cameroun (350;586). homotypic synonyms region checked to determine their status and disposition. Whether true Spongites Cape Verde Islands (598). fruticulosus occurs in the West African region is uncertain. Penrose (1991) Mauritanie (349). provides an account of S. fruticulosus in southern Australia. Babbini & SaoTome(211;350;586). Bressan (753: 246) state 'non 'fruticulosa' emendavit Penrose 1981; ICBN the eastern Atlantic '. . . in warm temperate and tropical parts of 1994, art. 62.4'. Specimens from the Cape Verde Islands identified (2 10: 214) Ocean . . .' (350;586). as Lithothamnion fruticulosum f. clavulata and as L. fruticulosum f. 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. crassiuscula also need to be checked to determine their status and disposi- Sahara]' (598). tion. - Tropical Africa (N. Gambia Congo river)' (598). Spongites wildpretii Afonso-Carrillo Note. Based on examinations of the lectotype from Cap Vert, Senegal (see Canaries (633;634;657;687;733;747). Woelkerling, 678: 23-24) and specimens from the Cape Verde Islands, absimile Foslie] Afonso-Carrillo, Chacana & Sanson (726) tramferredLithophyllum africanum [As Lithophyllum Canaries Foslie (199: 3) to Spongites as delimited by Penrose & Woelkerling (1988) (362;363). Cabioch nom. and Woelkerling (1988). This concept of Spongites encompassed the genus [AsNeogoniolithon absimile (Foslie & Howe) invalid] HydrolithonFoslie. Based on further studies, however, Penrose &Woelkerling Canaries (226;227;576;582;733). (1992) concluded that Spongites and Hydrolithon should be treated as distinct Note. Afonso-Carrillo (733) based Spongites wildpretii on a series of genera based on differences in tetrasporangial conceptacle anatomy (see also collections from the Canary Islands, including ones earlier referred to Afonso- Penrose & Chamberlain, 1993;Penrose, 1996a; Woelkerling, 1996/7). Lithophyllum absimile or the invalid name Neogoniolithon absimile (see to Keats Chamber- Carrillo (pers. comm.) suggested that the species probably belongs note under Spongites absimile). Subsequently, however, & not been that Hydrolithon as delimited by Penrose, but formal transfer has lain (754: 15, 18) and Chamberlain (702: 126) concluded Spongites of Keats effected. All specimens upon which published records Spongites africanum wildpretii is a heterotypic synonym of Hydrolithon samoense (Foslie) based need to identified and its homotypic synonyms from the West African region are & Y.M. Chamberlain based on examination of European specimens to the checked to determine their status and disposition. This also applies by Afonso-Carrillo but not listed by Keats & Chamberlain (754). types and other records (listed above) of Lithophyllum africanum f. withL. intermediumandL africanumf. truncatum, established concurrently Sporolithon f. africanum by Foslie (199: 3). The types of both Lithophyllum africanum truncatum is now in a distinct family of Corallinales, the intermedium (see Woelkerling, 678: 127; 730.) andL. africanumf. Sporolithon placed Comments on the incorrect use of the (see Woelkerling, 678: 226) also come from Cap Vert, Senegal. SporolithaceaeVerheij (1993). The record under the name Lithothamnion proboscideum from Sao Tome name Archaeolithothamnium for species referable to Sporolithon referred to and (Foslie, 197: 14) pertains to material Foslie (199: 3) subsequently are provided by Papenfuss (1968: 83), Woelkerling (1988: 220) Information to material Spongites africanum (as Lithophyllum). relating Moussavian & Kuss (1990: 932-934). identified as Lithothamnion ponderosum Foslie has been presented previ- africanum (Foslie) Afonso-Carrillo ously (John et al., 1994: 68). Sporolithon Canaries (11;582;726). Archaeolithothamnion (-ium) africanum Foslie] Spongites fruticulosus Kutzing [As Canaries [As Lithothamnion fruticulosum (Kutzing) Foslie] (6;70;139;191;205;212;227;361;363;365;366;493). Verde Islands (366). Canaries (6). Cape 124 W.J. WOELKERLING ET AL. Note. The combination Sporolithon africanum was first validly published Senegal (38B;38C;38D;55;59;555;556;722). by Afonso-Carrillo (II: 142). Wynne (1986: 2258), apparently unaware of Sierra Leone (30;350;586). the same the Afonso-Carrillo paper, subsequently proposed combination, Western Sahara (38B;38D). crediting Tomita, who had first used it in an unpublished thesis in 1976. The 'Atlantic Ocean (. . . African and American coasts, Canary Islands combination also appeared in an unpublished thesis of Oliveira in 1977. The holotype of Sporolithon africanum (from the Canary Islands) is 'Atlantique tempere et tropical (cotes d'Europe, d'Afrique . . .)' divided between PC (Woelkerling, 730) and TRH (Woelkerling, 678: 23). (190). Afonso-Carrillo (11: 142) examined the type but provided no details, and '...... (cotes ...... )....' (196). Townsend (1995) included comments on type material in a PhD thesis, but a Atlantique, tropical d'Afrique has to '. . . in warm and of the Atlantic ocean . . .' published account of type material in a modern context yet appear. temperate tropical parts Other published records of the species from the West African region also (350;586). of relevant voucher require verification through re-examination specimens. 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. al. Lemoine treated S. As noted by Afonso-Carrillo et (726: 133), (353: 146), Sahara]' (598). (as Archaeolithothamnium} as a heterotypic synonym of africamtm 'Widely distributed in tropical to temperate regions' (707). Lithophvlliim africanum Foslie (199: 3) [see account above of Spongites 'Oceano Atlantico (desde Gran Bretana hasta Sudafrica . . .)' (751). africanum (Foslie) Afonso-Carrillo, Chacana & Sanson]. [As Spyridia filamentosa Harvey] Spyridia aculeata (Schimper) Kutzing Canaries (44;254;306;40 1 ;439;547). See Spyridia hypnoides (Bory) Papenfuss. Sao Tome (261;263;265;269). 'Du sud de la Grande-Bretagne aux Canaries . . .' (89). Spyridia armata Kiitzing [As Spyridia filamentosa f. Rfriabilis (Clemente) J. Agardh] See Spyridia hypnoides (Bory) Papenfuss. Canaries (25). Spyridia clavata Kiitzing [As Conferva pallescens Bory] Angola (41;42;292;535). Canaries (90). Congo (249;250;292). [As Hutchinsia filamentosa (Wulfen) C. Agardh] Gabon (249;586). '. . . Atlantico ab Anglia ad Teneriffam . . .' (20). that of the Gambia (292;350;535;586). Note. Lawson & John (350;586) remarked many early reports of this from offshore islands in the Gulf of Guinea have been Ghana (292;350;377;586). species discounted by Steentoft (535) who believed them misidentifications of Mauritanie (624). Spyridia hypnoides (q.v.) They therefore placed all reports of S. filamentosa Sao Tome (251;265;292;316;318;323;350;535;586). from Sao Tome under S. hypnoides until re-examination and (if needed) Senegal (50;55;59;99;249;292;350;722). reaffirmation of the original determination is possible. Senegambia (27;61;133;296;410;535). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. Spyridia hypnoides (Bory) Papenfuss Sahara]' (598). Canaries (13;38B;38D;128A;226;227;306B;633;634;635;662;663; - Tropical Africa (N. Gambia Congo river)' (598). 751). [As Spyridia clavulata Kutzing] Cape Verde Islands (38B;38D;598;652;683;713). Angola (352). Gabon (586). [As Spyridia clavifera J. Agardh] Ghana (299;300;346;491;586). "... ad oras Senegambiae . . .' (25). Mauritanie (38B;38D). Principe (586). Spyridia clavifera J. Agardh Salvage Islands (38B;38D;598). See Spyridia clavata Kutzing. Sao Tome (586). Spyridia clavulata auct. Senegal (38B;38D;722). Africa of Mauritania; former W. See Centroceras clavulatum (C. Agardh) Montagne and Spyridia 'Subtropical [Senegal (N. Gambia); clavata Kutzing. Sahara]' (598). - Tropical Africa (N. Gambia Congo river)' (598). filamentosa Spyridia (Wulfen) Harvey 'Oceano Atlantico (desde las costas francesas hasta Senegal . . .' Note on authorities. De Toni (133: 1427-1429) placed Conferva palle- (751). scens Bory (90: 306, pl.V, figs 2A,B,C) in synonymy with Spyridia [As Spyridia hypnoides (Bory ex Belanger) Papenfuss] filamentosa. From the figures provided, there seems little reason to challenge Ghana (292;377). this. The problem then arising is that of the earliest specific epithet for the Gambia (296). taxon. Wulfen's Cryptogamia Aquatica, with the recognition of Fucus Mauritanie (349). filamentosus, was dated 1 803. Bory's work, dated the same year, states on the title page 'Germinal, An XI'. It will be necessary to establish the priority of Senegambia (296). these two works to assess correct nomenclature. Western Sahara (349). Annobon (Pagalii) (456;457). [As Spyridia hypnoides var. typica (Bory ex Belanger) Papenfuss] Ascension Island (37). Mauritanie (624). Canaries (8;13;16;38B;38C;38D;70;71;108;128A;133;177;19:226; [As Spyridia hypnoides (Bory in Belanger) Papenfuss] 227;229;230;235;236;237;295;304;306B;379;390;392;489;490;493; Canaries (745). 546;547;555;556;584;598;633;634;635;662;663;684;745;747;751;752). [As Spyridia aculeata (Schimper) Kutzing] Cape Verde Islands (598;639). Canaries (72;108;191;235;304;351;535;556). Ghana (290). Cape Verde Islands (37;41;100;183;191;259;535;556;639). Mauritanie (349;516;555;556;624). Guinea-Bissau (529). St Helena (644). Principe (535). Salvage Islands (38B;38C;38D;231;375;555;556;598). Salvage Islands (556). Sao Tome (25 1 ;265). SaoTome(93;535). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 125 Senegal (59;122;196;529;556). Cape Verde Islands (38; 191). '. . . Atlantique du Maroc au Senegal . . .' (196). Stichothamnion cymatophyllum B0rgesen [As Spyridia aculeata (Schimper) Kiitzing f. inermis] Canaries (16;71;128A;191;225;227;253;281;306B;583;598;633; Principe (93;535). 634). [As Spyridia aculeata (Schimper) Kiitzing f. typica, f. aculeata] [As Stichotamnion cymatrophilum B0rgesen] Sao Tome (93;535). Canaries (235). [As Spyridia hypnoides (Bory) Papenfuss var. disticha (B0rgesen) [As Stichothamnium cymatophyllum B0rgesen] G.W. Lawson & D.M. John f. inermis G.W. Lawson & D.M. John] Canaries (229). Canaries (292). Note. Feldmann (191) considered this alga to constitute a remarkable Gabon (294;350). palaeoendemic in the Canaries flora '. . . C'est, en effet, le seul genre Ghana (292;350;586). endemique parmi les algues des Canaries. Ses affinites possibles avec d'autres Senegal (292). genres de Rhodomelacees sont assez obscures et ne permettent pas de deceler [As Spyridia aculeata Kiitzing] son origine'. Angola (42). Streblocladia camptocladia (Montagne) Falkenberg Cape Verde Islands (38;42;49;145;259). Namibia (36B;348;707). Senegal (42). [As Orcasia pulla Simons] 'De Cadiz au Senegal . . .' (38;89). Angola (3 12A). 'In oceano atlantico tropico.' (320). Namibia (3 12A;5 12). '. . . et atlantico tropico' (318). 'Morocco to Senegal' (97). Streblocladia collabens Falkenberg [As Spyridia aculeata (Schimper) J. Agardh [or Kiitzing] var. disticha Cape Verde Islands (598). B0rgesen] 'Subtropical Africa [Senegal (N. of Gambia); Mauritanie; former W. Canaries (71;295). Sahara]' (598). Senegal (59;295;529). [As 'Streblocladia''] [As Spyridia aculeata (Schimper) Kiitzing var. hypnoides J. Agardh] Senegal (529). Senegal (55). Streblocladia corymbifera (C. Agardh) Kylin [As Spyridia aculeata (Schimper) Kiitzing var. hypneoides Kiitzing] Namibia (36B;707). Mauritanie (186;535;542). [As Polysiphonia corymbifera (J. Agardh) Harvey] Senegal (55;122;186;535). Namibia (167;348;523). 'S. Atlantic shores of Europe, Morocco to Senegal . . .' (97). 'Morocco to Senegal' (97) Streblocladia fasciculifera (Kiitzing) Falkenberg [AsSpyridia aculeata (Schimper) J. Agardh var. hypnoides J. Agardh] Namibia (348). Senegal (59). Streblocladia glomerulata (Montagne) Papenfuss [As Spyridia aculeata (Un. itin.) Zanardini] [As Streblocladia neglecta Schmitz] Cape Verde Islands (141 A). Senegal (122). [As Spyridia armata Kiitzing] 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; former W. Canaries (439). Sahara]' (598). Senegambia(318;320). [As Spyridia aculeata var. typica B0rgesen] Streblocladia neglecta F. Schmitz Canaries (71). See Streblocladia glomerulata (Montagne) Papenfuss. [As Spyridia filamentosa ] Suhria vittata (L.) J. Agardh Sao Tome (261;263;264;265). Namibia (36B;169;348;453;500;569;707;716). [As Spyridia insignis (J. Agardh) J. Agardh] . . .' '. . . im siidlichen Teile des atlantischen Oceans (502). Cape Verde Islands (37;138;191;598;625). meridionalis . . .' '. . . praesertim ad litoraAfricae vulgatissima (139). Senegal (408). [As Fucus vittatus L.] 'pantropical' (625). Ghana (271). '. . . meridionale . . .' (38). Afrique The record for Ghana Danish Guinea) is a - Note. (as probably Africa (N. Gambia river)' (598;625). 'Tropical Congo misdetermination, since, rather surprisingly, no material having been traced, Bindera J. [As insignis Agardh] the attribution is by nomenclatural equivalence alone. Original P.E. Isert Cape Verde Islands (528). material from Ghana, perhaps destroyed in a fire in 1807, requires re- if examination for certainty, although it is difficult to see how Suhria, present, Spyridia insignis (J. Agardh) J. Agardh could have been confused easily with other taxa. See Spyridia hypnoides (Bory) Papenfuss. Taenioma macrourum Thuret Spyridia sp. See Taenioma nanum (Kiitzing) Papenfuss. Angola (500). Canaries (66;214). Taenioma nanum (Kiitzing) Papenfuss Mauritanie (349). Bioko (346;350;586). Senegal (399). Canaries (17;341). . .' '. . . open shores in West Africa . (347A). Cape Verde Islands (639). Sierra Leone Stenogramme interrupta (C. Agardh) Montagne ex Harvey (295;350;432;586;598). Africa Gambia - river).' (598). Canaries (598;635). 'Tropical (N. Congo in warm and seas.' '. . . widespread temperate tropical Cape Verde Islands (37 ;1 15;172;598). probably [As Stenogramme interrupta Montagne] (350;586). 126 W.J. WOELKERLING ET AL. [As Taenioma macruorum Thuret] Tenarea corallinae P. & H. Crouan Canaries (1 1;227;489;584). See Lithophyllum corallinae (P. & H. Crouan) Heydrich. [As Taenioma macrourum Thuret] Tenarea hapalidioides (P. & H. Crouan) W.H. Adey & P. Adey Canaries (375:430:543). See Titanoderma hapalidioides (P. & H. Crouan) J.H. Price, D.M. Taenioma C. [As perpusillum Agardh] John & G.W. Lawson. Canaries (71). [As Taenioma perpusillum J. Agardh] Tenarea irregularis (Foslie) Me. Lemoine Sierra Leone (336:339). See Lithophyllum irregulare (Foslie) P. Huve ex Steentoft. The following are additional references: Taenioma perpusillum (J. Agardh) J. Agardh [As Tenarea irregularis (Foslie) Me. Lemoine] 1 Canaries ( 13;38B;38D;7 ; 108;226;227;304;589;634;635;684;747). Canaries (191;362;363). Cape Verde Islands (598). Sao Tome (362;363). Cote d'lvoire (287;288;295;296;350;586). [As Lithophyllum irregularis Foslie] Ghana (153;288;295;338;344;350;432;487;491;537;586). Sao Tome (206;212). Liberia (129;287;288;350;586). Salvage Islands (38B;38D;598). Tenarea polycephalum (Foslie) W.H. Adey . . .' '. . . Atlantique tropical et subtropical (184). See Titanoderma polycephalum (Foslie) Woelkerling, Y.M. Cham- seas . . .' Silva. '. . . probably widespread in warm temperate and tropical berlain & PC. (350:586). - Tenarea tortuosa (Esper) Me. Lemoine Tropical Africa (N. Gambia Congo river)' (598). Mauritanie (529). 'Virtually worldwide in tropical and subtropical seas . . .' (707). [As Lithophyllum tortuosum (Esper) H. Huve] [As Taenioma perpusillum J. Agardh] Canaries (107). Canaries (19 1;556). Mauritanie (349). Ghana (336). Note. According to Babbini & Bressan (753: 192), this species is an Salvage Islands (556). 'espece mediterraneene' (thus a mediterranean endemic species). Therefore, '. . . widespread in warmer seas.' (336). as also noted by Lawson & John (586: 2 1 0), these records appear doubtful but '. . . seems to be in warmer seas.' (63). widely spread need to be checked against the relevant specimens. Esper's epithet tortuosum Taenioma J. [As perpusilla (J. Agardh) Agardh] has been misapplied to a different species (see Huve, 272; Woelkerling et al., Canaries (684). 1985; Woelkerling, 730) commonly referred to Lithophyllum. Tenarea [As Taenioma perpusilla] tortuosa, the type and only known true species of Tenarea, is known with from the eastern Mediterranean 1988: Canaries (2 14). certainty only (Woelkerling, 109; Athanasiadis, 1995: 656). The earlier cross reference Note. Cribb (113) commented '. . . T. nanum has been distinguished from toLithophyllumcristatum f. crassa Hauck (John et al., 1994: 61) does not involve T. perpusillum (J. Agardh) J. Agardh mainly on the basis of its possession of Meneghini (Lloyd) records from the West African but rather the of two terminal hairs, T. perpusillum having three. However, the number of hairs any published region opinion Foslie that f. crassa, based on Melobesia crassa (see is variable in some reported populations of the two species, and Hollenberg (1898: 15) Lloyd 730), be a of Tenarea tortuosa. (1967) has questioned the justification of recognizing T. nanum as distinct Woelkerling, might heterotypic synonym from T. perpusillum? Thallium-Ionium claviferum J. Agardh Taenioma spp. Ghana (290;299;300;350;586;590). Mauritanie (624). '. . . in warm temperate and tropical seas' (350;586). - Tropical Africa (N. Gambia Congo river)' (598). Tayloriella tenebrosa (Harvey) Kylin [As Thamnoclonium sp.] Namibia (348;707). Ghana (292). Note: be with Thamnoclonium dichotomum J. Tayloriella virgata (C. Agardh) Papenfuss May conspecific (J. Agardh) See Scott et al. (1984) and Womersley (712). Angola (352). Agardh. Namibia (352;487). Thuretella schousboei (Thuret) F. Schmitz [As Polysiphonia virgata (C. Agardh) Sprengel] Canaries (8;38B;38D;227;598;745). Namibia (167:348). Salvage Islands (38B;38D;598). Note. Wynne (36B) has argued for the retention of Tayloriella virgata under Polysiphonia virgata (C. Agardh) Sprengel but as the above records Tiffaniella capitata (Bornet) Doty & Menez were omitted from our earlier publication (Lawson et al., 1995) they are Canaries (240;598;635;747;756). included here for completeness. Mauritanie (624?). '. . . O. Atlantico del N. hasta Canarias)' (747). Tenarea Bory (desde Europa [As Tiffaniella capitatum (Bornet) Doty & Menez] Woelkerling et al. (1985) concluded that Tenarea included only one Canaries (663). known species (T. tortuosa Bory) and that most other species [As Spermothamnion capitatum (Schousboe) Bornet] referred to the genus belonged to Lithophyllum or Titanoderma (see Canaries (71;189;191;227). entry for Titanoderma below). Further comments on Tenarea are '. . . Tingin Africae borealis . . .' (133). provided by Woelkerling (1988: 106-109) and Athanasiadis (1995). Note . For data on the genus and species of Tiffaniella, see Gordon (240). Tenarea adhaerens Me. Lemoine Tiffaniella gorgonea (Montagne) Doty & Menez See Neogoniolithon hirtum (Me. Lemoine) Afonso-Carrillo. Canaries (597;747). Tenarea confinis (P. & H. Crouan) W.H. Adey & P. Adey '. . . O. Atlantico Oriental y Occidental' (747). See Titanoderma confinis (P. & H. Crouan) J.H. Price, D.M. John & [As Tiffaniella gorgoneum (Montagne) Doty & Menez] G.W. Lawson. Canaries (240;598). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 127 Cape Verde Islands (240;597;598;652). evolutionary lines are present within Lithophyllum sensu lato (i.e. Salvage Islands (598). sensu Woelkerling & Campbell, 1992 and Woelkerling, 1996a) [As Callithamnion gorgonium Kiitzing] without complicating the matters of species placement and naming '. . . in atlantico ad Africanum . . .' (27). as well as specimen identification, it is possible to recognize two [As Callithamnion gorgoneum Montagne] subgenera, as has been done by some authors (e.g. Rosenvinge, Canaries (500). 1917; Lemoine, 363; Newton, 1931). 'In oceano atlantico ad orasAfricae. Specimen dedit amic. Montagne.' Titanoderma byssoides (Lamarck) Y.M. Chamberlain & Woel- (320). kerling Cape Verde Islands (37;408;410;500?;597). See Lithophyllum byssoides (Lamarck) Foslie [As Spermothamnion gorgoneum (Montagne) Hornet] The following are additional records: Canaries (38B;71;97;133;191;226;227;493). [As Lithophyllum byssoides (Lamouroux) Heydrich] Cape Verde Islands (713). Cape Verde 'islands (38B;71;133). [As Spermothamnion gorgonium Hornet] Mauritanie (6). 'Plante des Antilles et du Cap Vert, nouvelle pour les Canaries' Salvage Islands (38B). (547). Senegal (547). 'Plante des Antilles et du Cap Vert, nouvelle pour les Canaries' Titanoderma Nageli (547). Titanoderma is considered here to be a heterotypic synonym of Note. The neotype mentioned under Lithophyllum byssoides (John et Lithophyllum; further comments appear under Lithophyllum (John al., 1994: 60) has been superseded with the discovery of original Lamarck et al, 1994: see also & (1995: 270), 59); Braga Aguirre Verheij material, an account of which is provided by Woelkerling (729). (1994: 98) and Basso et al. (1996: 276). By contrast, Chamberlain Titanoderma confine & H. J.H. Price, D.M. John & (737: 204) has continued (see also Chamberlain, 1991 and Cham- (P. Crouan) G.W. Lawson berlain & Irvine, 736) to maintain Titanoderma 'pending more (as confinis) Note. In an earlier paper in this series, John et al. (1994: 64) treated this conclusive, probably genetic, data to determine their relationship . . taxon as a heterotypic synonym of Lithophyllum pustulatum, noting that .'. Babbini & Bressan (753: 150) follow Chamberlain & Irvine Chamberlain (1991, as Titanoderma) recognized it as a distinct variety of (736). Chamberlain (737: 204), however, has noted the difficulties in that species, and noting that all published records from the West African species such as Lithophyllum johansenii Woelkerling & assigning region required confirmation. The additional record of Chamberlain & to or Titanoderma as delimited in her stud- Campbell Lithophyllum Irvine (736: 105, as Titanoderma pustulatum var. confine) from the Canary ies; other examples are given by Campbell & Woelkerling (1990) Islands also requires confirmation. Earlier West African records are given by and Woelkerling & Campbell (1992). John et al. (1994: 64). Woelkerling (1996o: 229) reaffirmed the earlier As stated by Chamberlain (737: 204; see also Braga & Aguirre, conclusion of Woelkerling & Campbell (1992) that, in contrast to British (Chamberlain, 1991), southern Australian populations were : be identified in the populations 1 995 27 1 ), two distinct lines of development can and continuously variable, making it impossible to recognize distinct "Lithophyllum-Titanoderma complex': one in which the thallus highly varieties in that region. includes a layer of palisade cells and one in which no such layer is the thallus present. In most attached species with a palisade layer, Titanoderma corallinae (P. & H. Crouan)Woelkerling, Y.M. Cham- margin commonly does not immediately thicken while in species berlain & PC. Silva does lacking a palisade layer, immediate thickening commonly See Lithophyllum corallinae (P. & H. Crouan) Heydrich. occur. While such trends may be interpreted in evolutionary terms, The following are additional records: level results in investing these with formal taxonomic status at genus [As Titanoderma corallinae (P. & H. Crouan) Woelkerling, Y.M. a situation where some species cannot be placed with certainty in Chamberlain & PC. Silva] either one genus or the other. For some attached species, including Canaries (633;736). Lithophyllum johansenii and species mentioned by Campbell & 'British Isles to Canary Isles . . .' (649). (1990) and Woelkerling & Campbell (1992), known Woelkerling Titanoderma Y.M. Chamberlain with cystoseirae (Hauck) Woelkerling, infraspecific variation precludes generic placement certainty. PC. Silva informa- & For other species, known only from unattached specimens, See Lithophyllum cystoseirae (Hauck) Heydrich tion on the characters used by Chamberlain (737) for generic The following are additional records: delimitation is lacking altogether, thus making generic placement [As Dermatolithon cystoseirae (Hauck) H. Huve] purely conjectural. Of the eight British species placed by Chamber- Canaries (584). lain & Irvine (736) inLithophyllumsensu Chamberlain, for example, Mauritanie authors (367) the diagnostic characters of the genus as delimited by these '. . . au golfe de Guinee' (367). (immediate thickening behind a bistratose margin and a basal layer (Hauck) Heydrich] - of [As Lithophyllum cystoseirae of non-palisade cells also see key on p. 30 Chamberlain & Annobon (Pagalii) (455). Irvine, 736) are unknown for four (L. dentatum, L. duckeri, L. [As Titanoderma papillosum var. cystoseirae (Hauck) Me. Lemoine fasciculatum, L. hibernicum). ex G.W. Lawson & D.M. John] terms Investing apparent evolutionary trends in formal taxonomic Annobon (Pagalii) (586). to draw firm at generic level in cases where it is presently impossible Note. Woelkerling & Verheij (1995: 45) provide information on the not boundaries or in cases where data on species are missing or are lectotype. in either apparent in any known specimens serves no useful purpose the Titanoderma (Me. Lemoine) J.H. Price, D.M. John taxonomic or evolutionary terms, and it needlessly complicates geometricum identifi- & G.W. Lawson more practical matters of species placement and specimen to See Me. Lemoine. cation. As noted by Braga & Aguirre (1995: 271), it is possible Lithophyllum geometricum within a The following are additional records: recognize that two evolutionary lines are present single terms that two Verde Islands (370). genus. If one wishes to emphasize in taxonomic Cape 128 W.J. WOELKERLING ET AL. Titanoderma hapalidioides (P. & H. Crouan) J.H. Price, D.M. John [As Titanoderma pustulatum (J.V. Lamouroux) Nageli var. confine} G.W. Lawson Canaries (736). See Lithophyllum pustulatum (J.V. Lamouroux) Foslie Titanoderma Note. In an earlier paper in this series, John et al. (1994: 64) treated this sp. See taxon as a heterotypic synonym of Lithophyllum pustulatum, noting that all Lithophyllum spp. The is an additional record: published records from the West African region required confirmation. following Earlier West African records are given by John et al. (1994: 64). [As Dermatolithon]] Chamberlain (1991, as Titanoderma) and Chamberlain & Irvine (736, as Canaries (18). Titanoderma) recognized four distinct varieties of pustulatum, listing T. Trailliella intricata Batters hapalidioides as a heterotypic synonym of T. pustulatum var. macrocarpum. See In southern Australia, by contrast, Woelkerling & Campbell (1992, as Bonnemaisonia hamifera Hariot. Lithophyllum) and Woelkerling (1996a, as Lithophyllum) found that The following is an additional record: populations were highly and continuously variable, making it impossible to Canaries (598). recognize distinct varieties for that region, and they listed hapalidioides as a affinis De Toni heterotypic synonym of Lithophyllum pustulatum. Trematocarpus (J. Agardh) See Trematocarpus flabellatus (J. Agardh) De Toni Titanoderma J.H. D.M. John papillosum (Zanardini) Price, & Note. Trematocarpus affinis (J. Agardh) De Toni and T. flabellatus (J. are 1 G.W. Lawson Agardh) DeToni considered by Simons ( 983: 808) to belong to different See Lithophyllum papillosum (Zanardini ex Hauck) Foslie species. He is followed in this by Womersley (712). The following are additional records: Trematocarpus flabellatus (J. Agardh) De Toni [As Lithophyllum papillosum (Zanardini) Foslie] Namibia (348;707). Canaries (387). [As Lithophyllum (Dermatolithon) papillosum (Zanardini) Foslie] Trichogloeopsis pedicellata (M. Howe) I. A. Abbott & Doty Canaries (70). Canaries (38C;232B;598;635;745). Note. Afonso-Carrillo et al. (582) considered this taxon (as Goniolithon Tricleocarpa cylindrica (Ellis & Solander) Huisman & Borowitzka papillosum) to be doubtfully present in the Canaries; records therefore Canaries (663;734). necessitate confirmation, despite references given elsewhere. Note. See under Galaxaura cylindrica (Ellis & Solander) Lamouroux Titanoderma polycephalum (Foslie) Woelkerling, Y.M. Chamber- and G. oblongata (Ellis & Solander) Lamouroux. For further information, lain & PC. Silva see Huisman & Borowitzka (1990) and Huisman & Townsend (1993). See Lithophyllum polycephalum Foslie Tricleocarpa fragilis (L.) Huisman & Townsend The following are additional records: [As Tricleocarpa (Galaxaura) oblongata (Ellis & Solander) Huisman [As Goniolithon polycephalum (Foslie) Afonso-Carillo] & Borowitzka] Cape Verde Islands (11 ;597). Cape Verde Islands (7 13). [As Lithophyllum (Dermatolithon) polycephalum Foslie] Note. See also under Galaxaura cylindrica (Ellis & Solander) Lamouroux Canaries (70). andG. oblongata (Ellis & Solander) Lamouroux. InOtero-Schmitt(713) the Verde Islands Cape (362). authors are mistakenly given as '(Ellis et Solander) Huisman et Bornet'. Tenarea [As polycephalum (Foslie) Adey] Perez & Afonso-Carrillo (734) state that the occurrence of Tricleocarpa Canaries (11). fragilis (as Tricleocarpa oblongata (Ellis & Solander) Huisman & Borowitzka) [As Titanoderma polycephalum (Foslie) Woelkerling, Y.M. Cham- has to be confirmed. They did not discover this species in many samples from the Islands. consider berlain & PC. Silva] Canary However, they Galaxaura fragilis (Lamark) Lamouroux ex Decaisne to be with Canaries (633;737). synonymous Tricleocarpa cylindrica while Huisman & Townsend (1993) consider Tricleocarpa fragilis to be Cape Verde Islands (737). synonymous with, and having priority over, the combination Tricleocarpa Note. Chamberlain (737, as Titanoderma) has provided an account of oblongata (Ellis & Solander) Huisman & Borowitzska. holotype material in TRH. Vickersia baccata (J. Agardh) Karsakoff Titanoderma polyclonum (Foslie) Woelkerling, Y.M. Chamberlain Canaries (240;306B;489;499;598;635;684). & PC. Silva Cape Verde Islands (652;713). See Lithophyllum polyclonum Foslie Senegal (529;598). '. . . . . Canaries Titanoderma pustulatum (J.V. Lamouroux) Nageli Atlantique (. ...)... (33). See Lithophyllum pustulatum (J.V. Lamouroux) Foslie 'Lusit.-Africain- Medit.' (529). The following are additional records: [As Vickersia baccata (J. Agardh) Karsakoff emend. B0rgesen] Canaries [As Dermatolithon pustulatum (J.V. Lamouroux) Foslie] (13;38B;71;189; 191 ;226;227;490;634;745;747). Canaries (18). Salvage Islands (38B;231). '. . . [Titanoderma hapalidioides (P. & H. Crouan) J.H. Price, D.M. John O. Atlantico (desde Portugal hasta Cabo Verde) . . .' (747). & G.W. Lawson] [As Vickersia baccata J. Agardh] Canaries (227;362;375;582). Canaries (5). Salvage Islands (35B; 373). [As Vickersia canariensis Karsakoff] Canaries 1 1 1 [As Titanoderma pustulatum (J.V. Lamouroux) Nageli] ( 33; 139;3 ;493;499;538;547). Canaries (633;634;663;747). Vickersia canariensis Karsakoff '. . . Canary Islands, Senegal . . .' (649). See Vickersia baccata (J. Agardh) Karsakoff. [As Titanoderma pustulatum (J.V. Lamouroux) Nageli var. pustulatum} Vickersia sp. Canaries (736). Cape Verde Islands (639). Senegal (736). Ghana (377). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 129 Vidalia volubilis J. (L.) Agardh Wrangelia plebeja J. Agardh 1 1 1 1 1 Canaries(66;7 ;89; 33; 79; 9 1 ;226;227;390;392;439;448;5 7;584; See Wrangelia argus (Montagne) Montagne. 598;663). Wurdemannia miniata J. Feldmann & Hamel Senegambia (133;179;296). (Sprengel) Wurdemannia miniata J. ' [As (Duby) Feldmann & Hamel] '. . . an den atlantischen Kiisten von bis Spanien zum Senegal . Cameroun (350;586). (281;501). Canaries (584;598;635;720;747). '. . . Atlantique subtropicale (de Cadiz au Senegal)' (190). Cape Verde Islands (598;713;720). 'De Cadiz aux Canaries et au Senegal (Leprieur)' (89). Salvage Islands (598). 'From Cadiz southwards to Senegal' (71). Sao Tome Africa of (350;535;586). 'Subtropical [Senegal (N. Gambia); Mauritania; former W. - Tropical Africa (N. Gambia river)' (598). Sahara]' (598). Congo - [As Wurdemannia miniata (Draparnaud) J. Feldmann & Hamel] 'Tropical Africa (N. Gambia Congo river)' (598). Canaries (38B;177;188;191;194;227;489;535;745). Note. 1 1 According to Morris ( 99 ) the correct name \sOsmundaria volubilis Verde Islands (L.) R.E. Norris. See note under that name. Cape (38;38B;100;183;191). Sao Tome (535). Waldoia antillana W.R. '. . . cotes africaines de . . Taylor 1'Atlantique tropical et subtropical .' (194). Ghana '. . . cotes occidentales . (286;290;292;299;350;377;586). d'Afrique et aux Canaries . .' (184). . . '. in the . . tropical parts of Atlantic Ocean .' (350;586). [As Wurdemannia miniata (Draparnauld ex A. P. DeCandolle) - Tropical Africa (N. Gambia Congo river)' (598). Feldman & Hamel] Canaries (633;634). Wrangelia argus (Montagne) Montagne [As Wurdemannia setacea Harvey] Ascension Island (475). Canaries (38C;70;7 1 ;375;555;556). Cameroun (288;350;586). Cape Verde Islands (38;38C;145;555;556). Canaries 191 (38B;38C;38D;68;83;89;97; 124; ;240;351;407;417A; Salvage Islands (38C;555;556). 547;553;559;572?;598;633;634;635;662;663). [As Caulacanthus ustulatus (Martens) Kiitzing] Verde Islands Cape (37;38B;38C;38D;598;639;652;713). Sao Tome (251;263;264;265). Gambia (350;586). Note. Correct citation is Wurdemannia miniata (Sprengel) J. Feldmann Ghana (154;240;288;338;340;344;350;537;586;590). & Hamel, see Silva et al. (1987) and Silva et al. (746). Liberia (129;288;350;586). Wurdemannia setacea Harvey St Helena (644). See Wurdemannia miniata (Sprengel) J. Feldmann & Hamel. Salvage Islands (38B;38D;598). Togo (288;293;350;586). Wurdemannia sp. - Verde Islands 'Tropical Africa (N. Gambia Congo river)' (598). Cape (639) [As Wrangelia argus Montagne] Zanardinia marginata J. Agardh Ascension Island (474;475). See Galaxaura marginata (Ellis & Solander) Lamouroux. Canaries (60;68;83;97;139;227;375;457;547;745). Cape Verde Islands (191). Ghana (297 ;491). AN UPDATE OF CURRENT NAMES FOR NON- [As Wrangelia argus J. Agardh] Canaries GENICULATE CORALLINES REPORTED (439). FROM WEST AFRICA [As Wrangelia plebeja J. Agardh] Cape Verde Islands (38;150). In the second edition of the floristic account this [As Griffithsia argus Montagne] complementing series of critical assessments et John et Canaries (25 ;26;44;266;318;320;401). (Price al., 1986, 1988, 1992; al., 1994; Lawson et al., 1995; this Lawson & John (586: '. . . tropical and subtropical regions of the world.' (417A). paper), foreshadowed and nomenclatural in '. . . 196) re-alignments widespread on tropical and subtropical coasts . . .' (559). changes of coralline from West Africa as a . . species algae reported tropical '. widespread in warm temperate and tropical seas . . .' (350;586). consequence of renewed taxonomic interest in the group. Since Wrangelia penicillata (C. Agardh) C. Agardh 1 987, a number of changes in generic concepts have occurred, new Canaries (18;38B;38D;68;70;584;598;634;635;662;684;747;752). families and subfamilies have been described, and the status and Cape Verde Islands (598;639). disposition of various species have been reassessed. While the status Ghana (292;299;350;376;377;586). and disposition of many species reported from tropical west Africa Salvage Islands (598). still remain uncertain, and while most earlier records require confir- 'Mediterranean and warmer areas of the Atlantic Ocean' (269). mation, post- 1 987 studies on non-geniculate corallines have clarified '. . . widespread in warm temperate and tropical seas' (350;586). the status and generic disposition of a number of species and [As Wrangelia penicillata C. Agardh] infraspecific taxa based on types from or reported to occur in the Canaries ( 1 3;68; 177; 191 ;226;227;375;392;556). West African region. St Helena (644). The following list summarizes the current situation for non- Salvage Islands (38B;38D;556). geniculate Corallinales mentioned in previous parts of the West the '. . . Atlantic tropical et subtropical.' (190; 196). African critical assessment series, including floristic accounts of . The list is '. . John Atlantic Ocean (. . . African and American coasts, Canary Lawson & (350, 586). organized alphabetically by Islands...)' (177). specific epithet, with infraspecific taxa listed alphabetically within associated with and '. . . in dem warmeren Teilen der nordlichen Ha'lfte des atlantischen species. Generic names specific infraspecific taxa in the series are in after the relevant oceans . . .' (502). given parentheses epithet. ET AL. 130 W.J. WOELKERLING For each taxon, the following data are provided: references to the aequinoctiale (Lithophyllum, Porolithon) in the West African critical assessment series and the floristic name SERIES REFERENCES. Lawson & John, 350: 243; Lawson & John, accounts (Lawson & John, 350, 586); current placement/name of the 586: 215; John et al., 1994: 60; Lawson et al., 1995: 111. taxon; new records; and comments relating to that placement includ- based on identifications CURRENT PLACEMENT/NAME. Not determined; see comments. ing information on type material. Records are not at level (e.g. Choreonema sp., Mesophyllum sp.) only genus COMMENTS. Status and disposition uncertain; TRH holotype included. (Woelkerling, 678: 20) from Sao Tome not studied in detail in a reported/new records are presented for the following Recently modern context; placement in Porolithon in this series based on taxa: boreale (Foslie) Y.M. Chamberlain; H. crudatum Hydrolithon considerations explained under entry for genus (Lawson Penrose pragmatic (Bressan)Y.M. Chamber\ain;H.farinosum(LV. Lamouroux) etal., 1995: 111). & Y.M. Chamberlain; H. samoense (Foslie) Keats &Y.M. Chamber- Chamberlain Keats; L. lain; Leplophytum ferox (Foslie) Y.M. & africanum (Archaeolithothamnion, Sporolithon) lobatum Me. foveatum Y.M. Chamberlain & Keats; Lithophyllum SERIES REFERENCES. Price et al., 1986: 17; this paper. Lemoine; L. neoatalyense T. Masaki; Lithoporella melobesioides P. H. (Foslie) Fos\ie\ Lithothamnion corallioides(P. & H. Crouan) & CURRENT PLACEMENT/NAME. Sporolithon africanum (Foslie) Crouan; L sonderi Hauck; Mesophyllum engelhartii (Foslie) W.H. Afonso-Carrillo, according to Afonso-Carrillo (11: 142). Adey;M. erubescent (Foslie) Me. Lemoine ;M. lichenoides(J. Ellis) COMMENTS. Holotype from Canary Islands divided between PC Me. Lemoine; Phymatolithon /enormanJ//(Areschoug)W.H.Adey; (Woelkerling, 730) and TRH (Woelkerling, 678); conclusion of Pneophyllum fragile Kiitzing. Afonso-Carrillo (11) based on examination of TRH portion of absimile (Lithophyllum, Neogoniolithon, Spongites) holotype. SERIES REFERENCES. John et al., 1994: 60, 77; this paper. africanum (Lithophyllum, Porolithon, Spongites) CURRENT PLACEMENT/NAME. Spongites absimile (Foslie & M. SERIES REFERENCES. Lawson & John, 350: 244; Lawson & John, to Afonso-Carrillo but see Howe) Afonso-Carrillo, according (733), 586: 215; John et al., 1994: 60; Lawson et al., 1995: 112; this paper. comments. CURRENT PLACEMENT/NAME. Spongites africanum (Foslie) COMMENTS. Status and disposition of species uncertain; TRH Afonso-Carrillo, Chacana & Sanson, according to Afonso-Carrillo 678: from Jamaica examined Afonso- in this holotype (Woelkerling, 14) by et al. (726: 1 33), but see notes under main species entry paper. Carrillo (733) but placed with uncertainty in Spongites; see note to COMMENTS. Conclusion of Afonso-Carrillo et al. (726) based on Spongites absimile (this paper) for further information. study of TRH holotype (Woelkerling, 678: 23-24) from Cap Vert, accretum (Goniolithon, Lithophyllum, Neogoniolithon) Senegal. SERIES REFERENCES. Priceetal., 1988: 230; John etal., 1994:60,77. africanum f. intermedia (Lithophyllum) See under CURRENT PLACEMENT/NAME. Neogoniolithon accretum (Foslie & Spongites africanum. M. Setchell & L.R. Mason, according to Adey (1970: 8), but Howe) africanum f. truncata (Lithophyllum) see comments. See under Spongites africanum. COMMENTS. Status and disposition of species uncertain; conclu- amplexifrons (Lithophyllum, Lithothamnion, Melobesia, Pneo- sion of Adey (1970) requires verification via a modern, detailed 678: phyllum) study of the Florida US holotype in NY (Woelkerling, 14). SERIES REFERENCES. John et al., 1994: 60, 66, 71; Lawson et al., accretum f./var. canariense/canariensis (Lithophyllum, Neogonio- 1995: 105. lithon) CURRENT PLACEMENT/NAME. Pneophyllum amplexifrons (Harvey) SERIES REFERENCES. John et al., 1994: 60, 77 (under Y.M. Chamberlain & R.E. Norris), according to Chamberlain & Neogoniolithon accretum). Norris(1994). CURRENT PLACEMENT/NAME. Not determined; see comments. Additional records: Cape Verde Islands (625); pantropical (625). of Chamberlain Norris based COMMENTS. Status and disposition uncertain; holotype from Ca- COMMENTS. Conclusion & (1994) of & 1992: 98) nary Islands divided between PC (Woelkerling, 730) and TRH on study TCD lectotype (Woelkerling Campbell, (Woelkerling, 678) but not studied in detail in a modern context; from South Africa. placement under Neogoniolithon accretum (John et al., 1994: 77) in angolense (Lithothamnion) this series follows format conventions (John et al., 1994: 49) without implying synonymy with the type of the species. SERIES REFERENCE. John et al., 1994: 66. adhaerens (Tenarea) CURRENT PLACEMENT/NAME. Not determined; see comments. et 1994: 78 SERIES REFERENCES. John al., (under Neogoniolithon COMMENTS. Status and disposition uncertain; whereabouts of type hirtum); this paper. material uncertain; protologue (Romanes, 677) based on fossil from four localities. CURRENT PLACEMENT/NAME. Heterotypic synonym of specimens hirtum(Me. Lemoine in B0rgesen) Afonso-Carrillo, Neogoniolithon aninae (Lithophyllum) according to Afonso-Carrillo (11: 131). SERIES REFERENCE. John et al., 1994: 60. COMMENTS. Conclusion of Afonso-Carrillo (1 1) based on exami- PLACEMENT/NAME. aninae Foslie, accord- nation of holotype (from Canary Islands) in C. CURRENT Lithophyllum RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 131 ing to Adey (1970: 8), but see comments. COMMENTS. Status and disposition uncertain; TRH holotype (Woelkerling, 678: 23) from Sao Tome not studied in detail in a COMMENTS. Status and disposition uncertain; conclusion of Adey modern context; placement under Hydrolithon boergesenii in Price (1970) requires verification via a modern, detailed study of theTRH et al., ( 1 992) follows format conventions (see Price et al., 1 992: 1 23) holotype (Woelkerling, 678: 27) from the Cape Verde Islands; without implying synonymy with the type of the species. holotype fragments in PC (Woelkerling, 730). boreale (Hydrolithon) antarcticum (Lithothamnion) SERIES REFERENCES. Not previously reported. SERIES REFERENCE. John et al., 1994: 66. CURRENT PLACEMENT/NAME. Hydrolithon boreale (Foslie) Y.M. CURRENT PLACEMENT/NAME. Heterotypic synonym ofSynarthro- Chamberlain, according to Chamberlain (702: 1 16). phyton patena (Hooker f. & Harvey in Harvey) R.A. Townsend; see May & Woelkerling (1988: 68). NEW RECORDS. Canaries (687;702). COMMENTS. Delete from flora; no published record of occurrence COMMENTS. Conclusion of Chamberlain (702) based on study of from West African region found. TRH lectotype from Ireland. applicatum (Lithophyllum, Mesophyllum) bornetii (Leptophytum, Lithothamnion) SERIES REFERENCES. John et al., 1994: 60, 73, 78 (under Neogon- SERIES REFERENCES. John et al., 1994: 57, 66. iolithon hirturri). CURRENT PLACEMENT/NAME. See comments. CURRENT PLACEMENT/NAME. Heterotypic synonym of COMMENTS. Based on a detailed study of type material in PC Neogoniolithon hirtum(Me. Lemoine in B0rgesen)Afonso-Carrillo, (Woelkerling, 730) andTRH (Woelkerling, 678), Chamberlain ( 1 990) according to Afonso-Carrillo (11: 131), but see comments. placed the species mLeptophytum. Subsequently, Diiwel &Wegeberg based on exami- COMMENTS. Conclusion of Afonso-Carrillo (11) (1996) concluded from a study of relevant types that Leptophytum from Islands in nation of type material Canary C; lectotype, however, constitutes a heterotypic synonym of Phymatolithon. Babbini & under apparently not yet designated (Woelkerling, 730, Lithophyllum Bressan (753: 312), clearly without knowing the work of Diiwel & confirmation applicatum) and placement requires following Wegeberg (1996), considered the genus Leptophytum as insuffi- et 1994: lectotypification (John al., 78). ciently characterized, and the species as dubious and rare. Evidence from the type of bornetii presented by Chamberlain (1990) strongly atlantica (Lithoporella) suggests that the species belongs to Phymatolithon, but it has yet to SERIES REFERENCE. John et al., 1994: 65. be validly transferred into that genus (comments on invalid transfers to are under Lithothamnion bornetii; see CURRENT PLACEMENT/NAME. Heterotypic synonym of Lithopor- Phymatolithon provided Johnetal., 1994:66). ella melobesioides (Foslie) Foslie, according to Lemoine (37 1 : 44), but proposed synonymy not based on examination of type material brachycladum (Lithothamnion, Mesophyllum) (for data on type material, see Woelkerling, 678, 730). SERIES REFERENCES. Lawson & John, 350: 240; Lawson & John, Delete from no record of occurrence COMMENTS. flora; published 586: 210; John et al., 1994: 66, 73. from West African region found. CURRENT PLACEMENT/NAME. Mesophyllum brachycladum (Foslie) bisporum (Leptophytum, Lithophyllum, Lithothamnion, Phym- W.H. Adey, according to Adey (1970: 22), but see comments. atolithon) COMMENTS. Status and disposition of species uncertain; holotype John et 1994: Lawson et al., in with SERIES REFERENCES. al., 57, 60, 66; (Woelkerling, 678: 14) from St Helena Island mainly BM not studied in 1995: 102. fragments in PC andTRH (Woelkerling, 678, 730) but detail in a modern context. CURRENT PLACEMENT/NAME. Not determined; see comments. brassica-florida (Goniolithon, Lithothamnion, Melobesia, Neo- COMMENTS. Status and disposition uncertain; holotype from Ca- goniolithon) nary Islands in PC (Woelkerling, 730) with fragments in TRH not studied in detail in a modern context. 1 et 1994: (Woelkerling, 678); SERIES REFERENCES. Price et al., 1988: 23 ; John al., 67, 1 7 , 77, 78 (under Neogoniolithon mamillare). boergesenii (Goniolithon, Hydrolithon, Porolithon) Current name: brassica-florida (Harvey) Setchell & SERIES REFERENCES. Lawson & John, 350: 235; Lawson & John, Neogoniolithon to et al. et L.R. Mason, according Woelkerling (678: 324-326). 586: 206; Price et al., 1988: 230; Price et al., 1992: 131; Lawson a al., 1995: 112. COMMENTS. Conclusion of Woelkerling et al. (678) based on of the BM lectotype from South Africa. CURRENT PLACEMENT/NAME. Hydrolithon boergesenii (Foslie) study in Price et al. Foslie, according to Woelkerling (1992: 131). byssoides (Goniolithon, Lithophyllum, Titanoderma) Price et al., 1992) 1 et 1994: COMMENTS. Conclusion of Woelkerling (in SERIES REFERENCES. Price et al., 1988: 23 ; John al., 60; based on examination ofTRH 678: 40) from lectotype (Woelkerling, this paper. US Virgin Islands. CURRENT PLACEMENT/NAME. Lithophyllum byssoides (Lamarck) Porolithon) boergesenii f./var. africana (Goniolithon, Foslie, according to Woelkerling (729). SERIES REFERENCES. Price et al., 1992: 131 (under Hydrolithon COMMENTS. Conclusion of Woelkerling (729) based on a study of et 1995: 1 12. to come from the boergesenii); Lawson al., PC lectotype, said but not confirmed English CURRENT PLACEMENT/NAME. Not determined; see comments. Channel. 132 W.J. WOELKERLING ET AL. calcareum (Lithophyllum, Lithothamnion, Lythophyllum, canariense var. fasciata (Mesophyllum) Phymatolithon) SERIES REFERENCE. John et al., 1994: 74 (under Mesophyllum SERIES REFERENCES. John et al., 1994: 60, 67, 71; Lawson et al., canariense). 1995: 103. CURRENT PLACEMENT/NAME. Not determined; see comments. CURRENT PLACEMENT/NAME. Phymatolithon calcareum (Pallas) COMMENTS. Status and disposition uncertain; holotype from Ca- W.H. & D.L. McKibbin, to Woelkerling & Irvine Adey according nary Islands presumed to be in C but not studied in detail in a modern (1986a). context; placement under Mesophyllum canariense (John et al., 1994: 74) in this series follows format conventions (John et al., COMMENTS. Conclusion of Woelkerling & Irvine (1986a) based 1994: 49) without implying with the type of the species. on a detailed study of the designated neotype (from Falmouth synonymy in BM. harbour, England) capense/capensis (Lithophyllum, Lithothamnion) calcareum f. crassa (Lithothamnion) SERIES REFERENCE. John et al., 1994: 60, 67. SERIES REFERENCE. Lawson etal., 1995: 103 (underPhymatolithon CURRENT PLACEMENT/NAME. Not determined; see comments. calcareum). COMMENTS. Status and disposition uncertain; lectotype in CN CURRENT PLACEMENT/NAME. Lithothamnion calcareum f. crassa (Woelkerling & Verheij, 1995: 38) from South Africa not studied in (Philippi) Me. Lemoine, considered a heterotypic synonym of detail in a modern context; isolectotypes present in BM, L, and PC Lithophyllum racemus (Lamarck) Foslie, according to Basso et al. (Woelkerling & Verheij, 1995; Woelkerling, 19986). (1996:284-286). caribaeum (Lithophyllum, Neogoniolithon) COMMENTS. See comments for Lithophyllum racemus. Babbini & SERIES REFERENCE. John et al., 1994: 60, 77. Bressan (753: 124, 128) listed Lithophyllum crassum Philippi as a CURRENT PLACEMENT/NAME. caribaeum synonym of Lithophyllum racemus (Lamarck) Foslie (with a f. Neogoniolithon (Foslie) W.H. to but see comments. crassa (Philippi) Foslie) along with Phymatolithon calcareum f. Adey, according Adey (1970: 8), crassa Me. Lemoine. COMMENTS. Status and disposition uncertain; conclusion of Adey verification via a detailed of the TRH callithamnioides sensu Falkenberg (Melobesia) (1970) requires modern, study lectotype (Woelkerling, 678: 48) from the US Virgin Islands. SERIES REFERENCE. John et al., 1994: 71. confervicola (Melobesia, Pneophyllum) CURRENT PLACEMENT/NAME. Not determined; see comments. SERIES REFERENCES. John et al., 1994: 71; Lawson et al., 1995: COMMENTS. Application of name uncertain and apparently also 106. commonly misapplied (see comments in Chamberlain, 94: 351- CURRENT PLACEMENT/NAME. Pneophyllum confervicola (Kiitz- 352; John et al., 1994: 71; Chamberlain, 702: 116-117 under ing) Y.M. Chamberlain, according to Chamberlain (94: 385; 702: Hydrolithon boreale). Babbini & Bressan (753: 200) classified this 137). under Hydrolithon farinosum (P.V. Lamouroux) Penrose & Y.M. Chamberlain. Relevant collections of Falkenberg require re-investi- COMMENTS. Conclusions of Chamberlain (94;702) based on ex- gation as a basis for resolving uncertainties. amination of holotype from Italy in L (Woelkerling & Verheij, 1995: 41). canariense/canariensis (Lithophyllum, Lithothamnion, Meso- phyllum) confervicolum f. minutula (Pneophyllum) SERIES REFERENCE. Lawson et al., 1 995: 1 06 (under SERIES REFERENCES. Lawson & John, 350: 241 ; Lawson & John, Pneophyllum 586: 211; John et al., 1994: 60, 67, 74. confervicola). CURRENT PLACEMENT/NAME. of Pneo- CURRENT PLACEMENT/NAME. Mesophyllum canariense (Foslie) Heterotypic synonym confervicola (f. confervicola), according to Chamberlain Me. Lemoine, according to Reyes & Afonso-Carrillo (687). phyllum (19946: 140). COMMENTS. Conclusion of Reyes & Afonso-Carrillo (687) based COMMENTS. Conclusion of Chamberlain (702) based on examina- on examination of TRH portion (Woelkerling, 678) of holotype tion (Chamberlain, 94: 394) of TRH holotype (Woelkerling, 678: (from Canary Islands); major portion of holotype now known to be 151) from Norway. in PC (Woelkerling, 730). confine/confinis (Dermatolithon, Melobesia, Tenarea, Titano- canariense van difformis (Mesophyllum) derma) SERIES REFERENCE. John et al., 1994: 74 (under Mesophyllum SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 71; canariense). this paper. CURRENT see comments. PLACEMENT/NAME. Not determined; CURRENT PLACEMENT/NAME. Heterotypic synonym of Litho- phyllum pustulatum (Lamouroux) Foslie, according to Woelkerling COMMENTS. Status and disposition uncertain; holotype from Ca- & Campbell (1992: 79). Although others (Chamberlain, 1991; nary Islands presumed to be in C but not studied in detail in a Babbini & Bressan, 753: 181) list it as Titanoderma pustulatum var. modern context; placement under Mesophyllum canariense (John confine (P. & H. Crouan) Y.M. Chamberlain. et al., 1994: 74) in this series follows format conventions (John et al., 1994: 49) without implying synonymy with the type of the COMMENTS. Conclusion of Woelkerling & Campbell (1992) takes species. account of study of PC lectotype by Chamberlain (1991, as RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 133 Titanoderma); lectotype (Woelkerling, 730) presumed (Chamber- CURRENT PLACEMENT/NAME. Not determined. Babbini & Bressan to be from France. lain, 1991) (753: 258), however, listed it as Lithothamnion crispatum Hauck; see comments. conjuncta (Lithoporella, Mastophora) COMMENTS. Status and disposition uncertain; lectotype from Adri- SERIES REFERENCE. John et al., 1994: 66, 71. atic Sea in L (Woelkerling & Verheij, 1995: 44) vegetatively CURRENT PLACEMENT/NAME. Heterotypic synonym of concordant withLithothamnion, but absence of reproductive material Lithoporella melobesioid.es (Foslie) Foslie, according to Lemoine precludes certain generic placement; information on PC and TRH (37 1 : 44), but see comments. isolectotypes provided by Woelkerling (730). COMMENTS. Status and disposition uncertain; conclusion of crouanii (Lithophyllum) Lemoine (371) not based on study of lectotype (from Cape Verde SERIES REFERENCE. John et al., 1994: 61. Islands) inTRH (Woelkerling, 678); also treated as a distinct species of Lithoporella (Adey, 1970: 15). CURRENT PLACEMENT/NAME. Lithophyllum crouanii Foslie, ac- cording to Chamberlain et al. (1988). corallinae (Dermatolithon, Lithophyllum, Melobesia, Tenarea, Titanoderma) COMMENTS. Conclusion of Chamberlain et al. (1988) based on examination of lectotype from England in TRH (Woelkerling, 678: SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 60, 68). 72; this paper. cruciatum (Hydrolithon) CURRENT PLACEMENT/NAME. Lithophyllum corallinae (P. & H. Crouan) Heydrich, according toWoelkerling & Campbell ( 1 992: 41). SERIES REFERENCES, not previously reported. COMMENTS. Conclusion ofWoelkerling & Campbell (1992) based CURRENT PLACEMENT/NAME. Hydrolithon cruciatum (Bressan) on an examination of the lectotype (from France) in CO and the Y.M. Chamberlain, according to Chamberlain (702: 120). earlier detailed study of the lectotype by Chamberlain (1991: 66, as NEW RECORDS. Canaries (687;702). Titanoderma). COMMENTS. Conclusion of Chamberlain (702) requires confirma- corallioides (Lithothamnion) tion via a study of the TSB holotype from Italy; type not seen by SERIES REFERENCE. John et al., 1994: 67. Chamberlain (702). CURRENT PLACEMENT/NAME. Lithothamnion corallioides (P. & cystoseirae (Dermatolithon, Lithophyllum, Melobesia, Titano- H. Crouan) P. & H. Crouan, according to Chamberlain & Irvine derma) (701: 177). SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 61, NEW RECORD. Canaries (701). 72; this paper. COMMENTS. Conclusion of Chamberlain & Irvine (701) based on CURRENT PLACEMENT/NAME. Lithophyllum cystoseirae (Hauck) to John et al. the selection and study of the neotype; information on isoneotypes in Heydrich, according (1994: 61). PC provided by Woelkerling (730). COMMENTS. Conclusion of John et al. ( 1 994) based on data on the from in L & 1995: 45) corticiforme/corticiformis (Epilithon, Lithothamnion, Melo- lectotype Italy (Woelkerling Verheij, provided Huve and Athanasiadis besia) by (272) (1989). f./var. saxicola (Dermatolithon) SERIES REFERENCES. Price et al., 1986: 89; John et al., 1994: 67, cystoseirae 72. SERIES REFERENCE. John et al., 1994: 61 (under Lithophyllum CURRENT PLACEMENT/NAME. Heterotypic synonym of Melobesia cystoseirae). membranacea (Esper) J.V. Lamouroux, according to Chamberlain CURRENT PLACEMENT/NAME. Nom. nud.\ see comments. (94: 300, 306). COMMENTS. Name first coined by Huve (272: 234) and mentioned based on examina- COMMENTS. Conclusion of Chamberlain (94) by Lemoine (368: 6) but never validated with a description or in the Atlantic tion of the holotype (from an unnamed locality designation of a type specimen in accordance with Articles 36 and Ocean) in L; additional information on holotype provided by 37 of the ICBN (see Greuter, 1994). Woelkerling & Verheij (1995: 42). daedaleum (Lithophyllum) crassum (Lithothamnion) SERIES REFERENCE. John et al., 1994: 61. SERIES REFERENCES. John et al., 1994: 67; Lawson et al., 1995: CURRENT PLACEMENT/NAME. Lithophyllum daedaleum Foslie & 103 (under Phymatolithon calcareum). M. Howe, according to Adey (1970: 5), but see comments. CURRENT PLACEMENT/NAME. Lithothamnion crassum Philippi con- COMMENTS. Status and disposition uncertain; NY holotype sidered a heterotypic synonym of Lithophyllum racemus (Lamarck) (Woelkerling, 678: 27) from Puerto Rico not studied in detail in a Foslie, to Basso et al. (1996: 284-286) and Babbini & according modern context; information on isotypes and paratype at TRH, L, Bressan (753: 258). and PC provided by Woelkerling (678), Woelkerling & Verheij and COMMENTS. See comments for Lithophyllum racemus. (1995) Woelkerling (730). decussatum f. crispatum (Lithothamnion) planiuscula (Lithophyllum) SERIES REFERENCE. John et al., 1994: 61 (as "planiscula"). SERIES REFERENCE. John et al., 1994: 67. 134 WJ. WOELKERLING ET AL. CURRENT PLACEMENT/NAME. Not determined; see comments. expansa/expansum (Crodelia, Lithophyllum, Pseudolitho- phyllum) COMMENTS. Status and disposition uncertain; holotype from Mo- John et 1994: rocco in TRH (Woelkerling, 678: 173) not studied in detail in a SERIES REFERENCES. Price et al., 1986: 78; al., 62, 75 modern context; West African record questionable (John et al., 1 994: 63 (under Lithophyllum lobatum), (under Mesophyllum 112. 61). lichenoides); Lawson et al., 1995: PLACEMENT/NAME. of duckeri (Lithophyllum) CURRENT Heterotypic synonym Mesophyllum lichenoides, according to Woelkerling (1983b: 307) SERIES REFERENCE. John et al., 1994: 61. and Lawson et al. (1995: 1 12), but see comments. CURRENT PLACEMENT/NAME. Lithophyllum duckeri Woelkerling COMMENTS. Holotype from Sicily in L (Woelkerling & Verheij, a nom. nov. for Lithothamnion crasswn (1983a: 184), Philippi, 1995: 51) studied in detail by Woelkerling (19836: 307) who sug- considered a of Lithophyllum racemus heterotypic synonym gested conspecificity with Mesophyllum lichenoides', disposition of (Lamarck) Foslie, according to Basso et al. (1996: 284-286). plants referred to Pseudolithophyllum expansum (Philippi) Me. Lemoine sensu Lemoine discussed Furnari et al. On the COMMENTS. See comments for Lithophyllum racemus. by (1996). other hand, Babbini & Bressan (753: 130) listed this under ectocarpon (Lithothamnion, Mesophyllum) Lithophyllum frondosum (Dufour) Furnari, Cormaci & Alongi f. Babbini & Bressan, a name which has to SERIES REFERENCE. John et al., 1994: 67, 74. expansum (Me. Lemoine) be checked against the provisions of ICBN 1994 for acceptability CURRENT PLACEMENT/NAME. (Foslie) Mesophyllum ectocarpon (see Greuter, 1994) W.H. Adey, according to Adey (1970: 23), but see comments. expansum f. exigua (Lithophyllum) COMMENTS. Status and disposition uncertain; TRH lectotype SERIES REFERENCES. John et al., 1994: 75 (under Mesophyllum (Woelkerling, 678: 27) from Cap Blanc, Senegal not studied in detail lichenoides); Lawson et al., 1995: 112 (under Pseudolithophyllum in a modern context; placement in Mesophyllum somewhat in ques- expansum). tion (Adey, 1970: 23); information on isolectotype in L provided by Woelkerling & Verheij (1995: 51). CURRENT PLACEMENT/NAME. Not determined. Babbini & Bressan (753: 131) included this under Lithophyllum frondosum (Dufour) endophloea (Schmitziella) Furnari, Cormaci & Alongi without providing a statement on its See under Schmitziella endophloea Bornet ex Batters (see also taxonomic status. See comments. Woelkerling & Irvine, 1982). COMMENTS. Status and disposition uncertain; holotype from Al- engelhartii (Mesophyllum) geria in TRH (Woelkerling, 678) not studied in detail in a modern context; placement under Mesophyllum lichenoides in this series SERIES REFERENCES, not previously reported. follows format conventions (John et al., 1994: 49; Lawson et al., PLACEMENT/NAME. CURRENT Mesophyllum engelhartii (Foslie) 1995: 99) without implying synonymy with the type of the species. W.H. Adey, according to Woelkerling & Harvey (1993: 581). expansum f. involvens (Lithophyllum) NEW RECORD. Namibia (742). SERIES REFERENCE. John et al., 1994: 75 (under Mesophyllum based COMMENTS. Conclusion of Woelkerling & Harvey (1993) lichenoides). on study of TRH lectotype (Woelkerling (678: 84) from southern CURRENT PLACEMENT/NAME. Not determined. According to Australia. Babbini & Bressan (753: 138) f. involvens Vinassa = f. exigua Foslie erubescens (Lithothamnion, Mesophyllum) underLithophyllumfrondosum (Dufour) Furnari, Cormaci &Alongi, but provided no statement on its taxonomic status; see comments. SERIES REFERENCES. John et al., 1994: 67, 74. COMMENTS. Status and disposition uncertain; type material from CURRENT PLACEMENT/NAME. Mesophyllum erubescens (Foslie) the Mediterranean Sea (Vinassa, 1892) not studied in detail in a Me. Lemoine, according to Keats & Chamberlain (755: 175). modern context and whereabouts uncertain; placement under ADDITIONAL RECORDS. Cape Verde Islands (625); pantropical Mesophyllum lichenoides in this series follows format conventions (625). (John et al., 1994: 49) without implying synonymy with the type of the species. NEW RECORD. Senegal (755). expansum f. stictaeformis (Lithophyllum) COMMENTS. Conclusion of Keats & Chamberlain (755) based on examination of holotype from Brasil in TRH (Woelkerling, 678: SERIES REFERENCE. John et al., 1994: 75 (under Mesophyllum 85). lichenoides). esperi (Lithophyllum, Pseudolithophyllum) CURRENT PLACEMENT/NAME. Not determined. According to Babbini & Bressan (753: 131) this taxon can be included under SERIES REFERENCES. John et al., 1994: 61; Lawson et al., 1995: Lithophyllumfrondosum (Dufour) Furnari, Cormaci & Alongi. They 112. did not provide a statement on its taxonomic status. See comments. CURRENT PLACEMENT/NAME. Not determined; see comments. COMMENTS. Status and disposition uncertain; type material from COMMENTS. Status and disposition uncertain; species based on a the Mediterranean Sea (Areschoug, 1852: 517) not studied in detail series of collections from Canary Islands (Lemoine, 362: 63) but not in a modern context; placement under Mesophyllum lichenoides in lectotypified and original collections not studied in detail in a this series follows format conventions (John et al., 1994: 49) without modern context (John et al., 1994: 61). implying synonymy with the type of the species. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 135 farinacea (Melobesia) COMMENTS. Status and disposition uncertain; conclusion of Adey (1970) requires verification via a modern, detailed study of the TRH SERIES REFERENCES. Price et al., 1986: 92 (under Fosliella holotype (Woelkerling, 678: 96) from Florida, U.S.A.; holotype farinosa)', John et al., 1994: 72. fragment in PC (Woelkerling, 730). COMMENTS. Epithetfarinaceain the b'momia\Melobesiafarinacea foveatum (Leptophytum) Lamouroux an orthographic variant of Melobesia farinosa; see following entry. SERIES REFERENCES. Not previously reported. farinosa (Fosliella, Melobesia) CURRENT PLACEMENT/NAME. Not determined; see comments. SERIES REFERENCES. Lawson 350: Price et 1 & John, 234; al., 986: NEW RECORD. Namibia (743). 91; Lawson & John, 586: 205; John et al., 1994: 72. COMMENTS. Originally described as a species of Leptophytum CURRENT PLACEMENT/NAME. Hydrolithon farinosum (J.V. (Chamberlain & Keats, 743) based on a holotype from South Penrose & Y.M. to Penrose & Lamouroux) Chamberlain, according Africa deposited in L; Leptophytum now considered a heterotypic Chamberlain (1993). synonym of Phymatolithon (Duwel & Wegeberg, 1996) but not transferred to that and other NEW RECORD. Canaries (702). foveatum formally genus; type collections cited in protologue require further study to determine COMMENTS. Conclusion of Penrose & Chamberlain 1 based ( 993) generic placement. on examination of lectotype (from an unspecified locality in the Mediterranean) in CN. fragile (Pneophyllum) SERIES REFERENCE. Lawson et al., 1995: 106. farinosa f. callithamnioides (Fosliella) CURRENT PLACEMENT/NAME. ac- SERIES REFERENCES. Price et al., 1986: 91 (under Fosliella Pneophyllum fragile Kutzing, to Chamberlain (1983: and Penrose & farinosa); Lawson & John, 586: 205. cording 356) Woelkerling (1991:495). CURRENT PLACEMENT/NAME. Not determined; see comments. NEW RECORDS. Canaries (702;740). COMMENTS. Status and disposition uncertain; Fosliella farinosa f. COMMENTS. Conclusions of Chamberlain and Penrose & callithamnioides (Foslie) Y.M. Chamberlain based on Melobesia (1983) based on of L & farinosa f. callithamnioides Foslie (see Chamberlain, 94: 352 for Woelkerling (1991) study holotype (Woelkerling 1995: from an in the Mediterranean details); taxon apparently not lectotypified (Chamberlain, 94: 352); Verheij, 53) unspecified locality also in PC name in the sense of Chamberlain (94) subsequently suppressed Sea; holotype fragment (Woelkerling, 730). under boreale (Chamberlain, 702: 116-117). Hydrolithon fruticulosum/fruticulosus (Lithothamnion, Spongites) farinosa var. solmsiana (Fosliella, Melobesia) SERIES REFERENCES. John et al., 1994: 67; this paper. SERIES REFERENCE. Price et al., 1986: 91, 92 (under Fosliella CURRENT PLACEMENT/NAME. Spongites fruticulosus Kutzing, ac- farinosa). cording to Woelkerling (1985: 135) and Penrose (1991). CURRENT PLACEMENT/NAME. Not determined; see comments. COMMENTS. Conclusions of Woelkerling (1985) and Penrose based on of L & 1995: . di uncertain whereabouts of COMMENTS Status and sposition ; type (1991) study holotype (Woelkerling Verheij, an in the Mediterranean Sea. material uncertain; additional comments provided by John et al., 54) from unspecified locality (1994: 72, under Melobesia solmsiana), Chamberlain (94: 351, fruticulosum f. clavulata (Lithothamnion) under Fosliella farinosa f. callithamnioides), Taylor (1939, under under Fosliellafarinosa var. solmsiana) and Babbini & Bressan (753: 200, See Spongites fruticulosus. under Hydrolithon farinosum). fruticulosum f. crassiuscula (Lithothamnion) ferox (Leptophytum) See under Spongites fruticulosus. SERIES REFERENCE. Not previously reported. geometricum (Dermatolithon, Lithophyllum, Titanoderma) CURRENT PLACEMENT/NAME. Not determined; see comments. SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 62; NEW RECORD. Namibia (743). this paper. see comments. COMMENTS. Placed in Leptophytum by Chamberlain & Keats CURRENT PLACEMENT/NAME. Not determined; (743) based on study of TRH holotype (Woelkerling, 678: 92) from COMMENTS. Status and disposition uncertain; type material from South Africa; Leptophytum now considered a heterotypic synonym Canary Islands in C (unpublished data) not studied in detail in a of Phymatolithon (Duwel & Wegeberg, 1 996) butferox not formally modern context; species requires lectotypification (John et al., 1 994: transferred to that genus; type and other collections cited by Cham- 62). berlain & Keats (743) require further study to determine generic placement. gracile (Lithophyllum) REFERENCE. John et al., 1994: 62. floridanum (Lithothamnion, Mesophyllum) SERIES accord- SERIES REFERENCES. Lawson & John, 350: 241; Lawson & John, CURRENT PLACEMENT/NAME. Lithophyllum gracile Foslie, to 5), but see comments. 596: 211; John et al., 1994: 67, 74. ing Adey (1970: Status and conclusion of CURRENT PLACEMENT/NAME. Mesophyllum floridanum (Foslie) COMMENTS. disposition uncertain; Adey verification via a modern, detailed study of the TRH W.H. Adey, according to Adey (1970: 24), but see comments. (1970) requires 136 W.J. WOELKERLING ET AL. holotype (Woelkerling, 678: 108) from the Cape Verde Islands. CURRENT PLACEMENT/NAME. Neogoniolithon illitus(Me. Lemoine) Afonso-Carrillo, according to Afonso-Carrillo (11: 133), but see (Dermatolithon, Lithophyllum, Tenarea, Titano- hapalidioides comments. derma) COMMENTS. Conclusion of Afonso-Carrillo (11) based on exami- SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 62, nation of type material from Canary Islands in C, but lectotype 64 (under Lithophyllum pustulatum); this paper. apparently not yet designated and placement in Neogoniolithon CURRENT PLACEMENT/NAME. of Heterotypic synonym requires confirmation following lectotypification (John et al., 1994: to Lithophyllum pustulatum (J.V. Lamouroux) Foslie, according 78); syntype material also occurs in PC (Woelkerling, 730). Woelkerling & Campbell (1992: 79). incrustans (Lithophyllum, Lithothamnion) COMMENTS. Conclusion of Woelkerling & Campbell ( 1 992) takes SERIES REFERENCES. John et al., 1 994: 62, 65 (under account of study of CHE lectotype from France by Chamberlain Lithophyllum vickersiae), 67; Lawson etal., 1995: 1 13 (\\nferPseudolithophyllum : var. addi- ( 1 99 1 34, under Titanoderma pustulatum macrocarpum); this tional comments under Titanoderma hapalidioides in this paper. vickersiae); paper (under Spongites africanum). PLACEMENT/NAME. incrustans hapalidioides f./var. confinis (Lithophyllum) CURRENT Lithophyllum Philippi, according to Woelkerling (19836: 313-317). SERIES REFERENCE. John et al., 1994: 64 (under Lithophyllum pustulatum). COMMENTS. Conclusion of Woelkerling (19836) based on study of holotype from Sicily in L (Woelkerling & Verheij, 1995: 58). CURRENT PLACEMENT/NAME. Homotypic synonym of Melobesia confinis P. & H. Crouan and thus a heterotypic synonym of Litho- indicum (Lithothamnion) phyllum pustulatum (J.V. Lamouroux) Foslie, according to Woel- SERIES REFERENCE. John et al., 1994: 67. kerling & Campbell (1992: 79). Babbini & Bressan (753: 181), CURRENT PLACEMENT/NAME. Lithothamnion indicum Foslie, ac- however, consider it to belong to variety confinis of Titanoderma cording to Wilks & Woelkerling (1995: 558). pustulatum. COMMENTS. Conclusion of Wilks & (1995) based on COMMENTS. Conclusion of Woelkerling & Campbell (1992) takes Woelkerling study of TRH lectotype (Woelkerling, 678: 125) from Victoria, account of study of PC lectotype by Chamberlain (1991, as Australia. Titanoderma); lectotype (Woelkerling, 730) presumed (Chamber- to be from France. lain, 1991) irregulare/irregularis (Lithophyllum, Lithothamnion, Pseudo- hauckii (Lithophyllum) lithophyllum, Tenarea) SERIES REFERENCES. Lawson 350: Lawson & SERIES REFERENCE. John et al., 1994: 62, 79 (underNeogoniolithon & John, 245; John, mamillosum). 586: 217; John et al., 1994: 62, 67; Lawson et al., 1995: 113; this paper. CURRENT PLACEMENT/NAME. Avowed substitute name for Lithothamnion mamillosum Hauck, 1883 (non L. mamillosum CURRENT PLACEMENT/NAME. Not determined; see comments. Giimbel, 1871), according to Woelkerling & Verheij (1995: 57); COMMENTS. Status and disposition uncertain; holotype from Sao additional data provided by John et al. (1994: 79, under Tome in TRH (Woelkerling, 678: 130, under Lithothamnion irregu- Neogoniolithon mamillosum). lare Foslie) but not studied in detail in a modern context; additional COMMENTS. See entry for mamillosum below. comments provided by John et al. (1994: 62, under Lithophyllum irregulare). hirtum (Lithophyllum, Neogoniolithon) kaiserii (Lithophyllum) SERIES REFERENCE. John et al., 1994: 62, 78. SERIES REFERENCE. John et 1994: 63 CURRENT PLACEMENT/NAME. Neogoniolithon hirtum (Me. al., (as 'kaiseri'). to Afonso-Carrillo Lemoine) Afonso-Carrillo, according (11: 131), CURRENT PLACEMENT/NAME. Not determined; see comments. but see comments. COMMENTS. Status and disposition uncertain; species requires COMMENTS. Conclusion of Afonso-Carrillo (1 1) based on exami- typification and then type requires study in detail in a modern nation of type material from Canary Islands in C; lectotype, however, context (John et al., 1994: 63); syntype material present in TRH apparently not yet designated and placement in Neogoniolithon (Woelkerling, 678: 132). requires confirmation following lectotypification (John et al., 1994: 78). kotschyanum (Lithophyllum) hispanum (Lithothamnion) SERIES REFERENCE. John et al., 1994: 63. SERIES REFERENCE. John et al., 1994: 67. CURRENT PLACEMENT/NAME. Lithophyllum kotschyanum Unger, according to Verheij (1994: 100). CURRENT PLACEMENT/NAME. Invalid name, according to John et al. (1994:67). COMMENTS. Conclusion of Verheij (1994) based on examination of from Gulf of Bahrain in TRH (678: 1 33). COMMENTS. Original presentation (Gonzalez Henriquez, 235) holotype (Woelkerling lacks a description or diagnosis rendering name a nomen nudum and lejolisii (Fosliella, Heteroderma, Melobesia, Pneophyllum) thus invalid in relation to ICBN Art. 32.1 (Greuter, 1994). SERIES REFERENCES. Lawson & John, 350: 235; Price et al., 1986: illitus (Lithophyllum, Neogoniolithon) 92; Lawson & John, 586: 214; Price et al., 1992: 130; John et al., SERIES REFERENCE. John et al., 1994: 62, 78. 1994: 72; Lawson et al., 1995: 106. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 137 CURRENT PLACEMENT/NAME. Heterotypic synonym of CURRENT PLACEMENT/NAME. Not determined; see comments. Pneophyllum fragile KUtzing, according to Penrose & Woelkerling NEW RECORD. Canaries (744). (1991:496). COMMENTS. Status and disposition uncertain; type material from COMMENTS. Conclusion of Penrose & Woelkerling (1991) based Canary Islands in C (unpublished data) not studied in detail in a on study of lectotype from France in CHE. modern context; species requires lectotypification (John et al., 1994: lenormandii (Lithothamnion, Phymatolithon) 63). mamillare/mamillaris (Goniolithon, Lithothamnion, Melobesia, SERIES REFERENCES. John et al., 1994: 67; Lawson et al., 1995: Porolithon) 103. Current placement/name: Phymatolithon lenormandii Neogoniolithon, W.H. to Chamberlain & Irvine SERIES (Areschoug) Adey, according (701: REFERENCES. Lawson & John, 350: 241 ; Lawson & John, 224) and Duwel & Wegeberg (1996: 476). 586: 211; Price et al., 1988: 231; John et al., 1994: 67, 72, 78; Lawsonetal., 1995: 112. NEW RECORDS. Canaries (227;701). CURRENT PLACEMENT/NAME. Not determined; see comments. COMMENTS. Conclusions of Chamberlain & Irvine (701) and Additional record: Diiwel & Wegeberg (1996) based on examination of LD lectotype Cape Verde Islands (625); Pantropical (625). (Woelkerling, 1988: 219) from France. COMMENTS. Status and disposition uncertain; lectotype (Printz, 212: to 678: lenormandii f. squamulosa (Lithothamnion) pi. 47, legend fig. 15) apparently missing (Woelkerling, 144; John et al., 1994: 78) and thus not studied in detail in a modern SERIES REFERENCE. Lawsonetal., 1995: 103 (underPhymatolithon context. lenormandii). mamillosum (Goniolithon, Lithothamnion, Neogoniolithon) CURRENT PLACEMENT/NAME. Heterotypic synonym of SERIES REFERENCES. Price et al., 1988: 231; John et al., 1994: 67, Phymatolithon lenormandii (Areschoug) W.H. Adey f. lenormandii, 78. according to Chamberlain & Irvine (701: 225, 230), but see com- ments. Babbini & Bressan (753: 295), however, consider it to belong CURRENT PLACEMENT/NAME. Current placement for Lithotham- to a separate forma, squamulosa. nion mamillosum Hauck, 1 883 (non Giimbel, 1 87 1 ) not determined; see comments. COMMENTS. Conclusion of Chamberlain & Irvine (701) appar- ently not based on examination of TRH holotype from Norway COMMENTS. Status and disposition uncertain; lectotype at L (Woelkerling, 678: 206, under Lithothamnion squamulosum) and (Woelkerling & Verheij, 1995: 64) from the Adriatic Sea not studied thus requires verification. in detail in a modern context; see also entry for 'hauckii' above. lenormandii f. sublaevis (Lithothamnion) mamillosum f. microcarpa (Goniolithon) SERIES REFERENCE. John et 1994 : 78 SERIES REFERENCE. Lawsonetal., 1995: 103 (underPhymatolithon al., (under Neogoniolithon lenormandii). mamillosum). CURRENT PLACEMENT/NAME. Not determined; see comments. CURRENT PLACEMENT/NAME. Heterotypic synonym of Phymato- lithon lenormandii(Areschoug)W.H.Adey f. lenormandii, according COMMENTS. Status and disposition uncertain; TRH lectotype Irvine but see comments. to Chamberlain & (701: 225, 230), (Woelkerling, 678: 149) from the Cape Verde Islands not studied in detail in a modern context; placement under Neogoniolithon COMMENTS. Conclusion of Chamberlain & Irvine (701) appar- mamillosum (John et al., 1994: 78) follows format conventions ently not based on examination of TRH lectotype from England (John et al., 1994: 49) without implying synonymy with the type of (Woelkerling, 678: 211) and thus requires verification. the species. leptothalloideum (Lithophyllum, Pseudolithophyllum) marlothii (Lithophyllum) SERIES REFERENCES. John et al., 1994: 63; Lawson et al., 1995: SERIES REFERENCES. John et al., 1994: 63, 74 (under Mesophyllum 1 13. Current placement/name: Not determined; see comments. canariense). COMMENTS. Status and material disposition uncertain; type CURRENT PLACEMENT/NAME. Not determined; see comments. (whereabouts uncertain) from Annobon (Pagalu) (Pilger, 455) not Status and material studied in detail in a modern context. COMMENTS. disposition uncertain; syntype occurs in TRH (Woelkerling, 678) and PC (Woelkerling, 730) but lichenoides (Lithothamnion, Mesophyllum) lectotype not designated; additional comments provided by Cham- berlain (738: 154). SERIES REFERENCE. John et al., 1994: 67, 74. mediterranea (Litholepis) CURRENT PLACEMENT/NAME. Mesophyllum lichenoides (J. Ellis) Me. Lemoine, according to Woelkerling & Irvine (1986ft). SERIES REFERENCE. John et al., 1994: 59. In Babbini NEW RECORD. Canaries (701). CURRENT PLACEMENT/NAME. Not determined. & Bressan (753: 170) as Titanoderma mediterranea (Foslie) Woel- COMMENTS. Conclusion of Woelkerling & Irvine (1986ft) based kerling. See comments. on a detailed study of the BM neotype from England. COMMENTS. Status and disposition uncertain; TRH holotype lobatum (Lithophyllum, Mesophyllum, Pseudolithophyllum) (Woelkerling, 678) not examined in detail in a modern context; comments John et al. (1994: 59) and SERIES REFERENCES. John et al., 1994: 63, 75; Lawson et al., 1995: additional provided by by 113. Woelkerling (730). 138 WJ. WOELKERLING ET AL. melobesioides (Lithoporella, Mastophora) COMMENTS. Status and disposition uncertain; TRH holotype (Woelkerling, 678: 163) from the Canary Islands not studied in SERIES REFERENCE. John et al., 1994: 66, 7 1 . detail in a modern context; PC holotype fragments apparently CURRENT PLACEMENT/NAME. Lithoporella melobesioides (Foslie) missing (Woelkerling, 730). Foslie, to Turner & Woelkerling (1982a, b). according onkodes (Lithothamnion, Porolithon) NEW RECORDS. Cape Verde Islands (701), Senegal (701). SERIES REFERENCES. Lawson & John, 350: 244; Lawson & John, COMMENTS. Conclusions of Turner & Woelkerling (1982a, b) 586: 216; John et al., 1994: 67; Lawson et al., 1995: 1 12. based on detailed study of TRH lectotype (Woelkerling, 678: 148) CURRENT PLACEMENT/NAME. Hydrolithon onkodes (Heydrich) from S. Nilandu, Maldive Islands. Penrose & Woelkerling, according to Penrose & Woelkerling (1992: membranacea/membranaceum (Epilithon, Lithothamnion, 83). Melobesia) COMMENTS. Conclusion of Penrose & Woelkerling (1992) based 1986: SERIES REFERENCES. Priceetal., 89;Johnetal., 1994:67,72. on study ofTRH lectotype (Woelkerling, 678) from New Guinea and on earlier of by Penrose & Woelkerling ( 1 988); data CURRENT PLACEMENT/NAME. Melobesia membranacea (Esper) study lectotype on PC isolectotype provided by Woelkerling (730). J.V. Lamouroux, according to Chamberlain (1985) and Wilks & Woelkerling (1991). onkodes var. oligocarpa (Porolithon) Conclusions of Chamberlain and Wilks & COMMENTS. (1985) SERIES REFERENCE. Lawson et al., 1995: 112 (under Porolithon based on studies of CN Woelkerling (1991) neotype (Chamberlain, oligocarpum). 1985) from France; additional comments provided by John et al. CURRENT PLACEMENT/NAME. of Litho- (1994:72). Homotypic synonym phyllum oligocarpum Foslie. mildbraedii (Lithophyllum, Pseudolithophyllum) COMMENTS. See comments under entry for oligocarpum above. SERIES REFERENCES. Lawson & John, 350: 246; Lawson & John, orbiculare (Crodelia) 586: 217; John et al., 1994: 63; Lawson et al., 1995: 113. CURRENT PLACEMENT/NAME. Not determined; see comments. SERIES REFERENCE. Price et al., 1986: 79. PLACEMENT/NAME. orbiculare an COMMENTS. Status and disposition uncertain; whereabouts of type CURRENT Epithet orthographic variant of orbiculatum material uncertain; protologue (Pilger, 455) based on a collection (see below). from Annobon (Pagalu). COMMENTS. See comments for following entry. minutula (Fosliella, Melobesia) orbiculatum (Lithophyllum, Lithothamnion, Pseudolitho- SERIES REFERENCES. Price et al., 1986: 92; John et al., 1994: 72; phyllum) Lawson et al., 1995: 106 (under Pneophyllum confervicold). SERIES REFERENCES. John et al., 1994: 63, 68; Lawson et al., 1995: CURRENT PLACEMENT/NAME. Heterotypic synonym of Pneo- 113. phyllum confervicola (Kiitzing) Y.M. Chamberlain, according to CURRENT PLACEMENT/NAME. Lithophyllum orbiculatum (Foslie) Chamberlain (702: 137-141). Foslie, according to Chamberlain et al. (1991). COMMENTS. Conclusion of Chamberlain (702) based on studies COMMENTS. Conclusion of Chamberlain et al. (1991) based on (Chamberlain, 702, \994b) of TRH holotype (Woelkerling, 678) study of TRH lectotype (Woelkerling, 678) from Norway; the from Norway. binomial Crodelia orbiculare (Foslie) Kylin (in Price et al., 1986: neoatalyense (Lithophyllum) 79) pertains to Lithophyllum orbiculatum, but Kylin (281 : 208) used the spelling orbiculare and not orbiculatum. SERIES REFERENCES. Not previously reported. orotavicum (Goniolithon, Neogoniolithon) CURRENT PLACEMENT/NAME. Lithophyllum neoatalyense T. Lithophyllum, Masaki, to Chamberlain (737: 210). 1 according SERIES REFERENCES. Price et al., 1988: 23 ; John et al., 1994: 63, 79. NEW RECORD. Namibia (737). PLACEMENT/NAME. orotavicum COMMENTS. Conclusion of Chamberlain (737) based on study of CURRENT Neogoniolithon (Foslie) Me. andAfonso-Carrillo (1 1: HAK holotype from Japan. Lemoine, according toAdey (1970: 9) 133), but see comments. nephalidioides (Dermatolithon) COMMENTS. Conclusions of Adey (1970) and Afonso-Carrillo SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 64 (11) based on examination ofTRH portion of holotype (Woelkerling, (under Lithophyllum pustulatum). 678) from the Canary Islands; placement in Neogoniolithon as CURRENT PLACEMENT/NAME. Orthographic variant of Dermato- delimited by Penrose (1992, 1996c) and by Penrose & Chamberlain of lithon hapalidioides, a heterotypic synonym of Lithophyllum (1993), however, requires confirmation; data on PC portion pustulatum. Comments: See comments under entry for 'hapali- holotype provided by Woelkerling (678). dioides' above. pupil lost i in (Dermatolithon, Goniolithon, Lithophyllum, Titano- oligocarpum (Lithophyllum, Porolithon) derma) SERIES REFERENCES. John et al., 1994: 63; Lawson et al., 1995: SERIES REFERENCES. Price et al., 1986: 86; Price et al., 1988: 231; 1 12. Current placement/name: Not determined; see comments. John et al., 1994: 63; this paper. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 139 CURRENT PLACEMENT/NAME. Lithophyllum papillosum(Zanardmi Millepora polymorpha L., is a superfluous name for Phymatolithon ex Hauck) Foslie, according to Huve (272) and Babbini & Bressan calcareum (Pallas) W.H. Adey & D.L. McKibbin; details provided but see comments. (753: 307), by Woelkerling & Irvine (1986a); epithet polymorphum widely misapplied (Woelkerling & Irvine, 1986a) to plants referable to COMMENTS. Conclusion of Huve (272) based on study of L Phymatolithon purpureum (P. & H. Crouan) Woelkerling & L. Irvine lectotype (Woelkerling & Verheij, 1995) from the Adriatic Sea; (see entry for purpureum below). placement inLithophyllum as delimited by Woelkerling & Campbell (1992) and Woelkerling (1996), however, requires confirmation; polymorphum f. sublaevis (Phymatolithon) additional comments provided by John et al. (1994: 63) and SERIES REFERENCE. Lawson et al., 1995: 103. Woelkerling (1988: 216-217). CURRENT PLACEMENT/NAME. Superfluous name for Phymatolithon papillosum van cystoseirae (Dermatolithon, Titanoderma) polymorphum f. papillata Foslie, according to Woelkerling (678: SERIES REFERENCES. John et al., 1994: 61 (under Lithophyllum 211); also see comments. cystoseirae); this paper (under Titanoderma cystoseirae). COMMENTS. Status and disposition ofPhymatolithon polymorphum CURRENT PLACEMENT/NAME. Homotypic synonym of Litho- f. papillata Foslie uncertain;TRH lectotype (Woelkerling, 678: 1 68) phyllum cystoseirae (Hauck) Heydrich. from Helgoland, Germany not studied in detail in a modern context. COMMENTS. See comments under entry for 'cystoseirae' above ponderosum (Lithophyllum, Lithothamnion) and those by Babbini & Bressan (753: 307). SERIES REFERENCES. John et al., 1994: 62 (under Lithophyllum philippii (Lithothamnion, Mesophyllum) incrustans), 64, 68; this paper (under Spongites africanum). see SERIES REFERENCE. John et al., 1994: 68, 75. CURRENT PLACEMENT/NAME. Not determined; comments. CURRENT PLACEMENT/NAME. Not determined; see comments. COMMENTS. Status and disposition uncertain; TRH holotype (Woelkerling, 678: 168) from Sao Tome not studied in detail in a COMMENTS. Status and disposition uncertain; TRH lectotype modern context; additional comments provided by John et al. ( 1 994: (Woelkerling, 678: 171) from Italy not studied in detail in a modern 68). context; additional comments provided by John et al. (1994: 68). proboscideum (Lithophyllum) phyllactidium (Hapalidium) SERIES REFERENCES. John et al., 1994: 64; this paper (under SERIES REFERENCE. Lawson et al., 1995: 106 (under Pneophyllum Spongites africanum). confervicolum). CURRENT PLACEMENT/NAME. Lithophyllum proboscideum Foslie, CURRENT PLACEMENT/NAME. name for Phyllactidium Superfluous according to Adey (1970: 5), but see comments. confervicola Kutzing [basionym of Pneophyllum confervicola COMMENTS. Status and disposition uncertain; conclusion of Adey (KUtzing) Y.M. Chamberlain], according to Woelkerling & Verheij (1970) verification via a modern, detailed of the (1995:69). requires study lectotype from California, U.S.A. in TRH (Woelkerling, 678: 176); COMMENTS. Nomenclatural data & provided by Woelkerling comments on West African record provided in this paper under entry also see under above. Verheij (1995); entry 'confervicola' for Spongites africanum. polycephalum (Dermatolithon, Goniolithon, Lithophyllum, purpureum (Phymatolithon) Titanoderma) SERIES REFERENCE. Lawson et al., 1995: 103. SERIES REFERENCES. Price et al., 1986: 86; Price et al., 1988: 231 ; CURRENT PLACEMENT/NAME. Phymatolithon purpureum (P. & H. John et al., 1994: 64; this paper. Crouan) Woelkerling & L. Irvine, according to Woelkerling & Irvine CURRENT PLACEMENT/NAME. Lithophyllum polycephalum Foslie, (1986). according to Woelkerling & Campbell (1992: 22-23). COMMENTS. Conclusion of Woelkerling & Irvine (1986a) based 1 based COMMENTS. Conclusion ofWoelkerling & Campbell ( 992) on study of CO lectotype from France. on examination of TRH holotype (Woelkerling, 678: 174) from the pustulata/pustulatum (Dermatolithon, Lithophyllum, Melobesia, Cape Verde Islands; additional data on holotype provided by Cham- Titanoderma) berlain (737, as Titanoderma.). SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 64, polyclonum (Dermatolithon, Lithophyllum, Titanoderma) 72; this paper. SERIES REFERENCES. Price et al., 1986: 86; John et al., 1994: 64; CURRENT PLACEMENT/NAME. Lithophyllum pustulatum (J.V. this paper. Lamouroux) Foslie, according to Woelkerling & Campbell (1992: CURRENT PLACEMENT/NAME. Not determined; see comments. 78). 1 based COMMENTS. Status and disposition uncertain; TRH holotype COMMENTS. Conclusion ofWoelkerling & Campbell ( 992) in from France and from earlier of (Woelkerling, 678: 175) fragmentary and not studied in detail a on study of CN lectotype study modern context. lectotype by Woelkerling et al. (1985). polymorphum (Lithothamnion, Phymatolithon) pustulatum f. australis (Lithophyllum) et 1994: 64 SERIES REFERENCES. John et al., 1994: 68; Lawson et al., 1995: SERIES REFERENCE. John al., (under Lithophyllum 103. Current placement/name: Superfluous name; see comments. pustulatum). PLACEMENT/NAME. Not determined; see comments. COMMENTS. Phymatolithon polymorphum (L.) Foslie, based on CURRENT 140 WJ. WOELKERLING ET AL. COMMENTS. Status and disposition uncertain; TRH lectotype COMMENTS. Status and disposition uncertain; whereabouts of type (Woelkerling, 678: 35) from Canary Islands not studied in detail in material uncertain; additional comments provided by John et al. a modern context. (1994: 72, under Melobesia solmsiana), Chamberlain (94: 351, under Fosliella farinosa f. callithamnioides) and Taylor (1939, racemus (Lithophyllum, Lithothamnion) under Fosliella farinosa var. solmsiana). SERIES REFERENCES. John et al., 1994: 64, 67, 74 (under solmsii Mesophyllum brachycladum). (Melobesia) CURRENT PLACEMENT/NAME. Lithophyllum racemus (Lamarck) SERIES REFERENCES. Price et al., 1986: 92 (under Fosliella fari- Foslie, according to Basso et al. (1996). nosa); John et al., 1994: 72. etal. 1 select- COMMENTS. Conclusion of Basso ( 996) based on the CURRENT PLACEMENT/NAME. Invalid name, according to Woel- ion and study of a neotype (from Capri, Italy) housed at the Diparti- kerling (678). mento di Scienze della Terra, Istituto di Geologia e Paleontologia, COMMENTS. Original presentation lacks a description or diagnosis Universita di Milano, in Milano, Italy; heterotypic synonyms (ac- rendering name invalid (Woelkerling, 730); name possibly a variant cording to Basso et al., 1996) include Lithothamnion crassum of Melobesia solmsiana, but contrary to John et al. (1994: 72, under Philippi, Lithothamnion calcareum f. crassa (Philippi) Me. Lemoine, Melobesia solmsii), cannot be considered a homotypic synonym. and Lithophyllum duckeri Woelkerling; all West African records under these names to confirm identification as require checking solutum (Lithophyllum, Lithothamnion, Mesophyllum) misapplication of epithets likely in some or all cases; further infor- SERIES REFERENCE. John et al., 1994: 65, 68, 75. mation on the Lamarck species provided by Woelkerling (729) under the basionym (Millepora racemus). CURRENT PLACEMENT/NAME. Not determined; see comments. retusum (Lithophyllum) COMMENTS. Status and disposition uncertain; lectotype from Adri- SERIES REFERENCES. Lawson & John, 350: 237; Lawson & John, atic Sea in TRH (Woelkerling, 678: 203, under Lithothamnion 586: 207; John et al., 1994:65. fruticulosum f. soluta) not studied in detail in a modern context; additional comments provided by John et al. (1994: 68). CURRENT PLACEMENT/NAME. Lithophyllum retusum (Foslie) Foslie, according to Adey (1970: 5), but see comments. sonderi (Lithothamnion) COMMENTS. Status and uncertain, conclusion of disposition Adey SERIES REFERENCE. John et al., 1994: 68. (1970) requires verification via a detailed modern study of holotype CURRENT PLACEMENT/NAME. Lithothamnion sonderi from Sao Tome in TRH (Woelkerling, 678: 189); information on PC Hauck, ac- to Chamberlain isotype provided by Woelkerling (730). cording (750: 191). samoense (Hydrolithon) NEW RECORD. Canaries (750). SERIES REFERENCES. Not previously reported. COMMENTS. Conclusion of Chamberlain (750) based on detailed study of lectotype in L (Woelkerling & Verheij, 1995: 77) from CURRENT PLACEMENT/NAME. Hydrolithon samoense (Foslie) Keats Helgoland, Germany. & Y.M. Chamberlain, according to Keats & Chamberlain (754: 15). stictaeformis NEW RECORDS. Canaries (702;751). (Melobesia) COMMENTS. Conclusion of Keats & Chamberlain (754) based on SERIES REFERENCE. John et al., 1994: 73. study of TRH lectotype (Woelkerling, 678: 193) from Samoa. CURRENT PLACEMENT/NAME. Not determined; see comments. sauvageauii (Litholepis, Lithoporella, Melobesia) COMMENTS. Status and disposition uncertain; protologue SERIES REFERENCES. John et 1994: 72. al., 59, 66, (Areschoug, 1852: 517) based on mediterranean material not stud- ied in detail in a modern context; material in TRH CURRENT PLACEMENT/NAME. Lithoporella sauvageauii (Foslie) possible syntype 678: 207); additional material but not con- W.H. Adey, according to Adey (1970: 15), but see comments. (Woelkerling, presumed firmed to be in LD (John et al., 1994: 73). See also discussion in COMMENTS. Status and uncertain; conclusion of disposition Adey Babbini & Bressan (753: 130, under Lithophyllum frondosum). (1970) requires verification via a detailed modern study of holotype from the Canaries in TRH (Woelkerling, 678: 195). subtenellum (Goniolithon, Lithophyllum, Lithothamnion) simile (Lithophyllum) SERIES REFERENCES. Lawson & John, 350: 238; Lawson & John, 586: 208; Price et al., 1988: 231; John et al., 1994: 65, 68. SERIES REFERENCES. Lawson & John, 350: 237; Lawson & John, 586: John et 1994: 65. 208; al., CURRENT PLACEMENT/NAME. Lithophyllum subtenellum (Foslie) Foslie, to (1970: 6), but see comments and also CURRENT PLACEMENT/NAME. Lithophyllum simile Foslie, accord- according Adey Babbini & Bressan (753: 268, under Lithothamnion orbiculatum). ing to Adey (1970: 6), but see comments. COMMENTS. Status and conclusion of COMMENTS. Status and disposition uncertain; conclusion of Adey disposition uncertain; Adey verification via a detailed modern of (1970) requires verification via a detailed modern study of holotype (1970) requires study lectotype from France in TRH 678: from Sao Tome in TRH (Woelkerling, 678: 201). (Woelkerling, 215). solmsiana (Melobesia) tenuissimum (Lithothamnion, Phymatolithon) SERIES REFERENCES. Lawson & John, 350: 234; John et al., 1 994: SERIES REFERENCES. Lawson & John, 350: 242; Lawson & John, 72. Current placement/name: Not determined; see comments. 586: 212; John et al., 1994: 68; Lawson et al., 1995: 103. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 141 CURRENT PLACEMENT/NAME. Phymatolithon tenuissimum(Fos\\e) 1 1 6, 1 92) cite Tenarea tortuosa (Esper) Me. Lemoine as an accepted W.H. Adey, according to Adey (1970: 29), but see comments. species and as a synonym under Lithophyllum lichenoides. COMMENTS. Status and conclusion of disposition uncertain; Adey vickersiae (Lithophyllum, Lithothamnion, Pseudolithophyllum) (1970) requires verification via a detailed modern study of holotype SERIES REFERENCES. Johnetal., 1994: 65, 68; Lawson etal., 1995: from Sao Tome in TRH (Woelkerling, 678: 222); additional com- 113. ments provided by Lawson et al. (352: 103) and Babbini & Bressan under Lithothamnion (753: 268, tenuissimum). CURRENT PLACEMENT/NAME. Lithophyllum vickersiae Me. Lemoine, but see comments. thuretii (Choreonema) COMMENTS. Status and disposition uncertain; placement here in SERIES REFERENCE. Price et al., 1986: 69. Lithophyllum based on data on type material provided by Afonso- CURRENT PLACEMENT/NAME. Choreonema thuretii (Bornet) F. Carrillo (11: 139, as Pseudolithophyllum); protologue (Lemoine, Schmitz, according to Woelkerling (1987). 362: 42) based on seven collections from Canary Islands (all pre- sumably in C) but lectotype not yet designated or studied in detail in COMMENTS. Conclusion of Woelkerling ( 1 987) based on study of a modern context; additional comments provided by John et al. PC lectotype (Woelkerling, 730, under the basionym, Melobesia (1994:65). thuretii Bornet) from France. wildpretii (Spongites) tortuosa/tortuosum (Lithophyllum, Tenarea) See under Spongites wildpretii. SERIES REFERENCES. Lawson & John, 350: 238; Lawson & John, 586: John et 1994: this 210; al., 65; paper. ACKNOWLEDGEMENTS. We acknowledge with thanks assistance from many sources, of which the principal are: (i) Department of Botany, The CURRENT PLACEMENT/NAME. Tenarea tortuosa (Esper) Me. Natural History Museum, London, for provision of the necessary research Lemoine, according to Woelkerling et al. (1985). facilities to continue the series of papers of which the present is the terminat- COMMENTS. Conclusion of Woelkerling et al. (1985) based on a ing part; (ii) A grant (for GWL) towards this work from The Systematics of Natural detailed study of the FR lectotype from an unspecified locality in the Association; and (iii) Marian Short, Department Botany, The for careful work in this Mediterranean Sea; additional comments provided under entry for History Museum, London, her exceptionally editing Tenarea tortuosa in main part of this paper. Babbini & Bressan (753: paper. NUMERICAL LIST OF REFERENCES 2 Acuna Gonzales, 1970 41 Barton, 1897 1 1 5 Cullinane & Whelan, 1 982 5 Acuna Gonzales et al., 1970 42 Barton, 1901 117 Dangeard, 1948 6 Adey & Lebednik, 1967 44 Benftez, 1928 118 Dangeard, 1949 8 Afonso-Carrillo, 1980a 48 Bodard, 1965 122 Dangeard, 1952 9 Afonso-Carrillo, 19806 49 Bodard, 1966o 123 Dangeard, 1958 11 Afonso-Carrillo, 1984 50 Bodard, 19666 124 Dawson, 1956 13 Afonso-Carrillo & Gil-Rodriguez, 55 Bodard, 197 la 128A Delgado et al., 1986 1980 57 Bodard, 19716 129 De May et al., 1977 16 Afonso-Carrillo etal., 1979 58 Bodard, 1971c 131 DeToni, 1897 17 Afonso-Carrillo, Gil-Rodriguez & 59 Bodard & Mollion, 1974 133 DeToni, 1903 134 1905 Wildpret de la Torre, 1984 60 B0rgesen, 1916 DeToni, 18 Afonso-Carrillo, Gil-Rodriguez, 61 B0rgesen, 1917 138 DeToni, 1910 Haroun Tabraue, Villena Balsa & 63 B0rgesen, 1919 139 DeToni, 1924 140 De Toni & Forti, 1913 Wildpret de la Torre, 1984 66 B0rgesen, 1925 1888 20 Agardh, 1824 68 B0rgesen, 1927 141ADeToni&Levi, 1929 145 Dickie, 1874 21 Agardh, 1828 70 B0rgesen, 150 1877 25 Agardh, 1852 71 B0rgesen, 1930 Dickie, 153 Dickinson & Foote, 1950 26 Agardh, 1863 72 B0rgesen, 1931 1 54 Dickinson & Foote, 1 95 1 27 Agardh, 1876 83 B0rgesen, 1945 1926 30 Aleem, 1978 89 Bornet, 1892 167 Dinter, 169 1927 33 Ardre, 1970 90 Bory de St. Vincent, 1803 Dinter, 172 Dixon 1977 34 Ardre, 1980 93 Carpine, 1959 & Irvine, 177 Edelstein. 1964 36B Wynne, 1986 94 Chamberlain, 1983 1963 179 1901 37 Askenasy, 1888 97 Chapman, Falkenberg, 1920 182 Feldmann & Bodard, 1965 38 Askenasy, 1897 99 Chevalier, 183 1935 38B Audiffred & Weisscher, 1984 100 Chevalier, 1935 Feldmann, 184 Feldmann, 1937 38C Audiffred, 1985 105 Cinelli & Codomier, 1974 1938 38D Audiffred & Prud'homme van Reine, 107 Colman & Stephenson, 1966 186 Feldmann, 1985 1985 108 Cordeiro-Marino, 1978 187 Gabrielson, 188 Feldmann, 1939 39 Baardseth, 1941 113 Cribb, 1983 142 W.J. WOELKERLING ET AL. 189 Feldmann, 1941 297 John &Pople, 1973 448 Piccone, 1889 190 Feldmann, 1942 299 John et al., 1977 450 Piccone, 1900 191 Feldmann, 1946 300 John etal., 1980 451 Piccone, 1901 192 Feldmann, 1951 302 Johnston, 1969 453 Pilger, 1908 194 Feldmann & Hamel, 1934 304 Johnston, 1969/7 455 Pilger, 1919 196 Feldmann-Mazoyer, 1941 306 Johnstone & Croall, 1859 456 Pilger, 1920 197 Foslie, 1897 306B Jorge etal., 1986 457 Pilger, 1922 198 Foslie, 1900a 311 Karsakoff, 1896 474 Price & John, 1978 199 Foslie, 19006 312A Kensley & Penrith, 1980 475 Price & John, 1980 205 Foslie, 1906 316 Kutzing, 1843 477 Prud'homme van Reine, 1984 206 Foslie, 1907a 318 Kutzing, 1849 482 Reinbold, 1907 207 Foslie, 1907/7 320 Kutzing, 1862 487 Round, 1981 209 Foslie, 1908a 323 Kutzing, 1865 489 Santos Guerra, 1972 210 Foslie, 1908/7 329 Kylin, 1932 490 Santos Guerra et al., 1970 211 Foslie, 1909 336 Lawson, 1954 491 Sanusi, 1980 212 Printz, 1929 338 Lawson, 1956 493 Sauvageau, 1912 214 Fremy, 1936 339 Lawson 1957 496 Schmidt, 1924 217 Ganesan& West, 1975 340 Lawson, 1957/7 498 Schmidt, 1929 222 Gayral, 1966 341 Lawson, 1960 499 Schmidt, 1931 225 Gil-Rodriguez, 1980 344 Lawson, 1966 500 Schmidt & Gerloff, 1957 226 Gil-Rodriguez & Afonso-Carrillo, 346 Lawson, 1980a 501 Schmitz & Falkenberg, 1897 1980 347 A Lawson, 1985 502 Schmitz & Hauptfleisch, 1896-1897 227 Gil-Rodriguez & Afonso-Carrillo, 348 Lawson etal., 1990 512 Schneider, 1983 1981 349 Lawson & John, 1977 516 Seoane-Camba, 1960 229 Gil-Rodriguez &Wildpretde laTorre, 350 Lawson & John, 1982 517 Seoane-Camba, 1965 1980o 351 Lawson & Norton, 1 97 1 523 Simons, 1974 230 Gil-Rodriguez &Wildpretde laTorre, 352 Lawson etal., 1975 528 Sender, 1852 1980/7 353 Lemoine, 1911 529 Sourie, 1954 231 Gil-Rodriguez et al., 1978 354 Lemoine, 1912 532 South & Whittick, 1976 232B Gil-Rodriguez et al., 1985 356 Lemoine, 1915 535 Steentoft, 1967 235 Gonzalez Henriquez, 1976 359 Lemoine, 1924 537 Stephenson & Stephenson, 1972 236 Gonzales, 1977a 360 Lemoine, 1926 538 Svedelius, 191 1 237 Gonzales, 19776 361 Lemoine, 1928 542 Trochain, 1940 239 Goor, 1923 362 Lemoine, 1929a 543 Tseng, 1944 240 Gordon, 1972 363 Lemoine, 1929/7 546 Varo et al., 1979 249 Hariot, 1895 365 Lemoine, 1935 547 Vickers, 1897 250 Hariot, 1896 366 Lemoine, 1964 553 Weber-van Bosse, 1921 251 Hariot, 1908 367 Lemoine, 1965 555 Weisscher, 1982 252 Hariot, 1911 368 Lemoine, 1966 556 Weisscher, 1983 253 Haroun Tabraue et al., 1984 371 Lemoine, 1974 559 Wollaston, 1984 254 Harvey, 1846-1851 375 Levring, 1974 562 Wynne & Taylor, 1973 259 Hemsley, 1885 376 Lieberman et al., 1979 565A Yarishetal., 1985 261 Henriques, 1885 377 Lieberman etal., 1984 569 Anderson & Bolton, 1985 263 Henriques, 1886 379 Lopez Hernandez & Gil-Rodriguez, 570 Seagrief, 1984 264 Henriques, 1887 1982 572 Chang & Xia, 1963 265 Henriques, 1917 380 Lopez Hernandez et al., 1986 576 Afonso-Carrillo, 1985 266 Heydrich, 1894 387 May, 1912 582 Afonso-Carrillo et al., 1985 268 Hooker & Harvey, 1847 389 Mazza, 1903 583 Haroun Tabraue et al., 1985 269 Hoppe, 1969 390 Mazza, 1905-1925 584 Ribera Siguan et al., 1985 271 Hornemann, 1819 392 Menez & Mathieson, 1981 586 Lawson & John, 1987 271 A Huisman, 1985 397 Mildbread, 1922 589 Guiry, 1984 272 Huve, 1962 399 Mollion, 1976 590 John & Lawson (unpublished) 273 Irvine, 1983 401 Montagne, 1839-1841 596 Bailey & Harvey, 1 862 281 Kylin, 1956 402 Montagne, 1842 597 Prud'homme van Reine & Lobin, 1 986 286 John, 1972a 407 Montagne, 1856 598 Prud'homme van Reine (in lift., 10 287 John, 1972/7 408 Montagne, 1860 April 1987) 288 John, 1977 410 Murray, 1888-1889 624 Marcot-Coqueugniot, 1991 290 John, 1986 411 Naegele, 1960 625 Prud'homme van Reine & van den 292 John&Lawson, 1972a 417A Norris&Aken, 1985 Hoek, 1988 293 John&Lawson, 1972/7 430 Papenfuss, 1944 633 Pinedoetal., 1992 294 John&Lawson, 1974 432 Papenfuss, 1964 634 Elejabeitia et al., 1992 295 John & Lawson, 1977a 434 Papenfuss, 1968 635 Prud'homme van Reine (in litt., 296 John&Lawson, 1977/7 439 Piccone, 1884 Heincke Expedition, 1991) RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 143 639 Prud'homme van Reine & Pakker (in 699 Sanson& Reyes, 1995 733 Afonso-Carrillo, 1988 lift., 1991) 701 Chamberlain & Irvine, 19946 734 Perez & Afonso-Carrillo, 1993 644 Lawsonetal., 1993 702 Chamberlain, 19946 736 Chamberlain & Irvine, 1994a 646 Sanson etal., 1991 703 Evans etal., 1993 737 Chamberlain, 1996 648 Ballesteros et al., 1992 707 Stegenga etal., 1997 738 Chamberlain, 1994a 649 Irvine & Chamberlain, 1994 711 Maggs & Hommersand, 1993 740 Reyes & Afonso-Carrillo, 1995 652 Otero-Schmitt & Sanjuan, 1992 712 Womersley, 1994 742 Chamberlain & Keats, 1995 654 John & Lawson, 1991 713 Otero-Schmitt, 1994 743 Chamberlain & Keats, 1994 657 Afonso-Carrillo et al., 1992 714 Codomier, 1973 744 Reyes etal., 1988 662 Viera-Rodrfguez et al., 1987 716 Molloy, 1990 745 Reyes & Sanson, 1991 663 Prud'homme van Reine etal., 1994 717 Febles etal., 1995 746 Silvaetal., 1996 664 Harounetal., 1993 718 Ganesan, 1974 747 Guadeloupe Gonzalez et al., 1995 668 Athanasiadis, 1987 719 Maggs &Guiry, 19826 748 Soler-Onis et al., manuscript 677 Romanes, 1916 720 Pakker & Breeman, 1996 749 Soler-Onfs et al., 1995 678 Woelkerling, 1993 721 Martin etal., 1996 750 Chamberlain, 1992 683 Otero-Schmitt, 1993 722 Harper & Garbary, 1 997 75 1 Sanson & Reyes, 1996 684 Kristiansen et al., 1993 724 Reyes etal., 1993 752 Reyes & Sanson, 1996 685 Medina & Haroun, 1993 726 Afonso-Carrillo etal., 1993 753 Babbini & Bressan, 1997 687 Reyes & Afonso-Carrillo, 1993 729 Woelkerling, 1998a 754 Keats & Chamberlain, 1994o 695 Hardy & Seku, 1993 730 Woelkerling, 19986 755 Keats & Chamberlain, 19946 698 Rojas-Gonzales etal., 1994 732 Gabrielson & Hommersand, 1982 756 Barabara et al., 1992 Notes, i) The Corallineae (Ordo XII. pp. 506-576) was by J.E. Areschoug. ii) Some REFERENCES - versions of this second volume, Pars II, were issued as two separate texts Pars II: 1 being dated 1851 and Pars 11:2 1852. iii) There are also internal differences of Acuna Gonzalez, A. 1970. Algunos aspectos de la vegetacion submarina de las Islas numbering of pages between copies. BM copy is numbered straight through from Canarias. Vieraea[l]: 2-5. 337-720, including therein the Addenda [701-706] and Index [707-720]. In the Santos Guerra, A. & Wildpret de la Torre, W. 1970. Algunos aspectos de la copy from which the Cramer (1977) reprint was prepared, the Index [unnumbered, of vegetacion algal de la Playa de San Marcos, Icod, Tenerife. Cuadernos Botanica el 14 sides] is placed immediately after p. 700 and is followed by six sides [also Museo Canada 9: 30-36. unnumbered] of Addenda. Content of these unnumbered sides is exactly as the Adey, W.H. 1970. A revision of the Foslie crustose coralline herbarium. Kongelige numbered BM pages. 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Bulletin of RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 149 the British Museum (Natural History), Botany. 18: 195-273. Naturlichen Planzenfamilien .... I. Teil. 2. Abteilung: 149-150. Leipzig. 1 992. Seaweeds of the western coast of tropical Africa and adjacent & Hauptfleisch, P. 1 896- 1 897. Sphaerococcaceae In A. Engler & K. Prantl, Die islands: a critical assessment. IV. 3. Genera H-K. Bulletin Naturlichen . . . I. Teil. 2. Rhodophyta (Florideae) of pflanzenfamilien , Abteilung: 382-396. Leipzig. the British Museum (Natural History), Botany, 22: 123-146. Schneider, C.W. 1983. The red algal genus Audouinella Bory Nemaliales; H. 1 929. Foslie - Printz, (Ed.) M. 'Contributions to a Monograph of the Lithothamnia' . (Acrochaetiaceae) from North Carolina. Smithsonian Contributions on Marine Del Norske Kongelige Videnskabers Selskab Museet. Trondheim. Science 22: iii+[ 1 ]+25. van Prud'homme Reine, W. 1984. Marine algae of the Cape Verde Islands, historical Scott, F.J., Wetherbee, R. & Kraft, G.T. 1984. The morphology and development of and notes. preliminary Courier Forschungsinstitul Senckenberg 68: 135-137. some prominently stalked southern Australian Halymeniaceae (Cryptonemiales, 1987. Extractions of red report on algal distribution patterns, in lilt to J.H. Price, Rhodophyta) II. The sponge-associated genera Thamnoclonium Kiitzing and 10 April 1987. Codiophyllum Gray. Journal of Phycology 20: 286-295. 1 99 1 . reallocation Heincke Expedition, following examination of original Piccone Seagrief, S.C. 1984. A catalogue of South African green, brown and red marine algae. material from the Canary Islands, in litt. to D.M. John, 19 November 1991. Memoirs of the Botanical Survey South Africa 47: i-vi+1-71. Haroun, R.J. & Audiffred, P.A.J. 1994. A reinvestigation of Macaronesian Seoane-Camba, J. 1960. Nota sobre algunas especies de algas de la costa occidental seaweeds as studied by A. Piccone with remarks on those studied by A. Grunow. africana (sur de Cabo Blanco). Investigacion Pesquera 16: 91-103. Nova Hedwigia 58: 67-121. 1965. Estudios sobre las algas bentonicas en la costa sur de la Peninsula Iberica Hoek, C. van den 1988. Biogeography of Cape Verdean seaweeds. Courier (literal de Cadiz). Investigacion Pesquera 29: 3-216. Forschungsinstitut Senckenberg 105: 3549. Silva, P.C., Basson, P.W. & Moe, R.L. 1996. Catalogue of the benthic marine algae of & Lobin, W. 1986. Katalog der von den Kapverdischen Inseln beschriebenen the Indian Ocean. University of California Publications in Botany 79: 1-1259. Taxa von Algen (Algae: Chlorophyceae, Phaeophyceae & Rhodophyceae). Courier Menez, E.G. & Moe, R.L. 1987. Catalog of the benthic algae of the Philippines. Forschungsinstitut Senckenberg 81: 85-88. Smithsonian Contributions to Marine Science 27: 1-179. & Pakker, H. 1 99 1 . Collections from Cape Verde Islands, in litt. to D.M. John, 1 9 Simons, R.H. 1974. Algae (including diatoms and seaweeds). In J.H. Day, N.A.H. November 1991. Millard, & M.-L. Penrith, A guide to marine life on South African shores. 2nd ed.: Pueschel, C.M. 1 989. An expanded survey of the ultrastructure of red algal pit plugs. 239-261. Cape Town. Journal of Phycology 25: 625-636. 1983. The genus Trematocarpus (Sarcodiaceae, Rhodophyta) in southern Africa - Reinbold, T. 1 907. Die Meeresalgen der deutschen Tiefsee Expedition 1 898- 1 899. In and the exclusion of Sphaerococcus (Chondrus scutellatus. Bothalia 14: 803-808. C. Chun, Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Soler-Om's, E., Haroun, R.J. & Prud'homme van Reine, W.F. 1998. Mscr. Sebdenia Dampfer 'Valdivia' J 898-1 899. Bd. II, Teil II, Lieferung IV: 549-586+[8], pis VV- in Macaronesia. LVII. Jena. Viera Rodriguez, A. & Prud'homme van Reine, W.F. 1995. Sebdenia Reyes, J. & Afonso-Carrillo, J. 1993. Morphology and anatomy of Mesophyilum canariensis sp. nov. (Sebdeniaceae, Gigartinales, Rhodophyta), una nueva alga roja canariense (Corallinaceae, Rhodophyta) from the Canary Islands. Courier de profundidad delArchipielago Canario. Resumenes de comunicaciones. XlSimposio Forschungsinstitut Senckenberg 159: 27-132. Nacional de Botanica Criptogdmica, Santiago de Compostela, 18-21 de Septiembre 1995. Morphology and distribution of nongeniculate coralline algae de 1995: 15-16. (Corallinaceae, Rhodophyta) on the leaves of the seagrass Cymodocea nodosa Sender, O.W. 1852. Algae. In J.A. Schmidt, Beitrage zur Flora der Cap Verdischen (Cymodoceaceae). Phycologia 34: 179-190. Inseln. Mil Beriicksichtigung aller bisjetzt daselbst bekannten wildwachsenden und & Wildpret de la Torre, W. 1988. Structure of male conceptacles of kultivirten. Pflanzen Nach eigenen Untersuchungen und mil Benutzung der Lithophyilum lobatum (Corallinaceae, Rhodophyta). Actesdel Simposi International gewonnenen Resultate anderer Reisenden: 125-127. Heidelberg. de Botdnico Pius i Quer, 1988 l(criptogamia): 137-141. Sourie, R. 1954. Contribution a 1'etude e'cologique des cotes rocheuses du Senegal. & Sanson, M. 1991. Adiciones a la flora marina de la isla de El Hierro (Islas Memoires de I'lnstitut Francois d'Afrique Noire 38: l-342+[l]. Canarias). Vieraea 20: 71-81. Note. From the note on p. 117, it is clear that the algae were worked on mainly by J. 1996. Las algas epifitas en Cymodocea nodosa en El Medano, isla de Feldmann, but that Sourie took account of some of the views of Dangeard as Tenerife (Magnoliophyta, Cymodocea [.sic!]). Vieraea 25: 45-56. expressed in the latter's memoir on the Cap Vert (Dakar) peninsula algae. Since the & Afonso-Carrillo, J. 1993. Notes on some interesting marine algae new exact contribution of the various people involved is in doubt, we have left the from the Canary Islands. Cryptogamic Botanv 4: 50-59. reference in the name of Sourie, who seems to have exercised overall authorship. Ribera Siguan, M.A., Gomez Garreta, A. & Seoane-Camba, JA. 1985 ['1984']. South, G.R. & Tittley, I. 1986. A checklist and distributional index of the benthic Estudio biogeografico de la flora algologica bentonica marina de las Islas Baleares. marine algae of the North Atlantic Ocean. St. Andrews, Canada. Anales de Biologia Universidad de Murcia 2 (Seccion especial, 2): 147-159. & Whittick, A. 1976. Aspects of the life history of Rhodophysema elegans Rojas-Gonzales, B., Afonso-Carrillo, J. & Ibeas, C. 1994. New records of Rhodo- (Rhodophyta, Peyssonneliaceae). British Phvcological Journal 11: 349-354. melaceae (Rhodophyta) from the Canary Islands. Botanica Marina 37: 133-138. Steentoft, M. 1967. A revision of the marine algae of Sao Tome' and Principe (Gulf of Romanes, M.F. 1916. Note on an algal limestone from Angola. Transactions of the Guinea). Journal of the Linnean Society (Botany) 60: 99-146. 1 1997. the South Royal Society of Edinburgh 51: 581-584, pi. Stegenga, H., Bolton, J.J. & Anderson, R.J., Seaweeds of African Rosenvinge, L.K. 1917. The marine algae of Denmark. Part II. Rhodophyceae II. West Coast. Cape Town. (Cryptonemiales). Def Kongelige Danske Videnskabernes Selskabs Skrifter, 1 Raekke, Stephenson, T.A. & Stephenson, A. 1 972. Life between tidemarks on rocky shores. San Naturvidenskabelig og Matematisk Afdeling 7: 155-283, pis 3-4. Francisco. Round, F.E. 1981. The ecology of algae. Cambridge. Svedelius, N. 1 9 1 1 . Florideae. In A. Engler & K. Prantl, Die natiirlichen Pflazenfamilien Sanson, M. & Reyes, J. 1995. Morphological and geographical observations on four . . . Nachtrdge zum l.Teil, 2. Abteilung iiber die Jahre 1890 big 1910: 200-276. species of Ceramiaceae (Rhodophyta) new to the Canary Islands. Botanica Marina Leipzig. 34(6): 89-95. Taylor, W.R. 1939. Algae collected on the Presidential cruise of 1938. Smithsonian 2 1996. Sobre la morfologia de Spyridia filamentosa y Spyridia hypnoides en Miscellaneous Collections 98(9): 1-18. pis. las islas Canarias (Rhodophyta, Ceramiaceae). Vieraea 25: 37^4. Tomita, N.Y. 1976. Contribuicao ao conhecimento do genero Sporolithon 1 no Brasil. Ph.D. Universidade & Afonso-Carrillo, J. 1 99 . Contribution to the seaweed flora of the Canary (Corallinaceae, Cryptonemiales) Unpublished thesis, Islands: new records of Floridiophyceae. Botanica Marina 34(6): 527-536. de Sao Paulo. Brazil. Santos Guerra, A. 1972. Contribucion al estudio de la flora marina de la isla de La Townsend, R.A. 1995. The Sporolithaceae: It's place in the Florideophyceae. Unpub- Gomera. Vieraea 2: 86-102. lished Ph.D. thesis, University of Sydney, Australia. Acuna Gonzales, A. & Wildpret de la Torre, W. 1970. Contribucion al estudio Trochain, J. 1940. Contribution a 1'etude de la vege'tation du Senegal. Memoires de de la flora marina de la Isla de La Palma. Cuadernos Botanica El Museo Canario 9: I'lnstitut Franqais d'Afrique Noire 2: [l-6]+l 433+[63]. 20-29. Note. J. Feldmann clearly had a great deal to do with the main determinations on which the list 108-1 was based: since the extent to which the data were Sanusi, S.S. 1 980. A study on grazing as a factor influencing the distribution ofbenthie algal (pp. 10) is not and since there are other to the littoral algae. M.Sc. Thesis, University of Ghana, Legon. accepted or amended by Trochain clear, parts to have been attributable to we have Sauvageau, C. 1912. A propos des Cystoseira de Banyuls et Guethary. Bulletin de la text which seem definitely Trochain, accepted on the more correct Station Biologique d'Arcachon 14: 133-556. the latter as overall author. For individual comments species, citation would be J., in J.,' etc. Schmidt, O.C. 1924. Index algarum marinarum 1920-1923. Hedwigia 65: 1 1-27. authorship undoubtedly 'Feldmann. Trochain. 1944. Notes on the Taenioma. Madrono 7: 215-226. 1929. Beitrage zur Kenntnis der Meeresalgen der Azoren II. Hedwigia 69: 165- Tseng, C.K. algal genus 172. Turner, J.A. & Woelkerling, WJ. 1982a. Studies on the Mastophora-Lithoporella 21: 201-217. 1931. Die marine Vegetation der Azoren in ihren Grundzugen dargestellt. complex (Corallinaceae. development. Phycologia Bibliotheca Botanica 102: 1-116. 19826. Studies on the Mastophora-Lithoporella complex (Corallinaceae, II. and 21: 218-235. &Gerloff,J. 1957. Die marineVegetationAfrikas in ihren Grundzugen dargestellt. Rhodophyta). Reproduction generic concepts. Phycologia Willdenowia 1: 709-756. Varo, J., Ramirez, J. & Renteria, J. 1979. Estudio de la vegetacion bentonica del literal Acta Botanica Malacitana 5: 79-98. Schmitz, F. & Falkenberg, P. 1897. Rhodomelaceae. In A. Engler & K. Prantl, Die granadino. 150 W.J. WOELKERLING ET AL. Part IIIB. Verheij, E. 1993. The genus Spomlithon (Sporolithaceae fam. nov., Corallinales, Southern Australia D Gracilariales, Rhodymeniales, Corallinales and Canberra. Rhodophyta) from the Spermonde Archipelago, Indonesia. Phycologia 32: 1 84-196. Bonnemaisoniales: 237-283. 1994. Nongeniculate Corallinaceae (Corallinales. Rhodophyta) from the \99Sa. 3. Lamarck's Nullipores. In W.J. Woelkerling & D. Lamy, Non-geniculate coralline and the Paris Museum. and Spermonde Archipelago, SW Sulawesi, Indonesia. Blumea 38: 95-137. red algae Cryptogamie-ADAC Museum ' National d'Histoire Naturelle, Paris. In Vickers, A. 1 897(?)( 1 896' ].Contribution a la flore algologiques des Canaries. Annales press. 1 of at Science Naturelles (Botanique), 8, 4: 293-306. 998ft. 4. Type collections non-geniculate Corallinales housed the Laboratoire Note. The date is somewhat difficult to cite as there is confusion regarding the dates de Cryptogamie (PC). In W.J. Woelkerling & D. Lamy, Non-geniculate coralline red 1 and of various issues. It does seem possible that pre-prints were issued in 896 and this algae and the Paris Museum. Cryptogamie-ADAC Museum National d'Histoire is the date usually cited (see Lawson & Price. 1969: 345-346). Naturelle, Paris. In press. Viera-Rodriguez, M.A., Gil-Rodriquez, M.C., Audiffred, P.A.J., Prud'homme van & Campbell, S.J. 1 992. An account of southern Australian species ofLithophyllum Reine, W.F., Haroun-Tabraue, R. & Wildpret de la Torre, W. 1987. Contribution (Corallinaceae, Rhodophyta). Bulletin of the British Museum (Natural History), al estudio de la florula bentonica del islote de Montana Clara. Canarias. Vieraea 17: Botany, 22: 1-107. 271-279. Chamberlain, Y.M. & Silva, P.C. 1985. A taxonomic and nomenclatural Vinassa, P.E. 1892. Coralline mediterranee raccolte dal Prof Meneghini. Ani della reassessment of Tenarea, Titanoderma and Dermatolithon (Corallinaceae, Rhodo- Societd Toscana di Scienze Natural! Residente in Pisa. Pmcessi Verbal! 8: 58-60. phyta) based on studies of type and other critical specimens. Phycologia 24: Weber-van Bosse, A. 1921. Liste des algues du Siboga II Rhodophyceae Premiere 317-337. In A. 1993. An account of southern Australian of partie Protoflorideae, Nemalionales, Cryptonemiales. M. Weber, Siboga-Expeditie & Harvey, species Mesophyllum Australian 6: 571-637. . . . Monographie LIXb. Uikomsten op Zoologisch, Botanisch. Oceanographisch en (Corallinaceae, Rhodophyta). Systematic Botany 1982. Batters Geologisch Gebied . . . Livr. LXXXIX: [6]+187+392+[4]. & Irvine, L.M. The genus Schmitziella Bornet & (Rhodophyta): Weisscher, F.C.M. 1982. Marine algae from Ilheu de Fora (Salvage Islands). Boletim Corallinaceae or Acrochaetiaceae? British Phycological Journal 17: 275-295. do Museo Municipal do Funchal 34: 23-34. 19860. The neotypification and status of Phymatolithon (Corallinaceae, 1983. Marine algae from Selvagem Peguena (Salvage Islands). Boletim do Museo Rhodophyta). British Phycological Journal 21: 55-80. Municipal do Funchal 35: 41-80. 1986ft. The neotypification and status of Mesophyllum (Corallinaceae, Wilks, K.M. & Woelkerling, W.J. 1991. Southern Australian species of Melobesia Rhodophyta). Phycologia 25: 379-396. (Corallinaceae, Rhodophyta). Phycologia 30: 507-533. Penrose, D. & Chamberlain, Y.M. 1993. A reassessment of type collections of 1995. An account of southern Australian species of Lithothamnion non-geniculate Corallinaceae (Corallinales, Rhodophyta) described by C. (Corallinaceae, Rhodophyta). Australian Systematic Botany 8: 549-583. Montagne and L. Dufour, and of Melobesia brassica-florida Harvey. Phycologia Woelkerling, W.J. 1 983a. A taxonomic reassessment ofLithothamnium (Corallinaceae, 32: 323-331. Rhodophyta) based on studies of R.A. Philippi's original collections. British & Verheij, E. 1 995. Type collections of nongeniculate Corallinales (Rhodophyta) Phycological Journal 18: 165-197. in the Rijksherbarium, Rijksuniversiteit te Leiden (L), The Netherlands. Blumea 40: 1983ft. A taxonomic reassement of Lithophyllum Philippi (Corallinaceae, Rhodo- 33-90. phyta) based on studies of R.A. Philippi's original collections. British Phycological Wollaston, E.M. 1984. Species of Ceramiaceae (Rhodophyta) recorded from the Journal 18: 299-328. International Indian Ocean Expedition, 1962. Phycologia 23: 281-299. 1 985. A taxonomic reassessment ofSpongites (Corallinaceae, Rhodophyta) based Womersley, H.B.S. 1994. The marine benthic flora of Southern Australia Part HA. on studies of Kiitzing's original collections. British Phycological Journal 20: 1 23- 1 53. Canberra. 1987. The genus Choreonema in southern Australia and its subfamilial classifica- 1996. The marine benthic flora of Southern Australia Part IIIB Flora ofAustralia tion within the Corallinaceae (Rhodophyta). Phycologia 26: 1 1 1-127. Supplementary Series 5. Canberra. 1988. The coralline red algae: an analysis of the genera and subfamilies of Wulfen,F.X. 1803.CryptogamaAquatica.Arc/2/v/M>d/ Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. 21 Volume Seaweeds of the western coast of tropical Africa and adjacent No. 1 Historical and taxonomic studies in the genus Titanoderma islands: a critical assessment. IV Rhodophyta (Florideae) 4. (Rhodophyta, Corallinales) in the British Isles. Y.M. Chamber- Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F. lain. 199 1 . Pp. 1-80, 247 figs. 4235 Prud'homme van Reine and WJ. Woelkerling. Pp. 49-90. Studies on the Cretan flora 3. Additions to the flora of No. 2 Early collections of the Holy Thorn (Crataegus monogyna cv. Karpathos. N.J. Turland and L. Chilton. Pp. 91-100. 43.25 Biflora). A.R. Vickery. 1991. Pp. 81-83, 1 fig. A taxonomic study of the species referred to the ascomycete genus No. 2 Observations on the benthic marine algal flora of South Leptorhaphis. B. Aguirre-Hudson. 1991 . Pp. 85-192, 76 figs. Georgia: a floristic and ecological analysis. D.M. John, P.J.A. The typification and identification of Calymperes Pugh and I. Tittley. Pp. 101-1 14. crassilimbatum Renauld & Cardot (Musci: Calymperaceae). Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical Ellis. L.T. 1991. Pp. 193-194, 1 fig. 42.35 species (excluding Australia). D.J. Galloway. Pp. 1 15-160. Morphology and ecology of seedlings, fruits and seeds of Volume 22 Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp. 161-172. No. 1 An account of southern Australian species of Lithophyllum A of Bixa (Bixaceae), with reference to the leaf (Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. study particular undersurface indumentum as a diagnostic character. R.E. Campbell. 1992. Pp. 1-107, 63 figs. 41.25 Dempsey and N.C. Garwood. Pp. 173-180. 43.40 No. 2 Palynological evidence for the generic delimitation of Sechium and its allies. J.L. R. Lira-Saade & (Cucurbitaceae) Alvarado, Volume 25 J. Caballero. 1992. Pp. 109-121. No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. Pp. Seaweeds of the western coast of Africa and tropical adjacent 1-94. islands: a critical assessment. IV. Rhodophyta (Florideae) 3. William Roxburgh's St Helena plants. Q.C.B. Cronk. Pp. 95-98. Genera H-K. J.H. Price, D.M. John & G.W. Lawson. 1992. pp. 43.40 123-146. No. 2 Seaweeds of the western coast of Africa and Two new species of Solarium section Geminata (Solanaceae) tropical adjacent islands: a critical assessment. IV. (Florideae) 5. from Cerro del Torr in western Colombia. S. Knapp. 1992. Pp. Rhodophyta 147-152. Genera P. G.W. Lawson, WJ. Woelkerling, J.H. Price, W.F. Prud'homme Van Reine and D.M. John. 99-122. Fissidens ceylonensis Dozy & Molkenb. (Musci: Pp. A new of Fissidentaceae) and some allied taxa from southern India. L.T. species Odontorrhynchos (Orchidaceae, Spiranthinae) from Boliva. D.L. Szlachetko. Pp. 123-125. Ellis. 1992. Pp. 153-156, 2 figs. Linnaeus's of De New species of Piper (Piperaceae) from Central America. M. interpretation Prospero Alpine's plantis exoticis, with on the flora of Crete. N.J. Tebbs. 1992. Pp. 157-158. special emphasis Turland. 127-159. Studies on the Cretan flora 1. Floristic notes. N.J. Turland. Pp. Book review. M.G. Gilbert. P. 161. 43.40 1992. Pp. 159-164. Studies on the Cretan flora 2. The Dianthus juniperinus Volume 26 complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169. 1 of 41.25 No. A morphological study Chaetoceros species (Bacillariophyta) from the plankton of the Pacific ocean of Mexico. D.U. Hernandez-Becerril. 1996. Pp. 1-73. 43.40 Volume 23 No. 2 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. No. 1 Revision of Piper (Piperaceae) in the New World 3. The Myriandra to 28. Elodes. N.K.B. Robson. 1996. Pp. 75-217. taxonomy of Piper sections Lepianthes and Radula. M.C. 43.40 Tebbs. 1993. Pp. 1-50, 18 figs. Mounting techniques for the preservation and analysis of Volume 27 diatoms. S.J. Russell. 1993. Pp. 51-54. 1 fig. 43.25 No. 1 Notes on the diatom species Tetracyclus castellum (Ehrenb.) No. 2 New taxa of Gentiana from Western China and (Gentianaceae) Grunow with a description of Tetracyclus pseudocastellum nov. the T.-N. Ho and S.-W. Liu. 1993. 55- Himalayan region. Pp. sp. D.M. Williams. Pp. 1-5. 60. 2 figs. A new species of Calymperes (Musci: Calymperaceae) from New names and taxonomic notes on Gentianella combinations, Peninsular Malaysia. L.T. Ellis. Pp. 7-9. from South America and New Zealand. T.-N. (Gentianaceae) A phylogenetic conspectus of the tribe Hyoscyameae and S.-W. Liu. 1993. 61-66. Ho Pp. (Solanaceae). A.L. Hoare and S. Knapp. Pp. 1 1-29. Studies in Hypericum: validation of new names. N.K.B. A revision of Solarium section Pteroidea: Solanaceae. S. Robson. 1993. 67-70. Pp. Knapp and T. Helgason. Pp. 3 1-73. 1997. 43.40 Generic monograph of the Asteraceae-Anthemideae. K. No. 2 Systematics of Pogostemon (Labiatae) G.R. Bhatti and M. Bremer and C.J. Humphries. 1993. Pp. 71-177. 12 figs. 43.25 Ingrouille. Pp. 77-147. 1997 43.40 Volume 24 Volume 28 No. 1 Pre-Linnaean references for the Macaronesian flora found in No. 1 Morphology and ecology of seedlings, fruits and seeds of Leonard Plukenet's works and collections. J. Francisco-Ortega, Panama^ Vochysiaceae. N.C. Garwood. Pp. 1-16. A. Santos-Guerra and C.E. Jarvis. Pp. 1-34. A revision of the genus Mandragora (Solanaceae). S. Ungricht, Studies on the lichen genus Sticta (Schreber) Ach.: II. S. Knapp and J.R. Press. Pp. 17^*0. Typification of taxa from Swartz's Prodromus of 1788. D.J. The pteridophytes of Sao Tome" and Principe (Gulf of Guinea). Galloway. Pp. 35^*8. E. Figueiredo. Pp. 41-66. 1998. 43.40 CONTENTS 67 A revision of Brillantaisia (Acanthaceae) K. Sidwell 115 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 6. Genera [Q] R-Z, and an update of current names for non-geniculate Corallinales W.J. Woelkerling, G. W. Lawson, J.H. Price, D.M. John and W.F. Prud'homme van Reine jral History Museur BOTANY SERIES Vol. 28, No. 2, November 1998