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 (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Ms M.J. Short Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 7 16 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Email: intercept@andover.co.uk Internet: http://www.intercept.co.uk Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) © 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 KAREN S1DWELL 1 GENERALLY 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 of Hygrophila 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 INTRODUCTION The aim of this paper is to present a revision of Brillantaisia with a key for identification of species, addressing the following questions: 1 . How many species are there in Brillantaisia1. 2. Is the genus Brillantaisia monophyletic? 3. Are the subgeneric groups within Brillantaisia proposed by Lindau (1895«) monophyletic? 4. Is Hygrophila the sister group to Brillantaisia! 5. Is Hygrophila monophyletic? Brillantaisia P. Beauv. is a genus of mainly woody herbs and subshrubs from tropical Africa and Madagascar. Plants of Brillant- aisia generally have ovate leaves with a winged petiole, an open paniculate inflorescence of bilabiate purple flowers and capsules containing numerous seeds, each held on a hooked retinacula. At the outset of this study, there was no consensus concerning the delimi- tation of Brillantaisia, species boundaries within the genus or species relationships. The genus was thought to contain between 13 (Vollesen, pers. comm.) and 40 (Mabberley, 1987) species. The monospecific genus Plaesianthera (C.B. Clarke) Livera from Sri Lanka had recently been transferred to Brillantaisia as an Asian representative of the genus by Cramer (1991); however, this genus has been shown to be best placed within Hygrophila R. Br. (Sidwell, in press a) and is not discussed further in this paper. Prior to undertaking cladistic analyses, Brillantaisia is hypothesized as mono- phyletic (sensu Hennig, 1950, 1979), based on the presence of two posterior stamens, two staminodes and a laterally compressed upper corolla lip; the genus Hygrophila is hypothesized as the sister group to Brillantaisia based on previous classifications of Nees ( 1 847), Bentham (1876), Lindau (1895fl) and Bremekamp (1953, 1965), and the monophyly of the taxon Hygrophileae containing only Brillantaisia and Hygrophila, is provisionally accepted on the pres- ence of four aperturate pollen (Scotland, \992a, b, 1993; Furness, 1994) and a unique ndhF sequence (Scotland et al., 1995). Taxonomic history Brillantaisia was first described in 1 8 1 8 by Ambroise Marie Palisot de Beauvois from a collection made near Agathon in Benin, West Africa (Palisot de Beauvois, 1818). A single species, Brillantaisia owariensis P. Beauv., was described as having a four angled stem, violet flowers in a panicle and ovate-lanceolate toothed leaves with an acute tip and a winged petiole. Thirty years later, Nees von Esenbeck ( 1 847) overlooked the original publication of Brillantaisia and described two new genera of Acanthaceae from West Africa: Belantheria Nees and Leucoraphis Nees. Bentham (1849) recog- nized the oversight of Nees and transferred the two species of Leucoraphis to Brillantaisia as B. lamium (Nees) Benth. and B. vogeliana (Nees) Benth. and synonomized Belantheria belvisiana Nees with Brillantaisia owariensis. In 1853, Hooker addressed the already problematic issue of species delimitation in Brillantaisia. He agreed with Bentham (1849) that the three species described by Nees (1847) should be placed within Brillantaisia; however, after observation of a newly introduced garden plant from Sierra Leone (Lindley, 1853), Hooker (1853) concluded that the separation of those species on leaf margin and leaf shape characters was invalid as the character variation previously used to distinguish between the K. SIDWELL species could be seen on a single cultivated specimen. Hooker therefore united all the species recognized by Nees (1847) and Bentham ( 1 849) under the earliest name, Brillantaisia owariensis P. Beauv.' . Since that time, many new species of Brillantaisia have been described (see Sidwell, 1 997 for detailed discussion). Gustav Lindau was by far the most prolific describer of new species in the genus. Between 1893 and 1904 Lindau published descriptions of 25 new species of Brillantaisia. The first, largest and possibly most import- ant of these publications was Acanthaceae Africanae I (Lindau, 1893), in which, ten new species of Brillantaisia were described. Characters used by Lindau at the species level included habit (herb or shrub), leaf shape (ovate or obovate), leaf base shape (cordate or not cordate), leaf apex shape (acuminate or not), cystoliths (present or absent), whole plant indumentum (pubescent or glabrous), sepal length (equal or unequal) and number of ovules. By the time of his account of Acanthaceae in Engler and Prantl's Die Natiirlichen Pflanzenfamilien (Lindau, 1895a) Lindau had published fourteen names in Brillantaisia, and was the first and only worker to delimit subgeneric taxa within the genus. Lindau subdivided Brillantaisia into two sections: three species with spicate inflorescences were included in section Stenanthium Lindau, and nine species with paniculate inflorescences were mentioned in section Euryanthium Lindau. Section Euryanthium was further subdivided on leaf shape characters into Group A with lanceolate leaves (containing B. land- folia Lindau) and Group B with non-lanceolate leaves (containing all other species with a paniculate inflorescence). Brillantaisia is now generally considered to be more closely related to the genus Hygrophila R. Br. than to other members of the Acanthaceae because both genera possess a bilabiate corolla and numerous seeds. However, historically, the position of Brillantaisia within the Acanthaceae has been somewhat confused. Nees von Esenbeck ( 1 847) was the first to produce a worldwide monograph of the Acanthaceae. He refined his work on Asian Acanthaceae (Nees, 1 832), changing the rank of some taxa, and placed both Brillantaisia (as Belantheria and Leucoraphis) and Hygrophila in the subfamily Echmatacantheae (possessing retinacula), tribe Hygrophileae (co- rolla 5-parted; stamens 2 or 4; seeds numerous; fruits explosive), Subtribe 1 . This was the first grouping of Brillantaisia andHygrophila in the same taxon. However, in the same year Lindley (1847) listed Brillantaisia as of unknown affinity and noted that three eminent botanists working on the Acanthaceae - Brown, Nees and Meisner - were unable to agree on subdivision of the family. Lindley (1847: 679) noted that 'there are few natural Orders which now demand, in so eminent a degree, a searching investigation as that of Acanthads'. Bentham ( 1 876) subdivided the Acanthaceae into five tribes using a combination of characters (corolla aestivation, ovule number, seed shape and retinacula type). Brillantaisia andHygrophila were placed in tribe Ruellieae Benth. (aestivation contorted; seeds 2-many per locule; seeds laterally compressed; retinacula hooked), subtribe Hygrophileae (corolla two-lipped; filaments laterally connate). Baillon ( 1 89 1 ) recognized six series in Acanthaceae and considered Brillantaisia sufficiently distinct to be placed in the monogeneric series Brillantaisiees characterized by the presence of two fertile posterior stamens, a strongly bilabiate corolla and many seeds per capsule. 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 apparently mis-named by Hooker (1853). as 'Belantheria lamium' and 'Belantheria vogeliana', two combinations which had never been formally published prior to that work. The lumping of 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 'rippenpollen' with four equatorial apertures. Bremekamp (1965) also placed Brillantaisia with Hygrophila, in Acanthoideae- Ruellieae-Hygrophilineae, a group with bilabiate flowers in cymes, stamens united into a single group, style not held in place by a row or bundles of trichomes and subglobose, banded, usually four- aperturate pollen. As mentioned above, recent studies confirmed the monophyletic nature of the group consisting of these two genera based on a pollen type not found elsewhere in Acanthaceae (Scot- land, 1993) and a unique ndhF sequence (Scotland et al., 1995). Since the time of Nees von Esenbeck (1847) no monographic work has been undertaken for Brillantaisia and the prolific descrip- tion of species by Lindau (1893; 1894; 1895a, b, c, d; 1896; 1897; 1898; 1904) served to exacerbate difficulties in understanding the group. Several regional floras (Heine, 1966; Benoist, 1967; Champluvier, 1985; Vollesen, in prep, a & b) have addressed the problem of the number of species within Brillantaisia in Gabon, Madagascar, Rwanda and East Africa respectively. The authors of these works disagree in particular on the number of species in several morphologically distinct groups within Brillantaisia. Tax- onomists working on Brillantaisia have tended to split it into poorly defined groups and formally name many slightly differing forms of the great morphological variation within the genus. Many new species of Brillantaisia have been described, often from little material and based on characters that are highly variable within the genus. Identification of specimens has been inconsistent and a study cover- ing the entire range of geographical and morphological variation of the genus is essential to clarify this confusion. MORPHOLOGY Initially I considered it impossible to separate description of the morphological variation of Brillantaisia from discussion of the transformation of that information into comparative morphological data (characters and character states) for phylogenetic analysis. However, morphological data are analysed at two different levels in this study. Characters and character states of individual specimens are analysed when delimiting species of Brillantaisia, and charac- ters and character states of taxa are analysed when forming hypotheses of relations among those species. In order to emphasize the filtering of data undertaken when transforming observation and description into comparative data, a descriptive approach to discussion of characters is presented prior to discussion of character coding for cladistic analysis. A purely descriptive morphological section pro- vides an overview of the genus, directly comparable with many published monographic studies and some flora accounts for the Acanthaceae, recent examples of which include studies on Justicia (Graham, 1988), Justicia sect. Harnieria (Hedren, 1989), Justicia sect.Ansellia (Ensermu, 1990), Ruellia (Ezcurra, 1993), Strobilanthes (Wood, 1995), Dicliptera (Balkwill et al., 1996) and Phaulopsis (Manktelow, 1996). Brief discussion of morphological variation is given here and species that exhibit the range of variation of a particular organ are used as examples. Herbarium specimens rel- evant to the discussion are referenced by collector's name, number and herbarium acronym (e.g. Kibuwa 5191, UPS). The data were obtained through observation of herbarium speci- mens from BM*, BR*, C, DSM*, E*, ETH, FHO*, G, HBG, K*, LISC, M, MHU, MO*, NY, P*, PRE, S, SCA*, SRGH, W, WAG*, YA* and Z (abbreviations follow Holmgren et al., 1990). Visits have been made to the herbaria marked with an asterisk. Gross morpho- logical features were measured on mature vegetative and reproductive structures using a mm ruler or calipers which measure to 0.1 mm. Finer measurements were made using an 8 mm 0.1 graticule in a Leica Wild M8 dissecting microscope. Floral dissections were made after placing a flower into cold water and heating slowly until the water almost boiled. This usually took about 30 seconds and was sufficient to soften the delicate corolla. Boiling caused the corolla to disintegrate. Photographs were taken using a Nikon F-801s camera with FP4 black and white film and a Microtech MF80 fibre optic light source. Pollen preparations were acetolysed using a modifica- tion of Erdtman (1960) (see Sidwell, 1997). Light micrographs of pollen grains were taken using a Dialux 20EB photomicroscope. Scanning electron micrographs were taken at The Natural History Museum, London. Stubs were coated in gold-palladium 80:20% and viewed using an Hitachi S-800 SEM microscope with a working distance of 5-10 mm. Permanent slides of Brillantaisia wood were prepared according to Gourlay (pers. comm.): the stem was boiled for an hour, cut in transverse, tangential longitudinal and radial longitudinal sections on a sliding microtome and stained in safranin. The sections were passed through a dehydration series, washed three times in alcohol and mounted in Canada balsam. Habit Brillantaisia species range in habit from delicate herbaceous Brillantaisia land/alia Lindau, approximately 50 cm high, to the large woody species B. kirungae Lindau which reaches 7 metres tall. Within this range, Brillantaisia species are mainly erect, perennial 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 along branches that touch the ground, enabling them to spread vegetatively. The terms herb, shrub and tree have been loosely applied by collectors to describe the habit of Brillantaisia species. For example, B. madagascariensis ranges in habit from a 'soft succulent herb' (Mooney 607 1 , S) or a 'perennial herb with creeping and ascending stems' (Friis & Vollesen 613, C) to a 'shrub to 2m tall' (Kibuwa 5191, UPS). According to the model of plant architecture, described by Weberling (1989) after Troll (1964), plants can be divided into four distinct and clearly defined zones: main flores- cence; enrichment zone; inhibition zone and innovation zone. This model was used by Manktelow (1996) in her monograph of Phaulopsis and can be applied to species of Brillantaisia (Fig. 1). Figure 1 illustrates the architecture of Brillantaisia from the apex to the base of the plants as follows. The inflorescence is divided here into two distinct and more precisely defined parts: the terminal thyrse with lateral dichasial branches forms the main florescence and flowering lateral branches (coflorescences) which 'repeat the structure of the main axis to some extent' (Weberling, 1989: 225) form the enrichment zone. These two parts are separated from one another by the basal internode (bi on Fig. 1), and the whole flower- ing region is termed a synflorescence. Below the enrichment zone, where the axillary buds do not develop into coflorescences, is the inhibition zone, and below that, the region of vegetative growth is termed the innovation zone. The innovation zone is considered the region of production of new shoots after flowering in perennial plants. Brillantaisia lamium, B. vogeliana, B. owariensis and B. madagascariensis all exhibit the type of growth pattern described above and illustrated in Figure lA.Two species, B. lancifolia and/?. debilis Burkill have a main florescence and lack the enrichment zone (Fig. IB). In the annual species, B. pubescens T. Anderson ex Oliv. inhibition of the lateral shoots does not occur, lateral coflorescences occur to the base of the plant and the inhibition and innovation zones 70 K. SIDWELL Inhibition Innovation zone ' ' zone .a a § E •M .§ « g « s .1 00 S oo •& •- '£ 11 l! OQ a § 2 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) 3 1 . Fruit trichomes: absent (0), present ( 1 ) view trees. CLADOS (Nixon, 1992) was used to look at character distribution on the HENNIG86 trees. Two types of character transfor- mations were plotted onto the trees using CLADOS: either as unique occurrences (synapomorphies) in black or white or convergence or parallelism (homoplasy) in grey. The values of tree length, consist- ency index (CI) and retention index (RI) are provided. All characters are treated throughout as unweighted. Results The analysis produced 122 equally parsimonious trees, 87 steps long, CI=0.42 and RI=0.76.Tree 25/1 22 is presented in Figure 6 and the strict consensus of these trees is presented in Figure 7. Apart from a few collapsed nodes, the consensus tree (Fig. 7) is very similar to the tree given in Figure 6, and the major clades are the same in both trees. Although the consensus tree (Fig. 7) is accepted as a working hypothesis of the phylogeny ofBrillantaisia, the results are discussed here with reference to the characters shown on the tree in Figure 6. The results show that Brillantaisia and six species of Hygrophila both belong within a monophyletic group, the tribe Hygrophileae, based on a suite of pollen characters [21(1); 22(0); 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 species B. kirungae, B. owariensis and B. vogeliana). The outgroup to the Brillantaisia-Hygrophila clade is a group with non-bilabiate corollas [15(0)] containing the species Eremomastax speciosa, Mellera lobulata and Mimulopsis solmsii. The ((Brillantaisia- Hygrophila) (Eremomastax (Mellera-Mimulopsis))) clade is characterized by the presence of paniculate inflorescences [6(1)] and sepals which are free or fused only slightly at the base [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 01111 0?1?0 00010 10001 00101 11000 01 Dyschoriste verticillaris C.B. Clarke 11110 00100 10010 10071 10101 11710 00 Eremomastax speciosa (Hochst) Cufod. 10110 01100 00000 00011 00101 11000 01 Mellera lobulata S. Moore 10110 01100 11000 00001 10101 11000 01 Mimulopsis solmsii Schweinf. 10110 01100 11000 00000 00101 11000 01 Hygrophila auriculata (Schumach.) Heine 10010 00100 01001 10000 01010 00100 01 Hygrophila didynama (Lindau) Heine 11010 01001 11000 10110 01010 00100 11 Hygrophila laevis Lindau 11010 01100 01001 10070 01010 00100 70 Hygrophila linearis Burkill 11000 01000 10000 10013 01010 00100 10 Hygrophila pilosa Burkill 11010 01000 11000 10110 01010 00100 11 Hygrophila thwaitesii (Benth.) Heine -a 00010 01110 11000 10003 01010 00000 10 Hygrophila thwaitesii (Benth.) Heine-b 01010 onio 11000 10003 01010 00000 10 Brillantaisia debilis 10002 01000 00000 21102 01010 00100 10 Brillantaisia grottanellii-& 10012 00100 11100 21102 01010 00101 11 Brillantaisia grottanellii-b 10212 00100 11100 21102 01010 00101 11 Brillantaisia lamium. 10012 01100 00000 21102 01010 00100 11 Brillantaisia lancifolia-a 11001 01000 00000 21102 01010 00100 11 Brillantaisia lancifolia-b 11011 01000 00000 21102 01010 00100 11 Brillantaisia madagascariensis 10012 00100 01100 21102 01010 00111 11 Brillantaisia oligantha-SL 10012 11100 11000 21101 01010 00100 11 Brillantaisia oligantha-b 10112 01100 01100 21102 01010 00100 11 Brillantaisia owariensis-a 10112 01100 01100 21102 01010 00100 11 Brillantaisia owariensis-b 10113 01100 01100 21102 01010 00100 11 Brillantaisia pubescens. 00012 OHIO 11100 21111 01010 00100 11 Brillantaisia riparia-a 10012 OHIO 11000 21111 01000 00100 11 Brillantaisia riparia-b 10012 11110 11000 21111 01000 00100 11 Brillantaisia riparia-c 11012 11110 11000 21111 01000 00100 11 Brillantaisia riparia-d 11012 OHIO 11000 21111 01000 00100 11 Brillantaisia stenopteris-a 10012 11000 01100 21001 01010 00100 11 Brillantaisia stenopteris-b 10012 11000 11100 21001 01010 00100 11 Brillantaisia kirungae 10111 01101 01001 21102 01010 00100 11 Brillantaisia vogeliana. 10113 01100 11100 21102 01010 00100 11 Duosperma crenatum and Dyschoriste verticillaris form an unre- solved outgroup to all other species in this analysis. Within the Brillantaisia-Hygrophila clade, the twelve species of Brillantaisia form a monophyletic genus based on a laterally compressed upper corolla lip [16(1)], a membranous hinge at the apex of the corolla tube [15(2)] and a winged petiole with an entire or subentire margin [4(2)]. Within Brillantaisia, a clade containing eight spe- cies is delimited on the presence of staminodes greater than 2 mm long [19(2)]. The other four species within the genus all have minute staminodes and form a partially unresolved outgroup to that clade. Section Stenanthium of Lindau, recognized by him as consisting of all species of Brillantaisia with spicate inflores- cences (B. grottanellii and B. madagascariensis) is monophyletic in this analysis based on the persistence of the style in fruit [29( 1)], a character which maybe an artifact of the maturity of specimens examined for those species and requires reconsideration. Section Stenanthium of Lindau containing all other species of Brillantaisia is paraphyletic. Subgeneric taxa are not formally recognized in this account. The six species selected to represent Hygrophila form successive pairs of sister taxa to Brillantaisia. However, in this analysis, Hygrophila is paraphyletic and requires further study, including a more comprehensive cladistic analysis, before the precise sister group to Brillantaisia can be identified. Hygrophila is necessarily accepted as a paraphyletic genus for nomencla- tural convenience, but it should be recognized as a diverse group, in need of revision, and is best conceived as comprising all mem- bers of the tribe Hygrophileae that do not belong within Brillantaisia. Within the constraints of time and the Linnaean hierarchy, acceptance of well known groups which are paraphyletic is inevitable. Fig. 6 One of 122 equally parsimonious trees (tree 25/122) produced from analysis of the data matrix in Table 1. Synapomorphic characters are shown by the black and white bars and homoplasious characters by the grey bars. Character numbers are given above each bar and the state of that character is given below each bar. REVISION OF BRILLANTAIS1A 0 4 81 Du.crenatum 0 1 10202831 — IHHH}— Dy.verticillaris 1110 18 §— E.speciosa 1 15 1 HHh 20 6 13 -HH 1 o 0 0 1011 I — |— Me.lobulata 1 1 1 19 M.solmsii 1 6 2 1121222324252627 -0-f-HH-iHHHH 011010001 //. auricula! n H. laevis 3 7 1118 1030 -0-1- 1 1 1931 rUH 3 0 H. linearis 0 8 27 I — ff~ H.thwaitesii-a HhHH H.thwaitesii-b H.didynama 1718 -HH 1 1 H.pilosa 1 4 1516 HI • • I 0221 0 12 — 0~i~~ B.pubescens 0 1 23 — B.riparia-a 1 — §— B.riparia-d — 5. riparia-b — §— B.riparia-c — B.oligantha-a — B.oligantha-b 12 -IH 10 B. stenopteris-a — B.stenopteris-b — B.grotanellii-a 6 1229 19 0 1 1 10 HHH 1— B.grotanellii-b 1 1028 0 1 4 9 14 B. madagascariensis B.kirungae 1 1 1 I — B.owariensis-a 12 -IH — B.owariensis-b 10 1 — B.lamium 31 3 7 B. vogeliana 0 0 B.debilis 1 4 i — B.lancifolia-a -HH 3 1 Mh- B.lancifoliab 82 K. SIDWELL 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 Brillantaisia P. Beauv., Flore d'Oware 2: 67. t. 100, fig. 2 (1818); Burkill in Fl trap. afr. 5: 37 (1899); Durand & Durand, Syll.fl. congol: 416-418 (1909); Chevalier, Explor. hot. Afrique occ. franc. 1: 493 (1920); Robyns, Fl. pare nat. Albert (1947); Heine in Fl. W. trap. Afr. 2nd ed., 2: 405 (1963); Heine in Fl. Gabon 13: 28 (1966); Benoist, Fl. Madag.fam. 182: 28 (1967);Agnew, Upl. Kenya wild fls: 583 (1974); Troupin, Fl. pi. lign. Rwanda: 84 (1982); Champluvier in Fl. Rwanda 3: 444 (1985). Type species: Brillantaisia owariensis P. Beauv., Flore d'Oware 2: 68, t. 100: fig. 2(1818). Belantheria Nees in DC., Prodr. 11: 96 (1847). Type species: Belantheria belvisiana Nees in DC., Prodr. 11: 96 (1847). Leucoraphis Nees in DC., Prodr. 11: 97 (1847). Type species: Leucoraphis vogeliana Nees in DC., Prodr. 11: 97 (1847). Ruelliola Baillon in Bull. Mens. Soc. Linn. Paris 2: 852 (1890); Baillon, Hist. pi. 10: 427 (1891); Lindau in Nat. Pflanzenfam. 4(3b): 307 (1895). Type species: Ruelliola grevei Baillon in Bull. Mens. Soc. Linn. Paris 2: 852 (1890). Erect to prostrate herbs to erect, sturdy shrubs. Stems square, usually swollen at the nodes, glabrous to densely pubescent; cystoliths longitudinal, narrowing towards one end. Leaves opposite, decussate, petiolate, broadly ovate to ovate, occasionally elliptic or linear- lanceolate; leaf base cordate or truncate to cuneate; apex acute or acuminate; margin entire to irregularly toothed; glabrous to pubes- cent above and below; indumentum of multicellular, eglandular trichomes, evenly pilose to dense and villose above, mainly on midrib and primary lateral veins below; cystoliths usually visible, with a hand lens, above and below over entire leaf surface. Petiole usually winged, lamina decurrent in top half of petiole, though occasionally broadly winged to base; petiole wing margin entire, occasionally toothed towards apex. Inflorescence a variously modified dichasium forming either a terminal thyrse and/or lateral axillary thyrse with vegetative growth occasionally continuing above flowering axes; dichasial branching usually lax, becoming monochasial towards base of inflorescence, occasionally contracted forming spike with flowers appearing in verticillasters. Bracts on main inflorescence axis ovate to broadly ovate, foliaceous, equal in size to the uppermost leaves at base of inflorescence, becoming smaller towards the apex of the inflorescence; bractlets on lateral inflorescence branches linear to obovate. Calyx with five equal or unequal, linear to spathulate sepals, dorsal sepal longer and usually broader than the four lateral sepals; usually hirsute, with glandular and eglandular trichomes. Corolla two- lipped, purple, magenta, blue or violet, occasionally white; corolla tube cylindrical, often paler than corolla lobes, with two brown-yellow markings in throat; upper lip hooded, laterally com- pressed, two-lobed at apex, often glandular pubescent on outer surface; lower lip broad, ridged and bullate above, reflexed at edges, three-lobed at apex, occasionally with stiff trichomes on adaxial surface, two membranous lateral pouches at base of lower lip form a hinge with the apex of the corolla tube. Androecium with two posterior stamens fertile; filaments white, flattened, often sparsely pubescent towards the base; anthers sagittate, dorsifixed; two anterior stamens reduced to slender staminodes, often with membranous vestigial anther at apex, occasionally minute or absent. Gynoecium with slender, linear style, often sparsely pubescent towards base; stigma a single flattened lobe at the end of the style, second, lower lobe reduced to minute tooth; ovary two-locular on annular disc, usually pubescent, placentation axial; ovules numerous. Fruit a linear, convex, two-valved capsule, pointed at apex with deep longi- 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 slightly kidney shaped, flattened, each held on a hardened, hook- shaped retinacula, covered with adpressed trichomes which are hygroscopic and expand rapidly, producing mucous when wet. DISTRIBUTION. Throughout tropical Africa with two species ex- tending to Madagascar. HABITAT. The distribution of Brillantaisia species is centred in the Guineo-Congolian rain forests with species also occuring in montane rain forests and woodland throughout Africa. Plants of Brillantaisia 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 Africa Brillantaisia is commonly found in disturbed areas (e.g. roadsides, plantations, farmland). Brillantaisia is a genus of 12 species several of which are wide- spread and tend to be very variable morphologically, and is most diverse in the Guineo-congolian forests. Many species have previ- ously been described in the genus based on local variation which has been shown to be continuous by extensive study of specimens. The variation within each species is discussed in detail after each de- scription. A full list of specimens studied is provided in Sidwell (1997) and a constantly updated list is available from the author. Key to fertile specimens of Brillantaisia 1 . Inflorescence a contracted thyrse with short peduncles, forming a spike; flowers/fruit in verticillasters 2 Inflorescence a lax thyrse with elongated, clearly visible peduncles 3 2. Bracts with glandular and eglandular trichomes, linear-ovate; style often pubescent towards apex 1 1. B. grottanellii Bracts lacking glandular trichomes, ovate to broadly ovate; style pubes- cent only at base 12. B. madagascariensis 3. Bractlets obovate-rounded, shortly petiolate; corolla lips less than twice the length of the corolla tube; sepals spathulate; flowers with stiff, unicellular trichomes on inner surface of lower lobe 4 bractlets linear; corolla lips more than twice the length of the corolla tube; sepals linear or spathulate; flowers lacking unicellular trichomes on inner surface of lower lobe 5 4. Flowers small, 12-25 mm long, inflorescence highly branched; leaves 1.25-1.8 times longer than wide 1. B. pubescens Flowers large, 30-40 mm long; inflorescence branched 3-4 times; leaves 1 9-3.0 times longer than wide 2. B. riparia 5. Leaves lanceolate to elliptic, occasionally ovate, entire or barely cre- nate; inflorescence narrow terminal thyrse branching 1-2 times 9. B. lancifolia Leaves ovate to broadly ovate, entire or clearly toothed; inflorescence lax, open, terminal or lateral thyrse branching 2-many times 6 6. Fruit glabrous or with a few erect trichomes at the apex; leaves entire or subentire 7 Fruit pubescent; leaves toothed, or if leaves entire to subentire, inflores- cences slender, lateral, thyrse with apex usually reverting to vegetative growth 8 7. Leaves pubescent particularly towards petiole; fruit 22-30 x 2-5 mm, with 14-16 seeds per locule 8. B. lamium Leaves usually glabrous; fruit 18-22 x 1-2 mm, with 18-28 seeds per locule.... ... 10. B. debilis 84 K. SIDWELL 8. Fruit with 8-10 very rarely to 16 seeds per locule; woody shrub or small tree, leaves often drying black 6. B. kirungae Fruit always with more than 1 2 seeds per locule; herb to 2 m tall, not drying black 9 9. Flowers 15-25 mm long (rarely to 35 mm); fruit with 20-24 seeds per locule; leaves thin, rather papery when dry, often toothed at the top of the petiole wing 7. B. vogeliana Flowers 30-45(-50) mm long; fruit with 1 2-20 seeds per locule; leaves not thin and papery when dry, petiole wing usually entire 10 1 0. Lower corolla lip with clearly visible multicellular trichomes on upper surface 3. B. oligantha Lower corolla lip glabrous 1 1 1 1 . Leaf margin subentire or with small serrations; petiole wing tapering gradually to base of petiole: lower corolla lip strongly reflexed, centre of lower lip bent up to 90° 4. B. stenopteris Leaf margin usually strongly toothed; petiole wing tapering gradually to broad and toothed; lower corolla lip not not strongly reflexed .... 5. B. owariensis 1. H rill;i nt aisia pubescens T. Anderson ex Oliv. in Trans. Linn. Soc. London 29: 1 25 ( 1 875); Burkill in Fl. trop. afr. 5: 38 ( 1 899); Durand & Durand, Syll.fl. congol: 417 (1909); De Wild., Contr. FL Katanga 1: 143 (1913); Benoist in Cat. pi. madag. 13(1939); Vollesen in Opera Bot. 59: 79 (1980); Vollesen & Brummitt in Kew Bull. 36: 571 (1981). Type: Tanzania, Khutu, Kirengwe, Grant s.n. (K!-holotype). Fig. 8. Map in Vollesen & Brummitt (1981: 571). Brillantaisia rutenbergiana Vatke in Abh. Naturwiss, Vereine, Bremen 9: 131 (1885); Palacky, Cat. pi. madag. 3: 57 (1907). Type: Madagascar, Andranovaka, Rutenberg s.n. (P-holotype). Ruelliola grevei Baillon in Bull. Mens. Soc. Linn. Paris 2: 852 (1890); Baillon, Hist. pi. 10: 427 (1891); Lindau in Nat. Pflanzenfam. 4(3b): 307 ( 1 895).Type: Madagascar, Greve 26 (P!- holotype; P!-isotype). Brillantaisia anomala Lindau in Pflanzenw. Ost-afr. C: 366 (1895); Lindau in Bot. Jahrb. Syst. 24: 312-313 (1898); Lindau in Nat. Pflanzenfam. 4(3b): 296 (1895). Type: Mozambique, Villa Gouveia, de Carvalho s.n. (COI-holotype). B. pubescens var. rutenbergiana (Vatke) Benoist in Cat. pi. madag. 13(1939). Hygrophila pubescens (T. Anderson ex Oliv.) Benoist in Fl. Madag. fam. 182 1: 36 (1967) non Nees (1847). Icones: Anderson in Trans. Linn. Soc. London 29: pi. 125 pro parte (1875); Benoist in Fl. Madag. fam. 182. 1: fig. IV 14-18 (1967). Straggling annual, highly branched, aromatic herb, (0.3-)0.45-1 m tall, smelling minty or balsamic. Stems 2-5(-14) mm in diameter, covered in short glandular trichomes and longer fine, white eglandular trichomes. Leaves broadly ovate to ovate 50-1 10 x 85-160 mm, often caducous; leaf base cuneate to shortly attenuate, occasionally truncate; apex acute; margin entire, occasionally slightly crenate; indumentum of silky, eglandular trichomes above and below; lateral veins 7-9(-13) each side of midrib; petiole to 75 mm long, winged towards lamina, wing narrow, decurrent on petiole in upper Vi-Vi. Inflorescence highly branched open thyrse with lateral inflores- cences to base of stem, branches slender, rachis indumentum irregularly glandular-pubescent with longer eglandular trichomes. Bracts obovate to elliptic, to 35 x 45 mm, petiolate, irregular glandular pubescent on both surfaces; bractlets obovate-rounded, to 5x8 mm, persistent. Sepals unequal, spathulate, larger upper sepal 12-14 x 0.5 mm, 1 .5-2 mm wide at broadened apex, smaller lateral sepals 8-10(-13) x < 0.5 mm, pubescence of slender eglandular trichomes with shorter glandular trichomes, denser towards spathulate apex. Corolla purple to pale purple/pink occasionally with white upper lip, tube 5-10 mm long; upper lip 8-13(-15) mm long, 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 long; anthers 1-2.5 mm long; staminodes minute < 0.5 mm long or absent, glabrous. Gynoecium with style 8-9 mm long; stigma 1 mm long; ovary 2-3 mm long, glandular pubescent. Capsule 10-15 x 2-2.5 mm with 8-16 seeds per locule, yellow brown when dried, sparsely glandular pubescent with longer eglandular trichomes towards the apex. DISTRIBUTION. East and Central Africa, Malawi, Zambia, Zimba- bwe, Mozambique and Tanzania, west to Zaire, Madagascar. HABITAT. Sandy dry places, river beds, shade on forest floor or in savannah woodlands under, for example, Adansonia digitata and Faidherbia albida trees, or with Acacia, Albizia, Parkia, Lepis- anthes and Terminalia (Abdallah & Vollesen 95/176 K). Locally common; 0-1000 m. SELECTED COLLECTIONS DEMOCRATIC REPUPUBLIC OF THE CONGO: Upemba National Park, Kaswabilenga, 17 June 1948, 700 m, de Witte 03986 (BR, K). TANZANIA: Lindi, 1 1 September 1934, Schlieben 5323 (G, H, LISC, M, MO, PRE, S, US, Z); 2 km NW of Kingupira, 5 June 1975, Vollesen 2392 (C); 6 km along Kisiwani-Mnazi road, 800 m, 7 May 1995, Abdallah & Vollesen 95/176 (K). MOZAMBIQUE: Maringua, Dabi river, 25 June 1950, Chase 2547 (BM). MALAWI: Salima, Chipoka near Ilala Quarry, 25 May 1972, Salubeni 1803 (MO, SRGH); Chimpakati village, Mlunguzi River, 27 June 1987, Usi & Kaunda 606 (M AL). ZAMBIA: Petuake district, 5 September 1 947, Greenway & Brenan 8044 (PRE); NW of Mwern-na-ntipa, 7 August 1962, Tyres 339 (SRGH). ZIMBABWE: Binga district, September 1955, Davies 1426 (MO, SRGH); Urungwe district, 20 September 1981, Pope 1987 (MO, SRGH). 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 Corolla length to 25 mm Hinge at apex of corolla tube not well developed Leaf shape ovate branched 3-4 times 30-40 mm long well developed elliptic REVISION OF BRILLANTA1S1A 85 Fig. 8 Brillantaisia pubescens T. Anderson ex Oliv. A. Habit x 0.7; B. Lower leaf x 0.7; C. Bract and bractlets x 1 ; 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 genus. The highly branched inflorescence, small flowers and rounded persistent bracts are not found elsewhere \nBrillantaisia. Brillantaisia pubescens sensu lato was split into two subspecies by Vollesen & Brummitt (1981) on the basis of flower size. I consider the large flowered plants to be a different species (B. riparia) and treat B. pubescens s.s. as including only those plants with smaller flowers. Two specimens of B. pubescens have slightly larger flowers than most (Tinley 2639, SRGH; Thera & Kaunda 313, PRE); however, the leaf shape and venation are that of B. pubescens and this variation does not blur the boundary between B. pubescens and B. riparia. The original illustration of the small flowered B. pubescens by Fitch (Anderson 1875: pi. 125) almost certainly contributed to confusion over species delimitation as the type specimen was repre- sented as having many more, larger flowers than it actually has. It appears that large flowered specimens of B. riparia have been identified as B. pubescens by reference to the illustration. There are several characters by which plants of the two taxa can be clearly distinguished (Table 3). 2. Brillantaisia riparia (Vollesen & Brummitt) Sidwell, comb. nov. Type: Malawi, 13 miles N. of Kasungu, Dwangwa River, Pawek 3908 (K!-holotype). Fig. 9. Map in Vollesen & Brummitt ( 1 98 1 : 57 1 ). Brillantaisia pubescens var. riparia Vollesen & Brummitt in Kew Bull 36: 571 (1981); Vollesen in Opera Bot. 59: 79 (1980). Icon: Anderson in Trans. Linn. Soc. London 29: pi. 125 pro parte (1875). Annual viscid-pubescent, aromatic, rather woody herbs to 0.5-1 m tall. Stems erect or ascending; pilose with eglandular and glandular trichomes. Leaves oblong, elliptic to ovate 20-25(-52) x 39-59(- 95) mm, often fallen below; upper most leaves of main axis oblong, 5-8 x 10-15 mm; base gradually tapering, cuneate; apex acute, rounded at tip; margin entire to slightly crenate; glabrous to pilose; lateral veins 4-9(-l 1) each side of midrib; petiole 14-22 mm long, lamina decurrent on petiole forming very narrow wing sometimes to base. Inflorescence leafy-bracteate terminal thyrse, with lower lat- eral panicles, occasionally with vegetative growth continuing at apex; rachis covered in short simple trichomes with longer fine, non glandular and stalked glandular trichomes. Bracts as leaves; bractlets elliptic-obovate, rounded at apex, margin entire, glandular trichomes and slender white trichomes above and below. Sepals unequal, larger upper sepal linear-spathulate, 14-18 x 1 mm; smaller lateral sepals spathulate, 9-14 x 0.5-1 mm. Corolla purple to blue-purple; corolla tube 13-15x2 mm; upper lip 20-25 mm long, glabrous; lower lip 20-25 mm long, glabrous below, with straight erect stiff trichomes on centre of inner surface, trichomes white with a dark purple-black tip; apical lobes 2 x 2-3 mm, triangular, rounded to blunt or notched at apex. Androecium with filaments (10-)20-22 mm long; anthers 4-5 mm long; staminodes 3-4 mm long, pale, translucent, difficult to see with the naked eye, broadened slightly at apex sometimes to curved vestigial anther. Gynoecium with style to 30 mm long, stiff upward pointing trichomes towards base; stigma 2 mm long; ovary 3 mm long glandular pubescent. Capsule 14-19x2 mm with 8-12 seeds per locule, light brown, covered with simple short trichomes and longer, glandular trichomes. DISTRIBUTION. Mozambique and Malawi. HABITAT. Damp, shady places; 1000 m. SELECTED COLLECTIONS MOZAMBIQUE: Vila Cabral, 8 September 1942, Mendonqa 680 (LISC); Nampula, 3 October 1942, Mendon^a 1219 (LISC); between Namapa and Chiure, 19 August \948.Barbosa 1812 (LISC); Niassa, Vila Cabral, Meponda, 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 (MO, SRGH); Liwawazi river, 3 September 1986, Kaunda & Usi 429 (MAL); Liwonde, 9 September 1988, Banda & Kaunda 3452 (MO). Brillantaisia riparia was considered a variety of B. pubescens by Vollesen & Brummitt (1981). Due to the difference in flower size and shape, leaf shape, leaf venation and inflorescence structure, I consider B. riparia a separate species from B. pubescens. The two species are clearly very closely related yet are easily distinguished in the field. Brummitt (pers. comm.) noted that B. riparia should be compared with B. oligantha. Brillantaisia riparia and B. oligantha are superficially similar, both having lateral inflorescences, trichomes on the lower lip of the corolla and flowers of roughly equal size. However, there are several differences in gross morphology (Table 4) that clearly separate the two. 3. Brillantaisia oligantha Milne-Redhead in Mem. New York Bot. Card. 9: 20-21 (1954); Binns, Check List Herb. Fl. Malawi: 12 (1968). Type: Malawi, Nchisi Mountain, 30 January 1946, Brass 17042 (K!-holotype; BM!, MO!-isotypes). Fig. 10. Map 1. Viscid herb 0.3-1 .8 m tall. Stems 2-2.5 mm across, slightly winged on angles of stem, pubescent, with straggling eglandular and shorter multicellular glandular trichomes. Leaves ovate to broadly ovate, 36-65(-130) x 87-160(-210) mm; leaf base cuneate to attenuate; apex acuminate; margin subentire to shallowly crenate or dentate; 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 Corolla tube straight, long, narrow Lower corolla lip trichomes straight, single celled slender, curving inflated, short, broad, 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 petiole forming a slender wing almost to the base of the petiole. Inflorescence lateral few flowered thyrse to 45-75 mm long, branches very slender, branching 3—4 times, rachis pubescent. Bracts as leaves; bractlets narrowly obovate (0. 3-) 1.5— 4 x (0.8-)2-9 mm, glandular pubescent above and below. Sepals unequal; linear to spathulate, larger upper sepal 6-10 x 1 mm, smaller lateral sepals 5- 9 x 0.5 mm, pubescent. Corolla deep blue to purple, sometimes tinged brown on the upper lip, corolla tube 3 x 7.5 mm; upper lip 1 3- 17 mm long, sparsely pubescent outside; lower lip 14-18 mm long, strongly reflexed with lateral fold down centre and deep hinge at base, multicellular trichomes on inner surface; apical lobes to 2.5 x 3 mm. Androecium with filaments 10-15 mm long, pubescent at base; anthers 1-4 mm long; staminodes 1 mm long with small vestigial anther. Gynoecium with style 1-1.5 mm long, pubescent along the entire length; stigma 20-25 mm long; ovary 3 x 1 mm, covered in short glandular trichomes. Capsule 20-27 x 1 -2 mm with 10-12 seeds per locule, very sparse, short, glandular trichomes. DISTRIBUTION. Restricted to the Nchisi and Mughese mountain ranges in Malawi. HABITAT. Shady places in wet forest, or in dry evergreen or semi- evergreen forest, also along roadsides in these areas. Locally common; 1000- 1400m. SELECTED COLLECTIONS MALAWI: Nchisi Mountain, 29 July 1946, Brass 17021 (NY); 30 July 1946, Brass 17042 (BM, K, MO); 10 July 1960, Chapman 827 (BM); 1 September 1970, Salubeni 1485 (SRGH); Salubeni 1487 (MAL. PRE, SRGH): Chitipa district, Mughese, 1 2 September 1 977. Phillips 2836 (MO):ftw** 1 2984 (K, LISC, MAL, MHU, MO, SRGH). Brillantaisia oligantha is an easily distinguished species, with lat- eral axillary inflorescences only. The lower corolla lip is more strongly reflexed in B. oligantha than in other species of the genus and has multicellular trichomes on the adaxial surface. Previously thought to be restricted to the Nchisi mountains of Malawi, collec- tions from Mughese in Malawi are clearly the same species. Brillantaisia oligantha can appear morphologically similar to B. riparia, both species having entire leaves and fine white silky trichomes on vegetative parts, however, B. oligantlia has a much shorter, broader corolla tube, a more hooded upper lip, and a more slender, less branched inflorescence. This species is also closely related to B. stenopteris Sidwell, sp. nov. from Tanzania. However, the new species has larger flowers, lacks any trichomes on the lower lip of the corolla and is found in a different habitat to B. oligantha. 4. Brillantaisia stenopteris Sidwell, sp. nov. Type: Tanzania, Morogoro region, Kombola, 8 July, 1933, Schlieben 4068 (LISC!- holotype, MO!, PRE!-isotypes) Fig. 1 1. Map 2. Herba perennis usque ad 2 m alta. Folii lamina ovata vel latiovata 25-144 mm longa, 40-220 mm lata, apice acuminata, basi cuneata, margine obscure serrata, lamina 1 1-16 venatus, petiolo breviter decurrente; petiolo 15-90 mm longo. Inflorescentia ad 20 cm longa, cymosa, terminalis, interdum vegitativus ad apice. Corolla 24-30 mm longa, tubus circiter 8-9 mm longus; labium posticum 18-22 mm longum; labium anticum 17-23 mm longum, glabrum. Fructus incognitus. Species nova aflUnisBrillantaisia oligantha Milne-Redh. sed floribus majoribus et labium anticum sine pilis longis septatis differt. Herbs to 2 m tall. Stems erect, 2 mm in diameter, pubescent, trichomes white, eglandular with shorter glandular trichomes. Leaves ovate to broadly ovate 25-140 x 40-220 mm: leaf base cuneate; apex 90 K. SIDWELL acuminate; margin serrate, occasionally subentire; adaxial surface subglabrous to roughly pubescent, abaxial surface with fine non glandular trichomes; 11-16 lateral veins on each side of the midrib; petiole 1 5-90 mm long, lamina decurrent on the petiole forming a narrow wing, often to the base of the petiole. Inflorescence open panicle, sometimes becoming vegetative at the apex, to (4-)7-20 cm long; lateral branches slender, few flowered. Bracts as leaves or ovate, sessile; bractlets ovate-linear 7-12 x 1-3 mm. Calyx unequal, sepals slightly spathulate, larger upper sepal 10-12 x 1-2 mm, smaller lateral sepals 8-10 x 1 mm; covered in long non glandular and short glandular trichomes. Corolla purple, corolla tube 8-10x3 mm, upper lip to 18-22 mm long, glandular pubescent outside; lower lip 17-23 mm long, glabrous on inner surface. Androecium with filaments 25 mm long; anthers 4 mm long; staminodes 1 mm long, minute, lacking vestigial anther. Gynoecium with style 30 mm long; stigma 4 mm long; ovary 6 mm long, densely pubescent. Capsule not known. DISTRIBUTION. Restricted to the Morogoro region of Tanzania. HABITAT. Rain forest; 1000m. COLLECTIONS EXAMINED TANZANIA. Morogoro region, Kombola, 8 July 1933 Schlieben 4068 (LISC. MO, PRE); Kilosa district, 13 November 1987, Po$s et al. 8722 (UPS). Brillantaisia stenopteris is very closely related to B. oligantha from the Nchisi and Mughese mountains in Malawi. The two species have similar ovate, cuneate leaves with a long slender petiole wing. However, they can be easily distinguished due to the larger flowers and absence of trichomes on the inner surface of the lower corolla lip of B. stenopteris. The corolla tube of B. stenopteris is longer and thinner than that of B. oligantha, resembling the corolla tube of B. riparia, which is also closely related to B. stenopteris. 5. Brillantaisia owariensis P. Beauv., Flore d'Oware 2: 68 ( 1 8 1 8); Bentham in Niger Fl.: 477 (1849); Hooker in Bot. Mag.: 79: t. 4717 (1853); Durand & Schinz, Etudes fl. Congo 1:217 (1896); Burkill in Fl. trop. afr. 5: 40 (1899); Durand & Durand, Syll.fl. congoi: 417 (1909); Benoist in Bull. Soc. Bot. France 60: 335 (1913); Chevalier, Explor. hot. Afrique occ. franc . 1: 493 (1920); Hutchinson & Dalziel in FL W. trop. Afr. 2: 254 (1931); Heine in Fl. W. trop. Afr. 2nd ed., 2: 406 (1963). Type: Nigeria, Benin, Agathon, Palisot de Beauvois s.n. (Gl-lectotype; G!- isolectotypes). Belantheria belvisiana Nees in DC. Prodr. 11: 96 (1847). Type: as for B. owariensis. Brillantaisia patula T. Anderson in J. Proc. Linn. Soc. Bot. 7: 21 (1864); Burkill in Fl. trop. afr. 5: 41 (1899); Hiern in Cat. afr. pi. 1: 807 (1900); De Wild., Etudes fl. Bas- Moyen- Congo 1: 314 (1903-1906); Durand & Durand, Syll.fl. congoi.: 417 (1909); Benoist in Bull. Soc. Bot. France 60: 335 (1913); DeWild., Contr. Fl. Katanga 1: 143 & 2: 144 (1913); Mildbraed, Wiss. Erg. zweit. deut. Zentr.-Afr. Exped., Bot. (1922); Hutchinson & Dalziel in FL 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. trop. Afr. 2nd ed., 2: 406 ( 1 963); Heine in FL Gabon 13: 94, pi. 1 9, figs 1-6 (1966); Champluvier in Fl. Rwanda 3: 444-448 (1985). Type: Congo, Smith s.n. (K!-holotype; BM!, P!-isotypes). B. alata T. Anderson ex Oliv. in Trans. Linn. Soc. London 29: 125 (1875); S. Moore in J. Bot. 18: 197 (1880); Durand & Schinz, Etudes fl. Congo 1: 216-217 (1896); Durand & DeWild., in Bull. Soc. Roy. Bot. Belgique 36: 83 (1897); De Wild., Miss. Em. Laurent 1: 1 82 (1905); DeWild., Etudes fl. Bas- Moyen- Congo 1: 314 (1903-1906); De Wild., Contr. FL Katanga 1: 143 (1913). Type: Uganda, Unyoro, Speke & Grant 583 (K!-holotype). B. salviiflora Lindau in Bot. Jahrb. Syst. 17: 101 (1893); Burkill in 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. 92 336 (1913). Type: Togo, Bismarksburg, Buttner 341 (B|- holotype). B. nitens Lindau in Bot. Jahrb. Syst. 17: 102 (1893); Burkill in Fl. trap. afr. 5: 41 (1899); Lindau in Wiss Erg. deut. Zentr.-Afr. Exped., Bot. 2: 292 (191 1); Benoist in Bull. Soc. Bot. Fr. 60: 336 (1913); Mildbraed, Wiss. Erg. zweit. deut. Zentr.-Afr. Exped., Bot. (1922); Hutchinson & Dalziel in Fl. W. trap. Afr. 2: 254 (1931); Robyns, Fl. pare. nat. Albert. 2: 270-272 (1947); Heine in Fl. W. trap. Afr. 2nd ed., 2: 406 (1963); Blundell, Wildfl. Kenya: 104 ( 1 982); Champluvier in Fl. Rwanda 3: 448 ( 1 985); Blundell, Wild fl. E. Afr.: 389 (1987). Type: Cameroon, W. of Buea, Preuss 847 (Bt-holotype; K!-isotype). B. dewevrei De Wild. & Th. Dur. in Durand & De Wild., in Bull Soc. Roy. Bot. Belgique 38: 44-45 ( 1 899); Burkill in Fl. trop. afr. 5: 5 1 ( 1 899); De Wild. & Durand, Ann. Mus. Congo Ser. 3 Bot. 1 : 1 74- 175 (1901); De Wild. & Durand in Bull. Herb. Boissier 1: 833 (1901); Durand & Durand, Syll. fl. congol.: 416-417 (1909). Type: Congo, between Lukolela an Gombi, Dewevre s.n. (BR!- holotype). B. nyanzarum Burkill in Fl. trop. afr. 5: 39 ( 1 899); Robyns, Fl. pare. nat. Albert. 2: 272 (1947); Andrews, Fl. pi. Sudan 3: 171 (1956); Agnew, Upl. Kenya wildfls: 583 & 584 ( 1 974); Synnott in Comm. Forestry Inst. Occ. Papers 27: 68 ( 1985). Type: Kenya, Kavirondo, Scott-Elliot 6999 (K!-holotype). B. leonensis Burkill in Fl. trop. afr. 5: 41 (1899); Benoist in Bull. Soc. Bot. France 60: 335 (1913); Hutchinson & Dalziel in Fl. W. Trop. Afr. 2: 254 (1931). Syntypes: Sierra Leone, Scott-Elliot 3990a (K!), Windwood Reade s.n. (K!), Don s.n. (BM!). B. patula van welwitschii Burkill in Fl. trop. afr. 5: 41 (1899); Hiern in Cat. afr. pi 1: 807 (1900). Syntypes: Angola, near Sange, Quiapoza and Cuango Rivers, Welwitsch 5149 (BM!), Quibolo, Welwitsch 5182 (BM!, K!, P!). B. mahonii C.B. Clarke in Bull. Misc. Inform. 1906: 251 (1906); S. Moore in/ Bot. 45: 89 (1907); Lind, Comm.fl. pi Uganda: 140, fig. 82 ( 1962).Type: Uganda, Entebbe, Mahon s.n. (K!-holotype). B. bauchiensis Hutchinson & Dalziel in Fl. W. trop. Afr. 2: 253 (1931); Hutchinson & Dalziel Fl. W. trop. Afr. 2nd ed., 2: 407. Syntypes: Nigeria, Bauchi Plateau, Lely PI 29 (K!), Dent Young 195 (K!). Icones: P. Beauv., Flored'Oware 2: t. 100 fig. 2 (1818); Anderson in Trans. Linn. Soc. London29: pi. 124 ( 1 875); Hooker inBot. Mag.: 79: t. 4717 (1853) excluding glabrous fruit; Heine in Fl Gabon, Acanthacees: 13: pi. 19 figs 1-6 (1966); Agnew, Upl. Kenya wild fls: 584 (1974); Blundell, Wildfl. Kenya: pi. 264 (1982); Champluvier inf/. Rwanda 3: fig. 1 39: 2, 3 A-3D ( 1 985); Blundell, Wild./?. E. Afr.: pi. 849 (1987). Herbs or woody herbs to l-3(-4) m tall, viscid, often aromatic. Stems l-8(-15) mm in diameter, glabrous or pubescent towards apex. Leaves ovate to broadly ovate (35-)50-140(-210) x (25- )40-120(-185) mm; leaf base cordate to cuneate; apex acuminate sometimes markedly so; margin toothed, usually irregularly ser- rate, crenate towards apex with or without secondary toothing; eglandular pubescent above and below, sometimes densely pubes- cent; lateral veins 6-19 each side of the midrib; petiole (10-)30-90(-130) mm long, lamina decurrent in upper !/2-% length of petiole, occasionally entire length of petiole. Inflores- cence a many flowered terminal panicle (9-) 1 5-40(-60) cm long with 4-18 nodes; rachis glandular pubescent. Bracts ovate, (3- )20-50(-75) x (5-)15-25(-40) mm, usually falling early; bractlets linear oblong, to 20 x 5 mm, eglandular pubescent. Sepals subequal to unequal, linear, (5-)9-15(-22) x 1 mm long, rounded to slightly spathulate at the apex, glandular pubescent. Corolla K. SI DWELL pale purple to deep blue-purple often with yellow markings in throat; tube (5-)7-10(-14) x 2-6 mm long; upper lip (13-) 18- 30(-60) mm long, outer surface of upper lip glandular pubescent; lower lip (14-) 18-30(55) mm long, glabrous; apical lobes (l-)2- 5(-10) mm long. Androecium with filaments (20-)25-35(-40) mm long; anthers (2-)5-7(-10) mm long; staminodes (10-) 15- 25(-30) mm long with small vestigial anther at the apex. Gynoecium with style (15-)20-35(-45) mm long; stigma 2-4(-6) mm long; ovary 3-7 mm long, pubescent. Capsule (15-)18-26(- 33) x 2-3 mm with (10-)12-18(-24) seeds per locule; covered with eglandular and glandular trichomes. DISTRIBUTION. Throughout tropical Africa. HABITAT. Moist montane forest particularly in open shady places where competition is reduced in recently disturbed areas; 600-1600 SELECTED COLLECTIONS SIERRA LEONE: 18 November 1965, Adam 22109 (MO, P); 2 January 1966, Adam 22916 (MO, P); 27 November 1965, Adam 27084 (MO); Mt. Nimba, 14 December 1 966, Bos 2397 (WAG); 8°25'W7°32'N, 18 December 1967, Geerling & Bockdam 1859 (C, MO, WAG); LIBERIA: 10 January 1965, Adam 20549 (BR, K, MO, P, UPS); Grand Cape Mount Co., 21 December 1947, Baldwin 10778 (K, MO); 6 November 1947, Baldwin 101 82 (K, MO); Loffa country, 21 December 1966, Bos 2552 (K, WAG). IVORY COAST: 24 November 1909, Chevalier 22421 (P); 20 km de Man sur route de Danane, 23 November 1956, de Wilde 859 (WAG); MontTonkoui, pres de Man, 3 December 1985, Ake Assi 17170 (G). GHANA: Bauchi Plateau, 17 April 1955, Morton K416 (K, WAG); Kofordua, Dalziel 148 (C, E, K, M, MO, PRE, WAG). NIGERIA: Mandaga, Nambila Plateau, 4 January 1955, Latilo & Daramola FHI 28992 (FHO); Shasha Forest Reserve, Richards 3108 (BM, MO, NY); 1 1 December 1976, Ariwaodo 33. CAMEROON: 13 February 1927, Dalziel 8232 (US); Buea, Preuss 1029 (BM, M); Dschang, December 1938, Jacques-Felix 2600 (P); 31 km on Ebolowa-Ambam Road, 1 September 1974, de Wilde 7544 (WAG); Lake Oku, 6°13'N 10°28'E, Thomas 4381 (MO, YA); GABON: Woleu-Ntem Province, 14 November 1933, Le Testu 9395 (BR); CENTRAL AFRICAN REPUBLIC: 10 Decem- ber 1936, Eckendorfll (P); CONGO: Sangha, 2°N 13°55'E, 27 November 1991, Thomas 9005 (MO); DEMOCRATIC REPUPUBLIC OF THE CONGO: 12January 1914, Bequaert 1905 (BR); Kivu, Lac Mokoto, 27 July 1953, van cler Ben 648 (BR); Yangambi region, Yabahondo village, October 1952, Germain 8141 (BR, W, Z). RWANDA: Rwaza, SE de Ruhengeri, 23 February 1972, Auquier 2684 (BR). BURUNDI: 2 June 1969, Lewalle 3795 (BR). UGANDA: Bigera River, 3 August 1906, Bagshawe 1131 (BM). KENYA: December 1933, Dale 3208 (BR, FHO); January 1932, Brodhurst- HU1693 (Z).TANZANIA: 26January 1 99 1 , Ka \ombo 1072(MO).ANGOLA: 24 June 1958, Montiem, Santos & Murta 205 (PRE). Brillantaisia owariensis is the most widespread and variable of all species in the genus and has caused many problems of species delimitation for botanists over the years. After extensive herbarium studies, supplemented by field work in Cameroon and Tanzania, I consider it impossible to split this group into discrete taxa and recognize it as a very diverse 'oc/z/o-species' (sensu White, 1962: 79) and accept that, although unsatisfactory, this is the only viable option without further extensive studies in the field. Brillantaisia owariensis sensu meo encompasses all fairly robust, woody Brillantaisia plants with a paniculate inflorescence, toothed leaves with a winged petiole and pubescent fruit. Morphological variation within the group appears more pronounced in certain parts of Africa. In East Africa, for example, clear morphological differences can be observed between different isolated groups and two representative specimens may look nothing like one another when the extremes of morphological variation are observed. However, when morphologi- cal variation is studied accross the whole of Africa, character differences break down and distinct groups cannot be distinguished. REVISION OF BRILLANTAISIA 93 Variable characters include leaf shape, leaf base shape, leaf margin, number of lateral veins, calyx shape, inflorescence density, flower size, fruit length/width and fruit pubescence. The morphological variation within Brillantaisia owariensis is informally described below in terms of two main groups: the nitens group and the patula group. Plants which tend to differ from typical B. owariensis in possess- ing rather finely toothed leaves, a somewhat narrow petiole wing, larger number of lateral nerves and a relatively dense inflorescence could be assigned to the nitens part of Brillantaisia owariensis. Brillantaisia nitens was described by Lindau (1893) as being dis- tinct from all other Brillantasias due to the leaf pubescence, which is comparatively dense for the genus. Plants with characters of the nitens group tend to occur in the forests of Cameroon and further west in tropical Africa. Plants that are slightly more robust than typical Brillantaisia owariensis, with velvety-tomentose leaves, have previously been distinguished as Brillantaisia bauchiensis in Nigeria and B. leonensis in Sierra Leone. Specimens with longer calyx lobes and sub-persistent bracts were previously assigned tofi. mahonii. Specimens from East Africa that generally have the same morphological characters as those of the nitens group, but differ in the presence of more persistent bracts, have often been identified as B. nyanzarum. The patula group: The most distinct node of variation within Brillantaisia owariensis is seen on specimens that can be informally described as the patula group. These plants have a very open, lax, 'zigzag' inflorescence architecture and often have large flowers and a linear calyx. Specimens with these characteristics do not seem to be restricted to one area of Africa and have been collected from Guinea to Zaire and on Sao Tome. Most patula specimens have been collected from the Congo basin. Leaves of the patula 'node' often, but not always, have a distinct wing to the base of the petiole (Mocquery's 115, Z; Richards 3018, BM) which is occasionally amplexicaul. 6. Brillantaisia kirungae Lindau in Gotzen, Durch Afrika von Ost nach West, Sonderabdr.: 9 (1896); Burkill in Fl. trap. afr. 5: 42 (1899); Durand & Durand, Syll. fl congol. : 417 (1909); Robyns, Fl. pare nat. Albert 2: 269 (1947). Type: Zaire, Mount Kirunga, von Gotzen 48 (Bf-holotype). Fig. 12. B. ulugurica Lindau in Bot. Jahrb. Syst. 22: 112 (1897); Burkill in Fl. trop. afr. 5: 43 (1899); Brenan & Greenway, Checklist brit. empire 5 Tangan. terr. 2: 6 (1949). Syntypes: Tanzania, Uluguru Mts, Stuhlmann 8850 (B|) & Stuhlmann 9224 (B|) B. subulugurica Burkill in Fl. trop. afr. 5: 42 (1899); Binns, Check- list herb. fl. Malawi: 12 (1968). Type: Mozambique, Makua, Namuli Hills, Last s.n. (K!-holotype). B. grandidentata S. Moore in/ Bot. 45: 331 (1907). Type: Uganda, Toro, Fort Portal, Bagshawe 1270 (BM!-holotype). B. cicatricosa var. kivuensis Mildbr. in Bull. Jard. Bot. Etat 17: 86 (1943); Robyns, Fl. pare nat. Albert 2: 270 (1947). Type: Zaire, Lake Magera, de Witte 1434 (BR!-holotype; BR!-isotype). B. nitens sensu Agnew, Upl. Kenya wildfls: 583 (1974), non Lindau (1893). Icones: Champluvier in Fl. Rwanda 3: fig. 139, 1A-1D (1985); Fischer & Hinkel, Nat. env. Rwanda: fig. 68 (1992). Stout shrubby herbs 1-2 m tall to shrubs or small trees 2-5(-7) m tall with bole 1.4 m high and 12 cm diameter, viscid and sometimes aromatic. Stems erect, 5-20(-120) mm across, shortly puberulent. Leaves, broadly ovate 60-320 x 50-240 mm; leaf base cordate or truncate to attenuate or cuneate; apex acute to acuminate; margin irregularly toothed, teeth single or double, smaller becoming crenate towards the apex, rarely entire; puberulous; lateral veins 7-13 each side of the margin, lower laterals closer together; petiole (0-)10-140 mm long, lamina gradually decurrent on top '/2-% of petiole, upper pair of leaves at base of inflorescence sessile. Inflorescence a terminal thyrse 15-25(^0) cm long, lateral branches erect, at 30-50 degrees to the main rachis, flowers crowded at end of lateral branches; rachis shortly pubescent; trichomes, red-brown, non glan- dular, sometimes straggly purple trichomes. Bracts broadly ovate to ovate, occasionally elliptic, base slightly amplexicaul, apex rounded to acute; bractlets ovate(-linear), falling late or remaining in fruit. Sepals unequal, larger upper sepal linear, rounded at tip or slightly spathulate, (8-)10-14(-19) x l-3(-5) mm, often twice as wide as lateral sepals (clearly seen in bud), occasionally very large; smaller lateral sepals linear 8-15 x 1 mm; densely pubescent dorsally. Corolla pale blue to bright blue-purple or pink-purple, with darker markings on lower lip; corolla tube 6-10 x 3-6 mm, occasionally inflated; upper lip 19-35 x 6-10 mm, evenly covered in dense, eglandular and glandular trichomes; lower lip 19^40 x 10-18(-25) mm, ridged on upper surface; apical lobes triangular, 3-10 x 2-8 mm. Androecium with filaments 20-25(-30) mm long; anthers (3- )5-8 mm long; staminodes 8-12 mm, very slender often with well developed vestigial anther. Gynoecium with style 25^0 mm long; stigma 2 mm long; ovary 5-10 mm, covered in tangled glandular trichomes, sometimes also with non glandular trichomes. Capsule 30-40 x 2-5 mm with 8-10(-16) seeds per locule; black, glandular puberulous. Specimens often drying black. DISTRIBUTION. East and central Africa, from Kenya south to Zim- babwe and Mozambique, west to Uganda and Eastern Zaire. HABITAT. In gaps in mid altitude and montane forest, forming thickets, often along streams. Very local; 1000-3500 m. SELECTED COLLECTIONS DEMOCRATIC REPUPUBLIC OF THE CONGO: Kivu, 27 July 1959, Cambridge Congo Expedition 146 (BM, LISC, US). RWANDA: Pare des Volcans, 27 July 1974, Van der Veken 10350 (BR). BURUNDI: Rwegura, 29 May 1969, Lewalle 3640 (BR, G); Bubanza, 22 June 1980, Reekmans 9408 (K, MO, UPS). SUDAN: Imatong Mts, Gilo Village, 8 November 1980, Friis & Vollesen 58 (C). UGANDA: Ruwenzori Mts, Namwamba Valley, 17 January 1935, Taylor 3156 (BM, MO, NY, S). KENYA: 18 March 1977, Faden & Faden 77/919 (BR, US, WAG). TANZANIA: Rungwe, Kiwara River, 9 August 1 949, Greenway & Eggeling 8394 (PRE); Morogoro, Bunduki, 1 7 August 1 95 1 , Greenway & Eggeling 8594 (FHO); Nugwi River, 1 9August 1952, Carmichael 102 (FHO); Mbisi Mts, 17 June 1960, Leach & Brunton 10069 (SRGH); Ufipa, 6 August 1960, Richards 12983 (SRGH); Kilosa District, Ukaguru Mts, 7 August 1972, Mabberley 1403 (K), Mufindi Tea Estate, 17 August 1984, Bridson & Lovett 544 (MO). MOZAMBIQUE: Namuli Peaks, 26 July 1962, Leach & Schelpe 11470 (LISC, SRGH); MALAWI: Nchisi Mts, 4 September 1929, Bunt Davy 1228 (FHO); Kafwimba Forest, 4 July 1973, Pawek 6978 (C, PRE, SRGH, UPS). ZIMBABWE: Ngoruma reserve, 14 August 1962, Plowes 2261 (LISC, SRGH); Vumba, 20 September 1968, Mutter 802 (SRGH). Brillantaisia kirungae is the largest and most woody of the species of Brillantaisia and is easily recognized by the woody or shrubby habit; highly irregularly toothed leaves; large flowers in a crowded erect inflorescence; broad upper calyx lobe and large, few-seeded fruit. This species is rather variable in leaf shape and toothing (most collections lack the larger lower leaves, so measurements given above need refining after more field studies); the degree of contrac- tion of the inflorescence; density of the indumentum on both the inflorescence and fruit; and flower size, particularly the size of the lobes at the apex of the lower lip. Previous treatments of Brillantaisia kirungae have all divided it into more than one species. For example, 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 with subglabrous sepals and numerous seeds; B. kirungae with lanceolate bracts; B. ulugurica with large flowers and large lobes at the apex of the lower corolla lip and a new species, B. subulugurica which was considered distinct from B. ulugurica due to the shorter corolla tube. More recently, Champluvier (1985) recognized two species, B. kirungae and B. cicatricosa, separated on the size of the upper sepal. All of the characters used by previous authors to divide this species are continuously variable when the entire geographical variation of the group is accounted for. In his treatment of the genus for the Flora ofTropical East Africa and Flora Zambesiaca, Vollesen (in prep a & b) recognizes two species within this group, B. cicatricosa and B. ulugurica. He considers plants of B. cicatricosa to have irregularly large-toothed leaves; smaller lower corolla lip lobes; smaller seeds, to lack glandular trichomes on the fruit and to have a more northerly distribution from Kenya, Uganda, Burundi, Rwanda and Zaire. Brillantaisia ulugurica sensu Vollesen is recog- nized as distinct on the basis of smaller irregular teeth on the leaves; leaf margin becoming entire towards the apex; larger lobes on the lower corolla lip; larger seeds; both glandular and non glandular trichomes on the fruit; and a more widespread geographical range in Uganda, Burundi, Zaire, Tanzania, Malawi, Mozambique, Zimba- bwe and Zambia. Two characters used by Vollesen to distinguish B. cicatricosa from B. ulugurica are not overlapping or continuous in his species descriptions: seed size and fruit hair type. I have studied the two species across their entire range which includes specimens from Sudan and Angola. It appears that although some 'cicatricosa- like' specimens do have smaller flowers, deeply toothed leaves with a more cordate base, smaller seeds and no glandular tricomes on the capsule, these characters are continuously variable and there is no morphological data to suppport separatation of the two species. A new species from Tanzania was mentioned by Vollesen (in prep. a) as closely related to Brillantaisia ulugurica. Plants belonging to the new putative new species were recognized primarily by the presence of long, rather synthetic looking, purple, eglandular trichomes on the panicle and calyx, and by the larger capsule (no measurements given). Several intermediates between B. ulugurica sensu Vollesen, with short puberulent trichomes on the panicle and short glandular and eglandular trichomes on the calyx, and the purple haired sp. nov. cf. B. ulugurica have been studied (e.g. MOZAMBIQUE: Leach & Schelpe 1 1470; MALAWI: Chapman & Chapman 921 \ Mt Mulanje; Burn Davy 1228) and I consider the variation to represent one species. Within the single species recog- nized in this treatment, plants with a more lax inflorescence, more slender sepals and smaller flowers tend to occur further north-north west (SUDAN, Gilo Village, Friis & Vollesen 58 C; RWANDA, Pare des Volcans, Van der Veken 10350, BR; UGANDA, Kanaba Pass, Makerere College 29) whereas larger more robust plants are more commonly from Malawi and Mozambique. Typification of this species is somewhat problematic as men- tioned by Champluvier ( 1 985). I disagree with her decision to retain the epithet cicatricosa. Specimens of B. kirungae sensu meo have historically been named asfi. cicatricosa, B. kirungae, orfl. ulugurica, however, the type specimens of all three of these names were destroyed in Berlin. The protologue of the earliest name, B. cicatricosa Lindau, is somewhat ambiguous and does not fit the species descrip- tion above. Although the Latin diagnosis of B. cicatricosa mentions the large upper sepal of this species, several characters do not fit the group well, notably subglabrous sepals and 16-20 seeds in the capsule. The original description of B. cicatricosa Lindau men- tioned that the species was close to 5. pubescens T. Anders., however without a type specimen of B. cicatricosa the variation observed by Lindau when he made this seemingly misplaced statement cannot be checked. It may well be the case that Lindau based his observation on the illustration of Fitch in Anderson (1875) which has caused considerable confusion in Brillantaisia (see p. 86). Brillantaisia cicatricosa sensu Burkill ( 1 899) adds to the ambiguity of this name as he describes the species as having entire, glabrous leaves, a loose inflorescence, nearly glabrous sepals and 1 6-20 seeds, whereas the species recognized here has irregularly toothed pubescent leaves and a crowded inflorescence. A variety of B. cicatricosa Lindau was formally recognized by Mildbraed ( 1 943) on the presence of glandu- lar trichomes on the calyx. This taxon is illustrated in Flore du Rwan da (Champluvier, 1985) and clearly falls within B. kirungae as delimited here. However, I am not certain that Mildbraed saw the type specimens of B. cicatricosa or that he understood Lindau's concept of that species. Therefore, as the protologue offl. cicatricosa Lindau is ambiguous and the description of B. cicatricosa by Burkill does not fit this species, I treat this name as dubious, and use the second earliest name, B. kirungae, for this species. 7. Brillantaisia vogeliana (Nees) Benth. in Hooker, Niger FL: 477 (1849); Anderson inJourn. Linn. Soc. Bot. 7: 21 (1864); Burkill in Fl. trap. afr. 5: 40 ( 1 899); Benoist in Bull. Soc. Bot. France 60: 335 (1913); Hutchinson & Dalziel in Fl. W. trop. Afr. 2: 254 (1931); Exell, Cat.pl. S. Tome: 260(1944); Mildbraed, Wiss. Erg. zweit. deut. Zentr.-Afr. Exped., Bot.: 192 (1922); Robyns, Fl. pare. nat. Albert 2: 212 (1947); Heine in Fl. W. trop. Afr. 2nd ed., 2: 406 (1963); Heine inFl. Gabon 13: 92 (1966). Type: Fernando Po, Vogel 179 (K!-holotype; K!-isotype). Fig. 13. Leucoraphis vogeliana Nees, in DC., Prodr. 11: 97 (1847). Brillantaisia molleri Lindau \nBot. Jahrb. Syst. 17: 99 ( 1 893). Type: Sao Tome, Moller 33 (87) (Bt-holotype; COI!, K!-isotypes). B. preussii Lindau in Bot. Jahrb. Syst. 17: 100 (1893). Syntypes: Cameroon, Barombi, Preuss 320 (K!-isosyntype) Buea, Preuss 998 (BM!, K!-isosyntypes) B. soyauxii Lindau in Bot. Jahrb. Syst. 17: 101 (1893); Burkill in Fl. trop. Afr. 5: 39 (1899); Benoist in Bull. Soc. Bot. France 60: 335 (1913); non Heine in Fl. Gabon 13: 84 (1966). Type: Gabon, Munda, Sibange Farm, Soyaux 454 (Bt-holotype; E!, H!, US!,WAG!-isotypes). B. schumanniana Lindau in Bot. Jahrb. Syst. 17: 102-103 (1893). Type: Cameroon, Braun 47 (Bf-holotype). Icones: Heine in Fl. Gabon 13: pi. 18, figs 1-6 (1966); Heine, in Fl. W. trop. Afr. 2nd ed., 2: 406, fig. 300 (1963). Erect herbs to 0.2-1 m tall. Stems erect, 2-4 mm in diameter, branching near base, leaf petioles fused across each node, gla- brous or shortly pubescent with a fringe of trichomes at the nodes. Leaves ovate to broadly ovate 35-100(-l 10) x 45-120(-150) mm, papery, bright green, paler below, evenly pilose; leaf base truncate or cordate, occasionally shortly attenuate; apex slightly acumi- nate, rounded; margin irregularly serrated, dentate at base of lamina around the widest part of the leaf, crenate towards the apex of the leaf; lateral veins 7-13 each side of midrib, prominent; petiole (30-)40-120 mm, lamina decurrent on upper third of peti- ole, wing often toothed at top. Inflorescence a terminal lax thyrse (8-) 1 2-20(-30) cm long, with smaller lateral inflorescences in axils of lower leaves; rachis shortly glandular pubescent with slender non glandular trichomes. Bracts to 7 x 15 mm, often falling early; bractlets linear elliptic to 2 mm long, glabrous with fringe of irregular glandular trichomes. Sepals unequal, slightly spathulate, larger upper sepal 6-15 mm long, smaller lateral sepals 6-10(-12) mm long, viscid glandular pubescent. Corolla clear 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 paler, tube 2^4 x 5-10 mm, slightly shorter than the calyx lobes; upper lip 1 l-15(-25) mm covered with glandular and non glandular trichomes on outer surface, more so towards the apex; lower lip 10- 15(-25) mm long; apical lobes to 1 x 1 mm, central tooth broader than the laterals, glabrous. Androecium with filaments 12-17(-24) mm long, glabrous; anthers 2-4(-6) mm long; staminodes 6-10 mm long, flattened at tip to vestigial anther, glabrous. Gynoecium with style 15-20 mm long; stigma 1.5-3 mm long; ovary 3-4 mm long. Capsule (10-) 17-21 (-24) x 1-2 mm with 20-24 seeds in each locule, indumentum of slender white non glandular trichomes and short glandular trichomes. DISTRIBUTION. Throughout West Africa and on the islands of the Gulf of Guinea across Central African Repulic and Congo to Sudan, Uganda and Kenya. HABITAT. A weedy species common in wasteland, clearings, plan- tations and along roadsides; 60-1200 m. SELECTED COLLECTIONS IVORY COAST: Mont Momi, 29 October 1 966, AkeAssi 9 1 36 (G). GHANA: Atewa Range N. of Kibi, 3 November 1990, Manktelow, Steiner &Amponsah 89 (UPS). CAMEROON: N'Kolbisson, 3 November 1963. de Wilde & de Wilde-Duyfies 1 130 (K, MO, WAG, YA); Nkolbison, 8 km W. of Yaounde, Mount Minloua, 19 November 1964, Raynal 1 1964 (P); 17 km along Kribi- Lolodorf road, 22 September 1969, Bos 5387 (C, K, MO, PRE, UPS, WAG, YA); Chutes de Ntem, 40 km ESE of Campo, 10 December 1979, Letouzey 15340 (P); Sao Nicolau, 9 January 1980, de Wilde, Arends & Groenendijk 3 (WAG); crossing of road from S. Joao dos Angolares to Ribeira Peize, 12 January 1980, de Wilde, Arends & Groenendijk 95 (WAG); Kumba area, 13 October 1984, Thomas & Nemba 4048 (MO). BIOCCO: October 1911, Mildbraedl028 (H). CENTRAL AFRICAN REPUBLIC: 27 km S. of Nola, 29 November 1965, LeeuwenbergllQQ (BM, C, K, LISC, WAG). CONGO: Sulongo, 22 March 1959, Everard 6002 (PRE); Piste Meya - Mpassa, 28 April 1965, Farron 4085 (P). SAOTOME: Famosa, 5 January 1949, Espirito Santo 68 (BM); Monte Cape, 5 March 1968, Espirito Santo 4298 (LISC); road from Nora Moka to Manuel Jorge River, 1 5 October 1 993, Figueiredo & Arriegas 33 (FHO, K, LISC). GABON: Belinga Mines de Per, 25 June 1966, Halle 4058 (P); Boka-Boka, Mount Bengoue, 4 March 1979, Florence 1706 (P); Woleu-Ntem, 2 May 1986, Louis 2077 (WAG); 10 km from Belinga, 5 December 1986, Bos, van derLaan & Nzabi 10755 (K, US, WAG). DEMO- CRATIC REPUPUBLIC OF THE CONGO: Dyuma, Pare National Albert, 11 October 1955, de Witte 12827 (MO). SUDAN: Imatong Mountains, Talanga, 1 December 1980, Friis & Volleson 612 (C). UGANDA: Budongo forest, December 1935, Eggeling 3323 (K); Mabira forest, 8 November 1938, Loveridge25 (K). KENYA: Kakamega forest, 1 1 December 1956, Verdcourt 1683 (K). Brillantaisia vogeliana is often locally abundant as a weed along road and path sides and in clearings in secondary vegetation, small farms, plantations, etc. It is readily identified by leaves that are often cordiform, characteristically irregularly toothed and often with a toothed petiole; the highly branched inflorescence; rela- tively small flowers which are usually a clear mauve-violet and paler in the throat, and the pubescent fruit which have numerous seeds per locule. This species might be confused with B. owariensis, on the basis of the lax inflorescence and toothed leaves, and Vollesen tends to place East African specimens with rounded teeth and a wider inflorescence into that species. Plants of B. vogeliana differ, from those of B. owariensis, in possessing thinner, papery leaves with a more dentate, irregular leaf margin and pronounced dentate margin at the apex of the petiole wing; a more highly branched, slender inflorescence; smaller flowers and fruit with more seeds. A few specimens of B. vogeliana from the Congo basin have small flowers but larger leaves than typical B. vogeliana and seem to approach B. owariensis. White-flowered forms of this species are fairly common. Exell (1944: 261) noted the two colour forms and stated They seem to be identical except for the flower-colour and they always grow together, the purple- flowered form being rather more common'. 8. Brillantaisia lamium (Nees) Benth. in Hooker, Niger FL: 477 ( 1 849); Burkill in FL trop. Afr. 5: 38 ( 1 899); Hiern in Cat. afr. pi. 1: 807 (1900); Durand & Durand, Syll.fl. congol.: 417 (1909); Benoist in Bull Soc. Bot. France 60: 334 (1913); Chevalier, Explor. hot. Afrique occ. franc,. 1: 493 (1920); Mildbraed, Wiss. Erg. zweit. deut. Zentr.-Afr. Exped., Bot. 2: 90(1922); Hutchinson & 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 (1963); Heine inFl. Gabon 13: 88(1966).Type:A/we// 1842 (K!- lectotype) (P!-isolectotype). Fig. 14. Leucoraphis lamium Nees in DC., Prodr. 11: 97 ( 1 847). Type: As for Brillantaisia lamium. Brillantaisia palisotii Lindau in Bot. Jahrb. Syst. 17: 99 (1893); Henriques in Bol. Soc. Brot. 10: 146 (1893). Lindau in Nat. Pflanzenfam. 4(3b): 296, fig. 1 1 9 ( 1 895). Syntypes: Sierra Leone, Freetown, Preuss 16 (B); Sierra Leone, Freetown, Afzelius s.n. (UPS!); Togo, Bismarksburg, Buttner 231 (B), Burner 320 (B); Cameroon, Mungo, Bucholz s.n. (BM!); Sao Tome, Quintas 1235(88) (BM!, BR!, COI, Z!), Quintas 1 132(89) (COI); Angola, Luanda, Pogge 1123(7). B. eminii Lindau in Bot. Jahrb. Syst. 17: 103 (1893); Lindau in Pflanzenw. Ost-Afr. C: 366 (1895 d); Burkill in FL trop. Afr. 5: 38 (1899); Jex-Blake, Wildfl. Kenya: 103, fig. 84 (1948); Brenan & Greenway, Checklist brit. empire 5 Tangan. Terr. 2: 5 (1949); Andrews, FL pi. Sudan 3: 172 (1956); Synnott, Comm. forest. inst. occ. papers 27: 68 (1985). Syntypes: Tanzania, Bukoba, Stuhlmann 3664 (K!); Stuhlmann 3995 (Bt). B. owariensis sensu Hook, in Bot. Mag.: t. 4717, fig. 3 (1853); T. Anderson in Journ. Linn. Soc. Bot. 7: 339 (1863); Engler in Bot. Jahrb. Syst. 7: 339 (1886), non P. Beauv. (1818). B. subcordata De Wild. & T. Durand in: T. Durand & De Wild. Bull Soc. Roy. Bot. Belgique 38: 44-^5 (1899); De Wild. & Durand Ann. Mus. Congo 1: 175-176 (1901); DeWild. Miss. Em. Laurent 1: 182, t. 45 (1905); Durand & Durand, Syll. fl. congol.: 418 (1909). Type: Zaire, Bokakata, Dewevre 802 (BR!-syntypes). B. subcordata var. macrophylla DeWild. &T. Durand. in Contrib.fl. Congo 1, 2: 47 (1900). Type: Locality uncertain, village on the Congo river, Duchesne 13 (BR!-holotype). Icones: Lindau in Nat. Pflanzenfam. 4(3b): fig. 119A-G (1895); Heine, in FL W. trop. Afr. 2nd ed., 2: fig. 300 (1963); Heine, Fl Gabon: 13: pi. 17 figs 4-8 (1966). Perennial herbs (7-)20-100(-200) cm tall, creeping to erect, occa- sionally climbing with adventitious roots at nodes. Stems 4.5 mm in diameter, occasionally with longitudinal wings on angles, gla- brous to velvety-tomentose. Leaves ovate to broadly ovate (12-)25-80(-l 10) x (20-)45-100(-180) mm or rarely oblong and 55-90 x 20-30 mm, sparsely pilose to tomentose with white silky trichomes above, pilose to velvety-tomentose below; leaf base cordate, subcordate or truncate occasionally shortly attenuate; apex acute with short acuminate tip or obtuse and abruptly contracted to acuminate tip; margin entire, rarely subentire to slightly crenate; lateral veins 6-10(-13) each side of midrib; upper pair of leaves at base of terminal inflorescence sessile, petiole (0-)8-80(-l 10) mm long, unwinged or lamina decurrent close to apex of petiole. Inflo- rescence a terminal, open, branching thyrse, 12.5-35 cm long. 98 K. SIDWELL Fig. 14 Brillantaisia lamium (Nees) Benth. A. Habit x 0.7; B. Corolla dissection showing androecium x 2; C. Gynoecium x 2; D. Calyx x 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 inflorescence spreading almost at right angles to main axis, rachis usually sparsely covered in capitate glandular trichomes. Bracts ovate, adaxial surface pilose, often with slender multicellular trichomes, abaxial surface glabrous; bractlets rhomboid to linear, sessile, margin entire, adaxial surface covered with multicellular non-glandular trichomes, abaxial surface glabrous; becoming smaller and linear towards ends of lateral branches. Sepals sub-equal, linear, apex rounded, 4-7 mm long, green often tinged purplish; glabrous or with a few long glandular trichomes. Corolla deep bright blue- purple or violet occasionally pale lilac, one lip, often the upper; may be paler than the other; tube 5-8 x 2-4 mm, pale than lips, flushed inside with yellow, rarely completely white; upper lip (15-)22-30(- 35) mm long, outer surface finely covered in glandular multicellular trichomes; lower lip (13-)19-27(-30) mm long, inner surface occasionally sparsely pubescent; apical lobe to 2 x 3 mm.Androecium with filaments 12-35 mm long, usually white; anthers 4.5-6 mm long; staminodes exserted for 4-5 mm from corolla tube, small vestigial anther. Gynoecium with style to 40 mm long, sparsely pubescent at base, stigma 3-5 mm long; ovary 3-4 mm long. Capsule 22-30 x (2-)3-5 mm with 14-16 seeds per locule, blunt tipped, green, tinged red at apex when immature, brown-black when mature and/or dried; glabrous or with a few erect, non-glandular trichomes at the apex, covered in black gland dots. DISTRIBUTION. Throughout wet forest regions of west and central Africa, east to southern Sudan, through Zaire to northern Angola and through Uganda to northwest Tanzania. HABITAT. Forest clearings, shade beside rivers or streams, on lake shores or in swamps, often waterlogged/with roots in water, dis- turbed areas, roads, paths and plantations. In west Africa often very weedy. 'In moist exposed woody situations almost everywhere' (on Princes Island, Hiern, 1900: 807). Common; 200-1600 m. SELECTED COLLECTIONS SIERRA LEONE: Njala, 30 October 1928, Deighton 1419 (BM, K, MO). LIBERIA: Louisiana, 1 1 November 1966, Bos 2310 (WAG); 10 miles N. of Monrovia, September 1 970, Jansen 2207 (WAG). IVORY COAST: Sassandra, 1 8 June 1 963, de Wilde 280 (K, WAG, Z); Foret de YAPO, 8 km S. of Becedi- Bugnan, 6 August 1963, de Wilde 641 (WAG); 27 October 1976, Fabrigues 3306 (P). GHANA: Banyimade, 10 September 1956, Cudjoe, 161 (WAG). TOGO: Kpandu, 1924, Robertson 62 (BM). NIGERIA: Oban, 191 1, Talbot 981 (BM, K); Bauchi Plateau, August 1930, Lely 652 (MO); Cross River, 27 September 1985, Chile 27 (MO). CAMEROON: Bibundi, October 1891, Junger 248 (UPS); Bipindi, 1896, Zenker 1104 (BM); NW of Tibati, 9 September 1963, Letouzey 5666 (P); Bakaka forest, 27 August 1971, Leeuwenberg 8206 (H, LISC, MO, UPS, WAG); Moundjo, 27 October 1975, de Wilde 8522 (WAG). BIOCCO: Malabo, near Bae Basula, 13 September 1986, Carvalho 2445, (BM); Sooye", S. of Mamon, 16 February 1945, Roberty 6590 (G, Z). CENTRAL AFRICAN REPUBLIC: Koumbala, 25 October 1983, Fay 6055 (MO); Bambari, November 1923, Tisserant 1352 (P); Tisserant 3167 (BM, P). SAO TOME: 18 March 1968, Espirito Santo 4312 (LISC); Monte Cape, 14 August 1956, Monod 1 1681 (BM). DEMO- CRATIC REPUPUBLIC OF THE CONGO: 26 June 1914, Bequaert 4836 (BR). BURUNDI: Ruyigi, 3 May 1980, Reekmans 9060 (MO, SRGH). SUDAN: Imatong mountains, Talanga, 2 December 1980, Friis & Vollesen 6 1 9 (C); Imatong Mountains, Acholi Hills, November 1 98 1 , Howard UTT 1 6 (C). ETHIOPIA: Bidgood s.n. (K). UGANDA: shores of Lake Nabugabo, 1 3 November 1934, Taylor 1660 (BM); Malabigambo forest, 15 August 1950, Dawkins D6 1 6 (BM). KENYA: Itare river, October 1940, Copley B 1 1 88 (G); Nairobi, 12 October 1972, Hansen 713 (C). ANGOLA: Loanda, 1903, Gossweiler 131 (BM). Brillantaisia lamium is a widespread species that is readily recog- nized by the cordate leaves with an entire margin and unwinged 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. Fruit are characteristically glabrous but occasionally with a tuft of trichomes at the apex. East African specimens of Brillantaisia lamium are often far more robust and pubescent than West African representatives, which tend to be slender herbs; however, the variation appears to be continuous and there are no morphological and/or geographical data to support subdivision of the species. Vollesen (in prep, a) notes 'A similar pattern where eastern plants from higher altitudes have larger flow- ers than western lowland plants is known from other Acanthaceae, e.g. Anisosepalum alboviolaceum '. Brillantaisia lamium can form large weedy populations in areas of recently cleared vegetation or plantations of oil palm, cocoa or bananas, and in open grassy areas at higher altitude. In Cameroon slender low growing forms with very deep purple flowers were cultivated as an ornamental in small village compounds on Mount Cameroon. Collections from Kenya are of naturalized plants from Uganda and in swamp areas around Nairobi the species is now quite common. 9. Brillantaisia lancifolia Lindau in Engl. Bot. Jahrb. Syst. 17: 98 (1893);Burkill mFl. trop.Afr. 5: 40(1899); Benoist in Bull. Soc. Bot. France 60: 335 (1913); Heine in Fl. W. trap. Afr. 2nd ed., 2: 406 (1963); Heine in Fl. Gabon 13: 86 (1966). Type: Gabon, Sierra del Crystal, Mann 1688 (Bt-holotype, Kl-lectotype, P!- isotype). Fig. 15. Map 3. B. talbotiiS. Moore Cat. pi. TalbotOban distr.: 75 (1913); Hutchin- son & Dalziel in Fl. W. trap. Afr. 2: 254 (1931). Type: S. Nigeria, Oban Talbot 2000 (BM!, K!-syntypes). Icon: Heine, Fl Gabon, Acanthacees: 13: pi. 19 figs 1-3 (1966). Perennial herb to 0.3-0.5 m tall. Stems 2-3 mm in diameter, gla- brous to densely pubescent, petiole bases fused forming ridge across nodes. Leaves ovate-elliptic to lanceolate 25-130(-190) x 8-30(- 50) mm, smaller on axillary shoots, glabrous or very sparsely pubescent; base cuneate; apex acute; margin entire to very shallowly serrated or crenate, subentire towards apex, entire towards base; main lateral veins 5-8 on each side; petiole 5-22 mm long, unwinged, lamina decurrent at apex. Inflorescence a delicate, open, few-flow- ered panicle 50-150 mm long, rachis glandular pubescent. Bracts often caducous, linear-lanceolate, bractlets if present linear lanceo- late with rounded apex, sparsely pubescent. Sepals unequal, linear, tapering gradually towards the apex, larger upper sepal 6-10 mm long, smaller lateral sepals to 5-8 mm long, pubescent, with capitate glandular trichomes. Corolla deep blue to violet, often with brown marking in throat; corolla tube 5-10x2 mm, pale purple sometimes tinged green; upper lip 10-15(-20) x 2-3 mm, glandular pubescent, particularly of edge of upper lip; lower lip 10-18 x 6-8 mm, apical teeth rounded-triangular 1-2 x 1 mm. Androecium with filaments to 20 mm long, anthers 3-4 mm long; staminodes usually prominent, around half length of the corolla lips. Gynoecium with style 15-28 mm, stigma 1 mm, ovary 3 mm long, shortly glandular pubescent. Capsule 15 x 1-1.5 mm with c. 9-10 seeds per locule, brown when dry, evenly glandular pubescent. DISTRIBUTION. Restricted to a few montane regions in Nigeria, Cameroon and Gabon. HABITAT. On damp rocks, in dark ravines or at edges of waterfalls; 300-700 m. SELECTED COLLECTIONS NIGERIA: Oban, 1912, Talbot2WO (BM, K). CAMEROON: Ebone-Yabassi, 27 December 1967, Bamps 1632 (BR); Mount Cameroon, above Batoke, Thomas 2775 (MO). GABON: Cristal Mts, 21 January 1968, Halle & Villiers 4626 (P); Haute-Ngounye, 7 June 1926, Le Testu 5943 (BM, BR, P). Very few specimens have been collected of Brillantaisia lancifolia. However, from available material and without further opportunity for field study, I maintain this species as clearly distinct from all other species of Brillantaisia. Brillantaisia lancifolia has lanceolate leaves unlike any other species in the genus, the plants tend to be delicate and leafy, with numerous axillary vegetative shoots. The nodes are regularly spaced and the internodes rarely exceed 40 mm. The inflorescence is also very delicate with few flowers. Few of the specimens studied have fruit and additional counts of seeds per locule are needed. Two collections from Cameroon (Leeuwenberg 8994, WAG;Hepper 1 332, S) are intermediate in leaf shape between B. lancifolia and the less woody forms of B. owariensis; these specimens have ovate-elliptic leaves with a cuneate leaf base. How- ever, the flowers on these specimens are far larger than those of typical B. lancifolia and have been provisionally determined as part of the large and variable B. owariensis group. 10. Brillantaisia debilis Burkill in Fl. trap. Afr. 5: 39 (1899); Benoist in Bull. Soc. Bot. France 60: 335 (1913). Type: Cameroon, Efulen, Batanga, Bates 350 (Ki-lectotype, BM!, MO!, Z!-isotypes). Fig. 16. Map 4. Icon: Heine, Fl Gabon, Acanthacees: 13: pi. 16 figs 1-3 (1966). B. soyauxii sensu Heine in Fl. Gabon 13: 84, pi. 17, figs 1-3 (1966) non Lindau (1893); non Burkill (1899). 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- 1 10 x 25-58 mm, dull green, veins pale green to white; glabrous; lateral branches with smaller leaves 18^0 x 11-16 mm; leaf base cuneate; apex acuminate; margin entire to shallowly crenate; lateral veins 4-8 on each side of midrib; petiole 22^0(-80) mm, lamina decurrent towards base of petiole. Inflorescence a few flowered terminal panicle (50-)80-140(-180) mm long; rachis glabrous to glandular pubescent. Bracts ovate-obovate, 4-22 x 3- 1 0 mm, rounded at apex; bractlets 1-5 mm long, narrowly oblong-obovate, glabrous to glandular pubescent, pedicel 1-2 mm long. Sepals subequal, linear, rounded at apex, 5-10 x 0.5-1 mm, pale green, fringed with unequal, purple, glandular trichomes. Corolla violet to deep bright purple with pale green or yellow spots in throat; tube 2-3 x 5-12 mm; upper lip 20-28 x 7-10 mm long, outer surface of upper lip covered in short entangled, glandular trichomes, more so at apex; lower lip 19-25 mm long, glabrous above, sparsely glandular pubes- cent below; apical lobes triangular to 2 x 2 mm. Androecium with filaments whitish to 20 mm long; anthers 3-4 mm long; staminodes 5 mm long, densely glandular with a small vestigial anther at apex. Gynoecium with style 25 mm long; stigma 2 mm long; ovary to 4 mm long, glabrous. Capsule 18-22 x 1-2 mm with 18-28 seeds per locule, brown, glabrous. DISTRIBUTION. Cameroon and Gabon. HABITAT. Along rivers in primary forest in rocky, mossy places. 400-500(-900) m. SELECTED COLLECTIONS CAMEROON: Yaounde, Bitye, Bates 1004 (BM, MO); Lolodorf, 1896, Staudt 240 (G, K, NY, S, WAG); Akoakas, 15 February 1963, Raynal & Raynal 97 1 9 (P); near Molobo, 50 km S. of Batouri, 24 July 1 963, Letouzey 5472 (P); Kwoeinvom, 4 March 1964, de Wilde & de Wilde-Duyfies 2072 (BR. MO, SRGH, WAG, Z): 3 km N. of Lomie, 8 September 1965, Leeuwenberg 6560 (LISC, MO, WAG); Nvem, near Bitye. 20 October 1967, 102 K. SIDWELL Fig. 16 Brillantaisia debilis Burkill. A. Habit x 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 (P); N'Koemvon, 11 October 1974, de Wilde 7635 (WAG); Rocher de Akoakas, 27 March 1981, Meijer 15307 (MO); Cult. WAG, 20 December 1985, van derLaan 986 (WAG). GABON: near lake Moudiboubacoudou. 6 April 1927, Le Testu 6478 (BM); Le Testu 9135 (BM); Monts de Cristal, 3 February 1968, Halle & Villiers4891 (P); Akoga, 8 February 1968, Halle & Villiers 5047 (P); Cristal Mountains, 15 km NE of Asok, 21 August 1978, Breteler & de Wilde 1 87 (MO, WAG); Cristal Mountains, 2 1 January 1 983, de Wilde et al. 133 (WAG); W of Belinga, 18 July 1985, Bos, van der Laan & Nzabi 1 0696 (BR, WAG); 24 km SE of Medouneu, 5 February 1 986, Reitsma, Reitsma & Louis 1880 (NY, WAG); Wolou-Ntem province, Chantier Oveng near Mitzic, 6 May 1986, Louis 2161 (WAG). Plants of Brillantisia debilis have few flowers and are often very leafy, the vegetative parts of plants of this species are entirely glabrous and the leaves are often purple underneath. The inflores- cence may be glandular-pubescent on the rachis, peduncles, calyx and corolla. Many collections of this species have been mistakenly determined asfl. soyauxii. In Flare du Gabon, Heine ( 1 966) used the name B. soyauxii to describe a species which is glabrous except for the pedicels, calyx and, occasionally, the peduncles. The illustration of B. soyauxii in that work (Heine 1966: 87, 1-3) clearly fits the description of B. debilis given here. The type specimen of B. soyauxii has pubescent fruit, and a highly branched, many flowered inflorescence unlike B. soyauxii described in the Flore du Gabon (Heine 1966: 84-85) with few flowers and glabrous fruit. Other characters on the type specimen of B. soyauxii are inconsistent with the description of Heine (1966), including the larger, broader, spathulate upper calyx lobe and lower number of seeds in the fruit. Soyaux 454 is certainly a plant belonging to B. vogeliana. 11. Brillantaisia grottanellii Pichi-Sermoli in Miss. Stud. Lago Tana 7. Recherche Botaniche 1: 254-255 (1951); Cufodontis, Enum. pi. Aeth.: 930 (1964). Type: Ethiopia, Tucur Dinghia forest, Pichi-Sermoli 2099 (FT-holotype, BM!-photo). Map 5. Erect aromatic herb to l-2(-2.5) m tall. Stems 2-4 mm in diameter, rooting at nodes, covered in rather dense short, weak trichomes. Leaves ovate to broadly-ovate (35-)90-225(-250) x ( 10-)40-1 10(- 1 50) mm, olive green to dull dark green above, paler below, glabrous to pubescent above and below; base cuneate; apex acuminate; margin crenate to serrate, occasionally serrulate; 8-13 veins each side of the midrib; petiole (10-)20-70(-1 10) mm, lamina decurrent along !/2-% length of petiole. Inflorescence a terminal spike (60- )80-200(-340) mm long with smaller lateral spikes below, flowers in distinct verticillate whorls, rachis often visible between verticillasters. Bracts ovate to linear-lanceolate, entire or crenate, rounded at apex, densely pubescent with glandular and eglandular trichomes, bractlets linear, trichomes as for bracts. Sepals subequal, linear rounded at apex, 0.5-1 x 12-20 mm, densely covered in long glandular and eglandular trichomes. Corolla dark blue-purple to violet or bright blue-lilac; tube 8-1 3 x (2-)3-5 mm; upper lip 20-30 x 4-6 mm covered in short glandular trichomes, which are slightly longer at the apex of the petals; lower lip 22-30 x 14-20 mm; apical lobes to 7 mm long, acute. Androecium with filaments 35^40 mm long; anthers 6-7 mm long; staminodes 5-10 mm long Gynoecium with style 35 mm long pubescent to stigma, stigma 4 mm long. Capsule 20-22 x 2-3 mm with 8 seeds per locule, brown-black, covered with sparsely glandular trichomes. DISTRIBUTION. Ethiopia and possibly Sudan. HABITAT. Montane forest, beside streams in shade, in disturbed ground, occasionally collected from plantations; 1500-2000 m. SELECTED COLLECTIONS ETHIOPIA: Kaffa Prov., Beletta forest. 13 February 1957, Mooney 6725 (ETH): Wellega region near Dembidollo, Anfilo, 6 March 1957, Mooney 6890 (S): 7 km NW of Tippi airstrip, 14 January 1962, Meyer 7995 (US); 35 km W. of Lekemeti. 1 3 April 1 966. de Wilde & de Wilde-Duyfjes 1 0768 (MO, WAG); Kaffa Prov., Bonga, 9 January 1972. Ash 1454 (ETH, MO, UPS, WAG);. Kaffa Prov., 57 km from Jimma on Sheki-Gogeb river track, 8 104 K. SIDWELL Table 5 Six characters which differ between plants of Brillantaisia grottanellii and B. madagascariensis. Character B. groUanellii B. madagascariensis Bract shape Glandular trichomes on bract lanceolate-ovate present broadly ovate-ovate absent Trichomes on style along more than half length at base of style only of style Leaf margin Upper sepal shape Pollen ornamentation subentire-serrate linear-spathulate smooth entire or crenate linear irregularly spiny December 1972, Friis et al. 1677 (C); Mt. Karkarha, 18 February 1976, Ash 3402 (MO, UPS); Gofa region, 10 km from Sawla, 27 December 1983 Ensennu & Zerihun 649 (C. ETH); 43 km from MizanTeferi on road toTepi, 27 January 1984, Ensennu et al. 745 (ETH); 37-41 km on the track from Gore to Masha, 1 February 1984, Ensermu et al. 821 (ETH); 27 km from Bebeka coffee plantation on road to Guraferda, 7 November 1985 Puff & Ensermu 861107-4/6 (ETH); Wellega Region, Bedelle/Arjo road-Didessa road crossing, 16 January 1990, Friis et al. 6023 (ETH); Kaffa Prov., Beletta forest, 18 January 1990, Friis et al. 6041 (ETH); SUDAN: Torit Distr., near Katire, 9 February 1950, Jackson 1144 (BM, MO), identity uncertain. This species is clearly very closely related to Brillantaisia mada- gascariensis and, early in this study, was thought to be the same species. However, after detailed study of gross morphology and pollen characters, I recognize this species as distinct. Discussion with Dr Ensermu Kelbessa from Addis Ababa University who has studied these plants in the field convinces me that the madagas- cariensis-grottanellii group is best divided. Characters differentiating B. grottanellii from B. madagascariensis are shown in Table 5 and Figure 5. 12. Brillantaisia madagascariensis T. Anderson ex Lindau in Bot. Jahrb. Syst. 17: 103 (1893); Burkill inF/. trop.Afr. 5: 43 (1899); Palacky, Cat. pi. Madag. 3: 57 (1907); Chevalier, Explor. hot. Afrique occ.franq. 1: 493 (1920); Hutchinson & Dalziel in Fl. W. trap. Afr. 2: 254 (193 1); Brenan & Greenway, Checklist brit. empire 5 Tangan. Terr. 2: 5 ( 1 949); Heine in Fl. W. trap. Afr. 2nd ed., 2: 406 (1963); Cufodontis, Enum. pi. Aeth: 930 (1964); Benoist in Fl. Madag. fam. 182 1: 29 (1967); Agnew in Upl. Kenya wild/Is: 583 (1974); Champluvier in Fl. Rwanda 3: 446 (1985). Type: Madagascar, Betsileo, Nandihizana, Hildebrandt 3901 (Bt-holotype; BMI-lectotype; G!, M!, P!, W!, Z!- isolectotypes). Brillantaisia spicata Lindau in Bot. Jahrb. Syst. 20: 4 (1894); in Abh.. Preuss. Akad. Wiss. 50 & 54 (1894); in Wiss. Erg. deut. Zentr.-Afr. Exped., Bot. 2: 293 (1911). Syntypes: Tanzania, Usambara Mountains, Bangarra Lutindi, Hoist 3316 (COI, K!); Usambara Mountains, Gonja, Handei, Hoist 4216 (COI, G!, K!, M!, P!,W!, WAG!, US!, Z!). B. verruculosa Lindau in Bot. Jahrb. Syst. 22: 1 13 (1897); Burkill in FL trap. Afr. 5: 43 (1899); Mildbraed, Wiss. Erg. zweit. deut. Zentr.-Afr. Exped., Bot.: 192 (1922); Robyns Fl. pare. nat. Albert 2: 269(1947); Heine inF/. Gabonll: 89 (1966). Type: Cameroon, Yaounde, Zenker & Staudt 166 (Bf-holotype). B. bagshawei S. Moore in J. Bot. 46: 312-313 (1908). Type: Uganda, Bugoma forest, Bagshawe 1387 (BM!-holotype). B. majestica Wernham in J. Bot. 54: 229 (1969). Type: Cameroon, Buea, Mount Cameroon, Bates 817 (K!-holotype). Icones: Heine, Fl Gabon, Acanthacees: 13: pi. 18 figs 7-15 (1966); Benoist inF/. Madag. fam. 182, Acanthacees: fig. IV 8-13 (1967). Erect or scrambling much branched herb 0.2-2 m tall. Stems 2-5 (- 8) mm in diameter, shortly pubescent, more densely so at nodes, glabrous below. Leaves ovate to broadly ovate, 40-150(-180) x 30- 90(-130) mm, lower leaves becoming much larger, to 400 x 500 mm, usually sparsely pubescent, with short white or translucent trichomes on midrib and main veins, occasionally densely pilose; leaf base attenuate, occasionally truncate or cordate; apex acute- acuminate; margin entire, occasionally slightly crenate; 8-12(-14) main lateral veins each side of midrib; petiole 6-90 mm long, winged for 2/3 of length. Inflorescence a terminal spike with smaller lateral spikes below (20-)40-220(-350) mm long, flowers crowded into dense verticillate whorls, rachis densely pubescent, not usually visible between verticillasters. Bracts ovate to elliptic, apex rounded to acuminate, green with purple tinge, persistent. Sepals subequal, slender tapering to apex or linear and rounded at the apex, occasion- ally spathulate, larger upper sepal 10-30 x 1-2 mm, very slightly longer than the other four; lateral sepals 8-25 x 1-2 mm; green, red to purple at apex, densely pubescent on abaxial surface, trichomes white, less dense shorter trichomes on adaxial surface. Corolla variable shades of blue-purple, violet, magenta or white, with lateral yellow spots in the throat; corolla tube to 15 mm long; upper lip 15- 30 x 5-8 mm, glandular pubescent outside occasionally with long 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 with acute apex. Androecium with filaments 20-22 mm long, white; anthers 3-6(-8) mm long, creamy yellow or brown, pubescent along one side; staminodes 5-10 mm long. Gynoecium with style 25 mm long, often persistent when corolla has fallen, stigma 2.5 mm long; 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 eglandular trichomes. DISTRIBUTION. Throughout West Africa through the Congo basin to Ethiopia and south into Tanzania, also in Madagascar. HABITAT. Montane forest, often weedy in plantations, recently logged areas, clearings or pathsides; 500-1200 m. SELECTED COLLECTIONS LIBERIA: Yekepa, Yiti village, 24 January 1965, Adam 20734 (K, MO). BIOCCO: November 1911, Mildbraed 7126 (H). CAMEROON: 2°00'N 12°15'E, 2 January 191 1, Mildbraed 4\41 (H). GABON: Haute-Ngounye, 1924-1927, Le Testu 6431 (BM, MO); Lastoursville region, 1929-1931, Le Testu 8853 (BM, P). RWANDA: Nyungwe forest, near Rukuzi, 4 May 1972, Bouxin 1577 (BR). BURUNDI: Mabaya-Hua on Rwandan border, 22 June 1969 Lewalle 3783 (BR, G, SRGH); Mabayi valley, Bubanza, 16 October 1971, Reekmans 1 136 (BR). SUDAN: Torit District, Lotti forest, 2 January 1950, Jackson 1023 (BM); Talanga, 1 December 1980, Friis & Vollesen 613 (C). ETHIOPIA: Kaffa Prov., Abetu valley, near Belita, 18 October 1954, Mooney 6071 (ETH, K, S); 10 km E. of Bonga along Jimma road, 22 December 1965, de Wilde 9415 (C, DSM, ETH, MO, UPS, WAG); Kaffa Prov., Bonga, 20 November 1970, Friis, Hounde & Jacobsen 369 (C); IllubaborProv., 22 km S. of Gore, 19 December 1972, Friis et al. 1882 (C); UGANDA: Budongo Forest, 7 February 1935, Taylor 3327 (BM, MO); Mabira Forest, 8 November 1938, Love ridge 26 (MO). DEMO- CRATIC REPUPUBLIC OF THE CONGO: Kasongo-Kindu road, km 60, 29 June 1952, Germain 7826 (PRE); Valley de la Mbizi, near Burora, 10 May 1957, Gutzwiller 841 (BR); Kivu, W. of Kaluzi Mts, 26 April 1978, Lambinon 78/321 (BR). KENYA: Kakamega forest, 17 November 1975, Cunningham s.n. (LISC). TANZANIA: Mt Meru, Surra Forest, 10 Novem- ber 1969, Richards 24663 (C, M); Mshituni, above tea estate N. of Derema, 5 October 1979, Kibuwa 5191 (UPS). MADAGASCAR: August 1902, Perrier 9227 (P); near Doanyanala, 25 January-25 February 1949, Humbert 23090 (P); 6 December 1963, Rakotozafy 287 (P). Plants of this species are very distinctive in flower and fruit, with a narrowly spicate inflorescence, flowers in verticillasters, persistent bracts and linear, pubescent calyx lobes. Brillantaisia verruculosa has been upheld as a species by several recent workers, notably Heine (1966), solely on the basis of the floral colour being white rather than purple. White flowers are found in three other species of Brillantaisia and as this is the only character on which Heine split B. madagascariensis, I maintain this variation under the earliest name. NOMEN DUBIUM Brillantaisia cicatricosa Lindau in Bot. Jahrb. Syst. 20: 4 (1895) nomen dubium; Lindau in Pflanzenw. Ost-Afr. C: 366 (1895); Burkill in Fl. trap. Afr. 5:39 ( 1 899); Mildbraed, Deutch Zent. Afr. Exped. (1922); Robyns, Fl. pare, nat Albert 2: 270 (1947); Troupin, Fl. pi. lign. Rwanda: 84 (1982); Champluvier in Fl. Rwanda 3: 446 (1985); Fischer & Hinkel, Nat. Env. Rwanda: 81, fig. 68 (1992). Type: Zaire, W. slope of Ruwenzori Mountains, Buhtahu Valley, Stuhlmann 2301 (Bt-holotype). Problems with the application of this name are described in detail above (p. 95). Brillantaisia hirsuta T. Anderson in Journ Agric.-Hort. Soc. NS 1: 267 (1868). This name refers to a plant introduced to the Royal Botanic Gardens, Calcutta from Kew and which cannot now be traced. Anderson wrote of this species that 'It is common on the Map 5 The distribution of Brillantaisia grottanellii Pichi-Sermoli. 106 K. SIDWELL banks of streams in Western equatorial Africa. It differs from B. owariensis in its very prostrate habit, much smaller non decurrent leaves and in the smaller size of its flowers which are not glandular'. This description may refer to Brillantaisia vogeliana which has sparsely pubescent flowers, however, as this is the only information regarding the the identity of the plant in Calcutta, application of this name remains uncertain. Species transferred to Hygrophila Brillantaisia borellii Lindau in Bot. Jahrb. Syst. 33: 1 86 ( 1 904). This species was considered to be very different from all other Brillantaisia species due to the presence of four fertile stamens, very narrow leaves and the lack of an upper corolla lip. Brillantaisia didynama Lindau in Bot. Jahrb. Syst. 24: 3 1 3 ( 1 898). Considered by Lindau to be clearly distinct from all other species of Brillantaisia by virtue of having four fertile stamens and very distinctive pollen, Lindau maintained this species within the genus and placed it close to B. lancifolia due to the few-flowered inflorescence and lanceolate leaves. Brillantaisia thwaitesii Cramer in Kew Bull. 46: 335-338 (1991). Cramer transferred Plaesianthera thwaitesii (C.B. Clarke) Livera to Brillantaisia as he considered it close to Brillantaisia pube- scens. Recent studies (Sidwell, in press a) have shown that the species is best placed in Hygrophila until a greatly needed mono- graph of that genus is undertaken. Species transferred to Eremomastax Brillantaisia fulva Lindau in Bot. Jahrb. Syst. 24: 313-314 (1898). ACKNOWLEDGEMENTS. I would like to thank Robert Scotland and Kaj Vollesen for supervision throughout the course of my PhD study. Thanks also to John Baker, Ian Gourlay, Julie Hawkins, David Mabberley, Alison Strugnell and Rosemary Wise in Oxford; Alex Ball and Louisa Jones in the SEM unit at The Natural History Museum; everyone at Limbe Botanic Gardens and the National herbarium in Cameroon; Martin Cheek and Alan Paton at Kew and Axel Poulsen in Copenhagen. This research was funded by BBSRC research grant 93309762, with additional financial assistance from Wolfson College Oxford and the University of Oxford Hardship Fund. Funding to cover materials and tuition for all illustrations was received, with thanks, from the Linnean Society of London's A.G. Side fund for research in systematic biology. REFERENCES Ahmad, K.J. 1976. Epidermal studies in some species of Hygrophila and Dyschoriste (Acanthaceae). Journal of the Indian Botanical Society 55: 41-52. 1978. Epidermal hairs of Acanthaceae. Blumea 24: 101-1 17. Anderson, T. 1 875. Brillantaisia alula and Brillantaisia pubescens, In D. Oliver (Ed.), The Botany of the Speke and Grant Expedition. Transactions of the Linnean Society of London 29: 125-126. Baden, C. 1981. The genus Anisotes (Acanthaceae): a taxonomic revision. Nordic Journal of Botany 1: 623-664. Baillon, H. 1 89 1 . Acanthacees. In H. Baillon, Histoire des plantes: 10: 403-466. Paris. Balkwill, K. & Getliffe-Norris, F. 1985. Taxonomic studies in the Acanthaceae; the genus Hypoestes in southern Africa. South African Journal of Botany 51: 133-144. — Getliffe-Norris, F. & Balkwill,M-J. 1996. Systematic studies in the Acanthaceae: Dicliptera in southern Africa. Kew Bulletin 51: 1-61. Bateman, R.M., DiMichele, WA. & Willard, D.A. 1992. Experimental cladistic analysis of anatomically preserved arborescent lycopsids from the carboniferous of Euramerica: an essay on palaeobotanical phylogenetics. Annals of the Missouri Botanical Garden 79: 500-559. Benoist, R. 1967. Acanthacees. In H. Humbert (Ed.), Flore de Madagascar et des Comores: 182e famille. Paris. Bentham, G. 1849. Brillantaisia. In W. J. Hooker. (Ed.) Niger Flora: 477^478. London. 1 876. Acanthaceae. In G. Bentham, & W. J. Hooker (Eds), Genera plantarum: 2: 1060-1122. London. Blackmore, S. & Tootill, E. (Eds) 1984. The Penguin dictionary of botany. Middlesex. Bremekamp, C.E.B. 1944. Materials for a monograph of the Stobilanthinae (Acanthaceae). Verhandelingen der Nederkmdse Akademie van Wetenschappen Afdeeling Natuurkunde 41: 1-305. 1953. The delimitation of the Acanthaceae. Proceedings Koninkijke Nederlandse Akademie van Wetenschappen 56: 533-546. 1965. Delimitation and subdivision of the Acanthaceae. Bulletin of the Botanical Survey of India 1: 21-30. Bin-kill, L.H. 1899. Brillantaisia. In W.T. Thistleton-Dyer, (Ed.). Flora of Tropical Africa: 5: 37. London. Carlquist, S.& Zona, S. 1988. Wood anatomy of Acanthaceae: a survey. Aliso 12: 201- 227. Champluvier, D. 1985. Brillantaisia. In G. Troupin, (Ed.) Flore du Rwanda: Spermatophytes 3: 444-448. Tervuren. Chappill, J.A. 1989. Quantitative characters in phylogenetic analysis. Cladistics 5: 217-234. Cramer, L.H. 1991. Brillantaisia P. Beauv. (Acanthaceae), a new generic record for Asia. Kew Bulletin 46: 335-338. Cranston, P.S. & Humphries, C.J. 1988. Cladistics and computers: a chironomid conundrum? Cladistics 4: 72-92. Daniel T.F. 1984. Revision of Stenandrium (Acanthaceae) in Mexico and adjacent regions. Annals of the Missouri Botanical Garden 71: 1028-1043. De, A. I960. Cytological, anatomical and palynological studies as an aid in tracing affinity and phylogeny in the family Acanthaceae. IV. Transactions of the Base Research Institute Calcutta 31: 17-29. 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 of angiosperms: 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. 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. Ezcurra, C. 1993. Systematics of Ruellia (Acanthaceae) in southern south America. Annals of the Missouri Botanical Garden 80: 787-845. Farris, J.S. 1988. Hennig86: version 1.5. Published by the author, Jamaica Plains. Furness, C.A. 1994. The pollen morphology of Hygrophila and Brillantaisia (Acanthaceae: Ruellieae). Acta Botanica Gallica 141: 267-278. Gift, N. & Stevens, P.F. 1997. Vagaries in the delimitation of character states in quantitative variation - an experimental study. Systematic Botany 46: 1 12-125. Graham, V.A.W. 1988. Delimitation and infra-generic classification of Justicia (Acanthaceae). Kew Bulletin 43: 551-624. Grubert, M. 1 974. Studies on the distribution of myxospermy among seeds and fruits of angiospermae and its ecological importance. Acta Biologica Venezuela 8: 315- 551. Hauser, D.L. & Presch, W. 1991. The effect of ordered characters on phylogenetic reconstruction. Cladistics 7: 243-265. Hedren, M. 1989. Justicia sect. Harnieria (Acanthaceae) in tropical Africa. Acta Universitatis Upsaliensis Symbolae Botanicae Upsaliensis 29: 1-141. Heine, H. 1966. Acanthacees. In A. Aubreville (Ed.), Flore du Gabon: 13. Paris. Hennig, W. 1950. Grundziige einer Theorie der P hylogenetschen Systematik. Berlin. 1979. Phylogenetic systematics. Translated by D. D. Davis & R. Zangerl. Chicago. HUTU. W.P. 1900. Catalogue of the African plants collected by Dr Friedrich Welwitch in 1853-1861. London. Hobein, M. 1884. Uberden systematischen werth der cystolithen bei denAcanthaceen. Botanische Jahrbucherfiir Systematik. Pflanzengeschichte und Pflanzengeographie 5: 422-440. Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. 1990. Index Herbariorum. New York. Hooker, WJ. 1853. Brillantaisia owariensis P. Beauv. Curtis' s Botanical Magazine. 79: t.4717. Immelman, K.L. 1989. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae): palynology. Bothalia 19: 151-156. 1990. Studies in the southern African species of Justicia and Siphonoglossa (Acanthaceae): seeds. Bothalia 21: 49-59. Inamdar, J.A., Chaudhari, G.S. & Ramana Rao, T.V. 1990. Studies on the cystoliths of Acanthaceae. Feddes Repertorium 101: 4\l—424. REVISION OF BR1LLANTAIS1A Kippist, R. 1845. On the existence of spiral cells in the seeds of Acanthaceae. Transactions of the Linnean Society of London 19: 65-76. Lester, R.N. & Ezcurra, C. 1991. Enzyme etching treatment as an aid in the study of seed surface sculpture in Justicia and Ruellia (Acanthaceae). Botanical Journal of the Linnean Society 105: 285-288. Lindau, G. 1893. Acanthaceae Africanae I. Botanische JahrbUcher fiir Systematik, Pflanzengeschichte und Pflanzengeographie 17: 89-113. 1894. Beitrage zur systematik der Acanthaceen. Botanische JahrbUcher fur Systematik, Pflanzengeschichte und Pflanzengeographie 18: 36-64. 1895a. Acanthaceae. In A. Engler & K. Prantl (Eds), Die Natiirlichen Pflanzenfamilien: 4(3b): 274-354. Leipzig. 1895i. Brillantaisia anomala and B. cicatricosu. In A. Engler (Ed.), Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete. Geographische Verlagshandlung: 366. Berlin. 1 895c\ Acanthaceae Africanae 2. Botanische JahrbUcher fur Svstematik, Pflanzengeschichte und Pflanzengeographie 20: 1-76. 1895d. Brillantaisia ulugurica. In A. Engler (Ed.), Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete. Geographische Verlagshandlung: 9 1 . Berlin. 1896. Brillantaisia kirungae. In G. A. von Gotzen (Ed.), Verzeichniss deraufder Graf. v. Gotzen 'schen Expedition bei der Besteigung des Kirunga gesammelten pflanzen: 9. 1897. Acanthaceae Africanae 3. Botanische JahrbUcher fiir Systematik, Pflanzengeschichte und Pflanzengeographie 22: 1 12-127. 1898. Acanthaceae Africanae 4. Botanische JahrbUcher fur Svstematik, Pflanzengeschichte und Pflanzengeographie 24: 310-325. 1904. Acanthaceae Africanae 6. Botanische JahrbUcher fiir Svstematik, Pflanzengeschichte und Pflanzengeographie 33: 183-193. Lindley, J. 1847. The vegetable kingdom. London. 1853. Notices of new or rare garden plants. Journal of the Horticultural Society of London*. 128-130. Luckow, M. 1995. Species concepts: assumptions, methods and applications. System- atic Botany 20: 589-605. Mabberley, DJ. 1987. The plant book. Cambridge. McDade, L.A. 1 995. Species concepts and problems in practice: insight from botanical monographs. Systematic Botany 20: 606-622. Maddison, W.P. 1993. Missing data versus missing characters in phylogenetic analy- sis. Systematic Biology 42: 576-581. Manktelow, M. 1996. Phaulopsis (Acanthaceae) - a monograph. Acta Universiiatis Upsaliensis Symbolae Botanicae Upsaliensis 31: 1-184. Manning, J.C. & Getliffe-Norris, F. 1985. Taxonomic studies in the Acanthaceae: A reappraisal of the genera Duvemoia andAdhatoda in southern Africa. South African Journal of Botany 51: 475-484. Mayr, E. 1942. Systematics and the origin of species. New York. Mickevich, M.E & Lipscomb, D. 1 99 1 . Parsimony and the choice between different transformations for the same character set. Cladistics 7: 1 1 1-139. Mildbraed, J. 1946. Brillantaisia cicatricosa Lindau var. kivuensis Mildbraed van nov. Bulletin du Jardin Botanique de I'Etat 17: 86. Nees von Esenbeck, C.G. 1832. Acanthaceae. In N. Wallich (Ed.), Plantae Asiaticae Rariores: 3. London. 1847. Acanthaceae. In A. de Candolle (Ed.), Prodromus Systematis Naturalis Regni Vegetabilis: 11: 46-519. Paris. Nelson, G.J. & Platnick, N.I. 1981. Systematics and biogeography: cladistics and vicariance. New York. Nixon, K.C. 1992. CLADOS: version 1.38.1. New York. & Davis, J.I. 1991. Polymorphic taxa, missing values and cladistic analysis. Cladistics 7: 233-241. & Wheeler, Q.D. 1990. An amplification of the phylogenetic species concept. Cladistics 6: 211-223. 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- 209. Platnick, N.I., Griswold, C.E. & Coddington, J.A. 1991. On missing entries in cladistic analysis. Cladistics 7: 337-343. Pleijel, F. 1995. On character coding for phylogeny reconstruction. Cladistics 11: 309- 315. Punt, W., Blackmore, S., Nilsson, S. & Le Thomas, A. 1994. Glossary of pollen and spore terminology. LPP Contributions series No. I. Utrecht. Quicke, D.L.J. 1993. Principles and techniques of contemporary taxonomy. London. Radlkofer, L. 1883. Ueberdie systematischen Werth der Pollenbeschaffenheit bei den Acanthaceen. 5. B. buyer. Akademie der Wissenschaften., math-phys. Cl. 13: 256- 314. Raj, B. 1 96 1 . Pollen morphological studies in the Acanthaceae. Grana Palynologica 3: 1-108. Ramos, T. 1997. Tree Gardener: version 2.2. Sao Paolo. Rickett, H.W. 1 944. The classification of inflorescences. Botaw/a;/ Review 10: 1 87-23 1 . 1 955. Materials for a dictionary of botanical terms-Ill, Inflorescences. Bulletin of the Torrev Botanical Club 82: 419^45. 107 Si-hall nit, E. 1906. Beitrage zur anatomic der Acanthaceen-samen. Beihefte zum Botanist-hen Zentralblatt 19: 453-484. Scotland, R.W. 1991. A systematic analysis of Acanthaceae genera with contorted corollas. In S. Blackmore & S.H. Barnes (Eds), Pollen and Spores, Patterns of Diversification: Systematics Association special volume No. 44: 269-289. Oxford. — I992c/. Pollen morphology and taxonomic characters in Acanthaceae. Svstematic Botany 17: 337-340. 1992b. Systematics. similarity and Acanthaceae pollen morphology. Botanical Journal of the Linnean Society 109: 529-541. — 1 992c\ Chapter 2 - Character coding. In P.L. Forey, C.J. Humphries, I.J. Kitching, R.W. Scotland, D.J. Siebert & D.W. Williams (Eds), Cladistics: A Practical Course in Systematics: Systematics Association Publication No. 10: 14-21. Oxford. — 1993. Pollen morphology of Contortae (Acanthaceae). Botanical Journal of the Linnean Society 111: 471-504. — Endress, P.K., & Lawrence, TJ. 1994. Corolla ontogeny and aestivation in the Acanthaceae. Botanical Journal of the Linnean Society 114: 49-65. — Sweere, J.A., Reeves, P.A. & Olmstead, R.G. 1995. Higher-level systematics of Acanthaceae determined by chloroplast DNA sequences. American Journal of Botany 82: 266-275. Sell, Y. 1 969«. Les complexes inflorescentiels de quelquesAcanthacees etude particulere des phenomenes de condensation, de racemisation d'homogeneisation et dt troncature. Annales des Sciences Naturelles Botanique, Paris Series 12 10: 225-300. 1969/7. La dissemination des Acanthacees: Variations sur le type xerochasique fondamental. Revue Generate Botanique 76: 417^t53. Selvaraj, R. & Subramanian, D. 1983. Epidermal studies in some species of Acanthaceae. Journal of the Indian Botanical Society 62: 253-258. Sidwell, KJ. 1997. A revision of Brillantaisia P. Beauv. (Acanthaceae). Unpublished PhD. thesis. University of Oxford. In press a. The taxonomic position of the anomalous Sri Lankan species Brillantaisia thwaitesii (T. Anderson) Cramer (Acanthaceae) from Sri Lanka. Kew Bulletin. In press b. Monographing Brillantaisia P. Beauv. (Acanthaceae): How to delimit species and higher taxa? Proceedings of the 1 5'h AETFAT Congress. Harare. Singh, V. & Jain, D.K. 1975. Trichomes in Acanthaceae I: General structure. Journal of the Indian Botanical Society 54: 1 16-127. Stevens, P.F. 1984. Homology and phylogeny, morphology and systematics. System- atic Botany 9: 395^09. 1991. Character states, morphological variation and phylogenetic analysis: a review. Systematic Botany 16: 553-583. Thiele, K. 1993. The holy grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9: 275-340. Troll, W. 1964. Die Infloreszenzen: Typologie und Stellung im Aufbau des Vegetationskorpers I. Stuttgart. Vollesen, K. In prep a. Acanthaceae. The Flora of Tropical East Africa. In prep b. Acanthaceae. Flora Zambesiaca. & I? ruin mitt, R.K. 1981. Notes on Brillantaisia pubescens (Acanthaceae) with particular reference to discontinuity in flower size. Kew Bulletin 36: 569-573. Weberling, F. 1 989. Morphology of flowers and inflorescences. Cambridge. White, F. 1962. Geographic variation and speciation in Africa with particular reference to Diospyros. In D. Nichols (Ed.), Taxonomv and geography: Systematics Associa- tion Publication No. 4: 71-103. London. 1984 Unpublished fieldnotes. Department of Plant Sciences, University of Oxford. Wilkinson, M. 1 995. A comparison of two methods of character construction. Cladistics 11: 297-308. Wood, J.R.I. 1995. Notes on Strobilanthes (Acanthaceae) for the flora of Ceylon. Kew Bulletin 50: 1-24. EXSICCATAE Abdullah & Vollesen 95/176, pubescens (K). Achien 524, lamium (BR). Adam 2968, owariensis (K, MO); 4782, lamium (BR, K); 4830, lamium (MO); 5286, lamium (MO); 6910, lamium (MO); 7351, lamium (MO); 16325, lamium (K. P); 16453, lamium (K); 20464, lamium (K); 20549, owariensis (BR, K, MO, P. UPS); 20734, madagascariensis (K. MO); 22019, owariensis (MO, P); 22262, lamium (MO); 22916, owariensis (MO, P); 23291, lamium (MO, PRE); 241 16, lamium (MO); 24302. lamium (MO); 24663, lamium (MO); 24795, lamium (MO, PRE); 25258, owariensis (MO); 26520, lamium (MO); 26558, lamium (MO, PRE); 27084, owariensis (MO); 30140, lamium (MO). Adames 73, lamium (K); 694, lamium (K); 791, owariensis (K, UPS). Adebusuyi FHI 43972, lamium (K). Afzelius s.n., lamium (UPS). 108 K. SIDWELL Ake Assi 3244, lamium (G); 3333, owariensis (G); 5502, owariensis (G); 7186, lamium (G); 9136, vogeliana (G); 9243, lamium (G); 1221 1, owariensis (G, Z); 15745, lamium (G); 17170, owariensis (G). Alluard 309. owariensis (P); 390, owariensis (P). Anderson 1383. lamium (K). Andrada 1811, kirungae (LISC). Amr// 1842, lamium (K). Archibald 2589. kirungae (C). Arena's & Groenemlijk 3, vogeliana (WAG); 95, vogeliana (WAG). Ariwaodo 33. owariensis (MO): 77, owariensis (MO). As/? 1454. grottanellii (ETH, MO, UPS, WAG); 3402, grottanellii (MO, UPS). Auauier 2684, owariensis (BR). Bahault s.n. owariensis (P); s.n. kirungae (P). Bagshawe 1131, owariensis (BM); 1270, kirungae (US); 1342, owariensis (BM); 1381, madagascariensis (BM); 1506, owariensis (BM). BrtWuw 10182, owariensis (K, MO); 10778, owariensis (K, MO); 5993, /«wi/M/n (K, MO); 6175, /a/w/Mm (K); 6339, /am/w/n (K); 7070, /am/Mm (K, MO). ft/// 62. kirungae (MAL, SRGH). fiwwp.v 1632, lancifolia (BR); 2592, /a/n/wm (K). <& Kaunda 3452, riparia (MO). c$ Salubeni 1571, r/paria (MO, SRGH). Barbosa 1812. /-/panY/ (LISC); 2042, pubescens (LISC); 2395, pubescens (LISC). fiarfcr 1 131. /tfw/M/n (K); 1418, /a/nw/H (K). Barton (?) 103. /«;;//»/« (K). #«/« 216. vogeliana (BM, E, K, G, Z); 350, M//850, owariensis (WAG). La Crak 3122, pubescens (MO); 31 14, nparia (MO); 4083, kirungae (MO); 4600, kirungae (MO). Lamb 66, owariensis (K); s.n., lamium (UPS). Lambinon 78/321, madagascariensis (BR). Lane Poo/e 392, lamium (K). Lang Brown 9, kirungae (FHO); 81, owariensis (FHO). La/z7o 97, owariensis (K); FHI 43460, /amzwnz (K); FHI 67539, /aw/Mm (K). Lan/o <£ Daramola FHI 28926, owariensis (FHO, K); FHI 28969, owariensis (K); FHI 28992, owariensis (FHO). La?//o <* Fagbemi FHI 64782, /am/urn (K). Law/or <£ //a// 659, /am/Mm (K). Le TesfM 9, owariensis (BM); 1010, owariensis (FHO); 3541, owariensis (BM, P); 3557, /am/Mm (K); 5943, lancifolia (BM, BR, P); 6431, madagascariensis (BM, MO); 6478, ae/Wis (BM, P); 8853, madagascariensis (BM, P); 9135, debilis (BM); s.n., lamium (BM); 9395, owariensis (BR). Leac/z '/ze \\,lamium (K); 109, owariensis (K); 138, /am/Mm (K, P, S). Quintas 1235(88) /awiwrn (BM, Z). Raimano, Matos & Figueira 254, owariensis (LISC). Rakotozafy 287, madagascariensis (P). Ratabu 17, madagascariensis (MO). Rattray s.n., lamium (K). flayna/ 1 1964, vogeliana (P); 20477, kirungae (K). flayna/ cfc flayna/ 9719, aeW/w (P, YA); 10009, vogeliana (P, YA); 12167, owariensis (P, YA); 12966, owariensis (P, YA). Reekmans 1 136, madagascariensis (BR); 1986, kirungae (MO); 5237, kirungae (C, MO); 5918, /am/Mm (K, MO, PRE); 7028, /a/mum (K, MO); 8288, owariensis (MO, PRE, SRGH); 9060, /a/n/w/n (K, MO); 9408, kirungae (K, MO, UPS); 10009, kirungae (K, MO); 10017, kirungae (K, MO, PRE, UPS); 10073, kirungae (K, MO, UPS); 10604, kirungae (K, MO, PRE, S, UPS). Reitsma & Reitsma 1048, vogeliana (WAG). Reitsma, Reitsma & Louis 1880, deWfo (NY, WAG). Reynders 411, kirungae (PRE). Richards 3108, owariensis (BM, MO, NY); 12983, kirungae (SRGH); 24663, madagascariensis (C, M). Robertson 62, lamium (BM). Roberty 6590, /am/urn (G, Z); 6652, owariensis (G); 6946, /aw/urn (G, Z); 10695, owariensis (G, Z); 12613, /a/w/w/n (G, MO, Z); 13020, vogeliana (G); 15849, lamium (G); 15991, owariensis (G); 16148, owariensis (G); 16202, /am/Mm (G); 16210, owariensis (G, Z); 16412, owariensis (G). Robevazalia 1 1530, madagascariensis (P). Robson & Fanshawe 537, kirungae (LISC, SRGH). Robyns 151, owariensis (BR); 2098, owariensis (BR). Rodgers-Hall 1484, kirungae (DSM). Rosevear 41-29, /am/Mm (K); 68-31, owariensis (FHO); 82-29, /am/Mm (K). Saeger 1431, /am/Mm (C, MO). Sa/y 6312, madagascariensis (P). Salubeni 1485, oligantha (SRGH); 1487, oligantha (MAL, PRE, SRGH); 1489, kirungae; \8Q3, pubescens (MO, SRGH); 1 842, pubescens (MO). Salubeni & Kaunda 4523, pubescens (MAL). Salubeni & Tawakali 2334, riparia (MO). Samai 579, lamium (K). Sanford46\l, lamium (MO); 5812, lancifolia (K). Soft/We 953, /am/wm (G). Savory FHI 36599, owariensis (K). Savory <6 feay FHI 25015, owariensis (K); FHI 25056, owarten.s/.v (K); FH25017, owartenm (K). Saver 248, /am/ww (G, K, Z). Scaer/a 31 12, /am/wm (P); 3284, /aw/w/n (P). Schlieben 1358, kirungae (G, M, LISC, S, Z); 2767, kirungae (G, S, Z); 4068, stenopteris (LISC, MO, PRE); 4069, madagascariensis (G, H, LISC, M, P, S, Z); 4108, kirungae (G, LISC, M, S, Z); 5323, pubescens (G, H, LISC, M, MO, PRE, S, Z). Schmidt et al. 2090, /am/M/« (K). Scnne//6513, lamium (K). Schweich 2423, kirungae (LISC, PRE). Scott E///O? 3990, owariensis (BM); 3990a owariensis (K). Scurlock 17, lamium (K). Serer 97, c/ vogeliana (BR): 346, vogeliana; 716, owariensis (BR). Sharland 362, owariensis (K); 1 155, owariensis (K); 1202, owariensis (K); 1411, /aw/Mm (K); 1736, owariensis (K); 1802, S«arpe B8833, kirungae (G). S/zavv 26, owariensis (K). S/ra 670, lamium (P). Sma// 212, /am/wm (K); 243, /am/wm (K, P). Smeyers 32, kirungae (BR); 54, lamium). 12 K. SIDWELL Smith s.n., owariensis (K). Smythe 240, owariensis (K). Snowden 954, kirungae (G, MO); 1847, owariensis (G). SovaM.v 1 18, owariensis (M. W): 454, vogeliana (E, H, US). Stanton 38, owarienxis (EM, SRGH). Stauble 952, lamium (G). S/aurfr 240, dVW//.v (G, K. NY, S, WAG). Stauffer4f>, kirungae (PRE, UPS, Z). S/o/z 152. kirungae (G, M, MO, NY, S, W, Z). S/one 63, lamium (K); 86, lamium (K). Strid 2916, kirungae (S). Stubbings 200, lamium (K). Swarbrick 293, vogeliana (E, YA): TS2822, /am/Mm (K). .Svnse 31395. kirungae (BM, MO, S, US). Ta/fcof s.n., /awi/H/N (K, Z); 981, /am/ww (BM, K, Z); 993, owariensis (K); 1393, owariensis (MO); 1912, owariensis (MO); 2000, lancifolia (BM. K). Talbot & Talbot s.n., owariensis (K); 1270, owariensis (K). 7rt/on 1 172, lamium (BR). Tav/or s.n., madagascariensis (MO); 1660, lamium (BM, MO, S); 2477, owariensis (BM, MO); 2516, owariensis (BM); 3156, kirungae (BM, MO, S): 3203, owariensis (BM); 3284, owariensis (S); 3327, madagascariensis (BM, MO); 3384, owariensis (BM). TV/era c6 Kaunda 313. pubescens (PRE). Thijsen 276, lamium (WAG). TTioen 4829, pubescens (BR). Thomas, A.S. 1595. owariensis (BM); 1768, owariensis (BM); 2845, owariensis (MO); 2952, owariensis (MO, PRE, YA). rnoma.?, D.W 1722, /am/wwi (K); 2091, /am/H/n (K); 2096, /am/urn (S); 2320, /am/M/n (K); 2502, /a/w/wm (BM, MO, NY); 2609, vogeliana (MO, YA); 2762? /am/urn (W); 2775, lancifolia (MO); 2845, owariensis (MO); 2935. /am/Mm (K); 2952, owariensis (K, MO, PRE); D.W 3790, kirungae (MO); 4381. owariensis (MO, YA); 7149, owariensis (MO); 9005, owariensis (MO). Thomas, D.W. & Made od 5 1 06, owariensis (MO); 5145, owariensis (MO). Thomas, D.W cfc Mamto 4235, /a/n/wm (K, MO, PRE). Thomas, D.W., Mambo & Nemba 4987, owariensis (MO, NY). Thomas, D.W. & Nemba 4047, /am/H/n (MO); 4048, vogeliana (K, MO, YA). TTzomas, N.W. 102, owariensis (K); 2034, owariensis (K); 2285, owariensis (K). Thompson & Rawlins 1460, lamium (MO). 77zw/in <£ Mzoro 2904, kirungae (UPS); 3181, kirungae (MO, UPS). Tindall 29, owariensis (K). 7In/ey 2639, pubescens (LISC, MO, SRGH). Tisserant 1352, /am/ww (P); 2346, /am/Mm (BM); 2743, c/ vogeliana (BM, P); 3167, /am/urn (BM, P); 3245, owariensis (P); 3556, /amw/n (LISC, P). Tom? 486, r<>ar/a (LISC); 4586, kirungae; 35354, kirungae (LISC). Torre . Premiere liste des especes d'algues presentes sur la Pointe de Sarene (Senegal). Notes Africains 111: 81-89. 1971a. Halymenia senegalensis, nov. sp. [Algae], espece caracteristique de 1'infralittoral Sendgalais. Bulletin de I'lnstitut Fondamental d'Afrique Noire A, 33: 1-19. 19716. Sur un genre nouveau de Delesseriacees: Pseudobranchioglossum senegalense, algue de 1'infralittoral sendgalais. Bulletin de I'lnstitut Fondamental de I'Afrique Noire A, 33: 20-31. 1971c. Etude morphologique et cytologique d' Helminthocladia senegalensis (Rhodophycees. Nemalionale nouvelle a carpotetraspores et a cycle haplodiplophasique. Phycologia 10: 361-374. & Mollion, J. 1974. La vegetation infralittorale de la petite cote senegalaise. Bulletin Societe Phycologique de France 19: 193-221. B0rgesen, F. 1916. The marine algae of the Danish West Indies 2. Rhodophyceae [pt. 2]. Dansk Botanisk Arkiv 3(lb): 81-144. 1917. The marine algae of the Danish West Indies 2. Rhodophyceae [pt. 3]. Dansk Botanisk Arkiv 3(lc): 145-240. 19 19. The marine algae of the Danish West Indies 2. Rhodophyceae [pt. 5]. Dansk Botanisk Arkiv 3(lc): 305-368. 1925. Marine algae from the Canary Islands, especially from Teneriffe and Gran Canaria I. Chlorophyceae. Biologiske Meddelelser 5(3): 1-123. 1927. Marine algae from the Canary Islands, especially from Teneriffe and Gran Canaria III. Rhodophyceae. Part 1 Bangiales and Nemalionales. Biologiske Meddelelser 6(6): 1-97. 1929. Marine algae from the Canary Islands, especially from Teneriffe and Gran Canaria. III. Rhodophyceae. Part II Cryptonemiales, Gigartinales and Rhodymeniales. Les Melobe'siees by Mme Paul Lemoine. Biologiske Meddelelser 8(1): l-97+[9[. 1930. Marine algae from the Canary Islands, especially from Teneriffe and Gran Canaria III. Rhodophyceae. Part III Ceramiales. Biologiske Meddelelser 9(1): 1- 159. 1931. Some Indian Rhodophyceae, especially from the shores of the Presidency of Bombay. Bulletin of Miscellaneous Information Royal Botanic Gardens, Kew 1931(1): 1-24. 1945. Some marine algae from Mauritius. III. Rhodophyceae. Part 4 Ceramiales. Biologiske Meddelelser 19(10): 1-68. Bornet, E. 1 892. Les algues de P.-K.-A. Schousboe. Memoirs de la Societe National des Sciences Naturelles et Mathematiques de Cherbourg 28: 1 65-376. Note. Also published in Paris as a separate, with new prefatory pages, dated 1 892, and bearing two sets of pagination; the original as in the journal and a repagination from p. 1 to 216. Because of the prefatory pages, the original p. 165 becomes p. 5. Bory de St- Vincent, J.B.G.M. 1803. Essais sur les Isles Fortunees et ['antique Atlantide, ou precis de I'histoire generate de I'Archipel des Canaries. Paris. Boudouresque, C.-F., Perret-Boudouresque, M. & Knoepffler- Peguy, M. 1984. Inventaire des algues marines benthiques dans les Pyrenees-Orientales Mediterranees, France). Vie et Milieu 34: 41-59. Braga, J.C. & Aguirre, J. 1995. Taxonomy of fossil coralline algal species: Neogene Lithophylloideae (Rhodophyta, Corallinaceae) from southern Spain. Revue Palaobotanique et Palynologique 86: 265-285. Brummitt, R.K. & Powell, C.E. (Eds) 1992. Authors of plant names. Kew. Cabioch, J. 1972. Etude sur les Corallinacees. II. La morphogenese, consequences systematiques et phylogenetiques. Cahiers de Biologie Marine 13: 137-288. Campbell, S.J. & Woelkerling, W.J. 1990. Are Titanoderma and Lithophyllum (Corallinaceae, Rhodophyta) distinct genera? Phycologia 29: 14-125. Carpine, C. 1959. Aper$u sur les peuplements littoraux. In J. Forest, Campagne de la Calypso dans le golfe de Guinee et aux lies Principe, Sao Tome, Annobon (1956). Annales de I'lnstitut Oceanographique, Monaco 37: 1-244. Chamberlain, Y.M. 1983. Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. Bulletin of the British Museum (Natural History), Botany, 11: 291^63. 1985. The typification of Melobesia membranacea (Esper) Lamouroux (Rhodophyta, Corallinaceae). Taxon 34: 673-677. 1990. The genus Leptophytum (Rhodophyta, Corallinaceae) in the British Isles with descriptions of Leptophytum bometii, L. elatum sp. nov. and L. laeve. British Phycological Journal 25: 179-199. 1991. Historical and taxonomic studies in the genus Titanoderma (Rhodophyta, Corallinales) in the British Isles. Bulletin of the British Museum (Natural History), Botany, 21: 1-80. 1992. Observations on two melobesioid crustose coralline red algal species from the British Isles: Exilicrusta parva, a new genus and species, and Lithothamnion sonderi Hauck. British Phycological Journal 27: 185-201. 1994a. Pneophyllum coronatum (Rosanoff) D. Penrose comb, nov., P. keatsii sp. nov., Spongites discoideus D. Penrose et Woelkerling and S. impar (Foslie) Cham- berlain comb. nov. (Rhodophyta, Corallinaceae) from South Africa. Phycologia 33: 141-157. 19946. Mastophoroideae. In L.M. Irvine & Y.M. Chamberlain (Eds), Seaweeds of the British Isles. 1(2B): 113-158. London. 1996. Lithophylloid Corallinaceae (Rhodophyta) of the genera Lithophyllum and Titanoderma from southern Africa. Phycologia 35: 204—221. & Irvine, L.M. 1994a. Lithophylloideae Setchell. In L.M. Irvine & Y.M. Chamberlain (Eds), Seaweeds of the British Isles. 1(2B): 58-1 12. London. 19946. Melobesioideae Bizzozero. In L.M.Irvine & Y.M. Chamberlain (Eds), Seaweeds of the British Isles. 1(2B): 159-234. London. & Walker, R. 1988. A redescription of Lithophyllum crouanii (Rhodophyta, Corallinales) in the British Isles with an assessment of its relationships to L. orbiculatum. British Phycological Journal 23: 177-192. 1 99 1 . A redescription of Lithophyllum orbiculatum (Rhodophyta. Corallinales) in the British Isles and a reassessment of generic delimitation in the Lithophylloideae. British Phycological Journal 26: 149-167. & Keats, D.W. 1994. Three melobesioid crustose coralline red algae from South Africa: Leptophytum acervatwn (Foslie) comb, nov., Lfoveatum sp. nov. andLferox (Foslie) comb. nov. Phycologia 33: 1 1 1-133. 1995. The melobesioid alga Mesophyllum engelhartii (Rhodophyta, Corallinaceae) in South Africa. South African Journal of Botany 61: 134—146. & Norris, R.E. 1994. Pneophyllum amplexifrons (Harvey) comb, nov., a mastophoroid crustose coralline red algal epiphyte from Natal, South Africa. Phycologia 33: 8-18. Chang, C.F. & Xia, B.M. 1963. Polycavernosa, a new genus of the Gracilariaceae. Studies Marine Sinica 3: 1 19-126. Chapman, V.J. 1963. The marine algae of Jamaica Part 2. Phaeophyceae and Rhodophyceae. Bulletin of the Institute of Jamaica, Science Series 12(2): 1-195. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 145 Chevalier, A. 1920. Exploration botanique de I' Afrique occidentale Francaise. Tome I. Ennumeration des plantes recoltees avec une carle botanique, agricole etforestiere. Paris. 1935. Les lies du Cap Vert. Geographic, biogeographie, agriculture flore de 1'archipelago. Revue de Botanique Appliquee et d'Agriculture Tropicale 15: 733- 1090. Cinelli, F. & Codomier, L. 1974. Note floristique et repartition de Rhodophyce'es rares (KallymeniaceesetSebdeniacees)de la Mediterranee occidentale. Gio male Botanic a Italiano 108: 13-18. Codomier, L. 1973. Caracteres generaux et developpement des spores de Sebdenia dichotoma (J. Ag.) Berthold (Rhodophycees, Gigartinales). Phycologia 12: 97-105. Colman, J.S. & Stephenson, A. 1966. Aspects of the ecology of a 'tideless' shore. In H. Barnes, Some contemporary studies in marine sciences: 163-170. London. Cordeiro-Marino, M. 1978) '1977'). Rodoffceas bentonicas marinhas do Estado de Santa Catarina. Rickia 1: |6]+l-243. Cribb, A.B. 1 983. Marine algae of the Southern Great Barrier Reef Part I Rhodophyta. Australian Coral Reef Society, Handbook No. 2. Place of publication not given [presumably Brisbane]. ( 'ul I inane. J.P. & Whelan, P.M. 1982. The ecology and distribution of Stenogramme interrupta (C. Agardh) Montagne ex Harvey on the coast of Ireland. Proceedings of the Royal Irish Academy B, 81: 1 1 1-1 16. Dangeard, P. 1948. Sur la flore des algues marines du Maroc Occidental. Compte Rendu Hebdomedaire Seances de I'Academie des Sciences, Paris 227: 364—365. 1949. Les algues marines de la cote occidentale du Maroc. Botaniste 34: 89-198. 1952. Algues de la presqu'ile du Cap Vert (Dakar) et de ses environs. Botaniste 36: 193-329. 1958. Notice sur les travaux scientifiques (1931-1956) de M. Pierre Dangeard . . . Botaniste 42, Supplement: [2]+l-98+[4]. Dawson, E.Y. 1956. Some marine algae from the southern Marshall Islands. Pacific Science 10: 25-66. Delgado, E., Gonzales, M.N. & Jorge, D. 1986 [' 1984'].Contribucion al estudio de la vegetacion ficologica de la zona de Arinaga (Gran Canaria). Botanica Macaronesica 12-13: 97-110. De May, D., John, D.M. & Lawson, G.W. 1977. A contribution to the littoral ecology of Liberia. Botanica Marina 20: 41^-6. De Toni, G.B. 1897. Sylloge algarum omnium hucusque cognitarum ... 4. Sylloge floridearum . . . Sectio I - Familiae I-XI. Patavii. 1903. Sylloge algarum omnium hucusque cognitarum ... 4. Sylloge floridearum . . . Sectio III - Familiae V- VI. Patavii. 1905. Sylloge algarum omnium hucusque cognitarum ... 4. Sylloge floridearum . . . Sectio IV - Familiae I- II. Patavii.. 1910. Litteratura phycologica florae et miscellanea phycologica. Nuova Notarisia 21: 40-54. 1924. Sylloge algarum omnium hucusque cognitarum ... 6. Sylloge floridearum . . . Sectio V. Additamenta. Patavii. & Forti, A. 1913. Contribution a la flore algologique de la Tripolitaine et de la Cyrenai'que. Annales de I'Institut Oceanographique, Monaco 5(7): 1-56. & Levi, D. 1888. L algarum Zanardini. Venezia. Dickie, G. 1874. Enumeration of algae collected at the Cape- Verde Islands by H.N. Moseley, M.A., Naturalist to H. M.S. 'Challenger'. Journal of the Linnean Society (Botany) 14: 344-349. 1877. Supplemental notes on algae collected by H.N. Moseley, M.A.. of H. M.S. 'Challenger' from various localities. Journal of the Linnean Society (Botany) 15: 486-489. Dickinson, C.I. & Foote, VJ. 1950. Marine algae from the Gold Coast I. Kew Bulletin 5: 267-272. 1951. Marine algae from the Gold Coast: II. Kew Bulletin 6: 133-138. Dinter, K. 1926. Index, der aus Deutsch-Siidwestafrika bis zum Jahre 1917 bekannt gewordenen Pflanzenarten. XlX.Repertorium Specierum Novarum Regni Vegetabilis 22: 375-383. 1927. Index, der aus Deutsch-Siidwestafrika bis zum Jahre 1917 bekannt gewordenen Pflanzenarten. XXll.Repertorium Specierum Novarum Regni Vegetabilis 23: 363-371. Dixon, P.S. & Irvine, L.M. 1970. Miscellaneous notes on algal taxonomy and nomenclature, III. Botaniska Notiser 123: 474—487. 1977. Rhodophyta Introduction, Nemaliales, Gigartinales. In Seaweeds of the British Isles 1(1): xi+ 1-252. London. Diiwel, L. & Wegeberg, S. 1996. The typification and status of Leptophytum (Corallinaceae, Rhodophyta). Phycologia 35: 470-483. Note. Effective publication date: 26 September 1996 (stated on back cover of journal issue). Edelstein, T. 1964. On the sublittoral algae of the Haifa Bay area. Vie et Milieu 15: 1 77- 212. Elejabeitia, Y., Reyes, J. & Afonso-Carrillo, J. 1992. Algas marinas bentonicas de Punta del Hidalgo, Tenerife (Islas Canarias). Vieraea 21: 1-28. Evans, S.M., Gill, M.E., Hardy, EG. & Seku, F.O.K. 1993. Evidence change in rocky shore communities on the coast of Ghana. Journal of Experimental Marine Biology and Ecology 172: 129-141. Falkenberg, P. 1 90 1 . Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeresabschnitle. Fauna e Flora des Golfes von Neapel 26: 1-754. Febles, C.I., Arias,A., Gil-Rodriguez, M.C., Hardisson, A. & Sierra Lopez, A. 1995. Estudio in vitro de la actividad antimicrobiana de algas (Chlorophyta, Phaeophyta y Rhodophyta) recolectados en el literal de la isla de Tenerife. Annuario del Instituto de Estudios Canarios 34: 181-192. Feldmann, G. & Bodard, M. 1965. Une nouvelle espece de Botryocladia des cotes du Senegal. Bulletin de I'Institut Oceanographique, Monaco 65(1342): 1-14. Feldmann-Mazoyer, G. 1941. Recherches sur les Ceramiacees de la Mediterranee Occidentale. Alger. Feldmann, J. 1935. Algues marines des lies du Cap Vert recoltees par M. le Professeur Aug. Chevalier. In A. Chevalier, Les ties du Cap Vert. Geographic, Biogeographie, Agriculture flore de 1'Archipel. Revue de Botanique Appliquee et d'Agriculture Tropicale 15: 1069-1071. Note. This is also published as a separate with the original page numbers retained at the top of each page and a new sequence (pp. 1-358) at the bottom of the page. See also no. 100. 1937. Recherches sur la vegetation marine de la Mediterranee. La Cote des Alberes. Note. Originally published as Revue Algologique 10: 1-339. Printed 28 October 1937, but published with '1938' on title page of part The separate form was published with '1937' on title page and attributed inside as extracted from the Revue Algologique Tome X, Nov. 1937. The BM copy of the journal was received 22 June 1938. No textual differences exist between the two versions. 1938. Sur la repartition du Diplanthera wrightii Aschers. sur la cote occidentale d' Afrique. Bulletin de la Societe d 'Histoire Naturelles d 'Afrique du Nord29: 1 07- 112. 1939. Les Algues marines de la Cote des Alberes. IV - Rhodophycees. Revue Algologique 11: 247-330. Note. Includes Bangiales, Nemalionales, Gelidiales and Cryptonemiales. 1941. Les Algues marines de la Cote des Alberes. IV. Rhodophycees (suite). Revue Algologique 12: 77-100. Note. Covers Ceramiales. Travaux Algologiques 1 replaced volume 13 of the original series of Revue Algologique. 1942. Les Algues marines de la Cote des Alberes. IV. Rhodophycees (fin). Travaux Algologiques 1: 29-1 13. Notes. Covers Ceramiales. Travaux Algologiques 1 replaced volume 13 of the original series of Revue Algologique. 1 946. La flore marine des IlesAtlantides.Mwio(>e£ de la Societe de Biogeographie 8: 395^135. 195 1 . La flore marine de 1' Afrique du Nord. Compte Rendu Sommaire de Seance Societe de Biogeographie 28: 103-108. & Hamel, G. 1 934. Observations sur quelques Gelidiacees. Revue Generate de Botanique 46: 528-549. Foslie, M. 1897. On some lithothamnia. Kongelige Norske Videnskabernes Selskabs Skrifter 1897( 1 ): 1-20. 1898. Some new or critical lithothamnia. Kongelige Norske Videnskabernes Selskabs Skrifter 1898(6): 1-19. 1900a. New or critical calcareous algae. Kongelige Norske Videnskabernes Selskabs Skrifter 1899(5): 1-34. 1900/7. Five new calcareous algae. Kongelige Norske Videnskabernes Selskabs Skrifter. 1900(3): 1-6. 1902. New species or forms of Melobesieae. Kongelige Norske Videnskabernes Selskabs Skrifter 1902(2): 1-1 1. 1906. Den botaniske samling. Kongelige Norske Videnskabernes Selskabs Skrifter 1905(10): 17-24. 1907a. Algologiske notiser III. Kongelige Norske Videnskabernes Selskabs Skrifter 1906(8): 1-34. 1 907 b. Algologiske notiser IV. Kongelige Norske Videnskabernes Selskabs Skrifter 1907(6): 1-30. 1 908a. Algologiske notiser V. Kongelige Norske Videnskabernes Selskabs Skrifter 1908(7): 1-30. 1 908/7. Die Lithothamnien der Deutschen - Sudpolar Expedition 1 90 1 - 1 903 [Heft II]. In E. von Drygalski, Deutsche Sudpolar- Expedition 1901-1903 im Auftrage des Reichsministeriums des Innern, VIII. Band. Botanik: [2J+205- 2 19+[ 1 ]+pls 1 ]. Berlin. 1909. Algologiske notiser VI. Kongelige Norske Videnskabers Selskabs Skrifter 1909(2): 1-63. Fremy, P. 1936. Marine algae from the Canary Islands especially from Teneriffe and Gran Canaria IV. Cyanophyceae. Kongelige Norske Videnskabernes Selskabs Skrifter 12(5): 1^3. Furnari, G., Cormaci, M. & Alongi, G. 1996. Lithophyllum frondosum (Dufour) comb. nov. (Corallinaceae, Rhodophyta): the species to which Mediterranean 'Pseudolithoph\-llum expansum' should be referred. European Journal ofPhycology 31: 117-122. Gabrielson, P.W. l985.AgardhiellaveKUsNeoagardhiella(Solieri&ceae, Rhodophyta): another look at the lectotypification of Gigartina tenera. Taxon 34: 275-280. & Hommersand, M.H. 1982. The Atlantic species of Solieria (Gigartinales, Rhodophyta): their morphology, distribution and affinities. Journal ofPhycology 18: 31-45. 146 W.J. WOELKERLING ET AL. Ganesan, E.K. 1974. Studies on the marine algae of Venezuela V. Pseudogloiophloea Halliae. Journal of Phycology 10: 415-418. & West, J.A. 1 975. Culture studies on the marine red alga Rhodophysema elegans (Cryptonemiales, Peysonneliaceae). Phycologia 14: 161-166. Gayral, P. 1966. Les algues des cotes Francoises (Manche et Atlantique) Notions fondamentales sur I'ecologie, la biologic et la systematiques des digues marines. Paris. Gil-Rodriguez, M.C. 1980 ['1979']. Revision taxonomica-ecologia del genero Cvstoseira C. Ag. en el archipielago Canario. Vieraea 9: 1 15-148. Acebes Ginoves, J.R. & Perez de Paz, P.L. 1978. Nueves aportaciones a la flora ficologica de las islas Salvajes. In Anon., Contribution al estudio de la historia natural de las islas Salvajes: 45-72. Resultados de la Expedicion Cientifica 'Agamenon 76' (23 de febrero-3 de marzo de 1976). Santa Cruz de Tenerife. & Afonso-Carrillo, J. 1980. Adiciones a la flora marina y catalogo ficologico para la isla de Lanzarote. Vieraea 10: 59-70. 1981['1980'|. Catalogo de las algas marinas bentonicas (Cyanophyta, Chlorophyta, Phaeophvta \ Rhodophyta) para el Archipielago Canario. Tenerife. — Haroun Tabraue, R.J., Afonso-Carrillo, J. & Wildpret de la Torre, W. 1985. Adiciones al catalogo de algas marinas bentonicas para el Archipielago Canario. II. Vieraea 15: 101-112. & Wildpret de la Torre, W. 1980a. Contribution al estudio de la vegetation ficologica marina del litoral Canario. Tenerife [Encyclopedia Canaria]. 1980/7. Contribucion a la ficologia de la isla del Hierro. Vieraea 8: 245-260. Gonzalez Henriquez, M.N. 1976. Contribucion al estudio del epifitismo en Zostera marina L. (Zosteraceae) en la playa de Las Cameras (Gran Canaria). Botanica Macamnesica 2: 59-67. Gonzalez, N. 1977a. Estudio de la vegetacion litoral de la zona de Maspalomas. Botanica Macamnesica 4: 23-30. 1977/7. Estudio de la vegetacion bentonica litoral del noroeste de la Isla de Gran Canaria (Banaderos, San Felipe, Sardina, Las Nieves). Botanica Macamnesica 4: 85-104. Goor, A.C.J. van 1 923. Die Hollandischen Meeresalgen (Rhodophyceae, Phaeophyceae und Chlorophyceae) insbesondere der Umgebung von Helder, des Wattenmeeres und der Zuidersee. Verhandelingen der K. Akademie van Wetenschappen, Amsterdam.CTweede Sectie) 23(2): I-IX+[l]+l-232. Gordon, E.M. 1972. Comparative morphology and taxonomy of the Wrangelieae, Sphondylothamnieae, and Spermothamnieae (Ceramiaceae, Rhodophyta). Austral- ian Journal of Botany, Suppl. 4: [2]+l-180. Greuter, W. 1994. International Code of Botanical Nomenclature (Tokyo Code) adopted by the fifteenth International Botanical Congress, Yokohama, August- September 1993. Kbnigstein. [Regnum Vegetabile 131]. Guadeloupe Gonzalez, M.E. Gil-Rodriguez, M.C. & Fernandez Gonzalez, 1995. Flora y vegetacion marina Arrecife de Lanzarote. Fundacion C, sar Manrique, Lanzarote. Guiry, M.D. 1984. Structure, life history and hybridization of Atlantic Gigartina teedii (Rhodophyta) in culture. British Phvcological Bulletin 19: 37-55. Giimbel, C.W. 1871. Die sogenannten Nulliporen (Lithothamnium und Dactylopora) und ihre Betheiligung an der Zusammensetzung der Kalkgesteine. ErsterTheil. Die Nulliporen des Pflansenreichs (Lithothamnium). Abhandlungen der Ksniglich Bayerischen Akademie der Wissenschaften, Mathematisch-Physikalischen Classe, 11(1): 13-51, pis 1-2. Hardy, F.G. & Seku, F.O.K. 1993. Some notes on collecting sites and field records for marine algae in Ghana. The Phycologist 36: 2-7. Hariot, P. 1895. Liste des algues recueillies au Congo par M.H. Lecomte. Journal de Botanique, Paris 9: 242-244. 1896 ['1895']. Contribution a la flore algologique du Gabon et du Congo franfais. Compte Rendu de I' Association Franfaise pour I'Avancement des Sciences 24(2): 641-643. 1908. Les algues de San Thome (Cote occidentale d'Afrique). Journal de Botanique, Paris, Series 2, 1: 161-164. 1911 .Algues de Mauritanie recueillies par M. Chudeau. Bulletin Societe Botanique de France 58 [=Series 4, 11]: 438^45. Haroun, RJ., Prud'homme van Reine, W.F., Miiller, D.G., Serrao, E. & Herrara, R. 1993. Deep-water macroalgae from the Canary Islands: new records and biogeo- graphical relationships. Helgoldnder Meeresuntersuchungen 47: 125-143. Haroun Tabraue, R.J., Gii-Rodriquez. M.C., Afonso-Carrillo, J. & Wildpret de la Torre, W. 1984 ['1983']. Estudio del fitobentos del Roque de los Organos (Gomera). Catalogo floristico. Vieraea 13: 259-276. — & Wildpret de la Torre, W. 1 985 [ ' 1 984' ]. Vegetacion bentonica del Roque de Los Organos (Gomera). Anales de Biologia, Universidad de Murcia 2 (Seccion especial, 2): 107-117. Harper, J.T. & Garbary, DJ. 1997. Marine algae of northern Senegal: The flora and its biogeography. Botanica Marina 40: 129-138. Harvey, W.H. 1846-1851. Phycologia britannica ... 2, 3, Rhodospermeae ... 4. Chlomspermeae . . . [Synopsis nos. 280-388]. London. Hauck, F. 1883. Die Meeresalgen Deutschlands und Osterreichs. Part 5: 225-272. Leipzig. Note. The entire work (XXIV, 575 pp., 5 pis) appeared as a bound volume in 1885. Hemsley, W.B. 1885. II. - Report on the botany of the Bermudas and various other islands of the Atlantic and Southern Oceans. [First part] In C.W. Thompson & J. Murray, Report on the scientific results of the voyage of H. M.S. Challenger during the years 1873-76 under the command of Captain George S. Nares, R.N., F.R.S. and the late Captain Frank Tourle Thompson, R.N . . . , Botany - I: 1-137+[27|. London. Henriques, J.A. 1 885 [ ' 1 884' ] . Contribute, ao para o plantas colhidas por estudo da flora d'algumas possessoes portuguezas I F. Newton na Africa occidental. Boletim da Sociedade Broteriana 3: 129-140. 1886. Algae. In J.A. Henriques. Contribuigoes para o estudo da Flora d' Africa Flora de S. Thome. Boletim da Sociedade Broteriana 4: 217-221. 1887. Flora de S. Thome. In G.B. De Toni & D. Levi, Contributiones ad phycologiam extra-italicam. Notarisia 2: 381-383. Note. A complete extract from Henriques (1886) (263); the present text has been attributed to Henriques solely, as there appear to be no alterations in the algal text. 1917. Catalogo das especies de animals e plantas ate hoje encontradas no Una de S. Tome. Boletim da Sociedade Broteriana 27: 138-197. Heydrich, F. 1 894. Beitrage zur Kenntniss der Algenflora von Ost-Asien besonders der Insel Formosa, Molukken - und Liu-Liu-Inseln. Hedwigia 33: 267-306. Hollenberg, G.J. 1967. New marine algae from the Central Tropical Pacific Ocean. American Journal of Botany 54(10): 1 198-1203. Hooker, J.D. [& Harvey, W.H.] 1847. LV. Algae, L. In J.D. Hooker, The botany of the Antarctic voyage ofH.M. Discovery ships Erebus and Terror, in the years 1839-1843 ...,/. Flora Antarctica, Botany ofFuegia, The Falklands, Kuerguelen 's Land etc., Part II. Algae: 454-502. London. Hoppe, H.A. 1969. Marine algae as raw materials. In T. Levring, H.A. Hoppe & O.J. Schmid, Marine algae a survey of research and utilization. Botanica Marina Hand- books 1: 126-287. Hamburg. Hornemann, J.W. 1819. Anniversaria in memoriam Reipublicae Sacrae et Litterariae cum Universae, turn Danicae nostrae restauratae celebranda indicit Regiae Univer- sitatis Hauniensis Rector cum senatu academico. De Indole plantarum Guineensium [observationes]. Hauniae. Huisman, J.M. 1985. The Scinaia assemblage (Galaxauraceae, Rhodophyta): a re- appraisal. Phycologia 24: 403^118. & Borowitzka, M.A. 1990. A revision of Australian species of Galaxaura (Rhodophyta, Galaxauraceae), with a description ofTricleocarpa gen. nov. Phycologia 29: 150-172. & Townsend, R.A., 1993. An examination of Linnean and pre-Linnean taxa referable to Galaxaura and Tricleocarpa (Galaxauraceae, Rhodophyta. Botanical Journal of the Linnean Society 113: 95-101. Huve, H. 1962. Taxonomie, ecologie et distribution d'une Melobesiee Mediterraneene: Lithophyllum papillosum (Zanardini) comb, nov., nonLithophyllum (Dermatolithon) papillosurn (Zanard) Foslie. Botanica Marina 4: 219-240. Irvine, L.M. 1983. Seaweeds of the British Isles. 1(2 A). London. & Chamberlain, Y. 1994. Seaweeds of the British Isles. 1(2B). London. Johansen H.W. 1981. Coralline algae, a first synthesis. Florida. John, D.M. \912a. A new species of Botryocladia (Rhodophyceae, Rhodymeniales) from the Gulf of Guinea. Phycologia 11: 33-36. 1972/7. The littoral ecology of rocky parts of the north-western shore of the Guinea Coast. Botanica Marina 15: 199-204. 1 977 [ ' 1 976' ] . The marine algae of Ivory Coast and Cape Palmas in Liberia (Gulf of Guinea). Revue Algologique N.S. 11: 303-324. 1986. Littoral and sub-littoral marine vegetation. In G.W. Lawson (Ed.), Plant ecology in West Africa: systems and processes: 215-246. New York. & Lawson, G.W. 1972a ['1971']. Additions to the marine algal flora of Ghana I. Nova Hedwigia 21: 817-841. 1972/7. The establishment of a marine algal flora in Togo and Dahomey (Gulf of Guinea). Botanica Marina 15: 64-73. 1974. Observations on the marine algal ecology of Gabon. Botanica Marina 17: 249-254. 1977<7. The marine algal flora of the Sierra Leone Peninsula. Botanica Marina 20: 127-135. 1977/7. The distribution and phytogeographical status of the marine algal flora of Gambia. Feddes Reprium 88: 287-300. (unpublished). Additions to the marine algal flora of Ghana II. Note. Originally submitted to Nova Hedwigia and reached page proof, 21 Sept. 1972. Corrected and returned from Ghana, but no more heard of paper and never published. Revised by authors in 1973, 1974 and (finally before being abandoned) 1975. Much of the included information used subsequently elsewhere, but a few data not taken up and hence the present inclusion. 1991 . Littoral ecosystems of tropical Western Africa. In A.C. Mathieson & P.H. Nienhuis (Eds), Intertidal and littoral ecosystems. Ecosystems of the World 24: 297-322. Amsterdam. Price, J.H., Prud'homme van Reine, W.F. & Woelkerling, WJ. 1994. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 4. Genera L-O. Bulletin of the Natural History Museum London, Botany, 24: 49-90. Lieberman, D. & Lieberman, M. 1977. A quantitative study of the structure and RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 147 dynamics of benthic subtidal algal vegetation in Ghana (Tropical West Africa). Journal of Ecology 65: 497-52 1 . & Swaine, M.D. 1980. Strategies of data collection and analysis of subtidal vegetation. In J.H. Price, D.E.G. Irvine & W.F. Farnham (Eds), The shore environment. 1. Methods: 265-284. London. & Pople, W. 1973. The fish grazing of rocky shore algae in the Gulf of Guinea. Journal of Experimental Marine Biology and Ecology 11: 81-90. Price, J.H., Maggs, C. & Lawson, G.W. 1979. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. III. Rhodophyta (Bangiophyceae). Bulletin of the British Museum (Natural History), Botany, 7: 69- 82. Johnston, C.S. 1969o. Studies on the ecology and primary production of Canary Islands marine algae. Proceedings of the International Seaweed Symposium 6:213- 222. 1969/7. The ecological distribution and primary productivity of macrophytic marine algae in the eastern Canaries. Internationale Revue de Gesamten Hydmbiologie 54(4): 473-490. Johnstone, W.G. & Croall, A. 1859. The nature-printed British Seaweeds . . . 2. - Rhodospermeae. Fam. X-XIII. London. Jorge, D., Gonzales, M.N. & Delgado, E. 1986['I984']. Macrofitobentos del literal del Puerto de Las Nieves (Gran Canaria). Botanica Macamnesica 12-13: 1 1 1-122. Karsakoff, N. 1896. Sur deux Floridees nouvelles pour la flore des Canaries. Annales des Sciences Naturelles (Botanique) VIII, 4: 281-291. Keats, D.W. & Chamberlain, Y.M. 1994a. Three species ofHydrolithon (Rhodophyta, Corallinaceae): Hydrolithon onkodes (Heydrich) Penrose and Woelkerling, Hydrolithon superficial sp. nov., and H. samoense (Foslie) comb. nov. from South Africa. South African Journal of Botany 60: 8-21. 19946. Two melobesioid coralline algae (Rhodophyta, Corallinales), Mesophyllum erubescens (Foslie) Lemoine and Mesophyllum funafutiense (Foslie) Verheij from Sodwana Bay, South Africa. South African Journal of Botany 60: 1 75- 190. Kensley, B. & Penrith, M.-L. 1980. The constitution of the fauna of rocky intertidal shores of South West Africa. Part III. The north coast from False Cape Frio to the Kunene River. Cimbebasia ser. A, 5: 201-214. Kristiansen, A., Nielsen, R. & Pedersen, P.M. 1993. An annotated list of marine algae collected on Lanzarote, Canary Islands, January 1986. Courier Forschungsinstitut Senckenberg 159: 93-102. KiitzingF.T. 1841. Uberdie"PolypieresCalciferes"desLamouroux./n K.C.F.D.o.t.R. Fischer, Zu der offentlichen Prufung sammtlicher Calssen der Realschule zu Nordhausen welche den 2 April 1841 veranstaltet werden soil: 3-34. Nordhausen. Note. Published in the annual report of the Realschule zu Nordhausen for 1841. 1 843. Phycologia generalis oder Anatomie, Physiologic und Systemkunde des Tange. Lipsiae. 1849. Species algarum. Lipsiae. 1862. Tabulae phycologicae oder Abbildung der Tange 12: [4]+30+pls 1-100. Nordhausen. 1865. Tabulae phycologicae oder Abbildungen der Tange 15: [l]+36+pls 1-100. Nordhausen. Kylin, H. 1932. Die Florideenordnung Gigartinales. Acta Universitatis Lundensis II, 28(8): 1-88. 1956. Die Gattungen der Rhodophyceen. Lund. Lawson, G.W. 1954. Seaweeds from Sierra Leone. Journal of the West African Science Association 1(1): 63-67. 1956. Rocky shore zonation on the Gold Coast. Journal of Ecology 44: 153-170. 1957a. Some features of intertidal ecology of Sierra Leone. Journal of the West African Science Association 3: 166-174. 19576. Seasonal variation of intertidal zonation on the coast of Ghana in relation to tidal factors. Journal of Ecology 45: 831-860. 1960. The genus Taenioma in West Africa. New Phytologist 59: 361-366. 1966. The littoral ecology of West Africa. Oceanography and Marine Biology Annual Revue 4: 405^48. 1980a. Unpublished list (in litt.) of benthic marine algae from the intertidal and shallow subtidal of Fernando Poo (Bioko) collected during a field trip in December 1980. 1 9806. The Nigerian marine flora comes of age. Nigerian Field Society, Ibadan Field Notes No. 3: 1-12. 1985. Algae associated with mangroves in the Niger Delta area. In B.H.R. Wilcox & C.B. Powell (Eds), The mangrove ecosystem of the Niger Delta. Proceedings of a workshop: 56-67. Port Harcourt, Nigeria. & John, D.M. 1977. The marine flora of the Cap Blanc peninsula: its distribution and affinities. Botanical Journal of the Linnean Society 75: 99-1 18. 1982. The marine algae and coastal environment of Tropical West Africa. Beihefte Nova Hedwigia 70. Vaduz. 1987. The marine algae and coastal environment of Tropical West Africa (second edition). Beihefte Nova Hedwigia 93. Berlin. & Price, J.H. 1975. The marine algal flora of Angola: its distribution and affinities. Botanical Journal of the Linnean Society 70: 307-324. 1993. The marine algal flora of St. Helena: its distribution and biogeographical affinities. Courier Forschungsinstitut Senckenberg 159: 103-107. & Norton, T.A. 1971. Some observations on littoral and sublittoral zonation at Teneriffe (Canary Isles). Botanica Marina 14: 1 16-120. & Price, J.H. 1 969. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. I. Chlorophyta and Xanthophyta. Botanical Journal of the Linnean Society 62: 279-346. — Simons, R.H. & Isaac, W.E. 1990. The marine algal flora of Namibia: its distribution and affinities. Bulletin of the British Museum (Natural History), Botany, 20: 153-168. — Woelkerling, WJ., Price, J.H., Prud'homme van Reine, W. & John, D.M. 1995. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 5. Genera P. Bulletin of the Natural History Museum, Botany, 25: 99-122. Lemoine, P. 1911. Structure anatomique des Melobesie'es. Application a la classifica- tion. Annales de I'ln.ititut Oceangraphique, Monaco 2(2): 1-213+[12]. Note. Also issued in thesis form with identical pagination but a different title page. A summary appears in Bulletin de la Societe Botanique de France 58: 394—397 (P. Hariot, 191 1). Author listed as Mme P. Lemoine in the publication. 1912 ['191 1']. Catalogue des Melobesiees de 1'Herbier Thuret. Bulletin de la Societe Botanique de France 58: LI-LXV. 1915. Calcareous algae. Report on the Danish Oceanographical Expedition 1908-19 10 to the Mediterranean and adjacent seas. Biology K. 1,2: 1-30, pi. 1. 1924. Corallinacees du Maroc [I]. Bulletin de la Societe des Sciences Naturelles du Maroc 4: 1 13-134, pis 1-2. 1 926. Corallinacees du Maroc (II). Bulletin de la Societe des Sciences Naturelles du Maroc 6: 106-108. 1928. Une nouveau genre de Me'lobesiees: Mesophyllum. Bulletin de la Societe Botanique de France 75: 25 1-254. 1929c; ['1928']. LesAlgues calcaires (Melobesiees) des Canaries - leurs affinities. In Anon., Compte Rendu de la 52e Session, Association Francaise pour L'Avancement des Sciences ...La Rochelle 1928: 658-662. Paris. Note. The date of publication must be 1929. Footnote (1), p. 659, cites full reference data to Lemoine's own 1929 paper on Melobesiees in B0rgesen's 'Marine Algae from the Canary Islands . . . '. Also issued as an independently paginated reprint. 19296. Les algues calcaires (Melobesiees) des Lemoine M.(Mme P.). [Subfam. 1. Melobesieae]. In F. B0rgesen, Marine algae from the Canary Islands especially from Teneriffe and Gran Canaria III. Rhodophyceae Part II Cryptonemiales, Gigartinales and Rhodymeniales. Del Kongelige Danske Videnskabernes Selskab Biologiske Meddelelser 8(1): 19-68. 1935. Corallinaceae. In A. Chevalier, Les lies du Cap Vert. Geographic. biogeographie, agriculture flore de FArchipel. Revue de Botanique Appliquee et d'Agriculture Tropicale 15: 1071. 1964. Contribution a 1'etude des Melobesiees de 1'Archipel du Cap Vert. Proceed- ings of the International Seaweed Symposium 4: 234-239. 1965. Algues calcaires (Melobesiees) recueillies par le Professeur P. Drach (croisiere de la Calypso en mer Rouge, 1952). Bulletin Institut Oceanographique, Monaco 64(1 331): 1-20. 1966. Algues calcaires recueillies dans la Mer Rouge, en particulier dans le Golfe d'Eilat. Bulletin of the Sea Fisheries Researtch Station, Israel (Haifa) 42: 1-27, 1 pi. 1974. Contribution a 1'etude du genre Lithoporella (Corallinacees). Revue Algologique N.S. 11: 42-57, pis 4-6. Levring, I. 1974. The marine algae of the archipelago of Madeira. Boletim do Museu Municipal do Funchal 28(125): 1-1 1 1. Lieberman, M., John, D.M. & Lieberman, D. 1979. Ecology of subtidal algae on seasonally devastated cobble substrates off Ghana. Ecology 60: 1 151-1 161. 1984. Factors influencing algal species assemblages on reef and cobble substrata off Ghana. Journal of Experimental Marine Biology and Ecology 75: 1 29- 143. Lopez Hernandez, M. & Gil-Rodriguez, M.C. 1 982[ ' 1 98 1 ' ] . Estudio de la vegetaci6n ficologica del literal comprendido entre Cabezo del Socorro y Montana de la Mar, Giii'mar, Tenerife. Vteraea 11: 141-170. & Afonso-Carrillo, J. 1986. Sobre la presencia de Rhodophyllis divaricata (Stackhouse) Papenfuss (Rhodophyllidaceae Engler, Rhodophyta) en el Archipelago Canario. Anales Facultade Ciencias La Laguna Vol. especial 'Homenaje T Bravo'. Maggs, C.A. & Guiry, M.D. 1982a. The taxonomy, morphology and distribution of species of Scinaia Biv.-Bern. (Nemaliales, Rhodophyta) in north-western Europe. Nordic Journal of Botany 2: 517-523. 19826. The life history of Haematocelis fissurata Crouan frat. (Rhodophyta: Sphaerococcaceae). British Phycological Journal 17: 235. & Irvine, L.M., 1983. The life history in culture of an isolate of Rhododiscus pulcherrimus Crouan frat. (Rhodophyta) from Ireland. British Phycological Journal 18: 206. & Hommersand, M.H. 1993. Seaweeds of the British Isles. 1(4A). London. Marcot-Coqueugniot, J. 1991. A preliminary list of marine algae from the Bane d'Arguin (Mauritania). Botanica Marina 34: 195-199. Martin, M.J., Sanson, M. & Reyes, J. 1996. Morphology and anatomy of Papenfussiella kummo (Chordariaceae: Phaeophyta) from the Canary Islands. 148 W.J. WOELKERLING ET AL. Cryptogamie Algologie 17(3): 165-173. Masuda, M. & Ohta, M. 1981. Taxonomy and life history of Rhodophysema odonthuliue sp. nov. (Rhodophyta). Japanese Journal of Phycology 29: 15-21. May, D.I. & Woelkerling, W.J. 1988. Studies on the genus Synarthrophyton (Corallinaceae, Rhodophyta) and its type species, S. patena (J.D. Hooker et W.H. Harvey) Townsend. Ph\cologia 26: 50-71. May, W. 1912 ['1910-11']. Gomera die Waldinsel der Kanaren Reisetagebuch eines Zoologen. Verhandlungen des Naturwissenschaftelichen Vereins in Karlsruhe 24: 51-272. Note. The calcareous algae in this work are acknowledged as being determined by Heydrich; the rest are identifications by Reinbold. Mazza, A. 1903. La Schimmelmannia ornala Schousb. nel Mediterraneo. Nuova Notisaria 14b: 45-61. 1905-1 925. SaggiodialgologiaoceanicaM<0vaAfotari.H'a 16: 85-101, 129-141, 1905; 17: 1-13.41-56.81-101. 129-150. 1906; 18: 1-36,65-98, 126-152, 177- 195, 1907; 19: 1-24,49-66, 109-129, 153-170, 1908:20:6-18,65-86, 113-135, 1909; 21: 1-27,65-99, 125-152, 169-199, 1910; 22: 7-25, 1911:22:53-80, 1912; 23: 1-24, 57-78. 109-122, 1912; 24: 57-85, 1913; 157-174, 1914; 27: 1-53, 104- 155, 169-215, 1916;. 28: 176-239, 1917; Aggiunte al saggio di algologica oceanica (Florideae). Nuova Notarisia 30: 1-62, 1919:31:93-160, 1920:32: 1-48, 1921;33: 97-125, 1922. Note. Series continues, without relevant records, to 1925. Medina, M. & Haroun, R. 1993. Preliminary study on the dynamics of Cystoseira abies-marina populations in Tenerife (Canary Islands). Courier Forschungsinstitut Senckenberg 159: 109-112. Menez, E.G. & Mathieson, A.C. 1981. The marine algae of Tunisia. Smithsonian Contributions to Marine Science 10: i-viii+1-59. Mildbread, J. 1922. Wissenschafiliche Ergebnisse der Zweiten Deutschen Zentral- Afrika- Expedition 1910-1911 unter Fuhrung Adolf Friedrichs, Herzogs zu Mecklenburg 2 Botanik. Leipzig. Mollion, J. 1976 ['1975']. Etude quantitative d'une formation vegetale marine de 1'infralittoral supe'rieur au Senegal. Bulletin de I'lnstitut Fondamental d'Afrique Noire A. 37: 537-554. Molloy, FJ. 1990. Utilized and potentially utilizable seaweeds on the Namibian coast: biogeography and accessibility. Hydrobiologia 204/205: 293-299. Montagne, J.F.C. 1839-1841 ['1835-50']. Plantes cellulaires. In P. Barker-Webb & S. Berthelot. Histoire Naturelle des lies Canaries . . . 3(2), Phytographia Canariensis, Sectio ultima: I-XV+[1]+ 1-208. Paris. Note. For detailed consideration of the bibliography of this work, see W.T. Stearn in Journal of the Society for the Bibiography of Natural History, London I: 49-63 (1937). The correct date of publication is probably 1841; the introduction by Montagne is dated Paris, 1 January 1841. 1842. Troisieme centurie de plantes cellulaires exotiques nouvelles. Annales des Sciences Naturelles (Botanique) 18: 241-282. 1856. Svlloge Generum Specierumque Cryptogamarum quas in Variis Operibus Descriptas Iconibusque lllustratas . . . Paris. 1 860. Florula Gorgonea seu enumeratio plantarum cellularium quas in promontorio Viridi (Cap Vert) insulisque adjacentibus a diversis botanicis et imprimis Cl. Bolle. Annales Science Naturelles (Botanique) 4, 14: 210-225. Moussavian, E. & Kuss, J. 1 990. Typification and status of Lithothamnium aschersoni Schwager, 1883 (Corallinaceae, Rhodophyta) from Paleocene limestones of Egypt. A contribution to the synonymy and priority of the genera Archaeolithothamnium Rothpletz and Sporolithon Heydrich. Berliner Geowissenschaftliches Abhandlungen 120: 929-942. Murray, G. 1888-1889. Catalogue of the marine algae of the West Indian region. Journal of Botany, London 26: 193-196, 237-243, 303-307, 331-338, 358-363, 1888; 27: 237-242, 257-262, 298-305, 1889. Note. Re-paged reprints of the continuous text, pp. 1^6 [1888, pp. 1-28; 1889, pp. 28^6). Naegele, A. 1960. Note sur le peuplement algal de la presqu'ile du Cap- Vert. Notes Africaines 88: 118-119. Newton, L. 1931. A handbook of the British seaweeds. London. Norris, R.E. 1 99 1 .The structure, reproduction and taxonomy ofVidalia andOsmundaria (Rhodophyta, Rhodomelaceae). Botanical Journal of the Linnean Society 106: \-40. & Aken, M.E. 1985. Marine benthic algae new to South Africa. South African Journal ofBotanv 51: 55-65. Notoya, M. 1976a. On the influence of various culture conditions on the early development of spore germination in three species of the crustose corallines (RhodophytaXPreliminary Report). The Bulletin of Japanese Society of Phycology 24: 137-142. 19766. Spore germination in several species of crustose corallines (Corallinaceae, Rhodophyta). Bulletin of the Faculty of Fisheries Hokkaido University 26(4): 314- 320, pis 1-3. Oliveira, F.E.C. de, 1977. Algas marinhas bentonicas do Brasil. Unpublished Ph.D. thesis, Universidade de Sao Paulo, Brazil. Otero-Schmitt, J. 1993. Some local patterns of zonation of benthic marine flora and fauna in Sal, Santiago, S. Vicente and Brava (Cape Verde Islands). Courier Forschungsinstitut Senckenberg 159: 45-52. 1 994. Contribution to the knowledge of the Cape Verdean flora. Nova Hedwigia 59: 525-536. & Sanjuan, A. 1992. Epibiotic seaweeds of the Cape Verde Islands. Botanica Marina 35: 391-398. Pakker, H. & Breeman, A.M. 1996. Temperature responses of tropical to warm- temperate Atlantic seaweeds. II Evidence for ecotypic differentiation in amphi-Atlantic tropical-Mediterranean species. European Journal of Phycology 31: 133-141. Papenfuss, G.F., 1944. Structure and taxonomy ofTaenioma, including a discussion on the phylogeny of the Ceramiales. Madrono 7: 193-214. 1964. The development of the sexual organs and the cystcarp of Taenioma perpusillum. Journal of the Indian Botanical Society. 42A: 159-166. 1968. Notes on South African marine algae. V. Journal of South African Botanv 34: 267-287. Penrose, D. 1991. Spongite s fruticulosus (Corallinaceae, Rhodophyta), the type spe- cies of Spongites, in southern Australia. Phycologia 30: 438^188. 1992. Neogoniolithon fosliei (Corallinaceae, Rhodophyta), the type species of Neogoniolithon, in southern Australia. Phycologia 31: 338-350. 1996«. Genus Hydrolithon. In H.B.S.W. Womersley, The Marine Benthic Flora of Southern Australia D Part 111B. Gracilariales, Rhodymeniales, Corallinales and Bonnemaisoniales: 255-266. Canberra. 1996b. Genus Spongites. In H.B.S.W. Womersley, The Marine Benthic Flora of Southern Australia D Part IIIB. Gracilariales, Rhodymeniales, Corallinales and Bonnemaisoniales: 273-280. Canberra. 1996r. Genus Neogoniolithon. In H.B.S.W. Womersley, The Marine Benthic Flora of Southern Australia D Part IIIB. Gracilariales, Rhodymeniales, Corallinales and Bonnemaisoniales: 280-283. Canberra. & Chamberlain, Y.M. 1993. Hydrolithon farinosum (Lamouroux) comb, nov.: implications for generic concepts in the Mastophoroideae (Corallinaceae, Rhodophyta). Phycologia 32: 295-303. & Woelkerling, W.J. 1988. A taxonomic reassessment of Hydrolithon Foslie, Porolithon Foslie and Pseudolithophyllum Lemoine emend. Adey (Corallinaceae, Rhodophyta) and their relationships to Spongites Kiitzing. Phycologia 27: 159-176. 1991. Pneophyllum fragile in southern Australia: implications for generic concepts in the Mastophoroideae (Corallinaceae, Rhodophyta). Phycologia 30: 495-506. 1992. A reappraisal of Hydrolithon and its relationship to Spongites (Corallinaceae, Rhodophyta). Phycologia 31: 81-88. Perez, L. & Afonso-Carrillo, J. 1993. Estudios en las especies canarias de Galaxaura y Tricleocarpa (Galaxauraceae, Rhodophyta). Vieraea 22: 35-36. Piccone, A. 1884. Crociera del Corsaro alle hole Madera e Canarie del Capitano Enrico d'Albertis Alghe. Genova. 1889. Alghe della crociera del 'Corsaro' alle Azzorre. Nuovo Giornale Botanico Italiana Bolletino della Societa 21: 171-214 1900. Noterelle ficologiche. XI. Pugillo di alghe dell'isola S. Thiago (Capo Verde). Atti della Societa Ligustica di Scienze Naturali e Geograftche. Genova 11: 238-239. 1901. Noterelle ficologiche. XI. Pugillo di alghe dell'isola S. Thiago (Capo Verde). Nuovo Notarisia 12: 45-47. Note. Some data reported from Piccone (1900). Pilger, R. 1908. Kleinere Beitrage zur Kenntnis der Meeresalgen I. Hedwigia 48: 178- 183. 1919. Uber Corallinaceae von Annobon. Botanische Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzengeographie 55: 401-435. 1920 ['1920-21']. Algae Mildbraedianae Annobonenses. In A. Engler, Beitrage zur Flora von Afrika.XLVlll.BotanikaJaabucherfiir Systematik, Pflanzengeschichye und Pflanzengeographie 57: 1-14. Note. See also Pilger (1922). 1922. Algae. Corallinaceae. In J. Mildbraed, Wissenschafiliche Ergebnisse der Zweiten Deutschen Zentral-Afrika-Expedition 1910-1911 ... 11 Botanik: 157-158. Leipzig. /Vote. 'Algae' is a repeat of Pilger (1920) and 'Algae. Corallinaceae' of Pilger (1919). Pinedo, S., Sanson, M. & Afonso-Carrillo, L. 1992. Algas marinas bentonicas de Puerto de la Cruz (antes Puerto Orotava), Tenerife (Islas Canarias). Vieraea 21: 29- 60. Price, J.H. & John, D.M. 1978. Subtidal ecology in Antigua and Ascension: a comparison. Progress in Underwater Science [Report Underwater Association] N.S., 3: 111-133. 1980. Ascension Island, South Atlantic: a survey of inshore benthic macroorganisms, communities and interactions. Aquatic Botany 9: 251-278. & Lawson, G.W. 1 978. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment II. Phaeophyta. Bulletin of the British Museum (Natural History), Botany, 6: 87-182. 1986. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment IV. Rhodophyta (Florideae) 1 . Genera A-F. Bulletin of the British Museum (Natural History), Botany, 15: 1-122. 1988. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 2. Genera G. Bulletin of RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 149 the British Museum (Natural History), Botany. 18: 195-273. 1 992. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 3. Genera H-K. Bulletin of the British Museum (Natural History), Botany, 22: 123-146. Printz, H. (Ed.) 1 929. M. Foslie - 'Contributions to a Monograph of the Lithothamnia' . Del Kongelige Norske Videnskabers Selskab Museet. Trondheim. Prud'homme van Reine, W. 1984. Marine algae of the Cape Verde Islands, historical and preliminary notes. Courier Forschungsinstitul Senckenberg 68: 135-137. 1987. Extractions of report on red algal distribution patterns, in lilt to J.H. Price, 10 April 1987. 1 99 1 . Heincke Expedition, reallocation following examination of original Piccone material from the Canary Islands, in litt. to D.M. John, 19 November 1991. Haroun, R.J. & Audiffred, P.A.J. 1994. A reinvestigation of Macaronesian seaweeds as studied by A. Piccone with remarks on those studied by A. Grunow. Nova Hedwigia 58: 67-121. Hoek, C. van den 1988. Biogeography of Cape Verdean seaweeds. Courier Forschungsinstitut Senckenberg 105: 35—49. & Lobin, W. 1986. Katalog der von den Kapverdischen Inseln beschriebenen Taxa von Algen (Algae: Chlorophyceae, Phaeophyceae & Rhodophyceae). Courier Forschungsinstitut Senckenberg 81: 85-88. & Pakker, H. 1 99 1 . Collections from Cape Verde Islands, in litt. to D.M. John, 1 9 November 1991. Pueschel, C.M. 1 989. An expanded survey of the ultrastructure of red algal pit plugs. Journal of Phycology 25: 625-636. Reinbold, T. 1 907. Die Meeresalgen der deutschen Tiefsee - Expedition 1 898- 1 899. In C. Chun, Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer 'Valdivia' J 898-1 899. Bd. II, Teil II, Lieferung IV: 549-586+[8], pis VV- LVII. Jena. Reyes, J. & Afonso-Carrillo, J. 1993. Morphology and anatomy of Mesophyilum canariense (Corallinaceae, Rhodophyta) from the Canary Islands. Courier Forschungsinstitut Senckenberg 159: 27-132. 1995. Morphology and distribution of nongeniculate coralline algae (Corallinaceae, Rhodophyta) on the leaves of the seagrass Cymodocea nodosa (Cymodoceaceae). Phycologia 34: 179-190. & Wildpret de la Torre, W. 1988. Structure of male conceptacles of Lithophyilum lobatum (Corallinaceae, Rhodophyta). Actesdel Simposi International de Botdnico Pius i Quer, 1988 l(criptogamia): 137-141. & Sanson, M. 1991. Adiciones a la flora marina de la isla de El Hierro (Islas Canarias). Vieraea 20: 71-81. 1996. Las algas epifitas en Cymodocea nodosa en El Medano, isla de Tenerife (Magnoliophyta, Cymodocea [.sic!]). Vieraea 25: 45-56. & Afonso-Carrillo, J. 1993. Notes on some interesting marine algae new from the Canary Islands. Cryptogamic Botanv 4: 50-59. Ribera Siguan, M.A., Gomez Garreta, A. & Seoane-Camba, JA. 1985 ['1984']. Estudio biogeografico de la flora algologica bentonica marina de las Islas Baleares. Anales de Biologia Universidad de Murcia 2 (Seccion especial, 2): 147-159. Rojas-Gonzales, B., Afonso-Carrillo, J. & Ibeas, C. 1994. New records of Rhodo- melaceae (Rhodophyta) from the Canary Islands. Botanica Marina 37: 133-138. Romanes, M.F. 1916. Note on an algal limestone from Angola. Transactions of the Royal Society of Edinburgh 51: 581-584, 1 pi. Rosenvinge, L.K. 1917. The marine algae of Denmark. Part II. Rhodophyceae II. (Cryptonemiales). Def Kongelige Danske Videnskabernes Selskabs Skrifter, 1 Raekke, Naturvidenskabelig og Matematisk Afdeling 7: 155-283, pis 3-4. Round, F.E. 1981. The ecology of algae. Cambridge. Sanson, M. & Reyes, J. 1995. Morphological and geographical observations on four species of Ceramiaceae (Rhodophyta) new to the Canary Islands. Botanica Marina 34(6): 89-95. 1996. Sobre la morfologia de Spyridia filamentosa y Spyridia hypnoides en las islas Canarias (Rhodophyta, Ceramiaceae). Vieraea 25: 37^4. & Afonso-Carrillo, J. 1 99 1 . Contribution to the seaweed flora of the Canary Islands: new records of Floridiophyceae. Botanica Marina 34(6): 527-536. Santos Guerra, A. 1972. Contribucion al estudio de la flora marina de la isla de La Gomera. Vieraea 2: 86-102. Acuna Gonzales, A. & Wildpret de la Torre, W. 1970. Contribucion al estudio de la flora marina de la Isla de La Palma. Cuadernos Botanica El Museo Canario 9: 20-29. Sanusi, S.S. 1 980. A study on grazing as a factor influencing the distribution of bent hie littoral algae. M.Sc. Thesis, University of Ghana, Legon. Sauvageau, C. 1912. A propos des Cystoseira de Banyuls et Guethary. Bulletin de la Station Biologique d'Arcachon 14: 133-556. Schmidt, O.C. 1924. Index algarum marinarum 1920-1923. Hedwigia 65: 1 1-27. 1929. Beitrage zur Kenntnis der Meeresalgen der Azoren II. Hedwigia 69: 165- 172. 1931. Die marine Vegetation der Azoren in ihren Grundzugen dargestellt. Bibliotheca Botanica 102: 1-116. &Gerloff,J. 1957. Die marine VegetationAfrikas in ihren Grundzugen dargestellt. Willdenowia 1: 709-756. Schmitz, F. & Falkenberg, P. 1897. Rhodomelaceae. In A. Engler & K. Prantl, Die Naturlichen Planzenfamilien .... I. Teil. 2. Abteilung: 149-150. Leipzig. & Hauptfleisch, P. 1 896- 1 897. Sphaerococcaceae In A. Engler & K. Prantl, Die Naturlichen pflanzenfamilien . . . , I. Teil. 2. Abteilung: 382-396. Leipzig. Schneider, C.W. 1983. The red algal genus Audouinella Bory Nemaliales; (Acrochaetiaceae) from North Carolina. Smithsonian Contributions on Marine Science 22: iii+[ 1 ]+25. Scott, F.J., Wetherbee, R. & Kraft, G.T. 1984. The morphology and development of some prominently stalked southern Australian Halymeniaceae (Cryptonemiales, Rhodophyta) II. The sponge-associated genera Thamnoclonium Kiitzing and Codiophyllum Gray. Journal of Phycology 20: 286-295. Seagrief, S.C. 1984. A catalogue of South African green, brown and red marine algae. Memoirs of the Botanical Survey South Africa 47: i-vi+1-71. Seoane-Camba, J. 1960. Nota sobre algunas especies de algas de la costa occidental africana (sur de Cabo Blanco). Investigacion Pesquera 16: 91-103. 1965. Estudios sobre las algas bentonicas en la costa sur de la Peninsula Iberica (literal de Cadiz). Investigacion Pesquera 29: 3-216. Silva, P.C., Basson, P.W. & Moe, R.L. 1996. Catalogue of the benthic marine algae of the Indian Ocean. University of California Publications in Botany 79: 1-1259. Menez, E.G. & Moe, R.L. 1987. Catalog of the benthic algae of the Philippines. Smithsonian Contributions to Marine Science 27: 1-179. Simons, R.H. 1974. Algae (including diatoms and seaweeds). In J.H. Day, N.A.H. Millard, & M.-L. Penrith, A guide to marine life on South African shores. 2nd ed.: 239-261. Cape Town. 1983. The genus Trematocarpus (Sarcodiaceae, Rhodophyta) in southern Africa and the exclusion of Sphaerococcus (Chondrus scutellatus. Bothalia 14: 803-808. Soler-Om's, E., Haroun, R.J. & Prud'homme van Reine, W.F. 1998. Mscr. Sebdenia in Macaronesia. Viera Rodriguez, A. & Prud'homme van Reine, W.F. 1995. Sebdenia canariensis sp. nov. (Sebdeniaceae, Gigartinales, Rhodophyta), una nueva alga roja de profundidad delArchipielago Canario. Resumenes de comunicaciones. XlSimposio Nacional de Botanica Criptogdmica, Santiago de Compostela, 18-21 de Septiembre de 1995: 15-16. Sender, O.W. 1852. Algae. In J.A. Schmidt, Beitrage zur Flora der Cap Verdischen Inseln. Mil Beriicksichtigung aller bisjetzt daselbst bekannten wildwachsenden und kultivirten. Pflanzen Nach eigenen Untersuchungen und mil Benutzung der gewonnenen Resultate anderer Reisenden: 125-127. Heidelberg. Sourie, R. 1954. Contribution a 1'etude e'cologique des cotes rocheuses du Senegal. Memoires de I'lnstitut Francois d'Afrique Noire 38: l-342+[l]. Note. From the note on p. 117, it is clear that the algae were worked on mainly by J. Feldmann, but that Sourie took account of some of the views of Dangeard as expressed in the latter's memoir on the Cap Vert (Dakar) peninsula algae. Since the exact contribution of the various people involved is in doubt, we have left the reference in the name of Sourie, who seems to have exercised overall authorship. South, G.R. & Tittley, I. 1986. A checklist and distributional index of the benthic marine algae of the North Atlantic Ocean. St. Andrews, Canada. & Whittick, A. 1976. Aspects of the life history of Rhodophysema elegans (Rhodophyta, Peyssonneliaceae). British Phvcological Journal 11: 349-354. Steentoft, M. 1967. A revision of the marine algae of Sao Tome' and Principe (Gulf of Guinea). Journal of the Linnean Society (Botany) 60: 99-146. Stegenga, H., Bolton, J.J. & Anderson, R.J., 1997. Seaweeds of the South African West Coast. Cape Town. Stephenson, T.A. & Stephenson, A. 1 972. Life between tidemarks on rocky shores. San Francisco. Svedelius, N. 1 9 1 1 . Florideae. In A. Engler & K. Prantl, Die natiirlichen Pflazenfamilien . . . Nachtrdge zum l.Teil, 2. Abteilung iiber die Jahre 1890 big 1910: 200-276. Leipzig. Taylor, W.R. 1939. Algae collected on the Presidential cruise of 1938. Smithsonian Miscellaneous Collections 98(9): 1-18. 2 pis. Tomita, N.Y. 1976. Contribuicao ao conhecimento do genero Sporolithon (Corallinaceae, Cryptonemiales) no Brasil. Unpublished Ph.D. thesis, Universidade de Sao Paulo. Brazil. Townsend, R.A. 1995. The Sporolithaceae: It's place in the Florideophyceae. Unpub- lished Ph.D. thesis, University of Sydney, Australia. Trochain, J. 1940. Contribution a 1'etude de la vege'tation du Senegal. Memoires de I'lnstitut Franqais d'Afrique Noire 2: [l-6]+l— 433+[63]. Note. J. Feldmann clearly had a great deal to do with the main determinations on which the algal list (pp. 108-1 10) was based: since the extent to which the data were accepted or amended by Trochain is not clear, and since there are other parts to the text which seem definitely to have been attributable to Trochain, we have accepted the latter as overall author. For individual comments on species, the more correct authorship citation would undoubtedly be 'Feldmann. J., in Trochain. J.,' etc. Tseng, C.K. 1944. Notes on the algal genus Taenioma. Madrono 7: 215-226. Turner, J.A. & Woelkerling, WJ. 1982a. Studies on the Mastophora-Lithoporella complex (Corallinaceae. development. Phycologia 21: 201-217. 19826. Studies on the Mastophora-Lithoporella complex (Corallinaceae, Rhodophyta). II. Reproduction and generic concepts. Phycologia 21: 218-235. Varo, J., Ramirez, J. & Renteria, J. 1979. Estudio de la vegetacion bentonica del literal granadino. Acta Botanica Malacitana 5: 79-98. 150 W.J. WOELKERLING ET AL. Verheij, E. 1993. The genus Spomlithon (Sporolithaceae fam. nov., Corallinales, Rhodophyta) from the Spermonde Archipelago, Indonesia. Phycologia 32: 1 84-196. 1994. Nongeniculate Corallinaceae (Corallinales. Rhodophyta) from the Spermonde Archipelago, SW Sulawesi, Indonesia. Blumea 38: 95-137. Vickers, A. 1 897(?)( ' 1 896' ].Contribution a la flore algologiques des Canaries. Annales Science Naturelles (Botanique), 8, 4: 293-306. Note. The date is somewhat difficult to cite as there is confusion regarding the dates of various issues. It does seem possible that pre-prints were issued in 1 896 and this is the date usually cited (see Lawson & Price. 1969: 345-346). Viera-Rodriguez, M.A., Gil-Rodriquez, M.C., Audiffred, P.A.J., Prud'homme van Reine, W.F., Haroun-Tabraue, R. & Wildpret de la Torre, W. 1987. Contribution al estudio de la florula bentonica del islote de Montana Clara. Canarias. Vieraea 17: 271-279. Vinassa, P.E. 1892. Coralline mediterranee raccolte dal Prof Meneghini. Ani della Societd Toscana di Scienze Natural! Residente in Pisa. Pmcessi Verbal! 8: 58-60. Weber-van Bosse, A. 1921. Liste des algues du Siboga II Rhodophyceae Premiere partie Protoflorideae, Nemalionales, Cryptonemiales. In M. Weber, Siboga-Expeditie . . . Monographie LIXb. Uikomsten op Zoologisch, Botanisch. Oceanographisch en Geologisch Gebied . . . Livr. LXXXIX: [6]+187+392+[4]. Weisscher, F.C.M. 1982. Marine algae from Ilheu de Fora (Salvage Islands). Boletim do Museo Municipal do Funchal 34: 23-34. 1983. Marine algae from Selvagem Peguena (Salvage Islands). Boletim do Museo Municipal do Funchal 35: 41-80. Wilks, K.M. & Woelkerling, W.J. 1991. Southern Australian species of Melobesia (Corallinaceae, Rhodophyta). Phycologia 30: 507-533. 1995. An account of southern Australian species of Lithothamnion (Corallinaceae, Rhodophyta). Australian Systematic Botany 8: 549-583. Woelkerling, W.J. 1 983a. A taxonomic reassessment ofLithothamnium (Corallinaceae, Rhodophyta) based on studies of R.A. Philippi's original collections. British Phycological Journal 18: 165-197. 1983ft. A taxonomic reassement of Lithophyllum Philippi (Corallinaceae, Rhodo- phyta) based on studies of R.A. Philippi's original collections. British Phycological Journal 18: 299-328. 1 985. A taxonomic reassessment ofSpongites (Corallinaceae, Rhodophyta) based on studies of Kiitzing's original collections. British Phycological Journal 20: 1 23- 1 53. 1987. The genus Choreonema in southern Australia and its subfamilial classifica- tion within the Corallinaceae (Rhodophyta). Phycologia 26: 1 1 1-127. 1988. The coralline red algae: an analysis of the genera and subfamilies of Nongeniculate Corallinaceae. London. 1993. Type collections of Corallinales (Rhodophyta) in the Foslie Herbarium (TRH). Gunneria 67: 1-289. 1996a. Subfamily Lithophylloideae. In H.B.S. Womersley, The marine benthic flora of Southern Australia D Part IIIB. Gracilariales, Rhodymeniales, Corallinales and Bonnemaisoniales: 214-237. Canberra. 1996ft. Subfamily Mastophoroideae (excluding Hydrolithon, Pneophyllum, Spongites & Neogoniolithon). In H.B.S. Womersley, The marine benthic flora of Southern Australia D Part IIIB. Gracilariales, Rhodymeniales, Corallinales and Bonnemaisoniales: 237-283. Canberra. \99Sa. 3. Lamarck's Nullipores. In W.J. Woelkerling & D. Lamy, Non-geniculate coralline red algae and the Paris Museum. Cryptogamie-ADAC and Museum National d'Histoire Naturelle, Paris. In press. 1 998ft. 4. Type collections of non-geniculate Corallinales housed at the Laboratoire de Cryptogamie (PC). In W.J. Woelkerling & D. Lamy, Non-geniculate coralline red algae and the Paris Museum. Cryptogamie-ADAC and Museum National d'Histoire Naturelle, Paris. In press. & Campbell, S.J. 1 992. An account of southern Australian species ofLithophyllum (Corallinaceae, Rhodophyta). Bulletin of the British Museum (Natural History), Botany, 22: 1-107. Chamberlain, Y.M. & Silva, P.C. 1985. A taxonomic and nomenclatural reassessment of Tenarea, Titanoderma and Dermatolithon (Corallinaceae, Rhodo- phyta) based on studies of type and other critical specimens. Phycologia 24: 317-337. & Harvey, A. 1993. An account of southern Australian species of Mesophyllum (Corallinaceae, Rhodophyta). Australian Systematic Botany 6: 571-637. & Irvine, L.M. 1982. The genus Schmitziella Bornet & Batters (Rhodophyta): Corallinaceae or Acrochaetiaceae? British Phycological Journal 17: 275-295. 19860. The neotypification and status of Phymatolithon (Corallinaceae, Rhodophyta). British Phycological Journal 21: 55-80. 1986ft. The neotypification and status of Mesophyllum (Corallinaceae, Rhodophyta). Phycologia 25: 379-396. Penrose, D. & Chamberlain, Y.M. 1993. A reassessment of type collections of non-geniculate Corallinaceae (Corallinales, Rhodophyta) described by C. Montagne and L. Dufour, and of Melobesia brassica-florida Harvey. Phycologia 32: 323-331. & Verheij, E. 1 995. Type collections of nongeniculate Corallinales (Rhodophyta) in the Rijksherbarium, Rijksuniversiteit te Leiden (L), The Netherlands. Blumea 40: 33-90. Wollaston, E.M. 1984. Species of Ceramiaceae (Rhodophyta) recorded from the International Indian Ocean Expedition, 1962. Phycologia 23: 281-299. Womersley, H.B.S. 1994. The marine benthic flora of Southern Australia Part HA. Canberra. 1996. The marine benthic flora of Southern Australia Part IIIB Flora of Australia Supplementary Series 5. Canberra. Wulfen,F.X. 1803.CryptogamaAquatica.Arc/2/v/M>d/