0968-044C M^ • f Bulletin of The Natural History Museum Botany Series THE NATURAL HISTORY MUSEUM VOLUME 29 NUMBER 1 24 JUNE 1999 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. 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(Bot.) © The Natural History Museum, 1 999 Botany Series ISSN 0968-0446 Vol. 29, No. 1, pp. 1-79 The Natural History Museum Cromwell Road London SW7 5BD Issued 24 June 1999 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Land. (Bot.) 29(1): 1-46 Issued 24 June 1999 The moss family Calymperaceae (Musci Philippines \ X U6°lUr.N) THE IMAlunAL l^SJQ-KCM^ SEU M 17 JUN 1999 PRESENTED T 17XT HT 1?¥ T TO Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD BEN C. TAN Department of Biological Sciences, National University of Singapore, Singapore 119260 CONTENTS Introduction 1 Systematic account 2 Key to genera and species 2 Descriptions and illustrations of taxa 5 Excluded taxa 44 References 45 Systematic index 46 SYNOPSIS. A taxonomic account is presented of the nonleucobryoid genera in the moss family Calymperaceae occurring in the Philippines. Four genera are recognized: Calymperes (18 species), Chameleion (1 species), Mitthyridium (8 species) and Syrrhopodon (19 species). Keys to taxa are provided. Species newly recorded for the Philippines include Calymperes mangalorense Dixon & P. de la Varde, C. motleyi Mitt, ex Dozy & Molk., and C. subintegrum Broth. Mitthyridium fasciculatum var. cardotii (M. Fleisch.) A. Eddy is raised to the rank of subspecies (M. fasciculatum subsp. cardotii (M. Fleisch.) B.C. Tan & L.T. Ellis). INTRODUCTION The family Calymperaceae is a group of tropical mosses mainly confined to lowland and coastal forest communities. Its diversity and distribution worldwide have recently been reviewed and mapped by Reese (19876). Regional monographs of this family are available for southern India (Ellis, 1989), Malaysia (Reese & Mohamed, 1985; Reese, Mohamed & Mohamed, 1986; Mohamed & Reese, 1985), Borneo (Menzel & Schultze-Motel, 1990), Papua New Guinea (Reese, Koponen & Morris, 1986) and Australia (Reese & Stone, 1987, 1995). It is in the light of these new publications that a revision of the family was conducted for the Philippine archipelago based on the large collections made by the second author in recent years. There have been attempts to redefine the family Calymperaceae by the inclusion of several genera, such as Leucophanes and Exodictyon, in which the species possess leucobryoid leaves (see review in Reese, Koponen & Norris, 1986). Although the arguments presented are not agreeable to many, these new treatments of the family have much merit. However, in the present work we have chosen to include only the four genera with species that possess nonleucobryoid leaves, namely Calymperes, Syrrhopodon, Mitthyridium and Chameleion. The last one is a segregate genus of Syrrhopodon (Eddy, 1990). An early account of the Calymperaceae occurring in the Philip- pines can be found in Bartram (1939), who reported a total of three genera and 33 species. Iwatsuki & Tan (1979), in their checklist of Philippine mosses, report four genera and 35 species. This study recognizes four genera with 45 species. Three species are new records for the Philippine flora, namely, Calymperes mangalorense, C. motleyi and C. subintegrum. Except for C. mangalorense, which was known only from a few localities in India and Myanmar (Burma) before its discovery in the Philippines, these are wide- spread species in tropical Asia. Their presence in the Philippines is therefore not surprising. On the other hand, there are four taxa whose alleged presence in the country cannot be confirmed by the present study (see excluded taxa). The increase in the number of species from 33 to 45, and the data from the new collections made by the second author, represent a great improvement in our knowledge of the high diversity of the family Calymperaceae in the Philippines. Phytogeographically, the species of Philippine Calymperaceae can be grouped into the following six categories: 1. Pantropical - 8 species (c. 17%), Calymperes afzelii, C. erosum, C. lonchophyllum, Syrrhopodon gardneri and 5. parasiticus. 2. Palaeotropical to Oceania - 7 species (c. 15%), Calymperes serratum, C. taitense, Syrrhopodon armatus, S. involutus, S. spiculosus, S. trachyphyllus and 5. tristichus. 3. Indo-Pacific - 21 species (c. 46%), Calymperes aeruginosum, C. moluccense, C. motleyi, C. porrectum, Mitthyridium constrictum, M. flavum, M. repens, Syrrhopodon albovaginatus, S. confertus, S. croceus, S. japonicus and 5. muelleri. 4. Malesian-Pacific - one species (c. 2%), Mitthyridium subluteum. 5. South India, Indochina, SW China and Central Malesia disjunct - 4 species (c. 9%), Calymperes mangalorense, Chameleion peguense, Syrrhopodon flammeonervis and S. tjibodensis. 6. Malesia - 4 species (c. 9%), Calymperes robinsonii, C. subserratum, Mitthyridium wallisii and Syrrhopodon rufescens. 1. Philippine endemic - one species (c. 2%), Mitthyridium iwatsukianum. From the above analysis it is evident that the majority (c. 46%) of © The Natural History Museum, 1999 2 L.T. ELLIS AND B.C. TAN Philippine Calymperaceae are widely distributed from tropical Asia Key 1. Calymperes, Syrrhopodon and Chameleion to Oceania. About 17% are pantropical and 15% are palaeotropical in distribution. Only 9% (four species) are restricted to Malesia, one 1 Leaves linear, costa and marginal ribs covered with irregular nodules of species is Malesia and the western Pacific, and one species, small chlorophyllose cells in more or less transverse rows (Fig. 9b, c, f, Mitthyridium iwatsukianum, is locally endemic to Luzon island. 8) Calymperes strictifolium This overall distribution well reflects the predominantly coastal Leaves linear or not, lacking irregular nodules of small chlorophyllose distribution and preference for an epiphytic lifestyle in lowland cells 2 habitats among members of this family. Species of Calymperaceae 2 Leaves linear above hyaline base; cells immediately above hyaline must be efficiently wind dispersed to achieve such a broad range ,amina .„ ,eaf basg short,y subrectangulan smooth red with thick Evidence is seen in the frequent production of gemmae at the tips of pkted wa,ls distinctly largerthan the ceiis of the chlorophyllose lamina leaves; in many species some leaves become strongly modified for above (Fig. 18e) Syrrhopodon croceus the production of gemmae (see also Reese, 1987b). However, a few species, such as Calvmperes mangalorense, Chameleion peguense Reaves linear or not; cells above hyaline lamina not differentiated as .. ,. iU . described above 3 and Syrrhopodon tjibodensis, notwithstanding their gemma-pro- ducing capabilities, appear to have failed to disperse very far. They 3 Leaves soft, Ungulate to spathulate, with long, remote cilia on costa and have some limitation that has resulted in their present-day disjunc- leaf margin; dorsal and ventral surfaces of chlorophyllose lamina tive range in India, Indochina, SW China and a few island groups in smooth and flat (Fi§- 16g-») Syrrhopodon ciliatus central Malesia, where seasonally dry lowland rainforest predomi- Leaves variously shaped, lacking cilia, or if cilia present, then cells of nates. chlorophyllose lamina protuberant and/or papillose 4 4 Hyaline lamina often occupying two thirds or more of leaf blade and/or hyaline cells continuing distally from hyaline lamina to near leaf apex, SYSTF1VT ATTC1 ACCOUNT underlying the chlorophyllose lamina in 1 or 2 rows adjacent to costa — — (Fig. 18m, o) 5 Calymperaceae Kindb., Gen. Eur. N.-Amer. Bryin.: 11 (1897).Type Hyaline lamina occupying less than two thirds of leaf blade; eenus' Calvmneres Sw ex F Web chlorophyllose lamina without underlying rows of hyaline cells adja- cent to costa 7 Shoots erect or creeping, forming cushions, tufts or mats on trees, ,j.1T ..I r i-i.. i../"*, 5 Leaves 1.5-3.5 mm long with a short, but distinct chlorophyllose limb; rock and soil. Leaves mostly linear, ungulate or lanceolate. Costa . , margins denticulate from shoulders to apex strong, ending just below apex or excurrent, smooth, toothed, spmose Syrrhopodon confertus or ciliate; in nonleucobryoid genera usually composed of dorsal and ventral bands of stereids separated by 1-2 or more layers of guide Leaves 1-1 -5 mm long, lacking a distinct chlorophyllose limb; margins cells (in leucobryoid genera largely composed of large, empty largely entire (small teeth sometimes evident near leaf apex) 6 hyaline cells supporting networks or layers of small chlorophyllose 6 Leaves suberect to erecto-patent; peristome teeth papillose cells), superficial cells often differentiated. Lamina in basal region Syrrhopodon involutus of leaf composed of large, empty, thin-walled, porose hyaline cells; Leaves suberect to patent-recurved; peristome teeth finely stnate above basal region composed of small, thick-walled chlorophyllose c . do r f sc ns cells (sometimes incorporating hyaline cells), smooth, toothed, spinose or papillose. Leaf margin often with a marginal or 7 Distal extent of hyaline lamina poorly defined (showing a gradual intramarginal rib, entire, toothed, papillose, spinose or ciliate. Cla- transition from large hyaline cells in the base to small chlorophyllose vate to fusiform, multicellular, mostly uniseriate gemmae often cells in the distal leaf) (Fig. 8f); shoots usually acaulescent; leaves linear produced from a definite region of the costa (usually the apex), sometimes on specialized leaves. Dioecious. Sporophytes terminal; Hyaline lamina sharply defined with large, thin-walled hyaline cells seta smooth; capsule usually cylindrical. Peristome haplolepidous abutting small, thick-walled chlorophyllose cells (Fig. lOg); shoots (sometimes absent or reduced) and calyptra fugacious (Syrrhopodon, acaulescent or not; leaves linear or not 10 Chameleion and Mitthyridium), or peristome absent and calyptra g Leaves erect, bristle-like (wet or dry); margins above leaf base persistent (Calymperes). polystratose and largely entire Syrrhopodon aristifolius Kev to aenera and snecies LeaV6S variously incurled when dry> strap-like; margins above leaf base * unistratose and entire or, polystratose with well-developed, uni- to Note: This key includes four taxa which are highly likely to occur in multicellar, often double teeth 9 the Philippines, Calymperes crassinerve (Mitt.) Jaeg., Mitthyridium 9 Leaves <10-20 mm long; margins above leaf base polystratose, toothed junquilianum (Mitt.) H. Rob., Syrrhopodon prolifer var. albidus Calymperes serratum (Thwaites & Mitt.) S. Orban & W.D. Reese, and S. prolifer var. • /•/-« .\ o /~v u ' D « r r\ n Leaves c. 5 mm long; margins above leat base unistratose, entire tosaensis (Cardot) S. Orban & W.D. Reese. However, as the authors ^ , _ Calymperes subserratum are presently unaware of collections of these taxa from the study area they are not featured in the detailed descriptions of taxa in this ' ° Leaves Ungulate, less than 3 mm long; lamina on either side of hyaline paper region in leaf base formed by a broad band of rounded-quadrate/shortly subrectangular cells with thickened angles (Fig. 5k) 1 1 1 Plants with erect shoots, branched or simple; leaf margins various .... Leaves various; hyaline region in leaf base bordered on either side by ... Key 1 : Calymperes, Syrrhopodon, Chameleion narrow marginal (Fig. 24d) or intramarginal (Fig. 1 i) rib of thick- walled Plants with creeping main shoots and ascending branches; leaf margins largely formed (extending from leaf base to near apex) by a flattened, mostly unistratose* band of stereids Key 2: Mitthyridium *Polystratose in the Indonesian species Mitthyridium retusum (Besch.) W.D. Reese. CALYMPERACEAE IN THE PHILIPPINES 1 1 Cells of chlorophyllose lamina rounded, collenchymatous, mostly 8-20 x 7.5-15 |im; costa ending below leaf apex; gemmae produced from ventral surface of costal apex Calymperes motley! Cells of chlorophyllose lamina polygonal, not collenchymatous, mostly 7-12(-14) x 5-10(-12.5) u,m; costa usually excurrent with gemmae produced from all around the tip Calymperes tenerum 1 2 Hyaline leaf base with intramarginal band of thick- walled, linear cells (Figs li, 3b, h) 13 Hyaline leaf base with marginal rib of thick-walled linear cells (Figs 20m, 24d) 28 13 Costa in cross-section with 2 or more layers of guide cells (Fig. If).. 14 Costa in cross-section with a single layer of guide cells (Fig. 2e) .. 17 1 4 Leaves broadly linear, at apex abruptly narrowing into a linear proboscis (Fig. lOe) Calymperes taitense Leaves narrowly linear or strap-like, at apex tapering to a blunt to acute tip 15 15 Leaves strap-like, <9->15 mm long 16 Leaves bristle-like, up to 7.5 mm long Calymperes aeruginosum 16 Lamina abruptly narrowing into costa for a short distance above the hyaline base, giving leaf a petiolate appearance Calymperes robinsonii Lamina not narrowing into costa above hyaline base Calymperes lonchophyllum 17 Lumina in cells of chlorophyllose lamina not or hardly protruding dorsally or ventrally (walls papillose or not), subrectangular in cross- section (Fig. 3e) 18 Lumina in cells of chlorophyllose lamina protruding ventrally, thumb- nail- or shield-shaped in cross-section (Fig. 31) 20 18 Leaves dimorphic, with erect, bristle-like gemmiferous leaves and recurved, Ungulate nongemmiferous leaves Calymperes porrectum Leaves not dimorphic, all linear-lanceolate, lanceolate or strap-like 19 19 Leaves strap-like, <9->15 mm long; shoots usually acaulescent Calymperes lonchophyllum Leaves lanceolate to linear-lanceolate, 5-7.5 mm long; shoots with well-developed stems Calymperes fasciculatum 20 Costa excurrent; leaves hardly dimorphic, gemmae (when present) produced from all around the costal apex 21 Costa ending just below leaf apex; leaves dimorphic (gemmiferous and nongemmiferous); gemmae produced from the ventral surface of a modified costal apex 22 2 1 Rib bordering lamina in distal leaf intramarginal (bordered by 2-4 rows of chlorophyllose cells); cells of chlorophyllose lamina with one or two simple papillae on the ventral surface; distal-most cells of hyaline lamina acutely exserted, from above appearing to overlap adjacent cells of the chlorophyllose lamina Calymperes erosum Rib bordering lamina in distal leaf marginal; cells of chlorophyllose lamina with a single acute summit projecting from the ventral leaf surface; distal-most cells of hyaline lamina usually with flat ventral surfaces Calymperes mangalorense 22 Gemmiferous leaves lanceolate to Ungulate (when moist); blades of chlorophyllose lamina usually broader than costa, often abruptly nar- rowing into apical proboscis and becoming recurved below apex (Fig. 1 h) 23 Gemmiferous leaves linear, bristle-like; blades of chlorophyllose lamina often narrower than costa, tapering to just below apex (usually broaden- ing at apex) (Fig. 9i) 27 23 Marginal lamina bordering intramarginal rib in hyaline leaf base com- posed of 2->5 rows of small, subquadrate hyaline cells; proboscis in gemmiferous leaves narrow with lamina narrowly recurved, becoming plane but not, or barely, broadening out at apex; leaves commonly more than 4 mm long Calymperes afzelii Marginal lamina bordering intramarginal rib in hyaline leaf base com- posed of l-2(-3) rows of small, subquadrate to shortly subrectangular hyaline cells; proboscis in gemmiferous leaves with lamina broadening somewhat around the apex; leaves rarely more than 4 mm long .... 24 24 Nongemmiferous leaves up to 2 mm long; gemmiferous leaves strongly infolded when dry, with a cowl-like apex ... Calymperes crassinerve Nongemmiferous leaves usually more than 2 mm long; apex of hyaline lamina various in shape but rarely strictly truncate; gemmiferous leaves straight to curled when dry but not infolded, apices cowl-like or not . 25 25 Costa in gemmiferous and nongemmiferous leaves thick (with an inflated appearance), lacking stereids; leaf margins unistratose through- out Calymperes boulayi At least in nongemmiferous leaves costa lacking inflated appearance and incorporating stereids; leaf margins above hyaline base usually formed by a polystratose rib 26 26 Cells of chlorophyllose lamina 5- 1 2(- 1 4) (xm wide, thick- walled (thick walls particularly evident around shoulders of hyaline base where the cells are polygonal but possess rounded lumina), protruding acutely from the ventral leaf surface, often with thick, pleuripapillose summits; lamina at apices of gemmiferous leaves forming a broad, oval-concave 'collar' Calymperes moluccense Cells of chlorophyllose lamina 4-7.5 fim wide, protruding roundly to subacutely from the ventral leaf surface, not pleuripapillose or notably thickened; lamina at apices of gemmiferous leaves forming narrow, spoon-shaped 'collar' Calymperes graeffeanum 27 Superficial walls of cells in hyaline lamina of even thickness; in gemmiferous leaves lamina forming a narrow (often obscure) collar above the costal apex; in nongemmiferous leaves cells of chlorophyllose lamina subacutely to acutely protuberant from the ventral leaf surface (shield-shaped in cross-section); costa and margins without lamellae Calymperes subintegrum Superficial walls of cells in hyaline lamina with horizontal bands of thickening; in gemmiferous leaves lamina forming a funnel-shaped leaf apex; in nongemmiferous leaves cells of chlorophyllose lamina roundly protuberant from the ventral leaf surface (fingernail-shaped in cross- section); short lamellae (1-3 cells high) often apparent on ventral surface of costa and dorsal surfaces of marginal ribs Chameleion peguense 28 Leaf margin adjacent to apex of hyaline lamina entire 29 Leaf margin adjacent to apex of hyaline lamina with teeth, spines or cilia 35 29 Cells forming surface of marginal rib differentiated, in cross-section subquadrate to subrectangular or rounded 30 Cells forming surface of marginal rib not differentiated, composed of stereids 31 30 Leaves mostly patent to recurved with apices finely drawn out; leaf margins above mid-chlorophyllose limb strongly toothed Syrrhopodon tristichus Leaves erect to erecto-patent with blunt apices; leaf margins entire (sometimes weakly toothed near apex) Syrrhopodon muelleri 3 1 Leaves Ungulate or elongate triangular; often with filamentous gemmae produced just above the hyaline base from lateral cells in the ventral L.T. ELLIS AND B.C. TAN surface of the costa (Fig. 22g) 32 Leaves narrowly lanceolate or linear; gemmae not produced as described above 33 32 Cells of chlorophyllose lamina acutely protuberant from the ventral leaf surface, dorsally unipapillose (Fig. 22k); apex of hyaline lamina acute (Fig. 22g) Syrrhopodon parasiticus Cells of chlorophyllose lamina with compound papillae on dorsal and ventral surfaces (Fig. 24n); apex of hyaline lamina rounded- truncate (Fig. 24k) Syrrhopodon tjibodensis 33 Chlorophyllose lamina with acute, distally-pointing teeth arranged in transverse rows occurring at regular intervals (Fig. 16c, e) Syrrhopodon albovaginatus Chlorophyllose lamina lacking teeth arranged in distinct transverse 34 34 Leaf abruptly narrowing at apex of hyaline lamina, forming distinct 'shoulders' (Fig. 19c) Syrrhopodon flammeonervis Leaf gradually narrowing at apex of hyaline lamina, hardly forming 'shoulders' Syrrhopodon prolifer var. albidus 35 Marginal rib around mid-chlorophyllose limb with subquadrate to subrectangular or rounded superficial cells, some cells forming double or single, unicellular to multicellular teeth at regular intervals 36 Marginal rib around mid-chlorophyllose limb with linear superficial cells, entire or with each cell forming a distally pointing tooth 38 36 Cells of chlorophyllose lamina protruding acutely from the ventral leaf surface, often papillose dorsally and ventrally (Fig. 20t) Syrrhopodon gardneri Cells of chlorophyllose lamina flat to slightly roundly protuberant, smooth 37 37 Marginal rib lacking stereids; chlorophyllose lamina hardly constricted above hyaline leaf base; leaves less than 9 mm long; shoots with well- defined stem Syrrhopodon japonicus Marginal rib incorporating stereids; chlorophyllose lamina strongly constricted for a short distance above the hyaline leaf base; leaves 6- >30 mm long; shoots acaulescent Syrrhopodon loreus 38 Surface of costa largely formed by stereids, entire (except sometimes near leaf apex) (Fig. 24h) 39 Ventral and/or dorsal surfaces of costa composed of subquadrate to subrectangular or rounded cells (surface view), some cells often giving rise to teeth, spines or papillae (Fig. 23h, j) 40 39 Leaves with spinose teeth at apex Syrrhopodon prolifer var. tosaensis Leaves lacking spinose teeth at apex .... Syrrhopodon trachyphyllus 40 Superficial cells of marginal rib around mid-chlorophyllose limb each forming an acute, distally pointing tooth Syrrhopodon hispidissimus Marginal rib around mid-chlorophyllose limb entire, or only a few superficial cells forming teeth or spines 41 * 41 Leaves abruptly narrowing from subrectangular-elliptical hyaline leaf base into a linear chlorophyllose limb; margins spinose at 'shoulders', regularly toothed in distal chlorophyllose limb, from base of chlorophyllose limb to above mid-limb entire Syrrhopodon spiculosus Leaves ligulate, slightly tapering around apex of hyaline lamina; mar- gins often spinose orciliate from above mid-hyaline base to beyond mid leaf , not regularly toothed towards leaf apex Syrrhopodon armatus * An apparently undescribed variety of Syrrhopodon prolifer Schwagr., represented by a single Philippine collection, keys out here. See under excluded taxa. NB. Most of the genera in the family Calymperaceae that possess leucobryoid leaves are unlikely to be confused with the nonleucobryoid genera An exception is Exostratum L.T. Ellis. Exostratum blumei and E. sullivantii are known to occur in the Philippines. These species have a similar aspect to Syrrhopodon spiculosus and S. hispidissimus. However, unlike the latter, the leaves in Exostratum have a rather stiff appearance when wet or dry, in the costa there is a complete lack of stereids and the position occupied by guide cells mSyrrhopodon is occupied by chlorophyllose cells. Key 2. Mitthyridium 1 Leaves 1-3.5 mm long 2 Leaves mostly 3.5->5 mm long 9 2 Apices of leaves modified with lamina inrolled to form a cup or tube (proboscis) (Fig. 15b) 3 Apices of leaves plane, acute to rounded 5 3 Leaves up to 2.5 mm long, marginal band of thick-walled linear cells less than 50 urn wide at its broadest point; lamina at leaf apex loosely inrolled to form a tube, tube often flaring distally to form a narrow funnel Mitthyridium wallisii Leaf length various; marginal band of more than 100 urn wide at broadest point; modified leaf apices cup-shaped 4 4 Leaves tristichous, strictly ranked; cells of lamina smooth with evenly thickened walls, lumina seldom substellate; dorsal surface of costa near apex smooth Mitthyridium iwatsukianum Leaves seldom strictly ranked; cells of lamina papillose with incrassate, unevenly thickened walls enclosing substellate lumina (Fig. 12b); dorsal surface of costa near apex with spinose teeth Mitthyridium constrictum 5 Leaves tapering from base of chlorophyllose limb towards apex 6 Leaves with parallel sides, chlorophyllose limb not tapering until close to apex 8 6 Leaves mostly narrowly lanceolate with acute apices 7 Leaves with broadly flared shoulders, tapering to an obtuse apex Mitthyridium fasciculatum subsp. cardotii 7 Leaves 3-3.5 mm long; hyaline lamina occupying about a seventh of the leaf length Mitthyridium subluteum Leaves seldom more than 2 mm long; hyaline lamina occupying up to about a quarter of the leaf length Mitthyridium junquilianum 8 Leaves 1.5-2 mm long; apices acute or acuminate (Fig. 12d) Mitthyridium flavum Leaves 1-1.5 mm long; apices rounded-apiculate (Fig. 14b) Mitthyridium repens 9 Apices of leaves rounded-obtuse with marginal rows of lamina cells incrassate and smooth; apex of hyaline lamina truncate Mittyridium fasciculatum subsp. obtusifolium Apices of leaves acute or acuminate with marginal rows of laminal cells similar to those below; apex of hyaline lamina broadly acute 10 10 Leaves with broad, pronounced shoulders, tapering sharply to leaf apex; marginal band of linear, thick- walled cells usually extending from base to near apex of leaf; cells of chlorophyllose lamina (in surface view) formed by 2-4 rounded lobes, papillose with papillae easily seen in surface view Mitthyridium fasciculatum subsp. fasciculatum Leaves with barely defined shoulders (sometimes lacking), sides of chlorophyllose limb tapering gently towards leaf apex, or parallel and CALYMPERACEAE IN THE PHILIPPINES tapering abruptly near apex; marginal band of linear thick-walled cells extending from base to over two thirds the length of the chlorophyllose limb; cells of chlorophyllose lamina polygonal in surface view (not usually lobed), obscurely papillose, seen easily only in cross-section \ lit th\ i id in m papuanum Descriptions and illustrations of taxa Calymperes Sw. ex F. Web., Tab. Calyptr. operc. (1813). Type species: Calymperes lonchophyllum Schwagr. Shoots erect, simple or branched, forming mats or tufts. Leaves often dimorphic (gemmiferous and nongemmiferous), narrowly to broadly Ungulate, lanceolate or linear, consisting of a hyaline, semi- sheathing base that narrows slightly, sometimes abruptly, into a chlorophyllose limb, apices various. Costa ending below apex to excurrent. Cells of chlorophyllose lamina small, mostly isodiametric, smooth, papillose, and/or protuberant. Cells of hyaline lamina large, empty, mostly smooth, with round to irregular pores in superficial and transverse walls; border between hyaline and chlorophyllose lamina usually sharply defined. Marginal and/or intramarginal ribs frequently present, usually intramarginal in hyaline base. Gemmae produced in a radial group, often from the modified apices of specialized leaves, fusiform to clavate, sometimes filamentous and branched, multicellular, uniseriate, smooth or papillose. Dioecious. Perichaetia terminal (innovations often fertile). Seta normally exserted. Capsule cylindrical with a conical operculum, enclosed in a persistent calyptra. Peristome absent. Calymperes is most closely related to Syrrhopodon. The most important distinguishing feature is the presence in Calymperes of a persistent calyptra. It has been suggested that the calyptra in Calymperes plays a peristome-like role in the dispersal of spores (Edwards, 1980). In Syrrhopodon the calyptra is fugacious, falling from the capsule prior to spore dispersal. The differentiated leaf margin that occurs in many species in the Calymperaceae is often helpful in distinguishing between Calymperes and Syrrhopodon. In most species of Calymperes the hyaline lamina forming the leaf base possesses an intramarginal rib (in some species obscure or absent). In Syrrhopodon the hyaline lamina is often bordered by a marginal rib (in some species obscure or absent, rarely intramarginal). Calymperes occupies a broad range across the wet regions in tropical and subtropical belts (Reese, 1987a). Species are mostly corticolous and lowland in distribution (mostly below 800 m in the Philippine archipelago). Many are confined to tree trunks along the margin of humid forests, especially in coastal areas of small islands. There are about 24 species of Calymperes known from the Malesian region, of which 18 are reported here for the Philippines. A phytogeographical and floristic summary of the Philippine taxa is presented above in the introduction. Calymperes aeruginosum Hampe ex Sande Lac. in Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 13(2): 7 (1872). Type: Philip- pines, Basilan Island, near Maridanao, 5 January 1860, Semper s.n. (BM!-isotype). Fig. la-f. Calymperes mammosum Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 291 (1895). Type: Philippines, Cuming 2214 (BM!-holotype; BM!- isotype). Shoots reaching 1-1.5 cm high, usually dark green. Leaves mostly straight and bristle-like (Fig. la, b) (sometimes incurled when dry), erect to patent, up to c. 6 mm long, linear with a constriction in the lamina for a short distance above the hyaline base, broadening distally then narrowing gradually to a blunt, dentate apex. Costa strong, often occupying >25->90 % of the width of the leaf, ending just below apex in a bluntly pointed or slightly expanded truncate tip, largely smooth, in upper leaf sometimes with a few blunt, multicellular teeth; internally with 2 layers of guide cells sand- wiched between dorsal, median and ventral bands of stereids (Fig. If)- Chlorophyllose lamina sometimes hardly apparent along con- striction in leaf; cells mostly isodiametric to slightly longer than broad, with 4-6 sides or rounded, 6-12.5(-15) x 6-10(-12) |im, ventrally roundly to subacutely protuberant, dorsally flat. Hyaline lamina poorly to sharply defined. Leaf margin from around constric- tion in leaf to near apex plane to erect, formed by a thick polystratose rib, lacking stereids (Fig. If), superficial cells in surface view similar to those of the chlorophyllose lamina (slightly smaller) and some occasionally forming distant, blunt, (often multicellular) teeth; from constriction in leaf to below shoulders of leaf becoming unistratose, consisting of chlorophyllose lamina, denticulate; below shoulders of leaf to base often with an intramarginal band of long rectangular to linear, thick- walled cells, about 1-3 cells wide (Fig. Id) (frequently obscure), laminal margin denticulate, composed of shortly subrectangular, thin-walled, hyaline cells in l(-3) rows. Long, filamentous paraphyses produced in the axils of some leaves. Many leaves bearing gemmae on the ventral surface of the costal tip. HABITAT. On trunks and exposed roots of trees, and on basic rock, in damp, shady lowland rainforest. The specimen cited below was growing on soil on limestone. DISTRIBUTION. An Indo-Pacific species. Uncommon in the Philip- pines, but known also from Luzon and Mindanao. SPECIMEN EXAMINED. Palawan, St Paul Bay, St Paul Subterranean National Park, 25 May 1989, Tan 89-1413b (FH). Eddy (1990) notes plants of this species to be 'dark, blue-green to almost black'. However, this feature does not appear to be universal, as the recent collection cited above is green. Although yellowed by time, there is no remnant of dark colouration in the type ofCalymperes mammosum Besch. (dimming 2214, BM). Calymperes afzelii Sw. in Jahrb. Gewachsk. 1: 3 (1818). Type: Africa, Afzelius s.n. (BM!-isotype). Fig. Ig-m. Calymperes vriesii Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 307 (1895). Type: Sulawesi, Menahaye van Menado, de Vriese s.n. (Hb. Leyden, no. 4) (BM!-holotype). Shoots reaching >3.5 cm high, in yellowish green tufts or mats. Leaves up to 5.5 mm long, dimorphic: nongemmiferous leaves Ungulate; apex obtuse, broadly pointed, sometimes apiculate (Fig. Ig). Costa ending just below apex; above hyaline lamina rough with small, acute projections. Cells of chlorophyllose lamina mostly isodiametric with 4-6 sides, (3-)4-10(-12.5) x (3-K-7.5 ^m (Fig. Ik), roundly to acutely protuberant from the ventral leaf surface, sometimes with a simple papilla at the summit (Fig. 1m). Hyaline lamina sharply defined; cells in distal region often isodiametric, arranged in somewhat regular rows. Leaf margin from above hyaline base to near apex consisting of a polystratose marginal rib (usually strong), superficial cells subquadrate to shortly subrectangular, often forming small (unicellular) teeth, internally lacking stereids; in hyaline base with a narrow intramarginal band of linear, thick- walled cells (continuous with marginal rib in distal leaf), marginal lamina entire, composed of 2->5 rows of thin-walled, subquadrate to shortly subrectangular hyaline cells (Fig. li). Gemmiferous L.T. ELLIS AND B.C. TAN a,b,g c,h 1 mm 0.5 mm m - "° • d-f, i-m 50 jam Fig. 1 a-f. Calymperes aeruginosum Hampe ex Sande Lac. a-c: leaves (a, b: in ventral view, with c: detail of mid-leaf; d, e: cells of leaf in surface view (d: at margin in hyaline base, e: in chlorophyllose lamina; f: cross-section of chlorophyllose limb. g-m. Calymperes afzelii Sw. g: leaf in ventral view; h: apex of gemmiferous leaf (dorsal view); i-k: cells of leaf in surface view (i: at margin of mid-hyaline base, j: at margin in chlorophyllose limb, k: of chlorophyllose lamina); 1, m: cross-sections of leaf (1: costa, m: chlorophyllose lamina and marginal rib. a, c, d Drawn from Sarawak, Jermy 13734 (BM). b, e, f Drawn from Tan 89-1413b (FH). g-m Drawn from Tan 91-286 (BM). CALYMPERACEAE IN THE PHILIPPINES leaves similar to above but possessing a linear apical proboscis (Fig. Ih). Costa extending into proboscis. Lamina abruptly nar- rowing into proboscis and becoming tightly recurved, at leaf apex becoming plane and forming a narrow, denticulate margin around the costal tip. Gemmae produced from the ventral surface of the costal apex. HABITAT. On shaded trunks, exposed roots, decaying stumps and logs, sometimes on rock (the material cited below occurred on shale); occurring mostly in lowland areas. DISTRIBUTION. A pantropical species. A previous report of this species from the Philippines (Menzel & Schultze-Motel, 1990) does not cite a collection or specific locality. SPECIMENS EXAMINED. Balabac Island, Sitio Indalawan near Indalawan Village, 28 April 1 993, Tan 93-228 (BM). Palawan, Puerto Princesa, Barangay San Rafael, Batac Village, 1 May 1991, Tan 91-286 (BM). The leaves in Calymperes afzelii closely resemble smaller versions of those occurring in Calymperes taitense (Sull.) Mitt. However, the leaves of the latter always possess a narrow apical proboscis (poten- tially gemmiferous) and two rows of guide cells in the costa (a median layer and a smaller layer below the ventral surface). In C. afzelii at least some leaves have unmodified, flat apices, and there is a single (median) layer of guide cells in the costa (Fig 11). Calymperes boulayi Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 278 (1895). Type: Borneo, Korthals s.n. (BMMectotype (fide Ellis, 1988); L!, NY!-isotypes?). Fig. 2i-n. Calymperes dozyanum Mitt, sensu M. Fleisch.,Mwsc. Fl. Buitenzorg 1:266(1904). Shoots reaching 2(-3) cm high, forming yellowish green mats or tufts. Leaves dimorphic: nongemmiferous leaves up to 3.5 mm long, Ungulate to broadly Ungulate (Fig. 2i), with apices obtusely pointed. Costa ending below apex, with an inflated appearance, above hya- line base rough with small, acute projections, internally virtually lacking stereids. Chlorophyllose lamina often with laxly incurved margins, cells isodiametric to slightly longer than broad, mostly irregularly quadrate to hexagonal, 5-10(-13) x (3-)5-7.5 |im, ven- trally bluntly to acutely protuberant, dorsally flat, smooth or unipapillose. Hyaline lamina sharply defined. Leaf margin from around apex of hyaline lamina to leaf apex unistratose, entire or notched, rarely faintly denticulate toward leaf apex (Fig. 21); below apex of hyaline lamina to leaf base with a narrow, unistratose intramarginal band of thick-walled linear cells (sometimes weakly developed), marginal lamina consisting of a row of thin-walled, subrectangular hyaline cells, entire to faintly denticulate (Fig. 2k). Gemmiferous leaves reaching 4 mm long, linear to broadly Ungulate with an apical gemma-bearing proboscis (Fig. 2j). Costa as in nongemmiferous leaves but thicker. Lamina narrowing into probos- cis, usually becoming recurved, distally becoming plane and broadening into a concave, oval collar around the costal tip. Gem- mae produced from ventral surface of costal apex. HABITAT. On trunks of trees, logs and stumps, sometimes on rock, rarely on humus at the base of trees; mostly occurring in lowland areas. DISTRIBUTION. An Indo-Pacific species.As yet there are no records from Luzon. SPECIMENS EXAMINED. Mindanao, Zamboanga, \9\3,Binstead\42(BM). Palawan. Port Barton, 30 April 1991, Tan 92-312 (FH). The costa in both gemmiferous and nongemmiferous leaves of Calymperes boulayi has a rather inflated appearance and lacks stereids; marginal or intramarginal ribs are absent in the upper leaf, and at the apices of gemmiferous leaves a broad 'collar' of chlorophyllose lamina cups the gemmae (Fig. 2j). Together, these features distinguish this species from all others. Calymperes erosum Mull. Hal. in Linnaea 21: 182 (1848). Type: Surninam, near Paramaribo, Hb. Kegel 539 (PC!-isotype). Fig. 2a-h. Calymperes hampei Dozy & Molk., Bryol jav. 1: 48 (1856). Type: Java, Teysmann s.n. (BM!-isotype?). Calymperes sandeanum Besch. in Ann. Sci. Nat. Bot. s6r. 8, 1: 303 (1895). Type: Borneo, near Pontianak, van Oorschot s.n. (in Hb. Lacoste) (BM!-holotype). Shoots 8) rows of small, subquadrate, thick-walled, chlorophyllose cells; in hyaline base entire below, notched to denticulate distally; composed of l(-3) rows of narrowly rectangular, thin- walled, hya- line cells (Fig. 2b). Gemmiferous leaves with an excurrent costa; gemmae produced from all around the costal tip. HABITAT. On trunks, branches and exposed roots of trees, tree stumps, not infrequently on rock, sometimes on soil; mainly occur- ring in lowland areas. DISTRIBUTION. A nearly pantropical species. SPECIMENS EXAMINED. Balabac Island, Sitio Melville, 28 April 1993, Tan 93-232 (FH). Culion Island, Barangay Culion, 6-7 May 1992, Tan 92-328 (BM); Tan 92-381 (BM); Sitio Ugnisan, Leyson Rancho, 7 May 1992, Tan 92-377 (BM). Mindoro Island, between Bongabon and Pinamalayan, Feb- ruary-April 1941, Maliwanag 136 (BM, FH). For a comparison of Calymperes erosum with C. mangalorense Dixon & P. de la Varde see under account of the latter. Calymperes fasciculatum Dozy & Molk., Bryol. jav. 1: 50 (1856). Type: Java, Teysmann s.n. (L!-holotype; BM!-isotype). Fig. 3a-e. Calymperes johannis-winkleri Broth, in Mitt. Inst. Allg. Bot. Ham- burg 7(2): 122 (1928). Type: West Borneo, Bukit Raja, 1250 m, 9 December 1924, Winkler 3169 (BM!-isotype). Syrrhopodon hasagawae Tak. & Iwats. in J. Hattori Bot. Lab. 21: 240 (1959). Type: Japan, Kagshima Pref., Isl. Yaku, Odakumi, 14 L.T. ELLIS AND B.C. TAN a, i b-h, k-n n 1 mm 50 jam 0.5 mm Fig. 2 a-h. Calymperes erosum Miill. Hal. a: leaf (ventral view); b-d: cells of leaf in surface view (b: at margin of hyaline base, c: around apex of hyaline lamina, d: in chlorophyllose lamina and at margin); e-h: cross-sections of leaf (e: costa, f; chlorophyllose lamina, g: margin in chlorophyllose limb, h: lamina and margin near apex of hyaline lamina, i-n. Calymperes boulayi Besch. i: nongemmiferous leaf (ventral view); j: apex of gemmiferous leaf (ventral view); k, 1: cells of leaf in surface view (k: at margin of hyaline base, 1: at margin of chlorophyllose limb; m,n: cross-sections of leaf (m: costa, n: chlorophyllose lamina), a-h Drawn from Tan 92-328 (BM). i-n Drawn from Tan 92-312 (FH). April 1950, Hasagawa, Fukuhara & Fukui 42757 (NICH- holotype). Calymperes hasagawae (Tak. & Iwats.) Iwats. in 7. HattoriBot. Lab. 28:220(1965). Calymperes johannis-winkleri var. hasagawae (Tak. & Iwats.) Iwats. in J. Jap. Bot. 43: 476 (1968). Shoots reaching >5 cm high, in green tufts. Leaves erect to spreading (moist), mostly 5-7.5 mm long, lanceolate to linear-lanceolate, often narrowing abruptly a short distance above hyaline lamina (forming shoulders), apex narrowly acute, denticulate (Fig. 3a). Costa ending immediately below leaf apex in a blunt, toothed tip, mostly smooth. Cells of chlorophyllose lamina thick-walled, trans- versely to longitudinally elongate, with 4-6 sides, often rounded-elliptical, in distal leaf (8-) 10-1 7.5 x 10-15(-22.5) |im, smooth (Fig. 3d, e). Hyaline lamina extending from leaf base to about half way to shoulders, usually sharply defined. Leaf margins from leaf shoulders to near apex formed by a polystratose rib, i.e. a strand of stereid/substereid cells within a superficial layer of isodiametric cells similar to those of the lamina, some forming multicellular teeth (distant below, becoming more closely set to- wards leaf apex) (Fig. 3c); from shoulders to distal hyaline lamina undifferentiated, entire to notched; adjacent to hyaline lamina with an intramarginal band of thick-walled, linear cells in about 7 rows, marginal lamina entire to uneven, composed of thin- walled, shortly rectangular hyaline cells in a single row (Fig. 3b). Some leaves bearing gemmae on dorsal and ventral sides of costal apex; gemmae sometimes branched, sparsely papillose. HABITAT. On tree trunks, decaying wood and rock. DISTRIBUTION. An Indo-Pacific species. In the Philippines known previously only from Luzon. SPECIMEN EXAMINED. Panay Island, Antique, Mt Madyaas, 15-16 Janu- ary 1987, Price s.n. (FH). CALYMPERACEAE IN THE PHILIPPINES 1 mm 0.5 mm Fig. 3 a-e. Calymperes fasciculatum Dozy & Molk. a: leaf in ventral view; b-d: cells of leaf in surface view (b: at margin in hyaline base, c: at margin in chlorophyllose limb, d: in chlorophyllose lamina); e: cross-section of chlorophyllose limb. f-m. Calymperes graeffeanum Mull. Hal. f: leaf in ventral view; g: apex of gemmiferous leaf; h-j: cells of leaf in surface view (h: at margin in hyaline base, i: cells at margin in chlorophyllose limb, j: in chlorophyllose lamina); k-m: cross-sections of leaf (k: costa, 1: chlorophyllose lamina, m: marginal rib), a, d, e Drawn from Price s.n. (FH). b, c, e Drawn from Sarawak, Bell 2055 (BM). f-m Drawn from Tan 93-297 (FH). 10 L.T. ELLIS AND B.C. TAN Calymperes graeffeanum Mull. Hal. in/. Mus. Godeffroy 3(6): 64 (1874). Type: Western Samoa, Upolu, Graeffes.n. (BMl-isotype). Fig. 3f-m. Calymperes semperi Hampe in Besch. in Ann. Sci. nat. Bot. ser. 8, 1: 302 (1895).Type: Philippine Islands, Sempers.n. (BM!-holotype; L!-isotype?). Calymperes hyophilaceum Miill. Hal. ex Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 265, 287 ( 1 895).Type: Philippine Islands, Calumpit, Llanos s.n. (BM?, not found; B?, destroyed 1943?). Calymperes hyophilaceum var. robustum M. Fleisch., Muse. Fl. Buitenzorg 1: 265 (1904). Type: West Java, Rezidenz Krawang Bei Tjikao, 1899, Fleischer (Musci Frond. Arch. Ind. no. 263) (BM!, L!-isotypes). Shoots reaching l(-2) cm high, yellowish green. Leaves mostly 1 .5- 3.5 mm long, dimorphic: nongemmiferous leaves Ungulate to narrowly Ungulate, leaf apex obtuse to subacute (Fig. 3f). Costa ending below apex, above hyaline base rough with small, acute projections. Chlorophyllose lamina plane, erect or laxly incurved, cells drawn out ventrally as subacute to acute projections, dorsally flat or unipapillose, mostly 5-10 x 4-7.5 urn, mostly isodiametric with 4-6 sides (Fig. 3j, 1). Hyaline lamina sharply defined. Leaf margin from above apex of hyaline lamina to near leaf apex usually formed by a narrow, irregularly subdenticulate to denticulate polystratose rib (sometimes weak or absent), in surface view appear- ing as 2-3 rows of subquadrate chlorophyllose cells, sometimes incorporating stereids (Fig. 3i, m); adjacent to apex of hyaline lamina unistratose, entire to denticulate; from leaf base to below apex of hyaline lamina with a narrow, unistratose, intramarginal band of linear, thick-walled cells, marginal lamina unistratose, notched to denticulate, composed of 1-2 rows of shortly rectangular, thin-walled hyaline cells (Fig. 3h). Gemmiferous leaves lanceolate to narrowly Ungulate with an apical gemma-bearing proboscis, apex rounded or truncate (Fig. 3g). Costa thick, often with a slightly inflated appearance, ending below apex of proboscis, internally incorporating many wide-lumened substereid cells, normal stereids few. Chlorophyllose lamina narrowing gradually to abruptly into proboscis and becoming tightly recurved, broadening distally and becoming plane to form narrow margin around the costal tip. Gemmae produced from the ventral surface of the costal tip. HABITAT. On tree trunks and sometimes on rock, in lowland forest DISTRIBUTION. An Indo-Pacific species. Widespread in the Philip- pines. SPECIMENS EXAMINED. Luzon, Mt Makiling, 27 August 1931, Herklots P30c (BM); Bataan Province, Lomas River, Williams 815 (NY); Isabela, Palanan Wilderness, 19 May 1992, Tan 92-207 (BM); 21 May 1992, Tan 92- 181 (FH); Palawan: near Sitio Daan, Aborlan, 26 April 1992, Tan 92-259 (BM); St Paul Bay, St Paul Subterranean National Park, 25 May 1989, Tan 89-1417 (BM); Sabang Municipality, St Paul Underground River Subterra- nean Park, 3 May 1993, Tan 93-297 (FH); Iwahig, Balsahan, 4 April 1993, Tan 93-1 88 pro parte (FH); Barangay Puerto Princesa, Sitio Kalabayog, near Batac Village, 23-25 April 1993, Tan 93-222 (FH). Calymperes lonchophyllum Schwagr., Sp. muse, frond, suppl. 1(2): 333 (1816). Type: Central America, 'Guyanne', Richard s.n. (PC!-isotype). Fig. 4a-e. Shoots reaching <1.0->2.5 cm high, forming dense mats. Stems short to almost lacking. Leaves <9 >15 mm long, composed of a short, semi-sheathing, subelliptical hyaline base narrowing into a long-linear chlorophyllose limb (curled and contorted when dry, laxly suberect when moist); apex obtuse to acute, normally denticu- late (Fig. 4a). Costa ending in leaf apex to shortly excurrent, smooth; internally with l-2(-3) layers of guide cells (Fig. 4e). Chlorophyllose lamina occasionally transversely undulate, mostly unistratose, occasionally with bistratose streaks and patches, cells in surface view rounded, irregularly polygonal or quadrate to shortly subrectangular, mostly wider than long, <5-10(-12.5) x 7-15(-20) um, thick-walled, flat, smooth or pleuripapillose (Fig. 4d, e). Hya- line lamina usually sharply defined. Leaf margin in base (adjacent to hyaline lamina) with an intramarginal rib of thick-walled, linear cells (c. 3-6 cells wide), marginal cells in l-3(-6) rows, hyaline, thin-walled, irregular-elliptical, some in outermost row flask-shaped forming a bluntly denticulate margin (Fig. 4b); immediately above hyaline base formed by chlorophyllose lamina, unistratose, entire to denticulate; from a short distance above hyaline base to near leaf apex consisting of a robust polystratose rib, in section usually triangular, composed of small chlorophyllose cells and often incor- porating a central strand of stereids; entire below; above usually with one to two rows of distant teeth (arising from angles of rib, often paired) (Fig. 4c), becoming closer set and more pronounced toward the leaf apex, each composed of a short, single row of cells ending distally as an acute projection. Long axillary hairs sometimes present, median and distal cells about 4 times longer than wide. Gemmae sometimes produced from the ventral surface of the costal apex (occasionally also from well below the apex), yellowish brown with simple papillae. HABITAT. On tree trunks, sometimes on rock, in shaded rainforest. DISTRIBUTION. A widespread pantropical species. SPECIMENS EXAMINED. Luzon, Quezon National Park, Atimonan, 9 March 1986, Tan & Lipaygo s.n. (FH). Palawan, St Paul Bay, St Paul Subterranean National Park, 25 May 1989, Tan 89-1416 (BM); Puerto Princesa, Barangay Irawan, Mt Malinao, 4 May 1993, Tan 93-324 (FH); Barangay Apurawan, Sitio Daan, vicinity of Mt Tinik-basan, 27 April 1992, Tan 92-266 (BM). Calymperes serratum A. Braun ex Miill. Hal. and C. lonchophyllum are superficially very similar but may be separated by two key features. Firstly, in C. lonchophyllum the hyaline lamina (in most leaves) is very sharply defined, with the relatively large, thin-walled, empty cells at its apex abutting the small, thick-walled, green cells forming the base of the chlorophyllose lamina. In C. serratum the distal cells of the hyaline lamina intergrade gradually with those of the chlorophyllose lamina. The second key distinguishing feature was recognized by Akiyama & Reese (1993) and Reese & Stone (1995). Long axillary hairs occur in both species (particularly commonly in C. serratum). In C. lonchophyllum the median and distal cells of these hairs are about four times as long as broad; those in C. serratum are seldom more than twice as long as broad. Additionally, the chlorophyllose lamina in C. serratum is consist- ently smooth and unistratose, and the intramarginal rib in the hyaline leaf base is often weak to the point of absence. In C. lonchophyllum the chlorophyllose lamina is often papillose with bistratose patches, and the intramarginal rib in the hyaline leaf base is well-developed. Calymperes mangalorense Dixon & P. de laVarde in Arch. Bot. (Paris) 1(8-9): 164 (1927). Type: Southern India, Mangalore, Kanakanady, November 1925, Foreau 22 (BM!-holotype; PC!- isotype). Fig. 4f-k. Shoots reaching 2 cm high, forming yellowish green tufts or mats. Leaves up to 3.5(-4) mm long, hardly dimorphic; narrowly to broadly Ungulate, distal chlorophyllose lamina sometimes involute; leaf apex acute to obtuse, occasionally apiculate (Fig. 4f). Costa ending in apex to shortly excurrent, above hyaline base usually CALYMPERACEAE IN THE PHILIPPINES 11 1 mm 50 jam Fig. 4 a-e. Calymperes lonchophyllum Schwagr. a: leaf; b-d: cells of leaf in surface view (b: at margin in distal hyaline base, c: in marginal rib in chlorophyllose limb, d: in chlorophyllose lamina); e: cross-section of chlorophyllose limb. f-k. Calymperes mangalorense Dixon & P. de la Varde f: leaves; g-j: cells of leaf in surface view (g: at margin of distal hyaline base, h: around apex of hyaline lamina, i: in chlorophyllose lamina, j: at margin in chlorophyllose limb); k: cross-section of chlorophyllose lamina and marginal rib. a-e Drawn from Tan 89-1416 (BM). f-k Drawn from Tan s.n. (FH). rough with small, acute projections. Cells of chlorophyllose lamina subisodiametric with 4-7 sides, <5-ll(-12.5) x 5-10 (im (Fig. 4i), drawn out ventrally into rounded to acute projections, dorsally flat to unipapillose (Fig. 4k). Hyaline lamina sharply defined. Leaf margin beyond apex of hyaline lamina to near leaf apex, entire to uneven, formed by a polystratose rib of subquadrate to shortly subrectangular chlorophyllose cells (stereids sometimes present) (Fig. 4j, k), some- times becoming intramarginal around apex of hyaline lamina; from around distal hyaline lamina to leaf base with an intramarginal unistratose band of long rectangular to linear thick-walled cells (often continuous with marginal rib in upper leaf, sometimes poorly devel- oped), marginal lamina composed of 1-3 rows of thin- walled, subrectangular hyaline cells, entire to denticulate (Fig. 4g). Gemmiferous leaves with excurrent costa, gemmae produced from all around the costal tip. HABITAT. The specimen cited below occurred on the bark of a tree in open karst forest . DISTRIBUTION. Formerly known only from India and Burma. SPECIMEN EXAMINED. Coron Reef Island (Palawan Province), Coron Town, Lake Kayangan, off coast of Coron Municipality of Basuanga Island, 8 May 1992, Tan s.n. (FH) (new record). This species has been confused with Calymperes erosum. In the leaves of the latter there is an intramarginal rib in the limb; the distal rows of cells in the hyaline lamina protrude acutely from the ventral surface of the leaf, overlapping the cells immediately distal to them; and (in most specimens) at least a few of the ventrally protuberant cells forming the chlorophyllose lamina have bipapillose summits. The leaves of C. mangalorense have a marginal rib in the limb (Fig. 4k), lack protuberant hyaline cells (Fig 4h), and the ventrally protuberant cells forming the chlorophyllose lamina all possess unipapillose summits. Calymperes mangalorense closely resembles Calymperes graef- feanum in all features apart from the possession of leaves with an excurrent costa. 12 L.T. ELLIS AND B.C. TAN a,h, i 1 mm c-g,j-m 0.5 mm 50 Fig. 5 a-g. Calymperes tnoluccense Schwagr. a: nongemmiferous leaf (ventral view); b: apex of gemmiferous leaf (ventral view); c-e: cells of leaf in surface view (c: at margin of hyaline base, d: around apex of hyaline lamina, e: in chlorophyllose lamina); f, g: cross-sections of leaf (f: costa and chlorophyllose lamina, g: marginal rib and chlorophyllose lamina), h-n. Calymperes motleyi Mitt, ex Dozy & Molk. h: gemmiferous leaf (dorsal view); i: apex of nongemmiferous leaf; j, k: cells of leaf in surface view in (j: chlorophyllose lamina, k: margin of hyaline base); 1-n: cross-sections of leaves (1: costa (nongemmiferous), m: costa (gemmiferous), n: chlorophyllose lamina), a-g Drawn from Tan 92-186 (FH). h-n Drawn from Tan 93-240 (FH). Calymperes moluccense Schwagr. Sp. muse, frond, suppl. 2(1): 99 (1824). Type: Moluccas, Rawak, Gaudichaud 29 (15) (BM!- isotypes). Fig. 5a-g. Calymperes palisotii subsp. moluccense (Schwagr.) M. Menzel in M. Menzel & Schultze-Motel in Willdenowia 19: 489 (1990). Shoots reaching 2(-3) cm high, green above, often dark brown below, forming tufts or mats. Leaves 2.5-3.5 mm long, dimorphic: nongemmiferous leaves consisting of a narrowly lingulate chlorophyllose limb, extending (spreading when moist) from the slightly flared shoulders of a suberect hyaline base, distal chlorophyllose lamina often involute, leaf apex obtuse, sometimes apiculate (Fig. 5a). Costa ending below apex, above hyaline base rough with coronate-papillose projections. Cells of chlorophyllose lamina drawn out ventrally into acute, often coronate-papillose projections, dorsally smooth or unipapillose (Fig. 5f, g), thick- walled (especially in region adjacent to hyaline lamina (Fig. 5d)), mostly 5-13 x 5-12 |im, isodiametric with 4-6 sides or rounded. Hyaline lamina sharply defined, often differentiated: cells in rows nearer the margin narrow with unevenly thickened walls (Fig. 5c), in rows adjacent to the costa broad and evenly thin- walled. Leaf margin from above shoulders to near apex formed by a narrow, irregularly subdenticulate, polystratose rib, in surface view appearing as 2-3 CALYMPERACEAE IN THE PHILIPPINES 13 rows of subquadrate chlorophyllose cells, rarely incorporating stereids; at shoulders unistratose and denticulate, formed by small, thick-walled chlorophyllose cells; below shoulders with an intramarginal band of linear, thick-walled cells (sometimes continu- ous with marginal rib of upper leaf), marginal lamina unistratose, entire to denticulate, formed by 1-2 rows of subrectangular, thin- walled, hyaline cells (Fig. 5c). Gemmiferous leaves narrowly Ungulate to lanceolate with an apical, gemma-bearing proboscis (Fig. 5b). Costa stout. Lamina abruptly narrowed into proboscis and becoming tightly recurved; at apex, broadening and becoming plane to form a collar around the costal tip. Gemmae produced from ventral surface of costal tip. HABITAT. On the trunks and branches of trees, stumps and logs, sometimes on rock; occurring in lowland areas. DISTRIBUTION. Widespread in the Indo-Pacific. In the Philippines, in addition to the localities cited below, also known from Bohol and Mindanao. SPECIMENS EXAMINED. Culion Island (Palawan Province), near Sitio Ugnisan, 7 May 1992, Tan 92-382 (BM). Luzon, Isabela Province, Palanan Wilderness, 20 May 1 992, Tan 92-21 2 (FH); 21 May 1992, Tan 92-1 86 (FH); Bicobian Bay, 22 May 1992, Tan 92-177 (FH). Palawan (Palawan Prov- ince), Barangay Napsan, Sitio Tagkulit, Salakot Waterfall vicinity, 2 May 1993, Tan 93-265 (FH). Calymperes palisotii Schwagr. has been regarded by some authori- ties as a variety of C. moluccense. However, C. palisotii appears to be as distinct a species as any other in the genus (Ellis, 1987) and occurs mainly in the New World tropics, Africa and western SE Asia; it is extremely rare in Malesia and absent from Oceania. Calymperes moluccense has an Indo-Pacific distribution. In C. moluccense the leaves often have flared 'shoulders' where the hyaline lamina broad- ens before abruptly narrowing into the chlorophyllose limb; at the apices of gemmiferous leaves the chlorophyllose lamina forms a broad 'collar' around the gemmae (Fig. 5b); most cells in the chlorophyllose lamina and superficial cells in the costa protrude acutely and are often multipapillose on the protruding surface (Fig. 5f, g), those laminal cells adjacent to the distal hyaline lamina are usually strongly incrassate with a visible middle lamella (Fig. 5d); the cells of the hyaline lamina forming the rows adjacent to the intramarginal rib are often differentiated from those nearer the costa, being strikingly narrower with walls more unevenly thickened. In C. palisotii the leaves usually possess less pronounced shoulders (some- times not evident); the lamina at the apices of gemmiferous leaves forms a narrow band around the costal apex; the cells of the chlorophyllose lamina and the superficial cells in the costa are roundly protuberant and lack papillae on the protruding surface, those laminal cells adjacent to the distal hyaline lamina are not unusually thick-walled; and the cells of the hyaline lamina are evenly thin- walled and not differentiated, with a gradual decrease in cell size across the lamina from the costa to the intramarginal rib. Calymperes motley! Mitt, ex Dozy & Molk., Bryol. jav. 1: 48 (1856). Type: Borneo, Labuan Island, Tanjong, Motley s.n. (BM!- isotype). Fig. 5h-n. Shoots reaching 1.5 cm high, forming yellowish green tufts or mats. Leaves about 2 mm long (erect to patent when moist), dimorphic: nongemmiferous leaves obovate to Ungulate, distal chlorophyllose lamina sometimes involute, leaf apex obtuse to obtuse-apiculate. Costa ending below apex; in distal leaf ventral superficial cells protruding roundly, some dorsal superficial cells drawn out as blunt projections; internally incorporating some substereid cells. Cells of chlorophyllose lamina collenchymatous, rounded to subhexagonal, 8-20 x 7.5-15 |im (Fig. 5j), protruding roundly from the ventral leaf surface, dorsally flat to slightly convex and/or unipapillose (Fig 5n). Hyaline lamina sharply defined and enclosed on either side by broad marginal bands of small quadrate to shortly rectangular cells with differentially thickened transverse walls and/or angles (Fig 5k). Leaf margins unistratose, entire, slightly irregular. Gemmiferous leaves Ungulate with an apical, gemma-bearing, shortly suboblong probos- cis (Fig 5h). Costa often thicker than in nongemmiferous leaves, often incorporating substereid cells, extending into proboscis. Lamina abruptly narrowing into proboscis and becoming recurved, distally becoming plane to form a narrow, rounded margin around the costal tip. Gemmae arising from the ventral surface of the costal apex. HABITAT. On trees in lowland areas. The specimen cited below occurred in an open, windy mangrove forest of Avicenna. DISTRIBUTION. A widespread Indo-Pacific species. SPECIMEN EXAMINED. Balabac Island (Palawan Province), Sitio Melville, 28 April 1993, Tan 93-240 (FH) (new record). Calymperes motleyi and C. tenerum Mull. Hal. are similar in size, possess leaves with almost identical hyaline bases, and have some- times been confused with each other. Under the microscope they are immediately distinguishable. In C. motleyi, the large (8-20 x 7.5-15 urn), unipapillose, collenchymatous cells that form the chlorophyllose lamina (Fig. 5j) contrast stongly with the small (7-12.5 x 7-10(- 12.5) um) polygonal cells of the chlorophyllose lamina in C. tenerum (Fig. lOo). Apart from the key features, these species are also distinguishable by the attitude of the leaves on the stem when dry. The apices in C. motleyi tend to roll inwards; in C. tenerum the leaves when dry often twist spirally around the shoot, or curl variously. Calymperes porrectum Mitt, in J. Linn. Soc. Bot. 10: 172 (1868). Type: Samoa, Tutuila, Powell 10 (BM!-isotype). Fig. 6. Calymperes salakense Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 271, 302 ( 1 895). Type: Java, MontSalak, near Buitenzorg, 400 m, Kurz 154 (BM!-holotype). Calymperes scalare Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 303 (1895). Type: Philippines, Basilan Island, Semper s.n. (BM!- holotype). Shoots robust, reaching >6 cm high, in yellowish green tufts or mats. Leaves mostly 5-7 mm long, strongly dimorphic: nongemmiferous leaves consisting of a recurved (moist or dry), narrowly to broadly lanceolate limb extending from the (usually distinct) shoulders of a suberect hyaline base; leaf apex narrowly to broadly acute (Fig. 6b). Costa ending immediately below apex to shortly excurrent, above hyaline base superficial cells smooth or drawn out into small, acute projections. Cells of chlorophyllose lamina mostly isodiametric, subquadrate to rounded, <5-10(-12.5) x <5-10(-12.5)|im, smooth, flat or ventrally slightly protuberant (Fig. h, j). Hyaline lamina sharply defined. Leaf margin with an intramarginal rib extending from leaf base to near apex; from shortly above apex of hyaline lamina polystratose, a strand of stereid-like cells with superficial cells similar to those of the lamina (Fig. 6k); from above apex of hyaline lamina to leaf base becoming a band of linear, thick-walled cells in 4-5 rows. Marginal lamina unistratose, narrowing from shortly below apex of hyaline base towards leaf apex, from about midleaf apparent only as a series of single, large, multicellular teeth; below midleaf to mid-hyaline lamina uneven to dentate, composed of several rows of cells similar to those of the chlorophyllose lamina 14 L.T. ELLIS AND B.C. TAN a 1 mm c,d e-k 0.5 mm 50 Fig. 6 a-k. Calymperes porrectum Mitt, a, b: leaves (a: gemmiferous leaf, b: nongemmiferous leaf); c, d details of leaf (c: proximal hyaline base, and d: margin of chlorophyllose limb); e-h: cells of leaf in surface view (e: at margin in proximal hyaline base, f: at margin in distal hyaline base, g: at margin in chlorophyllose limb, h: in chlorophyllose lamina); i-k: cross-sections of leaf (i: costa, j: chlorophyllose lamina, k: margin in chlorophyllose limb, a-d Drawn from Semper s.n. (BM). e, f Drawn from North Borneo, Wood 1625 (BM). g-k Drawn from Papua New Guinea, Cheesman s.n. (BM). (Fig. 6g); from around mid-hyaline lamina to leaf base entire to denticulate, consisting of 1-3 rows of shortly rectangular to slightly irregular thin-walled hyaline cells (Fig. 6e). Gemmiferous leaves suberect, a linear chlorophyllose limb abruptly narrowing from a subelliptical hyaline base (Fig. 6a); leaf apex rounded to truncate, denticulate. Costa thick, occupying more than two thirds of the width of the chlorophyllose limb for most of its length, shortly excurrent and broadening slightly at apex; beyond hyaline base rough with subacute to acute projections. Chlorophyllose lamina narrowing gradually from above hyaline base to just short of leaf apex. Gemmae produced from all around costal tip. HABITAT. On tree trunks, rarely on soil, in moist, shaded forest; occurring from near sea level to around 900 m. DISTRIBUTION. An Indo-Pacific species. No local collections examined, apart from the type of Calymperes scalare Besch. Calymperes robinsonii B.C. Tan & W.D. Reese in W.D. Reese & B.C. Tan in Bull. Natl. Sci. Mus., Tokyo, Series B, 9: 30 (1983). Type: Philippines, Palawan Island, Mt Apis, Ebalo & Conklin 82803 (PNH-holotype). Fig. 7. Shoots acaulescent. Leaves <9->17 mm long, composed of a short, semi-sheathing, subelliptical hyaline base with a long-linear chlorophyllose limb (stiffly suberect when moist) (Fig. 7a); leaf blade immediately above hyaline base narrowing abruptly into costa, absent distally for a short distance (l-<2 mm) then re- emerging gradually into the chlorophyllose limb (i.e. forming a petiole) (Fig. 7c); apex subacute to obtuse, denticulate. Costa usu- ally ending in apex, smooth; internally mostly with 2—4 layers of guide cells (costa in region of petiole especially thick and with guide cells in several layers) (Fig. If). Chlorophyllose lamina sometimes undulate, mostly unistratose, sometimes with small bistratose patches, cells in surface view rounded, irregularly polygonal or CALYMPERACEAE IN THE PHILIPPINES 15 § b,c 1 mm 0.5 mm 50 Fig. 7 a-h. Calymperes robinsonii B.C. Tan & W.D. Reese a-d: leaf (a: semidiagram and details of b: apex, c: 'petiolate' region above hyaline base, d: distal hyaline base; e: cells at margin in leaf base; f-h: cross-sections of leaf (f: costa in distal chlorophyllose limb, g: chlorophyllose lamina, h: marginal rib in chlorophyllose limb), b, c Drawn from Tan 89-1426 (BM). d, f, g, h Drawn from Tan 93-224 (FH). e Drawn from Tan 89-1422 (BM). quadrate to shortly subrectangular, mostly wider than long, <5-10 x <5->l 2.5 um, thick- walled, flat, smooth to obscurely pleuripapillose. Hyaline lamina sharply defined, often bulging ventrally at distal extremity. Leaf margin from insertion to beyond mid-hyaline base similar to that in C. lonchophyllum with an intramarginal rib of thick-walled linear cells within a narrow marginal band of thin- walled hyaline cells (Fig. 7e); from distal hyaline base to petiole formed by a narrow band of chlorophyllose lamina, unistratose, denticulate; in petiole entire; from above petiole to near leaf apex consisting of a polystratose rib largely composed of small chlorophyllose cells, in cross-section triangular, lacking a central strand of stereids, distant multicellular teeth arising from angles of rib, (Fig. 7h). Groups of long, filamentous axillary hairs produced in some leaves. Gemmae sometimes produced from the leaf apex. HABITAT. On shaded tree trunks and occasionally on limestone outcrops. DISTRIBUTION. A western Malesian species. SPECIMENS EXAMINED. Luzon, Mt Arayat, 1896, Loher 1057 (BM). Palawan, St Paul Bay, St Paul Subterranean National Park, 25 May 1 989, Tan 89-1422 (BM); Tan 1426 (BM); Barangay Puerto Princesa, Sitio Kalabayog, near the Batac Village, '23, 25' April 1993, Tan 93-224 (FH). The abrupt and complete disappearance of the lamina just above the hyaline base and its equally abrupt reappearance shortly above will distinguish leaves of this species from those of any other of the long- leaved species of Calymperaceae occurring in the Philippines. A less extreme constriction of the lamina occurs in the leaves of Syrrhopodon loreus (Sande Lac.) W.D. Reese. However, in this species, the margins of the distal hyaline leaf base are replete with acute, thick- walled teeth (Fig. 2 It), and the costa possesses a single layer of guide cells (Fig. 21u). In C. robinsonii the hyaline base has an intramarginal rib, with a margin incorporating thin-walled hya- line cells (Fig. 7e); the costa includes 2—4 layers of guide cells (Fig. 7f). Calymperes aeruginosum also possesses linear leaves with a constricted lamina. This species has short leaves (c. 6 mm long) as compared to those in C. robinsonii (mostly 9-17 mm long). In the latter, the cells of the chlorophyllose lamina are flat ventrally, those of C. aeruginosum are ventrally protuberant. Calymperes serratum A. Braun ex Mull. Hal., Syn. muse, frond. 1: 527 (1849). Type: Java, Junghuhn s.n. (BM!, L!-isotypes). Fig. 8. Shoots reaching > 2 cm high, in dense green mats or tufts; stems very short, barely apparent. Leaves 10-20 mm long, linear, laxly erect to suberect (moist), with a subelliptical hyaline base; above hyaline base narrowing for a short distance then broadening slightly and gradually towards mid leaf; near apex narrowing to form an acute, dentate tip (Fig. 8a-d). Costa ending in apex to shortly excurrent, smooth; internally with 2(-3) layers of guide cells. Chlorophyllose lamina sometimes transversely undulate, cells transversely to longi- tudinally elongate, rounded-elliptical or with 4-6 sides, 5-1 0(- 12. 5) x 8-12.5 (im, smooth, flat to slightly protuberant (Fig. 8g, h, k). Hyaline lamina poorly defined, distal hyaline cells merging gradually 16 L.T. ELLIS AND B.C. TAN b,c,l 0.5 mm e-k 1 mm 50 Fig. 8 a-k. Calymperes serratum A. Braun ex Mull. Hal. a-d: leaf (a: semidiagram (ventral view), and details of b: apex, c: mid-leaf, d: lower leaf); e-i: cells of leaf in surface view (e: at margin of distal hyaline base, f: around apex of hyaline base, g: in proximal chlorophyllose lamina, h: in distal chlorophyllose lamina, i: at margin of chlorophyllose limb); j, k: cross-sections of leaf (j: costa, k: chlorophyllose lamina and marginal rib); 1: axillary hair, b-i Drawn from Herklots P14 (BM). j-1 Drawn from Elmer 10387 (BM). into chlorophyllose lamina above (Fig. 8f). Leaf margin a short distance beyond hyaline base formed by a polystratose rib, inter- nally sometimes incorporating a strand of stereids, superficial cells in surface view similar to those of lamina, many forming multicellu- lar, often distant, paired teeth (Fig. 8i, k); from proximal limit of marginal rib to below apex of hyaline lamina composed of chlorophyllose lamina, unistratose, entire to serrulate; from above mid-hyaline lamina to leaf base sometimes with a weak intramarginal band of linear thick-walled cells in about 3^4 rows, marginal lamina uneven to notched, consisting of 3-4 rows of subquadrate to shortly rectangular, thin-walled hyaline cells (Fig. 8e). Groups of long axillary hairs occur frequently, median and distal cells of hairs seldom more than twice as long as broad (Fig. 81). Gemmae pro- duced from dorsal and ventral sides of the costa at and near the leaf apex; gemmae sometimes branched, sparsely papillose. HABITAT. On tree trunks, mostly in lowland forest, but reaching over 1000 m. DISTRIBUTION. A palaeotropical species. In addition to localities in the Philippines cited below, also known from Mindanao. SPECIMENS EXAMINED. Luzon, Cordil\era,Semper s.n. (BM); Mt Makiling: 16 August 1931, Herklots P14 (BM, BM-K); 21 March 1982, Tan&Aguila 82-03 (BM); 6-9 December \9\2,Robinson 17121 (BM-K).Negros, Orien- tal Province: mountains in southern part of province, 1 May 1958, Brown 2859 (BM, BM-K); Cuernos Mountains, Dumaguete, June 1908, Elmer 10387 (BM, BM-K). The following combination of features should separate Calymperes serratum from other long-leaved species in the family, such as Calymperes lonchophyllum, C. robinsonii, Syrrhopodon aristifolius Mitt, and S. loreus (Sande Lac.) W.D. Reese: a) leaves curled when dry; b) hyaline lamina (in all leaves) poorly defined (i.e. with the cells of the hyaline lamina gradually merging into those of the chlorophyllose lamina); c) leaves with toothed, polystratose margins above the hyaline base; d) chlorophyllose lamina unistratose through- out; e) costa with 2(-3) layers of guide cells. The distinguishing features separating C. serratum from C. lonchophyllum are discussed in the account of the latter. CALYMPERACEAE IN THE PHILIPPINES 17 3. m n h v a, h, i 1 mm b,c d-g,j-n 0.5 mm 50 Fig. 9 a-g. Calymperes strictifolium (Mitt.) G. Roth a-c: leaf (a: in dorsal view, with details of b: distal region of hyaline base, c: chlorophyllose limb (dorsal view)); d-f: cells of leaf in surface view (d: at margin near leaf base, e: at margin above mid-hyaline base, f: at margin in chlorophyllose limb); g: cross-section of chlorophyllose limb, h-n Calymperes subintegrum Broth, h-j: leaves (h: nongemmiferous leaf in ventral view, i: gemmiferous leaf in dorsal view, with detail of j: gemmiferous apex in dorsi-lateral view); k-m: cells of leaf in surface view (k: at margin in hyaline base, 1: at margin in proximal chlorophyllose limb, m: in chlorophyllose lamina); n: cross-section of chlorophyllose lamina and margin, a-g Drawn from New Guinea, Eddy 6527 (BM). h-n Drawn from Alvarez Jr 0-781 138 (BM). Calymperes strictifolium (Mitt.) G. Roth in Hedwigia 51: 127 (1911). Fig. 9a-g. Syrrhopodon strictifolius Mitt, in Seem., Fl. vit.: 388 (1873). Type: Samoa, Tutuila, Powell s.n. (BM!-isotype). Syrrhopodon tuberculosus Ther. & Dixon in Dixon in J. Linn. Soc. Bot. 43: 303 (1916). Type: Borneo, Sekong, 22 April 1913, C.H. Binstead 84 (BM!-holotype). Calymperes tuberculosum (Then & Dixon) Broth., Nat. Pflanzenfam. 2nd ed., 10:240(1924). Shoots reaching 1.5-<2 cm high, in dark green mats. Leaves up to 5 mm long, obscurely dimorphic, linear-lanceolate, erect to erecto- patent (Fig. 9a). Costa usually ending immediately below leaf apex, above hyaline base often occupying over a third of the width of the leaf; from distal region of the hyaline base to leaf tip covered (ventrally and dorsally) in irregular, knob-like, multicellular nodules arranged in closely set transverse rows. Blades of lamina gradually narrowing towards leaf apex; at leaf apex forming an extremely narrow band around the costal tip (often eroded away); cells of chlorophyllose lamina isodiametric to about twice as long as broad, with 4-6 sides or rounded, 5-10(-12.5) x 5-7.5 |im; ventrally bluntly to acutely protuberant, sometimes crowned with papillae, dorsally flat or slightly convex. Hyaline lamina sharply defined. Leaf margin from around distal region of hyaline base to near leaf apex consisting of a thick polystratose rib with similar ornamenta- 18 L.T. ELLIS AND B.C. TAN tion to that of the costa, sometimes incorporating stereids (Fig. 9f, g); in hyaline base similar to that in C. afzelii, i.e. entire; possessing a narrow, unistratose intramarginal band of linear, thick-walled cells (continuous with marginal rib in distal leaf), marginal lamina com- posed of hyaline, thin-walled, shortly subrectangular cells arranged in about 2-5 rows (Fig. 9d). Some leaves producing gemmae from the ventral surface of the costal apex; narrow band of chlorophyllose lamina at the leaf apex tending to be pushed backwards as the gemmae develop forming a minute concave collar around the costal tip. HABITAT. On trunks and buttresses of trees, sometimes on logs or rocks; in damp, shady lowland rainforest. DISTRIBUTION. An Indo-Pacific species. In the Philippines re- ported from Batan, Luzon and Mindanao (see Tan & Iwatsuki, 1991). SPECIMEN EXAMINED. 'Philippine Islands', Cuming 2214b (BM, BM-K). Within the Calymperaceae, multicellular nodules (Fig. 9f) are unique to the leaves of Calymperes strictifolium. Calymperes subintegrum Broth, in J. Schmidt in Bot. Tidsskr. 24: 1 19 (1901). Type: Thailand, Koh Chang, near Klong Majum, [6 January 1900], Schmidt [14] (H!-holotype; BM!-isotype). Fig. 9h-n. Shoots reaching 1 cm high, forming yellowish green tufts. Leaves strongly dimorphic: nongemmiferous leaves Ungulate to lanceolate, upper lamina plane to involute, apex roundly subacute, 2-3(^0 mm long (Fig. 9h). Costa ending just below leaf apex; above hyaline base rough with simple, acute projections (more so on the ventral sur- face). Cells of chlorophyllose lamina isodiametric to slightly longer than broad with 4-6 sides, 7.5-15(-20) x 5-10(-12.5) urn (Fig. 9m), drawn out ventrally as acute, often unipapillose protuberances, dorsally smooth (Fig. 9n). Hyaline lamina sharply defined. Leaf margin above hyaline base consisting of a row of small, thick- walled, shortly rectangular cells, some projecting distally to form small teeth (Fig. 91); in hyaline base with a narrow, unistratose (rarely bistratose), intramarginal band of linear, thick-walled cells, 1-3 cells wide (sometimes poorly developed or absent), marginal lamina composed of a single row of shortly rectangular hyaline, thin-walled cells, some projecting distally to form notches or small teeth (Fig. 9k). Gemmiferous leaves erect, linear, 3-5(-8) mm long (Fig. 9i), costa incrassate, a little less than twice the thickness of that in nongemmiferous leaves. Blades of lamina from shortly beyond the hyaline base to the leaf apex narrower than costa, meeting above the costal tip to form a narrow, uneven to denticulate 'collar' (Fig. 9j). Gemmae produced from the ventral surface of the costal apex. HABITAT. On decaying wood and boulders in moist, shady forest. DISTRIBUTION. An Indo-Pacific species. Within the Philippines, in addition to the specimen from Luzon, cited below, this species has also been recorded from Palawan by Tan (1996). SPECIMEN EXAMINED. Luzon, Quezon National Park, Atimonan, 16 Sep- tember 1978, Alvarez Jr 0-781 138 (BM) (new record). The binomial, Calymperes subintegrum, was placed in synoymy with Calymperes schmidtii Broth, by Reese & Mohamed (1985). Ellis (1991) demonstrated that the holotype of the latter (Thailand, Schmidt 14, H) possesses features that are not shared with the type specimen of C. subintegrum, or those of any of its other proposed synonyms. The features of C. schmidtii are very similar to those of Calymperes tenerum Mull. Hal. Calymperes subintegrum is the earliest name for the species described above. Calymperes subintegrum is easily distinguished from other spe- cies with strongly dimorphic leaves occurring in the region. For example, Calymperes porrectum usually has recurved nongemmiferous leaves with a strongly developed intramarginal rib and large multicellular teeth along the margin. In C. subintegrum the leaves are not recurved, and the leaf margins above the hyaline base are weakly differentiated and possess small irregularly occurring, unicellular teeth. Chameleion peguense (Besch.) L.T. Ellis & A. Eddy possesses leaves in which the cells of the hyaline lamina have transverse bands of thickening, and the gemmiferous leaves possess a broad, cowl-like apex (Fig. lie). In contrast, C. subintegrum possesses leaves in which the cells of the hyaline lamina lack bands of transverse thickening, the gemmiferous leaves have a narrow, blunt apex, and the lamina forming the leaf apex is often obscure Calymperes subserratum M. Fleisch., Muse. Fl. Buitenzorg 1: 245 ( 1 904). Type: Java, Tjamea, 300 m, Fleischer s.n. (FH!-holotype). Fig. lOa-d. Calymperes clemensiae Broth, in Philipp. J. Sci. C. Bot. 8: 69 (191 3). Type: Philippines, Mindanao, Camp Keithley, Lake Lamao, June 1907, Clemens T' (FH!-isotype). Shoots <8 mm high, somewhat flattened into one plane, almost fan- like, forming mats. Stems very short, densely leaved, and often dense with rhizoids below. Leaves up to 5 mm long, not dimorphic, broadly linear, suberect to recurved, upper lamina often incurved to involute; leaf apex acute or apiculate, entire to denticulate (Fig. lOa). Costa ending immediately below, or in apex; superficial cells above hyaline base bluntly to subacutely protuberant (protuberances often more strongly pronounced towards leaf apex); internally with a single row of guide cells. Cells of chlorophylose lamina thick- walled, mostly isodiametric, with 4-6 sides or rounded, <5-10(-12.5) x <5-7.5 urn (Fig. lOb); dorsally subacutely to bluntly protuberant; ventrally flat to convex, smooth to unipapillose (Fig. lOd). Hyaline lamina poorly defined, hyaline cells gradually merging with chlorophyllose lamina. Leaf margin unistratose; in chlorophyllose lamina often erect to incurved, entire to subdenticulate (sometimes denticulate towards the leaf apex); in hyaline lamina unistratose, entire. Long, filamentous, uniseriate, hyaline paraphyses produced in axils of some leaves. HABITAT. On tree trunks, recorded at 300 m and 520 m (Reese & Streimann, 1994). DISTRIBUTION. Java, New Britain and Mindanao. No other local specimens examined. This species, maintained as distinct by Reese et al. (1986), was placed in synonymy with Calymperes serratum A. Braun ex Mull. Hal. by Eddy (1990) and Menzel & Schultze-Motel (1990). How- ever, Reese, Koponen & Norris (1986) and Reese & Streimann (1994) demonstrated that Calymperes subserratum is indeed a dis- tinct species. Its short leaves (up to 5 mm long), with unistratose margins (Fig. lOd) are easily distinguished from those of C. serratum, which are mostly 10-20 mm long, with polystratose marginal ribs above the leaf base. Calymperes taitense (Sull.) Mitt, in J. Linn. Soc. Bot. 10: 172 (1868). Fig. 10e-l. CALYMPERACEAE IN THE PHILIPPINES 19 a, e, m b-d, g-1, n-q 1 mm 50 0.5 mm Fig. 10 a-d. Calymperes subserratum M. Fleisch. a: leaf; b: cells of chlorophyllose lamina (dorsal surface); c, d: cross-sections of leaf (a: costa, b: chlorophyllose lamina and margin), e-1. Calymperes taitense (Sull.) Mitt, e, f: leaf (e: in ventral view, f: detail of margin in proximal hyaline lamina); g- h: cells of leaf in surface view (g: around apex of hyaline lamina, h: at margin of chlorophyllose limb (ventral view); i-1: cross-sections of leaf (i: costa in mid-hyaline base and j: in chlorophyllose limb, k: margin in hyaline base and 1: in chlorophyllose limb), m-q. Calymperes tenerum Mull. Hal. m: leaves; n, o: cells of leaf in surface view (n: near margin in hyaline base, o: in chlorophyllose lamina); p, q: cross-sections of leaf (p: costa, q: chlorophyllose lamina and marginal rib), a-d Drawn from Java, Fleischer s.n. (FH). e-1 Drawn from Tan 91-268 (BM). m-q Drawn from Tan 92-354 (FH). 20 L.T. ELLIS AND B.C. TAN Syrrhopodon taitense Sull., U. S. Exploring Expedition. Musci: 6 (1860 ['1859']).Type: Society Islands, Tahiti, 1838-1842, Wilkes Expedition s.n. (BM!-isotypes). Calymperes orientate Mitt, ex Besch. in Ann. Sci. Nat. Bot. sdr. 8, 1: 272, 296 (1895). Type: Borneo, Labuan, Motley s.n. (BM?, NY? - not found) Calymperes orientate var. polytrichoides M. Fleisch., Muse. Fl. Buitenzorg 1: 249 (1904). Type: Salak, bei Succamandri, 500 m, Fleischer s.n. (L [?] - holotype, fide Reese & Mohamed, 1985). Shoots reaching 2->6 cm high, dull green, forming tufts. Leaves mostly 6-7.5(-9) mm long, not dimorphic, a suberect, broadly subelliptical hyaline base with an erect to spreading (moist), linear- lingulate chlorophyllose limb ending in a linear proboscis (Fig. lOe). Costa extending into proboscis, ending immediately below apex, smooth; internally with two rows of guide cells (Fig. lOj). Chlorophyllose lamina abruptly narrowing into proboscis and be- coming narrowly recurved, distally becoming plane to form a blunt, dentate leaf apex (sometimes eroded away); cells mostly isodiametric, sometimes wider than long, with 4-6 sides or rounded, 5-12 x 5-7.5 Urn (Fig. lOg), ventrally roundly to subacutely protuberant, dorsally smooth (Fig. 101). Hyaline lamina sharply defined. Leaf margin from above hyaline base to near base of proboscis consisting of a thick polystratose rib (internally lacking stereids) with regular, often paired, multicellular teeth; in surface view cells isodiametric, subquadrate to irregularly rounded (Fig. lOh, 1); from proximal limit of marginal rib to leaf base with a narrow unistratose to polystratose intramarginal band of linear, thick-walled cells (continuous with marginal rib of upper leaf); marginal lamina shortly below apex of hyaline lamina unistratose, denticulate, consisting of small, thick- walled, chlorophyllose cells in about 8-14 rows; in mid to proximal hyaline base entire, unistratose, consisting of 2-5 rows of subquadrate to shortly subrectangular hyaline cells (Fig. lOf). Gemmae produced from the ventral surface of the costal tip. HABITAT. On trunks and exposed roots of trees and on rock; common in damp, shaded situations in lowland forest. DISTRIBUTION. A palaeotropical species reaching the Pacific is- land groups. Widespread in the Philippines. SPECIMENS EXAMINED. Leyte Island, Baybay, Mt Pangasugan, nearVISCA campus, 21-22 May 1984, Tan, Navarez & Raws 84-216 (BM). Luzon, Laguna, Mt Banahao, 5-7 March 1910, Robinson 9805 (BM); Oriental Mindoro, Sumagit Town, Bo. Magod, vicinity of Mt Wood along River Sapadao, 1 1-13 July 1983, Tan 83-123 (FH). Mindanao, Seno de Davao, 15 May 1890, Micholitz 5 (BM, BM-K). Palawan, western border of St Paul Subterranean National Park, 26 May 1989, Tan 89-1507 (BM); Puerto Princesa, Sitio Kalabayog, near Batac Village, 23-25 April 1993, Tan 93-220 (FH); Barangay San Rafael, trail to Batac Village, 1 May 1991, Tan 91-268 (BM); Barangay Apurawan, Sitio Daan, vicinity of Mt Tinik-basan, 27 April 1992, Tan 92-239 (BM); Tan 92-243 (BM); Aborlan, Sitio Daan, Barangay Aporawan, 27 April 1992, Tan 92-268 (FH). Panay Island, Capiz province, Libacao, May-June 1919, Martelino & Edano 35781 (BM). Polillo Island, August 1909, Robinson 9281 (BM). All well-developed leaves in specimens of Calymperes taitense possess a linear, apical proboscis (formed by the costa with narrow, recurved wings of chlorophyllose lamina). Other species of Calymperes and Syrrhopodon in the Philippines either possess dimorphic leaves (in which only some produce a special, gemma- bearing apex), or leaves without apical modification of this kind. Calymperes tenerum Mull. Hal. in Linnaea 37: 174 (1872). Type: India, Bengal, Calcutta, Kurz s.n.(BMI-isotype). Fig. lOm-q. Shoots reaching 1 cm high, forming green mats. Leaves often homomallously curled when dry, up to 3 mm long, hardly dimor- phic, obovate to lingulate with distal chlorophyllose lamina sometimes involute, leaf apex broadly acute (Fig. 10m). Costa usually excurrent, sometimes ending in apex, in distal leaf superfi- cial cells often forming rounded to acute projections. Cells of chlorophyllose lamina mostly 7-12.5x7-1 0(- 12.5) |im, isodiametric with 4 to 6 sides or rounded (Fig. lOo), ventrally drawn out as acute projections, dorsally flat or unipapillose (Fig. lOq). Hyaline lamina usually sharply defined, enclosed on either side by broad, unistratose, marginal bands of small quadrate to shortly rectangular cells with differentially thickened walls and/or angles (Fig. lOn). Leaf margins entire, beyond hyaline lamina sometimes unistratose but usually formed by a narrow polystratose rib of small isodiametric chlorophyllose cells, lacking stereids (Fig. 1 Oq). Gemmiferous leaves with excurrent costa; gemmae produced from all around the costal tip. HABITAT. On living and fallen trees and exposed tree roots, some- times on rock; occurring in lowland areas. DISTRIBUTION. A pan tropical species. Widespread in the Philip- pines. SPECIMENS EXAMINED. Lumbacan Island, October 1906, Elmer 5279 (BM). Luzon, Laguna, UPLB campus, 20 July 1985, Tans.n. (FH).Palawan, Barangay Apurawan, Sitio Daan, vicinity of Mt Tinikbasan, 27 April 1992, Tan 92-258 (FH); El Nido, 5 January 1992, Tan 92-354 (FH); Port Barton, 1992, Tan 92-315 (FH). Iwahig, Balsahan, 23 April 1993, Tan 93-188 pro parte (FH). Culion Island, Culion, Sister Convent Garden, 8 May 1992, Tan 92-347 (BM). Ursula Island, Barangay Rio Tuba, 29 April 1993, Tan 93- 257 (FH). The hyaline base in the leaves of Calymperes tenerum and C. motley i are almost identical. For a comparison of these species see under description of the latter. Chameleion L.T. Ellis & A. Eddy in A. Eddy, Handbook ofMales- ian Mosses 2: 250 ( 1 990). Type species: Chameleion cryptocarpos (Dozy & Molk.) L.T. Ellis & A. Eddy. Syrrhopodon section Heliconema Mitt, in J. Linn. Soc. Bot. 12: 112 (1869). Heliconema (Mitt.) L.T. Ellis & A. Eddy in L.T. Ellis in/ Bryol. 15: 728 (1989), horn, illeg. Shoots erect, simple or branched, often matted with rhizoids, form- ing mats or tufts. Leaves sometimes dimorphic (gemmiferous and nongemmiferous leaves), mostly lingulate, ligulate to oblong-lan- ceolate, consisting of hyaline, or partly hyaline base that usually narrows slightly into a chlorophyllose limb, apices truncate or rounded to broadly subacute, sometimes apiculate. Costa ending below apex, sometimes with short lamellae on ventral surface. Cells of chlorophyllose lamina small, mostly isodiametric, usually ven- trally protuberant, smooth or papillose. Cells of hyaline lamina large, empty, smooth, porose, walls often with transverse bands of thickening; border between hyaline and chlorophyllose lamina often poorly defined. Marginal and/or intramarginal ribs frequently present, marginal ribs in chlorophyllose limb sometimes with short lamellae. Gemmae produced, often in radial groups, sometimes from the apices of modified leaves. Perichaetia terminal. Seta very short. Capsule urn-shaped, immersed among the upper leaves of the shoot. Peristome syrrhopodontoid or absent. The name, Chameleion, was proposed by Ellis & Eddy in Eddy (1990) who also presented a short history of this segregate genus. As CALYMPERACEAE IN THE PHILIPPINES 21 a genus, the group resembles many small species of Calymperes in features of the gametophyte and Syrrhopodon in details of the sporophyte. There are three species recognized in this small tropical genus, only one of which, Chameleion peguense, has been recorded in the Philippines. This species is probably now extinct here as it has not been refound since its original collection from Mt Lumutan, Rizal Province in 1917. The mountain vegetation has subsequently be- come largely secondary forest, and grassland oflmperata cylindrica (L.) Ra'usch. The status of the genus Chameleion requires further research. The species included within the genus (C. cryptocarpos (Dozy & Molk) L.T. Ellis & A. Eddy, C. peguense and C. xanthophyllus (Mitt.) L.T. Ellis & A. Eddy) are united by the possession of capsules on very short setae that remain immersed among the leaves of the gametophyte. Chameleion xanthophyllus and C. cryptocarpos occur in tropical America and C. peguense has an Indo-Malesian distribu- tion. The leaves in the species of Chameleion are similar in form, but those of C. xanthophyllus lack some features shared by the leaves of C. peguense and C. cryptocarpos. For example, leaves in the latter two species possess hyaline cells in the basal lamina with unevenly thickened walls and series of large, subrectangular, transversely elongate, superficial pores; some cells on the ventral surface of the costa and those of the marginal rib in the chlorophyllose limb form short lamellae (only evident in well-developed specimens of C. peguense"). These features are absent from the leaves of C. xanthophyllus. Reese (1993) points out similarities between the leaves of C. xanthophyllus and Syrrhopodon rigidus Hook. & Grev. and some allied species (a group with exserted capsules, well accepted as belonging to the genus Syrrhopodon). He suggests that immersed capsules may have arisen more than once within the Calymperaceae and, therefore, undermines one of the principal justifications for recognizing the genus Chameleion. Similarities in the leaves of the species of Chameleion are said to be superficial. However, which features of these species of Syrrhopodon and Chameleion reflect their interrelationships requires greater clarification; a detailed cladistic analysis may help to resolve the problem. Presently, Chameleion appears to be worthy of retention. Chameleion peguense (Besch.) L.T. Ellis & A. Eddy in A. Eddy, Handbook ofMalesian Mosses 2: 250 (1990). Fig. 11. Calymperes peguense Besch. in Ann. Sci. Nat. Bot. ser. 8, 1: 269, 299 (1895). Type: Burma, Pegu, Yomah, Kurz 2928b (BM!-holotype). Calymperes ramosii Broth, in Philipp. J. Sci. 31: 281 (1926). Type: Philippines, Luzon, Province of Rizal, Mt Lumutan, July 1917, Ramos & Edano 29825 (BM!-isotypes). Heliconema peguense L.T. Ellis & A. Eddy in J. Bryol. 15: 730 (1989), horn, illeg. Syrrhopodon peguense (Besch.) W.D. Reese in /. Hattori Bot. Lab. 82: 243 (1997). Shoots 0.5->1.5 cm high, densely matted with rhizoids below, forming mats or tufts. Leaves strongly dimorphic: nongemmiferous leaves erect to patent (sometimes recurved), mostly 3-4 mm long, narrowly to broadly Ungulate, distal lamina often incurved; apex broadly obtuse to rounded or rounded-truncate, entire (Fig. lla). Costa ending just below apex; on the ventral surface near costal apex sometimes with one or two low lamellae (2-3 cells high); ventral superficial cells above the hyaline base subquadrate to shortly rectangular, smooth to roundly or subacutely protuberant, dorsal superficial cells similar but mostly longer relative to width and often with a shallow, simple papilla near end walls; internally with a single row of guide cells. Cells of chlorophyllose lamina isodiametric to slightly longer than broad, with 4-6 sides or rounded-elliptical, 5- 12.5(-15) x 5-10 urn (Fig. llf); ventrally roundly protuberant, dorsally flat to slightly convex or unipapillose (Fig. 1 Ih). Hyaline lamina often poorly defined, adjacent to costa sometimes incorpo- rating thick-walled, brownish yellow cells; largely composed of fragile hyaline cells, superficial walls with transverse bands of thickening (areas between thickened bands sometimes becoming pores) (Fig. 1 le). Leaf margins above hyaline base entire to irregu- larly and distantly toothed (teeth blunt), sometimes unistratose but usually incorporating an intramarginal polystratose rib with superfi- cial cells in surface view subquadrate to shortly subrectangular and often with an internal strand of stereids, dorsal surface of rib sometimes with one or two entire or distantly and bluntly toothed lamellae (1-3 cells high), marginal cells similar to laminal cells, forming a unistratose band 2-3 cells wide; in region around distal limit of hyaline lamina unistratose, entire to denticulate; in hyaline base incorporating an intramarginal, unistratose band of long rectan- gular to linear thick-walled cells (up to about 7 cells wide); marginal cells hyaline, shortly subrectangular, forming a single row (entire below, distally entire to denticulate). Gemmiferous leaves linear with a funnel-shaped apex, stiffly erect (curved when dry), exserted well above nongemmiferous leaves, c. 5 mm long (Fig. 1 \b, c). Costa strong (occupying over a third of the leaf width), ending below apex, distally rough with many superficial cells drawn out as acute projections; internally with 1-3 layers of guide cells between dorsal and ventral bands of substereids (Fig. 1 !/)• Chlorophyllose lamina usually involute, at apex forming a lax funnel-shaped collar above the tip of the costa; cells subquadrate to long subrectangular; dorsally and ventrally convex or drawn out as acute projections. Hyaline lamina narrow with cells similar to those in nongemmiferous leaves. Leaf margins above hyaline base similar to those in nongemmiferous leaves but lacking lamellae (some superficial cells of intramarginal polystratose rib acutely drawn out); in hyaline base intramarginal band of linear cells often obscure or absent. Gemmae produced in a radial mass from the ventral surface of the costal apex (within the 'laminal collar'). Perichaetia terminal. Seta >0.5 mm long. Capsule urn-shaped, >1 mm long. Peristome syrrhopodontoid with 16 papillose teeth. HABITAT. On trunks, branches and exposed roots of trees; occur- ring from near sea level to over 600 m. DISTRIBUTION. Disjunct from India, Indochina and Luzon Island in the Philippines. No other local material examined. Plants of this species are similar to those of some small species of Calymperes with dimorphic leaves (e.g. C. boulayi, C. subintegrum), but the leaves possess lamellae on the costa and leaf margins (in well-developed specimens), and the cells of the hyaline lamina have transverse bands of thickening. Sporophytes are very rare. Mitthyridium H. Rob. in Phytologia 32: 432 (1975). Type species: Mitthyridiumfasciculatum (Hook.& Grev.) H. Rob. Thyridium Mitt, in /. Linn. Soc. Bot. 10: 188 (1869), horn, illeg. Plants forming mats, primary shoots creeping, often with short ascending secondary branches. Leaves when dry variously curled, when moist mostly spreading; consisting of a suberect, often 22 L.T. ELLIS AND B.C. TAN a,b d-i 1 mm 0.5 mm 50 jLim Fig. 11 a-h. Chameleion peguense (Besch.) L.T. Ellis & A. Eddy a: nongemmiferous leaves in ventral view; b, c: gemmiferous leaf (b: in lateral view, with c: detail of gemmiferous apex); d-f: cells of leaf in surface view (d: at margin in hyaline base, e: in hyaline lamina, f: in chlorophyllose lamina); h: cross-sections of nongemmiferous leaf (g: costa, h: chlorophyllose lamina); i: cross-section of gemmiferous leaf through chlorophyllose limb, a-i Drawn from Ramos & Edano 29825 (BM). semiclasping hyaline base narrowing (sometimes abruptly) into a Ungulate or lanceolate to triangular-lanceolate chlorophyllose limb. Costa ending below apex, usually smooth, sometimes with small, sparse teeth; internally composed of dorsal and ventral bands of stereids with a single, median layer of guide cells, superficial cells sometimes differentiated. Cells of chlorophyllose lamina small, thick-walled, mostly isodiametric, polygonal to rounded or substellate, not usually protuberant, often multipapillose. Hyaline lamina well defined. Leaf margin mostly with a unistratose marginal rib of linear thick-walled cells extending from the base into the upper leaf. Calymperoid gemmae produced near and/or at apices of leaves (apices strongly modified in some species). Sporophytes terminal; seta smooth; capsule exserted, erect, cylindrical. Calyptra rostrate, fugacious. Peristome syrrhopodontoid. Mitthyridium is a small genus most closely related toSyrrhopodon, and is easily recognized by its creeping primary stems and broad leaf borders (in most species the leaf border is unistratose, an exception is the Indonesian species Mitthyridium retusum (Besch.) W.D. Reese in which the marginal rib is polystratose). The distribution of Mitthyridium is almost exclusively palaeotropical with only a single historical collection of a wide-ranging species recorded from South America (Reese, Mohamed & Mohamed, 1 986). Mitthyridium occurs exclusively at low altitudes. The species of Mitthyridium are highly variable and consequently have been differently interpreted by various authors. In this treatment, we have accepted a number of infraspecific taxa which reflect the morphological variations observed in specimens from the Philippines. There are 12-13 species of Mitthyridium reported from Malesia. Eight are described below for the Philippine archipelago, of which one, M. iwatsukianum B.C. Tan is a local endemic. Mitthyridium constrictum (Sull.) H. Rob. in Phytologia 32: 432 (1975). Fig. 12a-c. Calymperes constrictum Sull., U. S. Exploring Expedition. Musci: 6 (1860 ['1859'] ). Type: Hawaii ['Sandwich Island'] Wilkes Expe- dition s.n. (FH-lectotype). Shoots reaching c. 6 cm long, sparsely branched, densely leaved, forming stringy tufts and mats. Leaves sometimes in lax ranks, mostly 2.5-<3.0 mm long, subobovate with upper lamina involute, lower leaf suberect and clasping, upper leaf curled when dry, erect to spreading when moist; apex modified as a loosely tubular to funnel- shaped proboscis (tip mostly truncate) (Fig. 12a). Costa ending below leaf apex; dorsal surface near leaf apex scabrid with spinose teeth, otherwise smooth, composed of stereids/linear cells. Cells of chlorophyllose lamina 10-15(-17.5) x 7.5->12.5 |im, with very thick walls (abutting cell walls sometimes as broad as lumina), longer than broad to broader than long, irregularly polygonal to substellate, multipapillose below (papillae mostly thick and simple, rarely obscure) (Fig. 12b); towards leaf apex on dorsal leaf surface usually becoming unipapillose (papillae prominent, sometimes branched). Hyaline lamina sharply defined, occupying just less than a third of the leaf length. Leaf margins irregularly notched to CALYMPERACEAE IN THE PHILIPPINES 23 a a, d, f, h, k, 1, n 1 mm b, c, e, g, i,j, m, o 50 urn Fig. 12 a-c. Mitthyridium constriction (Sull.) H. Rob. a: leaf; b: cells of chlorophyllose lamina; c: cross-section of costa and chlorophyllose lamina, d, e. Mitthyridium flavum (Miill. Hal.) H. Rob. d: leaf; e: cells of chlorophyllose lamina, f, g. Mitthyridium cf.flavum! f: leaves; g: chlorophyllose lamina, h-j. Mitthyridium iwatsukianum B.C. Tan h: leaf in lateral view; i: cells of chlorophyllose lamina; j: cross-section of costa and chlorophyllose lamina, k-m. Mitthyridium junquilianum (Mitt.) H. Rob. k: leaf in ventral view, with 1: detail of distal chlorophyllose limb; m: cells of chlorophyllose lamina, n, o. Mitthyridium subluteum (Miill. Hal) H. Novak n: leaf in ventral view; o: cells of chlorophyllose lamina, a Drawn from Tahiti, Whittier 235 1 (BM). b, c Drawn from Louisades, Micholitz s.n. (BM). d, e Drawn from Tan 87-20 (FH). f, g Drawn from Sabah, Ellis 95-289 (BM). h-j Drawn from Tan & Tandang 82-371 (BM). k-m Drawn from Borneo, Motley s.n. (BM). n, o Drawn from Tan & Hernaez 87^49 (BM). denticulate from a short distance above leaf base to near apex; marginal rib very broad, 100-250(^00) |im wide at broadest point (adjacent to or slightly above distal hyaline lamina). Gemmae produced from ventral surface of costal apex (enfolded in incurled lamina of proboscis). HABITAT. On tree trunks and branches in lowland forest. DISTRIBUTION. An Indo-Pacific species. Uncommon in the Philip- pines. SPECIMEN EXAMINED. Luzon, Tayabas, Mt Binuang, May 1917, Ramos & Edano 28941 (FH) (duplicate in BM is M . fasciculatum). Mitthyridium constriction is superficially very similar to M. iwatsukianum B.C. Tan. In M. iwatsukianum the leaves lie in three very neat ranks along the stem and the cells of the chlorophyllose lamina are subquadrate to irregularly polygonal, smooth, with nar- row, evenly thickened walls. M. constrictum possesses leaves that lie in lax, untidy ranks and the cells of the chlorophyllose lamina are papillose with uneven walls which are often almost as broad as the lumina they define. 24 L.T. ELLIS AND B.C. TAN a, e, f b,g c,d 1 mm 0.5 mm 50 Fig. 13 a-d. Mitthyridiumfasciculatum (Hook. & Grev.) H. Rob. a, b: leaf ( a: in ventral view, with b: detail of apex); c, d: cells of leaf in surface view (c: in chlorophyllose lamina, d; in ventral surface of costa). e. M. fasciculatum subsp. cardotii (M. Fleisch.) B.C. Tan & L.T. Ellis e: leaf, f, g. M. fasciculatum subsp. obtusifolium (Lindb.) M. Menzel f, g: leaf (f: in ventral view, with g: detail of apex), a-d Drawn from Java, Eddy s.n. (BM). e Drawn from Tan, Navarez & Rams 84-218 (FH). f, g Drawn from Tahiti, Whittier 2172 (BM). Mitthyridium fasciculatum (Hook. & Grev.) H. Rob. \nPhytologla 32:43(1975). Mitthyridium fasciculatum (Hook. & Grev.) H. Rob. subsp. fasciculatum Fig. 13a-d. Syrrhopodon fasciculatus Hook. & Grev. in Edinburgh J. Sci. 3: 225 (1825). Type: Ternate Island, [C. Smith] s.n. (Hb. Dickson) (BMMectotype, fide Menzel & Schultze-Motel (1990)). Thy ridium fasciculatum (Hook. & Grev.) Mitt, in J. Linn. Soc. Bot. 10: 189(1868). Syrrhopodon leucoloma Mull. Hal. in Bot. Jahresber (Just.) 5: 86 (1883). Type: New Guinea, September 1875, Naumann s.n. (BM!-isotype). Syrrhopodon codonoblepharoides Mull. Hal. ex M. Fleisch., Muse. Fl. Buitenzorg 1: 228 (1904), nom. nud. Original material ?: Philippines, Cuming 2199 (in Hb. Hampe, BM!). Mitthyridium leucoloma (Mull. Hal.) H. Rob. inPhytologia 32: 433 (1975). Primary shoots reaching >8 cm, occasional ascending branches reaching >3 cm long, forming coarse stringy tufts and mats, densely leaved. Leaves 3.5-5 mm long, consisting of a subobovate, semisheathing hyaline base narrowing abruptly into a broadly lan- ceolate to Ungulate chlorophyllose limb (curled when dry, spreading when moist); leaf apex subacute to acute (Fig. 13a, b). Costa ending below leaf apex; dorsal surface composed of long rectangular to linear cells (some projecting distally); ventral surface above hyaline base composed of a (sometimes broken) layer of subquadrate to shortly subrectangular multipapillose cells, in hyaline base cells becoming slightly larger, smooth, thin-walled and hyaline. Chlorophyllose lamina strongly undulate; cells in surface view <5- 12 x <5-10 urn, isodiametric, subquadrate to irregularly polygonal or appearing to consist of 2-4 rounded lobes, multipapillose (papil- lae appearing to overlie lumina) (Fig. 13c). Hyaline lamina sharply defined with a broadly-pointed, almost truncate apex. Leaf margins irregularly denticulate (and papillose) from above mid-hyaline base to leaf apex; marginal rib extending from leaf base to near apex, broadest adjacent to distal hyaline lamina (c. 15->20 cells wide). Gemmae produced from near and at apex of costa. HABITAT. Generally a lowland species. In the Philippines Mitthyridiumfasciculatum has been collected from tree trunks and logs between 400 m and 800 m. CALYMPERACEAE IN THE PHILIPPINES 25 DISTRIBUTION. A palaeotropical species with doubtful reports from tropical America. Widespread in the Philippines. SPECIMENS EXAMINED. Philippines, Cuming s.n. (BM). Luzon, Mountain Province: June 1952, Mack 1395 (FH); Benguet, Mt Data, 26 March 1938, Santos 961 (BM, FH); Quezon Province, Atimonan: Quezon National Park, 9 March 1986, Tan & Lipaygo s.n. (FH); Malinao, 26 October 1937, Santos 857 (FH); Tayabas: Alabat, Mt Camagong, 22 October 1937, Santos 806 (FH); Mt Binuang, May 1917, Ramos & Edano 28941 (BM) (duplicate in FH is M. constrictum). Mindanao, Lanao, Dansalan, 29 September 1938, Zwickey 225b (FH). Mindoro Island, Lake Naujan, 15 April 1935, Bartlett 13561 (FH). Palawan, Puerto Princesa, Barangay Irawan, Mt Malinao, 4 May 1 993, Tan 93-302 (FH); Barangay Napsan, SitioTagkuit, Salakot waterfall vicinity, 2 May 1993, Tan 93-264 (FH). Nowak (1980), Eddy (1990), and Reese & Stone (1995) recognized Mitthyridium leucoloma as a distinct species. Menzel & Schultze- Motel (1990) placedM. leucoloma in synonymy wilhM. fasciculatum while Reese, Koponen & Morris (1986) placed it in synonymy with M. obtusifolium (Lindb.) M. Menzel. Reese & Stone (1995) distin- guish M. leucoloma from M. fasciculatum, and other taxa, by the possession of leaves with entire margins (sometimes weakly den- ticulate) and with the upper lamina more or less involute; there is a 'lack of flaring at the leaf shoulders' , and the cells at the apex of the hyaline lamina 'interfinger' with the proximal cells of the chlorophyllose lamina. In M. fasciculatum the leaves possess den- ticulate margins, an upper lamina not notably involute, flaring shoulders, and cells in the apex of the hyaline lamina not notably interfingering with the proximal cells of the chlorophyllose lamina. However, these apparently contrasting features may represent ex- tremes in a range of variation. In the isotype specimen of M. leucoloma (Naumann s.n., BM) there are many leaves with well- devoloped denticulate margins, an upper lamina not notably involute, and with somewhat flaring shoulders. A range of specimens, includ- ing the type of M. leucoloma, differ to varying degrees from the typical form of M. fasciculatum in the features outlined above. Whether these form a group distinct at the level of species remains equivocal. Nowak (1980) identifies two collections from the Philippines (one from Mindanao, the other unlocalized) as Mitthyridium leucoloma. Mitthyridium fasciculatum subsp. cardotii (M. Fleisch.) B.C. Tan & L.T. Ellis, stat. nov. Fig. 13e. Thyridium cardotii M. Fleisch., Muse. Fl. Buitenzorg 1: 228 (1904). Type: Java, Buitenzorg Botanical Garden, January 1 899, Fleischer (Muse. Frond. Archip. Indict Ser. II, exs. no. 73 (BM!-isotypes). Mitthyridium cardotii (M. Fleisch.) H. Rob. in Phytologia 32: 432 (1975). Mitthyridium fasciculatum var. cardotii (M. Fleisch.) A. Eddy, Handbook ofMalesian Mosses 2: 135 (1990). Differs from the type subspecies in having smaller shoots ( <4 cm long), branches (<1 cm long) and leaves (up to c. 2.5 mm long) with obtuse apices (Fig. 13e). HABITAT. On trunks of trees in lowland forest. DISTRIBUTION. Largely an Indo-Malesian subspecies, but appar- ently occurring as far west as the Seychelles. In addition to the localities in the Philippines cited below, this subspecies has also been reported from Leyte and Negros by Iwatsuki & Tan (1980). SPECIMENS EXAMINED. Leyte Island, Baybay, Mt Pangasugan, nearVISCA campus, 21-22 May 1984, Tan, Navarez & Rams 84-218 (FH). Luzon, Isabela, San Mariano, Barrio Disulap, Sierra Madre range, Dimahahabong Creek, 14 April 1991, 7a/z91-119 (BM). Reese, Koponen & Morris (1986) and Menzel & Schultze-Motel (1990) regard subspecies cardotii as a mere form of the type subspecies. However, Eddy (1990) proposed subspecies cardotii as a formal variety of Mitthyridium fasciculatum. The research of G. Jakab (pers. comm.) indicates that subspecies cardotii is not con- fined to Indo-Malesia, but also occurs in the Seychelles, where, hitherto, it has been treated as a distinct species, Mitthyridium micro-undulatum (Dixon) H. Rob. Mitthyridium fasciculatum subsp. obtusifolium (Lindb.) M. Menzel in M. Menzel & W. Schultze-Motel in Willdenowia 19(2): 502 (1990). Fig. 13f,g. Syrrhopodon obtusifolium Lindb. in Ofvers.Forh. Kongl. Vetensk.- Acad. 21: 605 (1865) ['1864']. Type: Tahiti, September 1852, Ponten s.n. (BMI-lectotype, BM!-isolectotype (fide Nowak (1980)). Thyridium obtusifolium (Lindb.) M. Fleisch., Muse. Fl. Buitenzorg 1: 228 (1904). Mitthyridium obtusifolium (Lindb.) H. Rob. in Phytologia 32: 434 (1975). Specimens of subsp. obtusifolium differ from those of the type subspecies in the possession of leaves with an obtuse apex (Fig. 13f, g), the rows of laminal cells forming the apical margin smooth and thicker walled than those below; the apex of the hyaline lamina tending to be truncate (rather than broadly acute) and the marginal rib generally broader. A collection from Luzon, Tan & Lipaygo s.n. (cited above under subsp. fasciculatum), possesses leaves with somewhat obtuse apices that approach halfway the characteristic form found in subsp. obtusifolium. In all other features, the leaves of this specimen are identical to those of subsp. fasciculatum. Un- equivocal evidence for the presence of this subspecies in the Philippines has yet to be discovered. Mitthyridium flavum (Mull. Hal.) H. Rob. in Phytologia 32: 433 (1975). Fig. 12d-g. Syrrhopodon flavus Mull. Hal. in Bot. Zeitung (Berlin) 13: 763 (1855). Type: Java, s.n. (B?-holotype, presumably destroyed 1943; isotypes not found). Thyridium flavum (Mull. Hal.) M. Fleisch., Muse. Fl. Buitenzorg 1: 232 (1904). Syrrhopodon luzonensis R.S. Williams in Bull. New York Bot. Card. 8: 338 (1914). Type: Philippines, Luzon, Lamao River, 90 m, March 1904, Williams 824 (FH!-isosyntype) . Thyridium luzonensis (R.S. Williams) Broth., Nat. Pflanzenfam. 2nd ed., 10:236(1924). Primary shoots prostrate, sometimes dense with rhizoids, <0.5->2.0 cm long, unbranched to dense with very short ascending branches. Leaves mostly 1.5->2 mm long, erect to spreading from a semi- sheathing hyaline base, lingulate, usually parallel-sided and abruptly narrowing at the apex to form a shortly pointed tip (Fig. 12d), sometimes gradually narrowing from hyaline base. Costa ending below apex, surface composed largely of stereids, some occasion- ally projecting at their distal ends to form minute teeth; lateral superficial cells sometimes shortly rectangular (at leaf apex some- times producing gemmae). Cells of chlorophyllose lamina in surface view 5-10 x 5-12.5 urn, slightly longer than broad to slightly 26 L.T. ELLIS AND B.C. TAN broader than long, irregularly polygonal or rounded, each with 1-4 simple papillae on the dorsal and ventral surfaces (papillae some- times obscure) (Fig. 12e). Hyaline lamina sharply defined, often with an acute apex. Leaf margins plane to undulate, with a narrow, unistratose rib of linear cells extending from the leaf base to distal leaf limb, 2-8 cells wide (broadest adjacent to distal hyaline lamina), sinuose to denticulate, rarely entire; margins toward leaf apex formed by cells of chlorophyllose lamina, entire to denticulate. HABITAT. On shaded tree trunks in forest. DISTRIBUTION. An Indo-Pacific species. Widespread in the Philip- pines. SPECIMENS EXAMINED. Leyte Island, Baybay, Mt Pangasugan nearVISCA campus, 21-22 May 1984, Tan, Navarez & Rams 84-217 (FH). Luzon, Bataan, Olongapo Naval Reservation, 25 May 1935, Bartlett 14114 (FH); Cagayan, hills at Sitio Babayuan, 30 October 1935, Bartlett 14926 (FH); Bartlett 14931 (FH); Isabela, Palanan Wilderness, 19 May 1992, Tan 92-205 (BM); Zambales, Candelaria, Bo.Tapaso, Mt Lanat, 15-16 March 1987, Tan 87-20 (FH). Mindoro Island, Mt Bulalacao, November 1939, Ebalo 231 (FH). Palawan, Mt Mantalingahan, between Nalpuan and Sandurapi, 26 April 1991, Tan 91-186 (BM); Aborlan, Barangay Apurawan, trail to Sitio Daan, 28 April 1992, Tan 92-297 (FH); Taytay, Lake Manguao (Lk Danao), 1 May 1992, Tan 92-311 (BM); Puerto Princesa, Barangay Irawan, Mt Malinao, 4 May 1993, Tan 93-316 (FH); Puerto Princesa, Sitio Kalabayog, near Batac Village, 23, 25 April 1993, Tan 93-221 (FH). Sibuyan Island, Magdiwang, BoTampayan, Mt Giting-Giting, 21 May 1987, Tan & Hernaez 87^55 (FH). The holotype of Mitthyridium flavum was presumably destroyed along with the major part of Muller's herbarium (B) in 1943. Should the apparent absence of isotype material be real the species may require neotypification. Miiller (1856), in his review of Dozy and Molkenboer's Bryologia Javanica, provides a strong indication of his concept ofM.flavum. Referring to the illustrations ofSyrrhopodon tenellum Dozy & Molk. he proposes that the latter species is conspecific with his earlier described Syrrhopodon flavus. These illustrations, and relevant material derived from Dozy and Molkenboer's herbarium (now in Hb. Hampe (BM) and Hb. Schimper (BM-K)), show features that agree with the concept ofM.flavum as presently understood (Eddy, 1990; Menzel & Schultze-Motel, 1990; Reese, Koponen & Norris, 1986). Most specimens can be identified as Mitthyridium flavum by their short leaves (>2 mm long) with parallel sides. The leaves in M. repens (Harv.) H. Rob. also have parallel sides, but are usually much smaller (1-1.5 mm long) and possess rounded-apiculate apices, not evident in the leaves of M. flavum (Fig. 14b). Mitthyridium iwatsukianum B.C. Tan in Mem. New York Bot. Card. 45: 453 (1987). Type: Philippines, Laguna, Cavinti Town, Bo. Lumot, near Sitio Ubali, 500-800 m, 24 October 1982, Tan & Tandang 82-371 (BM'.-isotype). Fig. 12h-j. Shoots reaching > 4 cm long, with sparse branches up to about 1 cm long (shoots and branches with a flattened appearance). Leaves strictly aligned in three ranks, mostly 2.5-3 mm long, subobovate, subrecurved from a clasping base, folded along costa; apex modified as a loosely tubular to funnel-shaped proboscis (Fig. 12h). Costa ending below leaf apex, smooth (dorsal surface of the costal apex lacking spinose teeth). Chlorophyllose lamina smooth; cells small, mostly 5-12.5(-15) x 5-12.5(-15) um, subquadrate to irregularly polygonal (never substellate), broader than long to longer than broad, with relatively narrow and evenly thickened walls (Fig. 12i). Hyaline lamina sharply defined, occupying just less than one third of the leaf length. Leaf margins irregularly notched to minutely den- ticulate from a short distance above the leaf base to near apex; marginal rib very broad, reaching 300-350 jam wide at its broadest point (near apex of hyaline lamina). Gemmae produced from ventral surface of costal apex (enfolded in incurled lamina of proboscis). HABITAT. On leaf sheath of Pinanga palm in lowland rainforest. DISTRIBUTION. Philippine endemic. Known only from the type collection. Eddy (1990) draws attention to the strong similarities between Mitthyridium iwatsukianum and M. constrictum. However, their differences, most strikingly those of the chlorophyllose lamina (Fig. 12b, i), presently justify the recog- nition of Mitthyridium iwatsukianum as a species. Similar diverse forms of chlorophyllose lamina occur as extremes of a range of variation in Mitthyridium wallisii; intermediates between the forms of lamina in M. iwatsukianum and M. constrictum have yet to be found. Mitthyridium papuanum (Broth.) H. Rob. in Phytologia 32: 434 (1975). Fig. 14i-o. Syrrhopodon papuanus Broth, in Oevfers. Forh. finska Vetensk.- Soc. 37: 156 (1895). Type: New Ireland, October 1893, Micholitz [94] (BM!-isolectotypes, fide Nowak (1980)). Thyridium papuanum (Broth.) M. Fleisch., Muse. Fl. Buitenzorg 1: 232(1904). Primary shoots reaching >4 cm long, with sparse ascending branches 3 cm long. Leaves 1-4 mm long, broadly lanceolate (gradually tapering from distal hyaline base to an acute apex) to lingulate (with parallel sides from base to near apex, at apex rapidly conracted into a subapiculate to apiculate, sometimes acuminate, point) (Fig. 14i, j); curved when dry, spreading when moist. Costa ending in apex; ventral and dorsal surfaces above midleaf largely formed by stereids, below midleaf formed by rectangular, thick- walled cells; on dorsal surface towards apex occasionally some cells giving rise to a small tooth from their distal ends. Chlorophyllose lamina undulate; cells mostly 5-12.5 x 5-10 um, quadrate to irregularly polygonal, or rounded, usually with evenly thickened walls, on dorsal and ventral surfaces with 1 or more simple papillae (papillae often obscure) (Fig. 14k). Leaf margin from shortly above leaf base to apex minutely and irregularly denticulate; marginal rib extending from leaf base, broadest adjacent to hyaline lamina, tapering to beyond midleaf (Fig. 141). Gemmae produced at leaf apex from shortly rectangular, lateral cells of costa. HABITAT. On tree trunks, mostly in lowland forest. DISTRIBUTION. A widespread Indo-Pacific species. SPECIMENS EXAMINED. Dalupiri Island, 31 October-5 November 1935, Bartlett 15843 (FH). Luzon, Mt Makiling, 27 August 1931, Herklots P28 (BM, FH); Isabela, Palanan Wilderness, 19 May 1992, Tan 91-222 (BM); 20 May 1992, Tan 92-213 (FH); Tayabas, Alabat, Mt Camagong, 21 October 1937, Santos 810 (FH). Reese, Koponen & Norris (1986) placed Mitthyridium papuanum in synonymy with M. luteum (Mitt.) H. Rob. and identified material from the Philippines as that species. Reese (1994) and Reese & Stone (1995) reinstated M. papuanum as a distinct species. The lectotype of Mitthyridium luteum, from Fiji (Milne 363, BM), is like a form of M. fasciculatum with relatively narrow, tapering leaves. The chlorophyllose lamina is composed of strongly papillose cells with incrassate walls. Many of the leaves in the type, and other specimens of M. papuanum, are parallel-sided and resemble giant forms of those occurring in Mitthyridium flavum. In M. papuanum, CALYMPERACEAE IN THE PHILIPPINES 27 the cells of the chlorophyllose lamina are relatively thin-walled and possess low papillae that, even in cross-section, are barely visible. Evidence from material presently available does not justify includ- ing M. papuanum in synonymy withM. luteum, and specimens from the Philippines, previously identified as M. luteum, are referable to M. papuanum. Mitthyridium luteum remains a good species based on Milne 363. Mitthyridium repens (Harv.) H. Rob. mPhytologia32: 434 (1975). Fig. 14a-h. Syrrhopodon repens Harv. in Hook., Icon. PL 1: t. 22, fig. 4 (1836); Harv. in Harv. & Hook, in J. Bot. (Hooker) 2: 7 (1840). Type: Peninsular Malaysia, Pinang ['Penang'], Wallich [H.I 204] (BM- K!-holotype?). Thyridium repens (Harv.) Mitt, in J. Linn. Soc. Bot. 10: 188 (1868). Primary shoots prostrate, forming mats, obscured by densely crowded, short, simple or innovating, ascending branches. Ascend- ing branches 1-7 mm high, densely leaved, sometimes densely matted with rhizoids below. Leaves <1-1.5 mm long, consisting of an erect, subrectangular hyaline base (slightly and gradually widen- ing from base to apex) narrowing slightly and gradually into an erect to spreading (moist), Ungulate chlorophyllose limb (tightly incurled when dry); leaf apex abruptly narrowing, broadly rounded or broadly rounded and subapiculate to apiculate, entire (Fig. 14a, b). Costa ending below leaf apex; surface near costal apex often composed of subrectangular cells (surface view), surface below apical region composed of stereids. Cells of chlorophyllose lamina in surface view 5-12.5(-15) x 5-12.5 Jim, longer than broad to broader than long, irregularly rounded to elliptical or appearing to consist of 2-4 (or more) rounded lobes; on dorsal and ventral surfaces each with several simple papillae (Fig. 14e, d). Hyaline lamina sharply defined with a somewhat truncate apex. Leaf margins plain to undulate (intermittently incurved); from below leaf apex to base consisting of a flattened band of stereids; from below leaf apex to around apex of hyaline lamina gradually broadening to c. 40-65 |im wide, regularly and sharply denticulate (Fig. 14c); below apex of hyaline lamina to leaf base slightly narrowing, entire to uneven. Gemmae produced towards and at leaf apex from ventral surface of costa. HABITAT. On trunks and buttresses of trees in lowland areas. DISTRIBUTION. An Indo-Pacific species. Philippine records are all from Luzon. SPECIMENS EXAMINED. Luzon, Quezon, Real, National Botanic Garden, 4 August 1985, Tan & Wijangco s.n. pro parte (BM); Bataan Province, Mt Mariveles, July 1904, Leiberg 1215 (FH). Mitthyridium repens has the smallest shoots of any species of Mitthyridium occurring in the Philippines. It often forms thin, dense mats resembling those of some diminutive species of Macromitrium and Schlotheimia in the family Orthotrichaceae. Mitthyridium subluteum (Mull. Hal.) H. Nowak in Bryophyt. Biblioth. 20: 144 (1980). Type:Samoa, Upolu, Graeffe s.n. (BM!- isolectotype). Fig. 12n-o. Codonoblepharum subluteum Mull. Hal. in J. Mus. Godeffroy 3(6): 67 (1874). Primary shoots prostrate, up to 2 cm long with sparse short branches. Leaves 3-3.5 mm long, lanceolate to linear-lanceolate with undulate margins (mostly 5-7 times longer than broad, slightly broadening from base to around distal extent of the short hyaline lamina, then tapering to an acute to narrowly acuminate apex) (Fig. 12n). Costa ending immediately below leaf apex; surface largely smooth (a few sparse teeth sometimes evident near leaf apex), above leaf base formed by stereids, in ventral leaf base formed by rectangular cells with mostly thin, porose walls. Chlorophyllose lamina occupying about six sevenths of leaf length, sometimes incurved above; cells 7.5-15 x 7.5-12.5 fim, irregularly polygonal, subelliptical or ap- pearing to consist of 2-4 rounded lobes; with walls unevenly thickened and several simple papillae (sometimes obscure) (Fig. 12o). Leaf margin denticulate from shortly above base to apex; marginal rib extending from base to near apex, often only 1-2 cells wide distally. Gemmae produced at leaf apex and from some dorsi- lateral cells of the costa for some distance below the leaf apex. HABITAT. In the Philippines collected from tree trunks up to 1100m. DISTRIBUTION. A Malesian species. Previously reported, but with no specimen citation, as occurring in Mindanao (Reese, Mohamed & Mohamed, 1986; Menzel & Schultze-Motel, 1990). SPECIMEN EXAMINED. Sibuyan Island, Magdiwang, Barangay Tampayan, Mt Giting-Giting, 21 May 1987, Tan & Hernaez 87-^49 (BM). The features of the collection from Sibuyan (Fig. 12n, o), cited above, accord especially well with those of the type of Mitthyridium subluteum. The latter has been widely accepted as representing a long-leaved (3-3.5 mm) form of M. junquilianum (Mitt.) H. Rob. in which the hyaline lamina often occupies a relatively small propor- tion (about one seventh) of the leaf length (Eddy, 1990; Menzel & Schultze-Motel, 1990; Reese, Koponen & Norris, 1986). Mohamed & Reese (1992) convincingly argue that M. junquilianum and M. subluteum are separate species. The present authors have not seen material from the Philippines that can safely be identified as M. junquilianum. The leaves of the isotype of M. junquilianum (Bor- neo, Motley s.n., BM) are relatively short (c. 2 mm) with the hyaline lamina often occupying about a quarter of the leaf length (Fig. 1 2k). Material recently collected in Sabah (e.g. Ellis 89-289, BM, UMS) possesses leaves that are not constant in form; some taper from around the apex of the hyaline base and closely resemble the leaves in the isotype of M. junquilianum (BM), whilst others (from the same shoot) are nearly parallel-sided and resemble the leaves of M. flavum (Fig. 12f). Consequently, this specimen is difficult to identify with certainty. The relationship between M. junquilianum and M. flavum requires further investigation. Mitthyridium wallisii (Mull. Hal.) H. Rob. in Phytologia 32: 435 (1975). Fig. 15. Syrrhopodon wallisii Mull. Hal. in Linnaea 38: 555 (1874). Type: Philippines, Luzon, Mahahai, 200 ft, 1870, Wallis s.n. (BM!- isotype; FH!-isotype). Shoots prostrate to ascending, in densely packed mats, with dense, ascending branches, mostly <0.5-1 cm long, densely leaved. Leaves mostly 1 .5-2.5 mm long, consisting of a semisheathing hyaline base narrowing into a Ungulate to narrowly triangular chlorophyllose limb (spreading when moist, incurled and crisped when dry), apex modified as a loosely tubular to narrowly funnel-shaped proboscis (often apiculate, sometimes emarginate) (Fig. 15a, b). Costa ending below apex (occasionally slightly excurrent as a mucro), surface formed by stereids, smooth (Fig. 15f). Cells of chlorophyllose lamina mostly 5-12.5 x 5-12.5 |im; longer than broad to broader than long, with irregularly polygonal to substellate lumina; thick- walled, smooth or papillose (papillae appearing to overlie cell walls) 28 L.T. ELLIS AND B.C. TAN m 9 k J g a, b 0.5 mm c-h, k-o 50 1 mm Fig. 14 a-h. Mitthyridium repens (Harv.) H. Rob. a, b: leaf (a: in ventral view, with b: detail of apex); c-e: cells of leaf in surface view (c: at margin above apex of hyaline lamina, d: of chlorophyllose lamina above apex of hyaline lamina, e: of distal chlorophyllose lamina); f-h: cross-sections of leaf (f: costa, g: chlorophyllose lamina, h: margin in chlorophyllose limb. i-o. Mitthyridium papuanum (Broth.) H. Rob. i, j: leaves (ventral view); k-m: cells of leaf in surface view (k: in chlorophyllose lamina, 1: of margin at mid-leaf, m: of margin near leaf apex); n, o: cross-sections of leaf (n: costa, o: chlorophyllose lamina), a-h Drawn from Tan & Wijangco s.n. pro parte (BM). i Drawn from Micholitz 94 (BM). j-o Drawn from Tan 91-222 (BM). (Fig. 15d, e). Leaf margin in hyaline base mainly entire; from distal hyaline base to near apex minutely and irregularly denticulate, in apical region entire to irregularly crenulate-denticulate; marginal rib extending from base of leaf to beyond midleaf, broadest adjacent to distal extent of hyaline lamina (c. 7-10 cells wide, <50 urn). Gemmae produced from ventral surface of costa at leaf apex (en- folded within incurled lamina of proboscis). HABITAT. On logs and tree trunks. DISTRIBUTION. A Malesian species, common in the Philippines. SPECIMENS EXAMINED. Luzon, Laguna, Cavinti Town, Sitio Calminoe, 25-26 February 1984, Tan & Trenbatt 84-50 (BM, FH); Quezon: Atimonan, 13 December 1980, Cadiz Fe Misa 12 (FH); Real, National Botanic Garden, 4 August 1985, Tan & Wijangco s.n. (FH); Tayabas, 17 September 1935, Pastrana 53 (FH); Guinayangan, February 1929, Doldulao 84421 (FH). Negros Island, Gimagon River, 6 January 1904, Copeland 73 (BM, FH); Dumaguete, November 1935, Chapman 41 (FH); May 1906, Whitford 1572 (BM). Panay Island, Capiz, October-November 1925, Edano 46254 (FH). Sibuyan Island, Magdiwang, Barangay Tampayan, Mt Giting-Giting, 21 May 1987, Tan & Hernaez 87^42 (BM). The cells of the chlorophyllose lamina m Mitthyridium wallisii vary widely in the thickness of their walls. At one extreme these cells possess relatively broad lumina and walls with unevenly thickened angles and sides (Fig. 15e); at the other extreme the walls appear broader than the lumina (Fig. 15d). All material of Mitthyridium wallisii from the Philippines, exam- ined for this study, is referable to the type variety. Mitthyridium wallisii var. crassum (Broth.) M. Menzel, with a largely Malesian distribution, possesses leaves reaching c. 4 mm long (approaching twice the length of most leaves seen in the type variety), which entirely lack the funnel-shaped apex present in some leaves in var. wallisii. Syrrhopodon Schwagr., Sp. muse, frond, suppl. 2: 110(1 824). Type species: Syrrhopodon gardneri (Hook.) Schwagr. Plants in tufts or mats, shoots erect, simple or branched, often CALYMPERACEAE IN THE PHILIPPINES 29 1 mm 0.5 mm c-g 50 Fig. 15 a-g. Mitthyridium wallisii (Mull. Hal.) H. Rob. a, b: leaf (a: in ventral view, with b: detail of gemmiferous apex; c-e: cells of leaf in surface view (c: at margin adjacent to apex of hyaline lamina, d, e: of chlorophyllose lamina); f,g: cross-sections of costa and chlorophyllose lamina, a-d, f Drawn from Wallis s.n. (BM). e, g Drawn from Tan & Wijangco s.n. (FH). densely matted with rhizoids below. Leaves when dry variously curled and twisted, when moist mostly erect to spreading or recurved; consisting of an erect to suberect, largely hyaline, semisheathing base extending (usually abruptly narrowing) into a ligulate, Ungulate or linear chlorophyllose limb. Costa ending just below apex to shortly excurrent, superficial cells linear to subrectangular, smooth or giving rise to teeth, spines, cilia or coronate papillose projections; internally composed of l(-2) layers of guide cells between dorsal and ventral bands of stereids. Cells of chlorophyllose lamina quad- rate, shortly rectangular, irregularly polygonal or rounded, broader than long to longer than broad, smooth and flat, or producing spines, teeth or papillose projections from dorsal and/or ventral surfaces of the leaf. Leaf margin often differentiated (nearly always so in the hyaline base); in proximal hyaline base usually consisting of a flattened rib of thick-walled linear cells, most often unistratose and entire; in distal leaf base often with spines, teeth or cilia; interrupted by chlorophyllose lamina, or continuing into upper leaf; in chlorophyllose limb obscure to strong, or absent, consisting of stereids and/or subrectangular, irregularly polygonal cells in 1- several layers; entire, denticulate, dentate, spinulose, or ciliate. In many species gemmae produced from the costa, most often from the costal apex (in a few species produced on specialized leaves). Sporophytes terminal; seta smooth; capsule exserted, erect, cylindri- cal. Peristome haplolepidous (lacking in some species), with 16 teeth (usually ridged and/or papillose); hyaline properistome often evident. Operculum rostrate; calyptra cucullate, fugacious. Syrrhopodon is the largest genus in the Calymperaceae and has the broadest latitudinal and altitudinal ranges. Several species have reached the temperate regions of Asia and North America (Reese, 19876). In the Philippines, the genus is primarily a corticolous moss of lowland rainforest, although several species occur on humid boulders. A few have successfully penetrated the montane or cloud forests above 1000 m. Syrrhopodon lacks autapomorphic characters, and consequently, has been thought to represent the ancestral or primitive form in the family Calymperaceae (Reese, 19876). Yet, the genus also has the largest number of endemics or locally derived species. There are about 30 Malesian species of Syrrhopodon, of which 19 are reported below for the Philippine archipelago. Syrrhopodon albovaginatus Schwagr., Sp. muse, frond, suppl. 2( 1 ): 1 3 1 ( 1 824). Type: Moluccas, Rauwack Island, Gaudichaud [14A (H. 1230)] (BM!-isotype). Fig. 16a-f. Syrrhopodon micholitzii Mull. Hal. in Paris, Index Bryol.: 1253 (1898), nom. nud. Original specimen: Philippines, Mindanao, Surigao, Jaganan, March 1896, Micholitz 171 (FH). Shoots slender, 3.5 cm high, in tufts or mats, with a scolopendroid appearance, sometimes dense with rhizoids below. Leaves close-set, mostly 3-3.5(>4.0) mm long; each consisting of an erect, semisheathing, narrowly elliptical hyaline base extending into a spreading to recurved, linear (slightly twisted) chlorophyllose limb with an acute to subacute dentate apex (Fig. 16a-d). Costa ending in apex, superficial cells shortly to long rectangular, above hyaline base some giving rise to subtransverse rows of acute teeth; internally with a single row of guide cells between dorsal and ventral layers of stereids. Cells of chlorophyllose lamina mostly isodiametric, rounded, quadrate, shortly rectangular or irregularly polygonal, smooth, mostly 7-12.5 x 5-10 |J.m, at short regular intervals along the leaf limb (on both dorsal and ventral surfaces) forming largely unbroken, oblique to transverse rows of acute, distally leaning teeth (Fig. 16e). Marginal rib continuous from leaf base to near apex, entire, above proximal hyaline base polystratose, composed of stereid cells, obscured in distal leaf where the margins of the chlorophyllose lamina become tightly incurved and apparently form a denticulate leaf margin. Sporophytes occasional, seta 5-7 mm long, capsule c. 1.5 mm long, erect, narrowly cylindrical; peristome teeth papillose; operculum with a long, erect, narrow, subulate beak. HABITAT. On decaying logs and tree stumps in lowland forest. DISTRIBUTION. A widespread Indo-Pacific species. Very common in the Philippines. SPECIMENS EXAMINED. Luzon, Laguna Province, Los Banos, Mt Makiling, June-July 1917, Elmer] 8457 (BM);Mt Makiling, 17 August \93l,Herklots P22 (BM); Siniloan, U.P. Quezon Land Grant, 3 January 1882, Baldovino 001 (BM); San Antonio, September-October 1912, Ramos 16670 (BM); Sierra Madre Range, 16 December 1985, Tan s.n. (FH); Aurora Province, 30 L.T. ELLIS AND B.C. TAN b-d,g e, f, h, i, 0.5 mm 50 Fig. 16 a-f. Syrrhopodon albovaginatus Schwagr. a-d: leaf (a: dorsal aspect, with details of b: apex of hyaline lamina, c: mid-chlorophyllose limb, d: apex); e: cells of chlorophyllose lamina in surface view; f: cross-section of chlorophyllose limb. g-i. Syrrhopodon ciliatus (Hook.) Schwagr. g: leaf in ventral view; h: cells of chlorophyllose lamina in surface view; i: cross-section of costa and chlorophyllose lamina, a-d Drawn from Tan 92-210 (BM). f Drawn from Tan 93-320 (FH). g-i Drawn from Micholitz 165 (BM). Sierra Madre Range, 23 October 1980, Alvarez 0-801367 (BM); Isabela Province, Palanan Wilderness, 20 May 1992, Tan 92-210 (FH). Mindanao, Sunlug, Seno de Davao, 12 May 1890, Micholitz 4 (BM); March 1896, Micholitz 171 (BM); Zamboanga Province, Basilan City, Lamitan, 4 May 1948, Santos 4075 (BM); 25 May 1949, Santos 4301 (BM). Mindoro, between Bongabon and Pinamalayan, February-April 1 94 1 , Maliwanag 3 1 2 (FH). Palawan, Puerto Princesa, Barangay Irawan, Mt Malinao, 4 May 1993, Tan 93-320 (FH); between Roxas and Port Barton, 1992, Tan 92-374B (FH). Panay, Capiz Province: Libacao, May-June 1919, Martelino & Edano 35768 (BM); April-May 1918, Ramos & Edano 30840 (BM). This species is unique in the possession of leaves with successive transverse rows of teeth projecting from the chlorophyllose lamina. Syrrhopodon aristifolius Mitt, in/ Linn. Soc. Bot. 10: 176 (1868). Type: Samoa, Upolu, 1000-2000 ft, May 1867, Powell^ (BM!- isotype). Fig. 17. Syrrhopodon subulatus Sande Lac. in Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 13(2): 5 (1872). Type: Sulawesi ['Celebes'], ex Hb. Sande Lac. s.n. (BM!-isosyntypes?). Syrrhopodon fallax Sande Lac. in Verh. Kon Ned. Akad. Wetensch. Amst., Afd. Natuurk. 13(2): 5 (1872). Type: Banca, Kurz s.n. (L- syntype); Borneo, Korthals s.n. (L-syntype). Shoots mostly 1-2 cm high, in bristly tufts. Stems short, often matted with rhizoids. Leaves long, flexuous, hair-like, mostly (<5-) 7-20 mm long, erect to patent or subfalcate to variously flexed from a broadly subelliptical base with long sloping shoulders; apex aristate (Fig. 17a-d). Costa strong, above leaf shoulders occupying most of the leaf; gradually narrowing distally into an often long excurrent, fine, entire arista; superficial cells above leaf base quad- rate to shortly rectangular, smooth and flat; internally with l-2(-3) layers of guide cells between dorsal and ventral layers of stereids (Fig. 17h). Chlorophyllose lamina occupying leaf shoulders, merg- ing proximally with the hyaline lamina; above leaf shoulders continuing linearly (seldom more than 5-6 cells wide on either side of the costa); beyond midleaf towards apex tapering into costa; unistratose but sometimes with bistratose patches; cells long to shortly rectangular or elliptical, isodiametric to longer than broad, smooth, sometimes slightly convex ventrally, mostly 10-20 x 7.5- 12.5 )im (often more elongated in leaf shoulders, some reaching >30 |im long) (Fig. 17g). Hyaline lamina merging into chlorophyllose lamina below shoulders of leaf, not usually sharply defined (Fig. 17f). Leaf margins from above leaf shoulders towards apex formed by a polystratose rib (in some leaves continuing beyond distal limit of chlorophyllose lamina and becoming continuous with the costa), lacking stereids, superficial cells in surface view quadrate to shortly rectangular, smooth, some forming distant, often multicellular teeth; CALYMPERACEAE IN THE PHILIPPINES 31 1 mm c, 0.5 mm e-h, k, 1 50 Fig. 17 a-h. Syrrhopodon aristifolius Mitt, (typical form) a-d: leaf (a: semidiagrammatic representation, with details of b: hyaline base, c: mid-leaf, d: apices); e-g: cells of leaf in surface view (e: at margin of hyaline base, f: around apex of hyaline lamina, g: in chlorophyllose lamina); h: cross-section of leaf in chlorophyllose limb. i-1. S. aristifolius (short-leaved form) i, j: leaf (i: in ventral view, withj: detail of apices); k: chlorophyllose lamina in surface view, 1: chlorophyllose limb in cross-section, a-h Drawn from Solomon Islands, Lee 803 (BM). i-1 Drawn from Peninsular Malaysia, Ellis s.n. (BM). around leaf shoulders unistratose, usually entire, rarely subdenticulate; in proximal leaf base unistratose, entire, composed of small hyaline cells, sometimes with walls thicker than those of adjacent hyaline cells (marginal band of linear thick-walled cells rarely apparent) (Fig. 17e). Seta c. 1 1 mm long; capsule c. 1.5 mm long; operculum with a long subulate beak. HABITAT. On tree trunks in lowland rainforest. DISTRIBUTION. An Indo- Pacific species. Bartram (1939) cites three Philippine collections (as Syrrhopodon fallax Sande Lac. and 5. subulatum Sande Lac.) from Mindoro and Mindanao. No other local specimens examined. As with most species in the Calymperaceae, plants of Syrrhopodon aristifolius are highly plastic. An atypical, under-developed form, with leaves consistently <5-7 mm long (Fig. 17i), occurs in lowland tropical rainforest. This form is all but absent from herbaria, yet extensive populations have been sampled in Sabah and the Malayan Peninsula. It remains to be detected in the Philippines. Superficially this form has some resemblance to Calymperes subserratum, but leaves in the latter possess unistratose margins and much smaller cells in the chlorophyllose lamina (<5-10(-12.5) x 5- 7.5 um) which are ventrally protuberant. The leaves in 5. aristifolius have polystratose margins and larger, mostly flat cells in the chlorophyllose lamina (10-20 x 7.5-12.5 urn). Syrrhopodon armatus Mitt, in J. Linn. Soc. Bot. 7: 151 (1863). Type: West Africa, Bagroo River, 1861, Mann s.n. (NY-syntype; BM!-isosyntype); West Africa, banks of the Nun, September 1860, Mann [549] (NY-syntype; BM!-isosyntype). Fig. 19h-n. Syrrhopodon fimbriatus Mull. Hal. inLinnaea 37: 151 (1872), horn. illeg. Syrrhopodon fimbriatulus Mull. Hal. in J. Mus. Godeffroy 3(6): 52 (1874). Type: Australia, Brisbane River, 1864, Dietrich s.n. (B- holotype, presumably destroyed; isotypes not found, fide Reese & Bartlett(1982)). Syrrhopodon larminatii Broth. & Paris in Rev. Bryol. Lichenol. 28: 125 (1901). Type: Vietnam, Hanoi, Lao Cai, Larminat s.n. (H- BR-holotype). Syrrhopodon tsushimae Cardot in Bull. Herb. Boissierser. 2,7:716 32 L.T. ELLIS AND B.C. TAN (1907). Type: Japan, Faurie 1637 (PC?-holotype). Syrrhopodon philippinensis E.B. Bartram in Philipp. J. Sci. 68: 83 (1939). Type: Philippines, Luzon, Zambales Prov., hills between San Marcelino and Mount Pinatubo, Bartlett 14219 (FH!- holotype). Shoots 0.4-1 cm high, densely leaved, matted with rhizoids below, forming dense, yellowish green mats. Leaves l-2(-3) mm long, linear, erect to patent (incurled-crisped when dry), consisting of a narrowly elliptical hyaline base tapering gradually into a chlorophyllose limb; leaf apex broadly obtuse to almost truncate, cuspidate (cusp sometimes dorsally inclined) (Fig. 19h, i). Costa (prominent dorsally) ending below apex in a blunt, dentate point; dorsally (usually from mid-hyaline base to apex) and ventrally (usually from above hyaline base to apex) with short to long, erect to distally-curving, acute spines. Chlorophyllose lamina laxly incurved, rarely slightly undulate; cells longer than broad to broader than long, with 4-6 sides or rounded-elliptical, 5-15 x 5-10 u.m (Fig. 191), ventrally and dorsally drawn out as single, often tall, sometimes curving, subacute projections (Fig. 19n). Hyaline lamina sharply defined. Leaf margins from proximal chlorophyllose limb to beyond midleaf (occasionally to near apex) usually consisting of a thin, largely entire strand of stereids (sometimes weak, intermittent or absent) (Fig. 19k); in hyaline base composed of a narrow, flattened strand of linear, thick- walled cells (<3-5 cells wide, usually continu- ous with marginal strand in distal leaf); adjacent to and beyond distal region of hyaline lamina giving rise to a row of acute spines and/or cilia (Fig. 19j). Gemmae sometimes produced from ventral surface of costal apex. Sporophytes rare. Seta c. 5-6 mm long; capsule cylindrical, c. 1 mm long; peristome teeth strongly papillose. HABITAT. Corticolous DISTRIBUTION. A Palaeotropical species. In the Philippines con- fined to Luzon Island. Asian and Australasian plants of this species were previously referred to Syrrhopdonfimbriatulus Mull. Hal. Reese & Stone (1995) recog- nized this taxon to be conspecific with Syrrhopodon armatus, earlier described from Africa. Syrrhopodon armatus is very similar toSyrrhopodon trachyphyllus Mont. The latter has slightly more robust shoots and leaves than S. armatus but the main distinguishing feature is the form of the chlorophyllose lamina. In 5. trachyphyllus the cells of the chlorophyllose lamina are 10-15(-17.5) x 8-12.5 |im, each with a crown of papillae projecting from the dorsal and ventral surfaces (Fig. 24i). Those in 5. armatus are 5-15 x 5-10 |im, with a single papilla projecting from the dorsal and ventral surfaces (Fig. 19n). Syrrhopodon ciliatus (Hook.) Schwagr., Sp. muse, frond, suppl. 2(1): 114, 130(1824). Fig. 16g-i. Weissia ciliata Hook., Musci Exot. 2: 7, 171 (1820). Type: Ternate Island, Hb. Dickson (BM!-holotype?; BM!-isotype). Shoots mostly <0.5-l(-4) cm high, soft, pale green, often densely covered with red rhizoids below. Leaves delicate, suberect to reflexed, 2.5 cm, densely leaved, forming pale, dense mats and cushions. Leaves erect to patent, lanceolate, 1 .5-3 mm long, consist- ing of a short, linear-lanceolate chlorophyllose limb extending from a narrowly suboblong hyaline base (usually occupying over half the leaf length); leaf apex usually drawn out into a fine, denticulate, acute point (Fig. 18k, 1). Costa narrow (especially in leaf base), ending in apex to excurrent; superficial cells long and narrow, ventral surface smooth, dorsal surface above hyaline base smooth to spinose (spines most frequently arising from cells adjacent to lamina); internally (above base) consisting of a layer of guide cells (2-3 cells wide) separating dorsal and ventral bands of stereids. Lamina at leaf shoulders often recurved. Cells of chlorophyllose lamina obscured on ventral and dorsal surfaces of leaf by tall projections crowned with compound papillae. Hyaline lamina often occupying up to two thirds of leaf length, sharply defined but narrowly tapering into the chlorophyllose limb between the costa and the narrow wings of the proximal chlorophyllose lamina (Fig. 18m). Leaf margins from shoulders to near leaf apex composed of stereids or substereids in a thin, polystratose rib, at shoulders notched to denticulate, above denticulate with single or double teeth; from below shoulders to leaf base entire, consisting of a flattened, narrow band of thick-walled, linear cells. Sporophytes rare, not seen in Philippine material. HABITAT. On tree trunks in lowland rainforest. DISTRIBUTION. An Indo-Pacific species. Bartram (1939) cites Phil- ippine specimens from Luzon, Negros and Panay (as Syrrhopodon amoenus Broth.). SPECIMEN EXAMINED. Luzon, Isabela Province, San Mariano, Barrio Disulap, Dimahahabong Creek, 14 April 1991, Tan 91-140 pro parte (FH). The leaves of Syrrhopodon confertus are distinguishable from those of related species, such as 5. involutus Schwagr. and S. rufescens Hook. & Grev., by the possession of a short but well-defined chlorophyllose limb. In the latter two species, the chlorophyllose CALYMPERACEAE IN THE PHILIPPINES 33 g n b,k c,d,l 1 mm e-j, m-p 0.5 mm 50 Fig. 18 a-j. Syrrhopodon croceus Mitt, a-d leaf (a: diagram of whole leaf, with details of b: base, c: region above base, d: apex); e-g: cells of leaf in surface view (e: above hyaline lamina, f: at margin in chlorophyllose limb, g: in chlorophyllose lamina); h-j cross-sections of leaf (h: costa, i: marginal rib in chlorophyllose limb, j: chlorophyllose lamina), k-p. Syrrhopodon confertus Sande Lac. k-1: leaf (k: in dorsal view, with details of 1: apex in ventral view, m: apex of hyaline lamina in dorsal view); n-p: cross-sections of leaf (n: near base, o: above mid-leaf, p: near apex), a-j Drawn from Tan 92-173 (BM). k, m Drawn from Peninsular Malaysia, Burkill 2112 (BM). 1, n-p Drawn from Sabah, Ellis 143-366 (BM). lamina is confined to the apex of shortly pointed leaves, otherwise composed of hyaline cells. Syrrhopodon croceus Mitt, in J. Linn. Soc. Bot. Suppl. 1: 41 (1859). Fig. 18a-j. Calymperidium croceum (Mitt.) M. Fleisch., Muse. Fl Buitenzorg 1: 219 (1904). Type: Singapore, Wallich s.n. [3640] (BM!- isosyntype). Shoots in dark green to reddish brown tufts, 2-5 cm high, often densely matted with rhizoids below. Leaves <5-10 mm long, stiff, straight or slightly curving, with an erect, elliptical semisheathing base narrowing gradually into an erect to patent, linear-ligulate chlorophyllose limb, ending in a blunt, denticulate apex (Fig. 1 8a, b, d). Costa ending just below leaf apex to excurrent (tip dentate and often, especially in gemmiferous leaves, slightly expanded), in distal leaf-limb scabrid on both dorsal and ventral surfaces, proximally becoming smooth. Hyaline lamina clearly defined, usually confined to the lowermost third to half of leaf base, hyalocysts often yellow tinged; distal leaf base occupied by bright orange-red, incrassate, long rectangular to linear, porose cells (Fig. 18e); distally these becoming smaller and paler, gradually merging into the chlorophyllose lamina of the lower leaf-limb. Chlorophyllose lamina often incurved to near leaf apex, composed of shortly subrectangular to subelliptical cells with walls sometimes thickened at the angles, 34 L.T. ELLIS AND B.C. TAN mostly 10-20 x 7.5-10mm, each projecting acutely from the ventral leaf surface (usually terminating in a sharp papilla), dorsally unipapillose (Fig. 18g, j). Margins of leaf base entire, adjacent to hyaline lamina formed by a broad, flat, orange-red band of incrassate, porose, linear cells (sometimes cells in outermost row of band shortly rectangular, thin-walled and hyaline); distally becoming indistinct, merging with the upper basal lamina; around and shortly above shoulders of leaf becoming clearly defined once more, com- posed of shortly to long subrectangular, incrassate cells in 5-8 rows, some in marginal row giving rise to long, acute teeth (Fig. 1 8c); from above shoulders to near leaf apex becoming a thick, polystratose rib composed of stereids with a broken superficial layer of shortly rectangular cells, some forming distant, large, single or double (sometimes triple) teeth (Fig. 18f, i). Gemmae sometimes produced from the ventral surface of the costal apex. Seta 12-17.5 mm long; capsule cylindrical, 1 .5^<2 mm long. HABITAT. On logs, tree trunks, and limestone rock in shade. DISTRIBUTION. An Indo-Pacific species, very common in the Phil- ippines. SPECIMENS EXAMINED. Basilan, Semper s.n. (in Hb. Hampe, BM). Leyte, Baybay, Mt Pangasugan, 21-22 May 1984, Tan, Navarez & Raros 84-235 (BM). Luzon, Isabela Province, San Mariano, Sierra Madre Range, Barrio Disulap, Dimahahabong Creek, 14 April 1991, Tan 91-140 pro parte (FH); ibid, Sitio Digoyo, 17 April 1991, Tan 91-170 (BM); Palanan Wilderness, 21 May 1992, Tan 92-173 (BM); Laguna Province, Cavinti, Bo. Lumot, Ubali River, near Sitio Ubali, 24 October 1982, Tan & Tandang 82-376 (FH); Quezon Province, Real, National Botanical Garden, 24 December 1981, Tan 81^46 (Muse. Philipp. Fasc. 2 no. 29) (BM). Palawan, Puerto Princesa, Barangay Irawan, Mt Malinao, 4 May 1993, Tan 93-308 (FH). Sibuyan, Magdiwang, Barangay Tampayan, Mt Giting-Giting, 1 May 1987, Tan & Hernaez 87^73 (BM); ibid, 21 May 1987, Tan & Hernaez 87-445 (BM). This species is highly variable. There are many atypical forms in which the leaves may be very short and in which distinctive features, such as the orange-red cells in the distal leaf base or the teeth along the margin of the lower leaf limb, may be obscure or completely missing. Syrrhopodon flammeonervis Mull. Hal. mLinnaea 38: 557 (1 874). Type: Philippines, Luzon, 1871, Wallis s.n. (FH!, H-isotypes). Fig. 19a-g. Syrrhopodon flammeonervis var. robustus Dixon in J. Linn. Soc. Dot. 50: 81 (1935). Type: Sarawak, Ulu Tinjar, Sungei Balapau, G. Laiun, c. 1100 m, 2 November 1932, Oxford Expedition [Richards] 2381 (BM!-holotype). Shoots 2->6 cm high, in pale to dull green tufts. Leaves 3->6 mm long, consisting of an erect, clasping, narrowly oblong hyaline base abruptly narrowing into an erect to spreading, linear chlorophyllose limb that is V -shaped in cross-section (loosely and irregularly curved and twisted when dry); apex blunt, dentate (Fig. 19a, b). Costa ending immediately below apex; towards and at apex dorsal surface with large, acute teeth, proximally cells in rows adjacent to the lamina sometimes forming acute to spinose teeth, otherwise smooth (dorsal and ventral surfaces largely composed of stereids). Chlorophyllose lamina at leaf apex forming a narrow, spinose margin around the costal apex, ultimate cell often in the form of a large, dorsally slanting, spinose tooth (Fig. 19b); below apex cells isodiametric to longer than broad, with 4-6 sides or irregularly rounded-elliptical, 8-17(-25) x 7.5- 10(- 12.5) |im (Fig. 19e), dorsally and ventrally drawn out into coronate-papillose projections (towards leaf apex projections sometimes becoming more spinose) (Fig. 19f). Hyaline lamina sharply defined. Leaf margin sometimes incurved above and/or recurved at leaf shoulders; from around apex of leaf shoulders to near leaf apex formed by a thick strand of stereids (Fig. 19g), largely entire but near apex some superficial cells forming teeth; from shoulders to leaf base composed of a flattened band (continuous with rib in upper leaf) of thick- walled, linear cells in about 3-7 rows (sometimes cells of outermost row with thin walls) (Fig. 19d); in shoulders uneven to denticulate, below shoul- ders entire. Sporophytes not seen. HABITAT. Eddy (1990) reports this species as occurring 'on bark and acidic rocks in moist rainforest between 1000 and 2000 m alt.'. DISTRIBUTION. SW China, Indochina, Philippines, Borneo. In the Philippines known from the islands of Luzon, Palawan and Negros. SPECIMENS EXAMINED. Luzon, Mt Makiling, 16 August 1931, Herklots P18b (BM). Palawan, Mt Manalsal, May 1929, Edano 80869 (BM). Syrrhopodon gardneri (Hook.) Schwagr., Sp. muse, frond, suppl. 2(1): 110(1824). Fig. 20i-u. Calymperes gardneri Hook., Musci Exot. 2:146 (1819). Type: Ne- pal, Gardner [1205] (BM!-holotype, BM!-isotype). Syrrhopodon curranii Broth, in Philipp. J. Sci. C, 5: 142 (1910). Type: Philippines, Luzon Island, Benguet Province, December 1908, H.M. Curran 15636 (BM!-isotype). Shoots 0.5->5 cm high. Leaves mostly 3-5 mm long, erect to patent, narrowly to broadly linear-ligulate from an erect, elliptical hyaline base; ending in an obtusely pointed, denticulate apex (Fig. 20i-k). Costa ending just short of leaf apex in a dentate tip; in upper leaf many superficial cells forming acute, forward-pointing teeth, often with one or two small papillae; especially towards leaf apex distant, subtransverse rows of relatively larger teeth sometimes occur (Fig. 20q). Chlorophyllose lamina often broadly incurved to below leaf apex; cells quadrate to shortly rectangular or subelliptical (mostly isodiametric or longer than broad), 7-17.5(-20) x 7-10 |j,m (Fig. 20p), projecting acutely from the ventral leaf surface, each projec- tion bearing one or more small papillae, dorsally uni- or pleuri-papillose (Fig. 20t, u). Hyaline lamina well-defined (Fig. 201). Marginal ribs in proximal leaf base entire to dentate, flat, composed of long, narrow, thick-walled cells (Fig. 20m); in distal leaf base vanishing into chlorophyllose lamina; from around mid- hyaline base to above shoulders rib lacking, margin denticulate to subciliate (Fig. 20n); above shoulders polystratose, erect or incurved, composed of quadrate to shortly rectangular cells (stereids absent), double or single teeth occurring at intervals along the margin (Fig. 20o). Gemmae sometimes produced from the ventral surface of the costal apex. Seta dull orange, 4-5 mm long; capsule cylindrical, 1- 1.3 mm long. HABITAT. On tree trunks in rainforest. Most frequent at 1 000-2000 m, occurring at higher altitudes than many taxa in the Calymperaceae. DISTRIBUTION. A nearly pan tropical species. In the Philippines confined to Luzon. SPECIMEN EXAMINED. Luzon, Benguet Province, Buguias, Lake Tabayog, 25 October 1985, Tan & Hernaez 85-127 (BM, FH). The leaves of Syrrhopodon gardneri are superficially similar to those of S.japonicus (Besch.) Broth, but are usually shorter. In most specimens of S. gardneri the leaves seldom exceed 5 mm long, the cells of the chlorophyllose lamina project acutely from the ventral leaf surface and dorsally and ventrally are usually papillose CALYMPERACEAE IN THE PHILIPPINES 35 a 1 mm b, c, h, i 0.5 mm d-g,j-n 50 Fig. 19 a-g. Syrrhopodon flammeonervis Miill. Hal. a-c leaf (a: lateral view, with details of b: apex, and c: distal hyaline base); d, e: cells of leaf in surface view (d: at margin in mid-hyaline base, e: in chlorophyllose lamina); f-g: cross-sections of leaf (f: chlorophyllose limb, g: margin of chlorophyllose limb), h-n. Syrrhopodon armatus Mitt, h, i: leaf (h: ventral view, with detail of i: distal chlorophyllose limb in ventral view); j-1: cells of leaf in surface view (j at margin in distal hyaline base, k: at margin in mid-chlorophyllose limb, 1: in chlorophyllose lamina (dorsal view)); m, n: cross- sections of leaf (m: costa, n: chlorophyllose lamina), a-g Drawn from Edano 80869 (BM). h-n Drawn from North Borneo, Binstead 35 (BM). (Fig. 20t, u). In most specimens of 5. japonicm the leaves are up to 7-8 mm long, the cells of the chlorophyllose lamina lack papillae and are more roundly protuberant (Fig. 2 In), often with the greater protuberance from the ventral leaf surface. Syrrhopodon hispidissimus Dixon in J. Malayan Branch Roy. Asiat. Soc. 6: 23 (1928). Type: Sumatra, Siberoet Island, Septem- ber 1924, Boden Kloss 10581b (BM!-holotype). Fig. 20a-g. Syrrhopodon perarmatus E.B. Bartram in Farlowia 1: 42 (1943), horn, illeg. (non 5. perarmatus Broth.). Syrrhopodon hispido-ciliatus E.B. Bartram in Farlowia 1: 504 (1944). Type: Papua New Guinea, Palmer River, 2 miles below junction, Black River, July 1936, Brass 7161a (FH!-holotype). Shoots delicate, <0.3-1.5 cm high, forming pale green, soft mats or tufts. Leaves consisting of a linear, erect to patent chlorophyllose limb narrowing from a suberect, subelliptical hyaline base, < 1.5-3 mm long; apex subacute to acute, dentate (Fig. 20a-c). Costa ending below apex; dorsal and ventral surfaces above hyaline base with close-set, transverse-oblique rows of long, acute, distally leaning, unicellular spines (may exceed 35 |im long) (Fig. 20g); internally with a single row of guide cells. Chlorophyllose lamina <5-c.lO cells wide on either side of costa; cells mostly isodiametric to slightly longer than broad, 5- 1 2.5 x 5-7 Jim; each with a thick, often 36 L.T. ELLIS AND B.C. TAN JM T^«WK«? t W3F a,b, i,j 1 mm c-g, m-u 0.5 mm 50 fim Fig. 20 a-h. Syrrhopodon hispidissimus Dixon a, b leaves, with details of c: apex, and d: region around apex of hyaline lamina; e-g: cells of leaf in surface view (e: at margin of distal hyaline base, f: at margin of chlorophyllose limb, and g: in costa); h: diagrammatic cross-section of leaf through chlorophyllose limb. i-u. Syrrhopodon gardneri (Hook.) Schwagr. i-1: leaves (i, j in ventral view, with details of k: apex (ventral view), and 1: distal region of hyaline lamina); m-q: cells of leaf in surface view (m: at margin of hyaline lamina, n: at margin adjacent to apex of hyaline lamina, o: at margin in chlorophyllose limb (ventral view), p: of chlorophyllose lamina, q: of costa (ventral view)); r-u: cross-sections of leaf (r, s: costa, t, u: chlorophyllose lamina), a, c, d Drawn from Ebalo 737 (FH). b Drawn from Sumatra, Boden Kloss 10581b (BM). e, f, g Drawn from Papua New Guinea, Brass 7 161 a (FH). i, k, m, r, t Drawn from Tan & Hernaez 85-127 (BM). j, 1, n-q, s, u Drawn from Curran 15636 (BM). CALYMPERACEAE IN THE PHILIPPINES 37 tall crown of papillae protruding from the dorsal and ventral surfaces (in surface view papillae obscuring the shape of the underlying cells). Leaf margins from above shoulders of leaf to near apex formed by a thin polystratose rib of transversely aligned, long, narrow cells that are drawn out distally as acute, forward pointing teeth (in transverse ranks of 2-4) (Fig. 20f); from shoulders to leaf base consisting of a flattened band of long, narrow, thick-walled cells (continuous with rib of upper leaf), at shoulders producing a row of cilia (cilia often exceeding 80 urn long) (Fig. 20e), entire below. Sporophyte unknown. HABITAT. On exposed roots, tree trunks and buttresses in shaded situations. DISTRIBUTION. An uncommon Indo-Pacific species. In the Philip- pines the species has been found in Mindanao and probably also occurs in Luzon. SPECIMEN EXAMINED. Mindanao, Zamboanga Province, Muralong Moun- tain near Kabasalan, 27 November 1940, Ebalo 737 (FH). Syrrhopodon hispidissimusis sometimes confused with S. spiculosus. In the latter, the margin of the leaf above the hyaline base is formed (except near the leaf apex) by a rib of stereids which is smooth and entire (Fig. 23h); in S. hispidissimus most cells at the surface of the marginal rib give rise to sharp, distally-pointing teeth (Fig. 20f). Syrrhopodon involutus Schwagr., Sp. muse, frond, suppl. 2(1): 1 17 (1824). Type: Moluccas, Rawak Island, Gaudichaud 11 (26) (BM!-isotype). Fig. 21a-g. Syrrhopodon revolutus Dozy & Molk. inAnn. Sci. Nat. Bot. ser. 3, 2: 315 (1844). Type: Java and Borneo, Korthals s.n. (L-holotype; BM!-isotype). Trachymitrium borneense Hampe in Nuovo Giorn. Bot. Ital. 4: 280 (1872). Type: Borneo, Sarawak, Beccari 46 (BM!-holotype). Syrrhopodon borneensis (Hampe) A. Jaeger, Gen. Sp. muse. 1: 316 (1873). Leucophanella borneense (Hampe) M. Fleisch., Muse. Fl. Buitenzorg 1: 197 (1904). Leucophanella revoluta (Dozy & Molk.) M. Fleisch., Muse. Fl. Buitenzorg 1: 198 (1904). Shoots l-2(->3) cm high, freely branched, densely leaved, in densely packed, pale green cushions, often loosely matted with red rhizoids below. Leaves 1-1.5 mm long, erect to erecto-patent, lanceolate, above often involute or V-shaped in cross-section; leaf apex obtuse to shortly pointed, entire to denticulate; largely com- posed of hyaline lamina (Fig. 2 la, b). Costa thin, ending in or just below apex; dorsal surface smooth below, nearing leaf apex often scabrid, teeth mostly formed by cells in rows adjacent to the lamina; ventral surface smooth, rarely toothed near apex; internal structure similar to that in Syrrhopodon confertus, consisting of a layer of guide cells (2-3 cells wide) separating dorsal and ventral bands of stereids (ventral band sometimes poorly developed) (Fig. 21d-f). Chlorophyllose lamina occupying leaf apex, proximally narrowly tapering down either side of the hyaline lamina, ceasing prior to midleaf; cells mostly quadrate to shortly rectangular, or irregularly rounded-elliptical, mostly 5->15 x 5->10 urn (Fig. 21c); dorsally and/or ventrally flat or protruding subacutely, smooth or unipapillose (Fig. 21e, g). Hyaline lamina occupying about two thirds to more than four fifths of the leaf length, sharply defined; cells large, thin- walled, shortly subrectangular, seldom in more than about 5-6 rows on either side of the costa. Leaf margins often recurved above; from leaf base to near apex entire, consisting of a narrow, smooth, unistratose to bistratose strand of stereids (Fig. 21c, d, e). Seta red, smooth, <6->10 mm long; capsule erect, shortly cylindrical, <1 mm long, yellowish brown with a red rim, lid long rostrate, reddish brown; peristome teeth <100->150 um long, papillose. HABITAT. On trunks and exposed roots of trees and decaying logs. DISTRIBUTION. A palaeotropical species. Common in the Philip- pines. SPECIMENS EXAMINED. Catanduanes, 14 November- 1 December 1917, Ramos 30610 (BM, BM-K); Luzon, Loher 1056 (BM-K); Baguio, 1910, Sanchez 6 (BM); Benguet, Micholitz 153 (BM-K); Laguna, San Antonio, Ramos 16672 (BM). Mindanao, March 1896, Micholitz 179 (BM, BM-K); Agusan Norte, Cabadbaran, Mt Hilog-Hilong, 31 May-2 June 1984, Tan & Navarez 84-492 (FH). Mindoro, between Bongabon and Pinamalayan, 5 February-5 April 1941, Maliwanag 313 (FH). Syrrhopodon japonicus (Besch.) Broth., Nat. Pflanzenfam. 2nd ed., 10: 233 (1924). Fig. 21h-o. Calymperes japonicus Besch. in J. Bot. (Morot) 12: 296 (1898). Type: Japan, Nagasaki, March 1895, Faurie 15454 (BM!- holotype). Calymperes datense E.B. Bartram in Philipp. J. Sci. 68: 98 (1939). Type: Philippines, Luzon, Benguet subprovince, Mount Data, 8000 ft, Hodden 116 (FHI-lectotype, fide Mohamed & Reese, 1985). Shoots <2-5(-6) cm high, often matted with red rhizoids below. Leaves 7-8(-9) mm long, linear-lanceolate, bristle-like, patent to recurved from an erect, narrowly elliptical hyaline base (shoulders of leaf often not very pronounced), at apex tapering to an acute, dentate tip (Fig. 21h, i). Costa strong, excurrent, sometimes long- excurrent; superficial cells mostly flat and smooth but often forming teeth near costal apex; internally with a single row of guide cells between dorsal and ventral bands of stereids (Fig. 21m). Cells of chlorophyllose lamina quadrate to subrectangular, mostly 1-3 times longer than broad, 6-12.5(-15) x 5-8 urn (Fig. 211), projecting roundly to subacutely from the ventral leaf surface, dorsally flat to slightly protuberant, smooth (Fig. 2 In); around leaf shoulders usu- ally grading gradually into areas of larger, subquadrate to subrectangular incrassate cells (often with transverse walls and corners preferentially thickened) that grade proximally into the large, thinner walled cells of the hyaline lamina in the leaf base (sometimes hyaline cells thick-walled to base of leaf) (Fig. 21j). Margins from above shoulders to near leaf apex formed by a polystratose rib (sometimes weak), superficial cells subquadrate to rounded, double (sometimes triple) teeth occuring at short intervals, lacking stereids (Fig. 21k, o); in shoulders and upper leaf base unistratose and undifferentiated, denticulate; in proximal leaf base usually entire, formed by a flattened rib composed of long, narrow thick-walled cells (cells in marginal row often with slightly thinner walls than those in inner rows), vanishing distally. Sporophytes apparently rare, not seen in Philippine material. Setae. 15 mm long; capsule 1.5-2 mm long. HABITAT. In montane rainforest; on tree trunks, buttresses, moist humus and logs, usually in shade. DISTRIBUTION. Japan, China, Indo-China, Malesia. Widspread in the Philippines. 38 L.T. ELLIS AND B.C. TAN a I a, h, i, q, r_ b, s c-g,j-o,t-v 50 1 mm 0.5 mm Fig. 21 a-g. Syrrhopodon involutus Schwagr. a: leaf, with b: detail of apex in ventral view; c: cells of chlorophyllose lamina above hyaline lamina; d-g: cross-sections of leaf (d: near base of leaf, e: above mid-leaf, f: costa near apex, g: chlorophyllose lamina), h-o. Syrrhopodon japonicus (Besch.) Broth. h, i: leaves; j-1: cells of leaf in surface view (j: around apex of hyaline lamina, k: at margin in mid-leaf, 1: in chlorophyllose lamina); m-o: cross-sections of leaf (m: costa, n: chlorophyllose lamina, o: marginal rib in chlorophyllose limb), p-v. Syrrhopodon loreus (Sande Lac.) W.D. Reese p-s: leaf (p: diagram of whole leaf, with details of q; hyaline base and proximal chlorophyllose limb, r: apex, and s: distal hyaline base); t: cells at margin of leaf adjacent to apex of hyaline lamina; u, v: cross-sections of leaf (u: costa and chlorophyllose lamina, v; marginal rib in chlorophyllose limb), a-g Drawn from Tan & Navarez 84-492 (FH). h Drawn from Tan, Navarez & Amoroso 84-365 (BM). i, m-o Drawn from Gruezo 5798 (FH). j-1 Drawn from Madden s.n. (FH). p-t Drawn from Tan 91-108 pro parte (FH). u, v Drawn from Tan & Wijengco 85-105 pro parte (BM). CALYMPERACEAE IN THE PHILIPPINES SPECIMENS EXAMINED. Mindanao, Bukidnon Province, trail to summit of Mt Kitanglad, 25-27 May 1984, Tan, Navarez & Amoroso 84-365 (BM, FH). Negros, Cuerno de Negros, 28 January 1937, Magdamo 1 (FH); Magdamo 12 (FH). Palawan, Mt Mantalingahan Range, Padparan, trail to Mt Inang Baboy Peak, 27 April 1979, Gruezo 5798 (FH). Syrrhopodon loreus (Sande Lac.) W.D. Reese in Phytologia 56: 306(1984). Fig. 21p-v. Calymperes loreum Sande Lac. in Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 13(2): 7 (1873). Type: Borneo, Labuan, Motley s.n. (L-holotype). Calymperes setifolium Hampe in Besch. in Ann.. Sci. Nat. Bot. sen 8, 1: 304 (1895). Type: Philippines, Basilan, Semper s.n. (BM!- holotype). Calymperes ebaloi E.B. Bartram in Farlowia 1: 505 (1944). Type: Philippines, Zamboanga Prov., Mindanao, Lilimbrog Mountain, near Kabasalan, Ebalo 664 (FH!-holotype). Shoots 3-5 cm high, with a short, obscure stem. Leaves 6->30(- >40) mm long, hair-like, erect, falcate or variously curving from a narrowly elliptical hyaline base; chlorophyllose limb notably con- stricted for a short distance above hyaline base; towards apex tapering to a long, fine, denticulate tip (Fig. 21p-s). Costa strong, excurrent or ending in leaf apex, largely smooth but often sparsely toothed near apex, superficial cells shortly rectangular, internally guide cells forming a single row between dorsal and ventral bands of stereids (Fig. 21u). Chlorophyllose lamina narrow, often undulate; from shoulders of leaf tapering proximally into the leaf base along either side of the hyaline lamina and gradually merging into the broad marginal ribs in the lower part of the leaf base; around shoulders of leaf cells towards margins slightly elongated, with the marginal row forming teeth or short spines (Fig. 21s, t); distally, from shoulders of leaf tapering or abruptly narrowing, virtually disappearing into costa, then broadening gradually above into long, linear limb (constricted region of lamina sometimes exceeding twice the length of the leaf base) ; towards apex tapering into the long, fine leaf tip; cells subquadrate to shortly rectangular (mostly longer than broad) or rounded, 5-15(-17) x 5-10(-12.5) [im, dorsally flat and smooth, ventrally flat to roundly protuberant (Fig. 21u). Hyaline lamina usually well-defined. Marginal ribs in proximal leaf base broad and flat, entire, composed of long, narrow, thick-walled cells, vanishing distally into chlorophyllose lamina; absent around leaf shoulders and below region of constricted lamina; above constric- tion narrow, polystratose, composed largely of a band of stereids adjacent to longitudinal rows of shortly to long rectangular chlorophyllose cells (sometimes forming a partial layer around the band of stereids) (Fig. 21g); some chlorophyllose cells forming large distant teeth, towards the leaf apex teeth becoming more closely set and sometimes double. Sporophytes reddish brown; seta 12-13 mm long; capsule 1.5-<2 mm long. HABITAT. Mostly occurring on tree trunks in lowland rainforest. DISTRIBUTION. An Indo-Pacific species. In the Philippines known from Luzon and Mindanao. SPECIMENS EXAMINED. Luzon, Isabela Province, Sitio Digoyo, 17 April 1991, 7an91-108 proparte (FH); Quezon Province, Real, National Botanical Garden, 4 August 1985, Tan & Wijangco 85-105 pro parte (BM). Syrrhopodon loreus is distinguishable from other species in the Calymperaceae with long leaves by the possession (in combination) of the following features: a) a sharply defined hyaline lamina (Fig. 39 21s); b) acute, thick-walled marginal teeth adjacent to the distal hyaline lamina (Fig. 21s, t); c) the constriction of the chlorophyllose lamina just above the hyaline base (Fig. 21q); and d) a single row of guide cells in the costa (Fig. 21u). Syrrhopodon muelleri (Dozy & Molk.) Sande Lac., Bryol. jav. 2: 224(1870). Fig. 22a-f. Calymperidium muelleri Dozy & Molk., Bryol. jav. 1: 51 (1856). Type: Java, Holle s.n. (BM!-isotypes?). Shoots up to >1.5 cm high, forming stiffly bristled tufts and mats. Stems very short, obscure. Leaves stiffly erect to erecto-patent, 6- >12 mm long, consisting of a linear chlorophyllose limb arising from a narrowly subelliptical hyaline base (limb sometimes spirally twisted when dry); apex obtuse (rarely acute), entire or with one or two small teeth (Fig. 22a, b). Costa ending in apex, usually smooth, above hyaline base sometimes some superficial cells projecting from their distal ends; internally with a single layer of guide cells (Fig. 22e); in upper leaf subtriangular in cross-section. Cells of chlorophyllose lamina quadrate to shortly rectangular or rounded- elliptical, 5-12.5 x 5-10 nm, obscured by a crown of low papillae or smooth, not protuberant (Fig. 22d, f)- Leaf margin from base to near apex formed by a thick, entire, polystratose rib; above leaf shoulders terete in cross-section, consisting of a strand of stereids largely enclosed by a superficial layer of chlorophyllose cells (subrectangular in surface view) (Fig. 22d, f); below shoulders becoming flattened, superficial cells linear or undifferentiated. Gemmae sometimes produced from the ventral surface of the costal apex. Seta 1 1-14 mm long; capsules 1-1.4 mm long, reddish brown. HABITAT. On tree trunks in lowland to mid-montane (c. 800 m) rainforest. DISTRIBUTION. An Indo-Pacific species. SPECIMENS EXAMINED. Catanduanes, 14 November 1917, Ramos 30609 (BM); Luzon, Isabela Province, Sitio Digoyo, 17April 1991, 7an91-108 pro parte (FH); Quezan Province, Real, National Botanical Garden, 4 August 1985, Tan & Wijangco 85-105 pro parte (FH); Quezon Province (Tayabas), Tagcauayan, Foxworthy & Ramos 13097 (BM); Mindanao, 1880, Montana s.n. (BM); Montana 192 (BM); Palawan, Mt Mantalingahan, between Nalpuan and Sandurapei, 26 April 1991, Tan 91-225 (BM); Palawan Prov- ince, Puerto Princesa, Barangay Irawan, Mt Malinao, 4 May 1993, Tan 93-336 (FH). Sibuyan, Magdiwang, Bo. Tampayan, Mt Giting-Giting, 21 May 1987, Tan & Hemaez 87-456 (FH). Syrrhopodon muelleri is distinguished from all other Malesian species by its erect, linear, largely entire leaves, with thick, subcylindrical marginal ribs composed of superficial chlorophyllose cells enclosing a core of stereids. Syrrhopodon parasiticus (Brid.) Besch. in Ann. Sci. Nat. Bot. se"r. 8,1:298(1895). Fig. 22g-k. Bryum parasiticum Brid., Muscol. recent. 2(3): 54 (1803). Type: Jamaica, Swartz s.n. (BM!-isotypes). Syrrhopodon wiemansii M. Fleisch., Muse. Fl. Buitenzorg 1: 204, 210 (1904). Type: Java, Tjipannas, Fleischer s.n. (FH, H- isotypes?). Calymperopsis wiemansii (M. Fleisch.) M. Fleisch. inBiblioth. Bot. 80:5(1913). Calymperopsis parasitica (Brid.) Broth., Nat. Pflanzenfam. 2nd ed., 10:235(1924). 40 L.T. ELLIS AND B.C. TAN g 1 a,g 1 mm b,c,l d-f, h-k 0.5 mm 50 jam Fig. 22 a-f. Syrrhopodon muelleri (Dozy & Molk.) Sande Lac. a-c: leaf (detail a: from base to above mid-leaf, b: apex, c: distal hyaline base; d: cells of chlorophyllose lamina with marginal rib in surface view; e, f: cross-sections of leaf (e: costa, f: chlorophyllose lamina with marginal rib), g-k. Syrrhopodon parasiticus Brid. g: leaf in ventral view with gemmae; h, i: cells of leaf in surface view (h: at margin in mid-leaf, i: in chlorophyllose lamina (dorsal view)); j, k: cross-sections of leaf (j: costa, k: chlorophyllose lamina and marginal rib. a-f Drawn from Tan 91-225 (BM). g-k Drawn from Sumatra, Jacobson 1 1393b (BM). Shoots 0.5-1.5 cm high, often densely matted with rhizoids below, forming mats. Leaves 3-4 mm long, consisting of an erect suboblong hyaline base narrowing slightly into a suberect to patent-reflexed (moist), Ungulate to lanceolate chlorophyllose limb; leaf apex broadly acute, entire to subdenticulate (Fig. 22g). Costa ending in apex; mostly smooth with a surface largely composed of stereids; dorsal surface from just above hyaline base to midleaf (or beyond) often composed of quadrate to shortly rectangular, subacutely protuberant cells (sometimes forming only one or two rows along either side of the ventral surface in this region) (Fig. 22j). Chlorophyllose lamina often incurved; cells isodiametric to slightly longer than broad, with 4-6 sides or rounded-elliptical, 5-12.5(-15) x 5-8(-10) M.ITI (Fig. 22i); ventrally acutely to subacutely protuberant (often tipped with a small papilla), dorsally unipapillose (Fig. 22k). Hyaline lamina sharply defined, with a long acute apex penetrating the chlorophyllose lamina. Leaf margins entire to subdenticulate; above hyaline base consisting of a thin, intermittent to continuous strand of stereids or substereids (Fig. 22h, k); in hyaline base consisting of a broad rib of linear, thin- to thick-walled cells (not continuous with strand in chlorophyllose limb but merging distally into the chlorophyllose lamina). Gemmae long, filamentous, uniseriate (Fig. 221); produced from rows of cells on either side of the ventral surface of the costa from slightly above the hyaline base to well beyond midleaf (Fig. 22g). Sporophytes rare, not seen in Philippine material. HABITAT. On trunks of trees. DISTRIBUTION. A nearly pantropical species. Tixier (1967) cites a Philippine specimen from Baguio in Luzon (as Calymperopsis wiemansii (M. Fleisch.) M. Fleisch.). No other local specimens examined. Syrrhopodon rufescens Hook. & Grev. in Edinburgh J. Sci. 3: 227 (1826). Type: Singapore, Wallich 2211 (BM!-holotype). Fig. 23a-e. Leucophanella rufescens (Hook. & Grev.) M. Fleisch., Muse. Fl. Buitenzorg 1: 200 (1904). Shoots l->2 cm high, densely leaved, forming pale green cushions, often loosely matted with red rhizoids below. Leaves 1.5->2 mm long, suberect to patent-recurved, lanceolate, V-shaped in cross- CALYMPERACEAE IN THE PHILIPPINES 41 a,g 0.5 mm b-e, h-j 50 jim 1 mm Fig. 23 a-e. Syrrhopodon rufescens Hook. & Grev. a, b: leaf (a: dorsal view, with b: detail of proximal chlorophyllose lamina); c-e: cross-sections of leaf in c: lower leaf, d: mid-leaf, and e: near apex, f-j Syrrhopodon spiculosus Hook. & Grev. f, g: leaf (f: ventral view, with g: detail of apex); h, i: cells of leaf (ventral surface) (h: in costa and blade of chlorophyllose limb, i: at margin of distal hyaline lamina); j: cross-section of chlorophyllose limb, a-e Drawn from Elmer 16843 (BM). f Drawn from Thailand, Kerr456 (BM). g-j Drawn from Peninsular Malaysia, Spare 1726 (BM). section, narrowing from above midleaf to an acute apex; largely composed of hyaline lamina (Fig. 23a). Costa thin, ending in apex, composed of a median layer of guide cells (about two cells wide) between dorsal and ventral bands of stereids, superficial cells differ- entiated (Fig. 23c-e), in distal leaf shortly subrectangular in surface view, many giving rise to teeth, toward leaf base elongated and smooth. Chlorophyllose lamina occupying less than one fifth of leaf length, narrowly to broadly tapering down either side of hyaline lamina, ceasing prior to midleaf, cells mostly quadrate to shortly rectangular, irregularly rounded-elliptical, mostly 6-12.5(-17.5) x 6-10 fjm (Fig. 23b); dorsally and ventrally flat or protruding suba- cutely, often with an apical papilla. Hyaline lamina sharply defined, cells large, thin-walled, seldom more than about 5-6 rows on either side of the costa. Leaf margins often recurved for a short distance above midleaf; from leaf base to apex entire, consisting of a narrow, smooth unistratose or bistratose strand of stereids. Seta red, smooth, c. 6->8 mm long; capsule erect, shortly cylindrical, c. 1 mm long, yellowish brown with a red rim; operculum long-rostrate; peristome teeth c. 150 urn long, usually with papillose longitudinal striations. HABITAT. On trees in lowland forest. DISTRIBUTION. A Malesian species. SPECIMEN EXAMINED. Luzon, August 1916, Elmer 16843 (BM, BM-K). Eddy (1990) points out the strong similarity between Syrrhopodon rufescens and 5. involutes. At least one Philippine specimen (Elmer 16843) possesses the features apparently definitive of 5. rufescens and is separable from the locally collected material of 5. involutus. However, the relationship between these taxa needs further study using fresh material with peristomes in a good conditon.. Syrrhopodon spiculosus Hook. & Grev. in Edinburgh J. Sci. 3: 226 (1825). Type: Singapore, Wallich [83 (H.1207)] (BM!-holotype; BM!-isotype). Fig.23f-j. Shoots mostly l-3(-4) cm high. Leaves 2->4 mm long, consisting of an erect, narrowly elliptical, semisheathing hyaline base rapidly narrowing distally into a patent to spreading linear chlorophyllose limb with a blunt, dentate apex (Fig. 23f, g). Costa ending just below 42 L.T. ELLIS AND B.C. TAN apex; above hyaline leaf base on dorsal and ventral surfaces with long, distally pointing, acute spines (often in subtransverse rows) (Fig. 23h). Cells of chlorophyllose lamina isodiametric to longer than broad, with 4-6 sides or rounded-elliptical, mostly 5-> 1 2 x 5-8 (im; drawn out dorsally and ventrally as erect, coronate-papillose projec- tions (towards leaf apex often becoming more spinose and curving distally) (Fig. 23h, j). Hyaline lamina sharply defined. Leaf margins towards and at apex dentate; from above shoulders of leaf to a short distance below apex often incurved, consisting of an entire poly stratose rib of stereids (Fig. 23h, j); from around shoulders of leaf to base composed of a flattened band of thick- walled, linear cells (continuous with rib in upper leaf), at shoulders giving rise to a row of long, acute spines (Fig. 23i), below shoulders entire. Sporophytes rare. Seta > 7 mm long; capsule cylindrical, c. 1-1 .5 mm long. HABITAT. On decaying logs, and trunks and twigs of trees in damp, shaded situations at low altitude. DISTRIBUTION. A palaeotropical species with sporadic distribu- tion. Reported for the Philippines by Bartram (1939), as a single collection from Panay. No other local specimens examined. Syrrhopodon tjibodensis M. Fleisch., Muse. Fl. Buitenzorg 1: 209 ( 1904).Type: Java, Tjibodas, 1450 m, September 1899, Fleischer (Musci Frond. Arch. Ind., ser. VI, exs. no. 261) (BM!-isotype). Fig. 24j-n. Syrrhopodon bartlettiiE.B. Bartram inPhilipp. J. Sci. 68: 82 (1939). Type: Philippines, Luzon, Mountain Prov., Baguio, 25 March 1935, Bartlett 13331 (FH!-holotype). Calymperopsis tjibodensis (M. Fleisch.) M. Fleisch. inBiblioth. Bot. 80:5(1913). Shoots 0.5->1.5 cm high, densely leaved, forming dense mats. Leaves erect to spreading (moist), mostly 2->3.5 mm long, consist- ing of a shortly subrectangular hyaline base that narrows slightly distally into a broadly linear-lanceolate chlorophyllose limb (often curved (inner edge ventral) and/or longitudinally folded, i.e. V- shaped in cross-section) (Fig. 24j); apex narrowly to broadly acute or apiculate, entire or with 1-2 small teeth. Costa ending in apex, smooth, dorsal and ventral surfaces formed by stereids (Fig. 24n). Cells of chlorophyllose lamina isodiametric to slightly longer than broad, with 4-6 sides or rounded-elliptical, mostly 7.5-15 x 7.5-1 1 fim; dorsally and ventrally slightly protuberant and strongly pleuripapillose (papillae often obscuring outlines of cells) (Fig. 24m, n). Hyaline lamina sharply defined with a somewhat truncate apex (Fig. 24k, 1). Leaf margins entire, adjacent to apex of hyaline lamina often narrowly recurved (Fig. 241); from below base of chlorophyllose limb to near apex consisting of a narrow rib of stereids (Fig. 24n); from between distal and mid-hyaline base to insertion composed of a flattened band of linear, thick-walled cells (about 3-5 cells wide, continuous with rib in upper leaf). Gemmae (fusiform-clavate) often produced from the ventral surface of the costa shortly above the apex of the hyaline lamina (Fig. 24k). Sporophytes apparently unknown. HABITAT. On trunks of trees. DISTRIBUTION. China, Indonesia (Java), Philippines. Bartram (1939) cites a specimen from Luzon (as Syrrhopodon bartlettii, see above). SPECIMEN EXAMINED. Palawan, St Paul Bay, St. Paul Subterranean National Park, 25 May 1989, Tan 89-1361 (FH). Syrrhopodon tjibodensis is very similar to S. parasiticus, but is most easily distinguished by the possession of pleuripapillose cells form- ing the chlorophyllose lamina (Fig. 24m, n). In 5. parasiticus these cells are ventrally protuberant (sometimes with a single papilla) and dorsally unipapillose (Fig. 22i, k). Syrrhopodon trachyphyllus Mont., Syll. Gen. Sp. Crypt. 47 ( 1 856). Type: Singapore, Gaudichaud, Hb. Montagne (BM!-isotypes). Fig. 24a-i. Syrrhopodon semperi Mull. Hal. in Linnaea 38: 557 (1874). Type: Philippines, Luzon, 1861, Semper s.n. (BM!-isotype). Shoots <0.5-1.5 cm high, forming dense, low mats. Stems some- times thickly matted with rhizoids towards base. Leaves about 2-3 mm long, consisting of a short, slightly tapering, broadly linear- rectangular, suberect to patent (when moist) chlorophyllose limb extending from a suberect, shortly oblong hyaline base; apex ob- tusely pointed to subcucullate (Fig. 24a-c). Costa ending just below or in apex, smooth; formed by dorsal and ventral strands of stereids separated by a unistratose layer of guide cells, superficial cells not differentiated (ie. surface formed by stereids) (Fig. 24h). Cells of chlorophyllose lamina mostly isodiametric to slightly longer than broad, with 4-6 sides, 10-15(-17.5) x 8-12.5 |im (Fig. 24g); ventrally protruding subacutely and crowned with papillae; dorsally flat to slightly convex, with a crown of papillae (Fig. 24i). Hyaline lamina sharply defined. Leaf margins at leaf apex denticulate; above hyaline base to near leaf apex often erect to incurved, with a thin (often intermittent), usually entire polystratose rib composed of stereids (Fig. 24i); adjacent to apex of hyaline lamina becoming unistratose, often weak, sometimes giving rise to acute teeth or spines (Fig. 24e, f); in leaf base entire, formed by a flattened band of linear, thick-walled cells (Fig. 24d). Sporophytes not seen. HABITAT. Mostly on tree trunks, sometimes on rock. DISTRIBUTION. A palaeotropical species. In the Philippines, ap- parently confined to Luzon. No local specimens examined, apart from the type of Syrrhopodon semperi. For a comparison of this species with Syrrhopodon armatus see under the description of the latter. Eddy (1990) illustrates two specimens with different features as Syrrhopodon trachyphyllus. One has features which conform to the above description and match those of the type specimen, the second possesses costae with a superficial layer of papillose chlorophyllose cells and cells in the chlorophyllose lamina apparently narrower than those described above. The material on which the latter illustra- tion is based may not represent 5. trachyphyllus and plants with its features have yet to be recorded for the Philippines. Syrrhopodon tristichus Nees ex Schwagr., Sp. muse, frond, suppl. 4: 31 Ib (1842). Type: Java, Hb. Nees (BM!-isotypes). Fig. 24o-t. Syrrhopodon macrotristichus Broth, in Leafl. Philipp. Bot. 2: 652 (1909). Type: Philippines, Luzon, Lucban, Tayabas Province, May 1907, Elmer 7713 (BM!-isotype, NY-isotype). Syrrhopodon victorianus E.B. Bartram in Be mice P. Bishop Mus. Occ. Pap. 19(11): 222 (1948). Type: Fiji, Viti Levu, Mount Victoria, Greenwood 1159 (FH-holotype). Shoots 2-9 cm long, slender, occasionally branched, often matted CALYMPERACEAE IN THE PHILIPPINES 43 b, c, k, 1, p, q d-i, m, n, r-t 1 mm 0.5 mm 50 urn Fig. 24 a-i. Syrrhopodon trachyphyllus Mont, a-c: leaf (a: in ventral view, with details of b: apex in ventral view with gemmae, and c: of lower leaf); d-g: cells of leaf in surface view (d: at margin in proximal hyaline base, e, f: at margin adjacent to apex of hyaline lamina, g: in chlorophyllose lamina); h, i: cross-sections of leaf (h: costa, i: chlorophyllose lamina and marginal rib), j-n. Syrrhopodon tjibodensis M. Fleisch. j-1: leaf (j: in ventral view, with detail of region around apex of hyaline lamina k: in ventral view with gemmae, and 1: in dorsal view showing recurved margin); m: cells of chlorophyllose lamina in surface view; n: cross-section of chlorophyllose limb. o-t. Syrrhopodon tristichus Nees ex Schwagr. o-q leaf (o: in lateral view, with details of p: apex, q: distal hyaline base); r, s: cross-sections of leaf (r: costa, s: chlorophyllose lamina and marginal rib); t: cells of chlorophyllose lamina in surface view, a-d, f-i Drawn from Sarawak, Bell 2008 (BM). e Drawn from Semper s.n. (BM). j-n Drawn from Tan 89-1361 (FH). o-t Drawn from Price s.n. (FH). 44 L.T. ELLIS AND B.C. TAN a 1 mm Fig. 25 a-e. Syrrhopodon prolifer var.? a: leaf; b, c: details of leaf surface (b: in chlorophyllose lamina, c: at margin of chlorophyllose lamina); d, e: cross- sections of chlorophyllose limb (d: costa, e: margin), f. Syrrhopodon prolifer var. papillosus (Mull. Hal.) W.D. Reese f: cross-section of costa in chlorophyllose limb. g. Syrrhopodon prolifer var. albidus (Thwait. & Mitt.) S. Orban & W.D. Reese g: cross-section of chlorophyllose limb. h. Syrrhopodon prolifer var. tosaensis (Cardot) S. Orban & W.D. Reese h: cross-section of costa in chlorophyllose limb, a-e Drawn from Tan & Hernaez 87-507 (BM). f Drawn from Venezuela, Funcke & Schlim 358 (BM). g Drawn from Sri Lanka, Thwaites 53 (BM). h Drawn from Japan, Iwatsuki, Sharp & Sharp 1044(BM). with rhizoids below. Leaves in three lax ranks (obscured by flexing of leaf limbs when dry), 4-1 1 mm long, bristle-like, mostly patent to recurved from an erect, long, narrowly oblong hyaline base; apex dentate, long and finely drawn out (Fig. 24o, p). Costa ending just short of leaf apex; superficial cells shortly to long rectangular, mostly smooth and flat, or with slightly projecting distal ends, sometimes forming teeth close to leaf apex; internally with a single row of guide cells between dorsal and ventral layers of stereids (Fig. 24r). Chlorophyllose lamina incurved to above mid-limb, forming a channel; composed of quadrate to subrectangular or irregularly rounded to elliptical, thick-walled cells, isodiametric or longer than broad, 8-17(-20) x 6-10 mm (Fig. 24t), each with a crown of low papillae on the dorsal and ventral surfaces (sometimes poorly devel- oped) (Fig. 24s). Marginal ribs strong, continuous from leaf base to shortly below apex, a polystratose band of stereid or substereid cells; in proximal leaf base flattened and entire; around shoulders of leaf becoming thicker and more rounded in section (commonly reaching 30-38 mm wide); from above shoulders towards leaf apex gradually becoming thinner and more flattened (Fig. 24s); distally (from about mid-limb) giving rise to acute, forward pointing, often double teeth, towards leaf apex these are usually more closely set and more commonly single. Sporophytes uncommon; seta 6-7 mm long; capsule 1.5-1 .8 mm long. HABITAT. Mostly occurring on tree trunks in rainforest. DISTRIBUTION. A palaeotropical species. SPECIMENS EXAMINED. Basilan, Sempers.n. (in Hb. Hampe, BM). Luzon, Rizal Province, April 1914, Loher 15164 (BM). Mindanao, Agusan Norte Province, Mt Hilong-Hilong, May-June 1984, Tan & Navarez 84-503 (BM). Mindoro,Baco,MtHalcon, 17April 1987,7an87-l 17(BM); ISApril 1987, Tan 87-136 (BM). Negros, Negros Oriental Province, Dumaguete, Cuernos Mts, June 1908, Elmer 91 6% (BM); May 1908, Elmer 9^6 (BM). Palawan, Motley s.n. (in Hb. Hooker, BM-K). Panay, Mt. Madyaas, 15-16 January 1987, Price s.n. (FH). Sibuyan, northern slope of Mt Citing Citing, near Mayo's Peak, 22 May 1987, Tan & Hernaez 87-509 (FH). The leaves of Syrrhopodon tristichus have a superficial resemblance to those ofS.flammeonervis Mull. Hal. In the latter the surface of the costa is undifferentiated, composed of stereids (Fig. 19f); in S. tristichus the costa possesses a superficial layer of rectangular chlorophyllose cells (surface view). Excluded taxa Mitthyridium luteum (Mitt.) H. Rob. - See under M. papuanum. Mitthyridium undulatum (Dozy & Molk.) H. Rob. - There are no reliable records of M. undulatum from the Philippines. The speci- mens cited under this name by Bartram (1939) have subsequently been redetermined, mostly as Mitthyridium fasciculatum. Three other taxa, Calymperes crassinerve (Mitt.) A. Jaeger*, Mitthyridium junquilianum (Mitt.) H. Rob.* and Syrrhopodon prolifer var. albidus (Thwait. & Mitt.) S. Orban & W.D. Reese* have had the Philippines either mentioned in the discussion of a species range, or included in maps showing the geographical range of a species (Mohamed & Reese, 1985; Reese, 1987a, 19876; Reese, Mohamed & Mohamed, 1986). There is no mention of a definitive Philippine specimen in these publications. Although the distribution pattern of these species suggests a high probability of their occur- rence in the Philippines, we have opted to exclude them in this treatment owing to the lack of a voucher specimen documenting their presence in the country. Syrrhopodon prolifer Schwagr. var. ? - Syrrhopodon prolifer, with its many varieties, was revised by Orban & Reese (1990). A collec- tion from Sibuyan Island, Tan & Hernaez 87-507 (BM), resembles a small form of the neotropical endemic S. prolifer var. papillosus CALYMPERACEAE IN THE PHILIPPINES 45 (Mull. Hal.) W.D. Reese. Shoots of Tan & Hernaez 87-507 have erect to recurved, green, linear leaves, about 3-5 mm long. The cells forming the ventral surface of the costa above the hyaline base are subquadrate to shortly subrectangular or rounded (in surface view) with compound papillae (Fig. 25d); cells in the chlorophyllose lamina (mostly 10-18(-22.5) x 10-12.5(-15) um) are similarly crowned on their dorsal and ventral surfaces with compound papil- lae (Fig. 25b, e). The leaf margins adjacent to the apex of the hyaline lamina are entire to distantly denticulate. Syrrhopodon prolifer var. papillosus possesses erect to recurved, linear leaves, up to >7 mm long with a reddish brown costa. The cells forming the ventral surface of the costa above the hyaline base (Fig. 25f), and those forming the chlorophyllose lamina, are similar to those in Tan & Hernaez 87-507 (except the cells of the chlorophyllose lamina have slightly thicker walls). The leaf margins adjacent to the apex of the hyaline lamina are most often entire. Syrrhopodon prolifer var. albidus (Thwait. & Mitt.) S. Orban & W.D. Reese and 5. prolifer var. tosaensis (Cardot) S. Orban & W.D. Reese have an Asian distribution. In these varieties (and most others) the ventral surface of the costa is smooth (except at the leaf apex) and composed of stereids (Fig. 25 g, h). Leaves of var. tosaensis are about 2—4 mm long, generally smaller than those in Tan & Hernaez 87- 507, but possess similar laminal papillae. According to Orban & Reese (1990), in var. tosaensis, the leaf margins adjacent to the apex of the hyaline lamina are consistently toothed. In var. albidus the leaves are 3-6 mm long, with entire leaf margins. The cells of the chlorophyllose lamina are similar in size and ornamentation to those of Tan & Hernaez 87-507, but are relatively thick-walled. Tan & Hernaez 87-507 cannot be readily identified as any of the presently recognized varieties of Syrrhopodon prolifer. However, the authors are reluctant to describe a new variety on the basis of a single specimen. ACKNOWLEDGEMENTS. We are grateful to the directors and staff of FH, H, L, NY and PC for the loan of specimens, and thank Dr A.J. Harrington (BM) and Professor W.D. Reese (LAP) for their valuable comments on the manu- script for this paper. REFERENCES Akiyama, H. & Reese, W.D. 1 993. Taxonomic studies of mosses of Seram and Ambon (Moluccas, East Malesia) collected by Indonesian-Japanese botanical expeditions. VI Calymperaceae. Journal of the Faculty of Science, University of Tokyo, Section in, 15(3): 199-218. Bartram, E.B. 1939. Mosses of the Philippines. Philippine Journal of Science 69(1- 4): 1^25. Eddy, A. 1990. A handbook ofMalesian mosses. 2. London. Edwards, S.R. 1980. Spore discharge in Calymperes. Journal of Bryology 11: 95-97. Ellis, L.T. 1987. Taxonomic notes on Calymperes. Journal of Bryology 14: 681-690. 1988. Taxonomic notes on Calymperes II. Journal of Bryology 15: 127-140. 1989. A taxonomic revision of Calymperes in southern India and neighbouring islands. Journal of Bryology 15: 697-732. 1991. Calymperes schmidtii Broth, in J. Schmidt and C. subintegrum Broth, in J. Schmidt, two distinct species from Malesia. Journal of Bryology 16: 589-593. Iwatsuki, Z. & Tan, B.C. 1979. Checklist of Philippine mosses. Kalikasan, The Philippine Journal of Biology 8(2): 179-210. 1980. Noteworthy Philippine mosses at the Hattori Botanical Laboratory (NICH), Japan. Kalikasan, The Philippine Journal of Biology 9(2-3): 267-282. Menzel, M. & Schultze-Motel, W. 1990. The bryophytes of Sabah (North Borneo) with special reference to the BRYOTROP transect of Mount Kinabalu. XI Calymperaceae (Bryopsida). Willdenowia 19: 475-542. Mohamed, H. & Reese, W.D. 1985. Syrrhopodon (Musci: Calymperaceae) in Malay- sia and adjacent regions. Bryologist 88: 223-254. 1992. Syrrhopodon stonae, new to Malaya, and notes on other Malayan Calymperaceae. Bryologist 95(1): 88-91. Miiller, C. 1856. Review of: Dozy, F & Molkenboer, J.H. 'Bryologia javanica (Fascicle VI: 1855, Fascicle VII-X: 1856)'. Botanische Zeitung 15: 530-535. Nowak, H. 1980. Mitthyridium in Ozeanien. Bryophytorum Bibliotheca 20: 1-236. Orban, S. & Reese, W.D. 1990. Syrrhopodon prolifer (Musci: Calymperaceae): a world view. Bryologist 93(4): 438 444. Reese, W.D. 1987a. Calymperes (Musci: Calymperaceae): world ranges, implications for patterns of historical dispersion and speciation, and comments on phylogeny. Brittonia 39: 225-237. 1 987b. World ranges, implications for patterns of historical dispersal and speciation, and comments on phylogeny of Syrrhopodon (Calymperaceae). Memoirs of the New York Botanical Garden 45: 426-445. 1993. Calymperaceae. Flora Neotropica 58: 1-102. 1994. The subgenera of Mitthyridium (Musci). Journal of the Hattori Botanical Laboratory 75: 41^4. & Bartlett, JJt 1982. Syrrhopodon fimbriatulus C. Mull., and the family Calymperaceae (Musci), new to New Zealand Island Territories. Journal of Bryology 12: 209-214. Koponen, T. & Norris, D.H. 1986. Bryophyte flora of the Huon Peninsula, Papua New Guinea. XIX. Calymperes, Syrrhopodon, Mitthyridium (Calymperaceae, Musci). Acta Botanica Fennica 133: 151-202. & Mohamed, H. 1985. A synopsis of Calymperes (Musci: Calymperaceae) in Malaysia and adjacent regions. Bryologist 88: 98-109. & Mohamed, A.D. 1986. A synopsis of Mitthyridium (Musci: Calymperaceae) in Malaysia and adjacent regions. Bryologist 89: 49-58. & Stone, I.G. 1987. New records of Australian Calymperaceae and keys to Australian species of Calymperes, Mitthyridium and Syrrhopodon. Journal of Bryology 14(3): 487^94. 1995. The Calymperaceae of Australia. Journal of the Hattori Botanical Laboratory 78: 1-40. & Streimann, H. 1994. Calymperes subserratum (Musci), new to eastern Malesia, with notes on C. serratum and C. subulatum. Bryologist 97(1): 80-82. Tan, B.C. 1996. Biogeography of Palawan mosses. Australian Systematic Botany 9: 193-203. & Iwatsuki, Z. 1991. A new annotated Philippine moss checklist Harvard Papers in Botany 3: 1-64. Tixier, P. 1967. Le genre Calymperopsis (C. M.) Fleisch. (Calymperaceae). Revue Bryol. Lichenol., N.S. 35(1^): 265-302. * These taxa are covered in the key. 46 L.T. ELLIS AND B.C. TAN SYSTEMATIC INDEX Accepted names are in roman and synonyms in italic; new names are in bold. Bryum parasiticum Brid. 39 Calymperes Sw. ex Web. 2, 5 Calymperes aeruginosum Hampe ex Sande Lac. 3, 5, 6 Calymperes afzelii Sw. 3, 5, 6 Calymperes boulayi Besch. 3, 7, 8 Calymperes clemensiae Broth. 1 8 Calymperes constrictum Sull. 22 Calymperes crassinerve (Mitt.) A. Jaeger 2, 3, 44 Calymperes datense E.B. Bartram 37 Calymperes dozyanum sensu M. Fleisch. 7 Calymperes ebaloi E.B. Bartram 39 Calymperes erosum Mull. Hal. 3, 7, 8 Calymperes fasciculatum Dozy & Molk. 3, 7, 9 Calymperes gardneri Hook. 34 Calymperes graeffeanum Mull. Hal. 3, 9, 10 Calymperes hampei Dozy & Molk. 7 Calymperes hasagawae (Tak. & Iwats.) Iwats. 8 Calvmperes h\ophilaceum Mull. Hal. ex Besch. 10 var. robustum M. Fleisch. 10 Calymperes japonicus Besch. 37 Calymperes johannis-winkleri Broth. 7 var. hasagawae (Tak. & Iwats.) Iwats. 8 Calymperes lonchophyllum Schwagr. 3, 5, 10 Calymperes loreum Sande Lac. 39 Calymperes mammosum Besch. 5 Calymperes mangalorense Dixon & P. de la Varde 3, 10 Calymperes moluccense Schwagr. 3, 12 Calymperes motley i Mitt, ex Dozy & Molk. 3, 12, 13 Calymperes orientale Mitt, ex Besch. 20 var. polytrichoides M. Fleisch. 20 Calymperes palisotii subsp. moluccense (Schwagr.) M. Menzel 12 Calymperes peguense Besch. 2 1 Calymperes porrectum Mitt. 3, 13 Calymperes ramosii Broth. 2 1 Calymperes robinsonii B.C. Tan & W.D. Reese 3, 14, 15 Calymperes salakense Besch. 13 Calymperes sandeanum Besch. 7 Calymperes scalare Besch. 1 3 Calymperes schmidtii Broth. 18 Calymperes semperi Hampe in Besch. 10 Calymperes serratum A. Braun ex Mull. Hal. 2, 15, 16 Calymperes setifolium Hampe in Besch. 39 Calymperes strictifolium (Mitt.) G. Roth 2, 17 Calymperes subintegrum Broth. 3, 17, 18 Calymperes subserratum M. Fleisch. 2, 18, 19 Calymperes taitense (Sull.) Mitt. 3, 18, 19 Calymperes tenerum Mull. Hal. 3, 19, 20 Calymperes tuberculosum (Ther. & Dixon) Broth. 17 Calymperes vriesii Besch. 5 Calymperidium croceum (Mitt.) M. Fleisch. 33 Calymperidium muelleri Dozy & Molk. 39 Calymperopsis parasitica (Brid.) Broth. 39 Calymperopsis tjibodensis (M. Fleisch.) M. Fleisch. 42 Calymperopsis wiemansii (M. Fleisch.) M. Fleisch. 39 Chameleion L.T. Ellis & A. Eddy in A. Eddy 2, 20 Chameleion cryptocarpos (Dozy & Molk.) L.T. Ellis & A. Eddy 20 Chameleion peguense (Besch.) L.T. Ellis & A. Eddy 3,21,22 Codonoblepharum subluteum Mull. Hal. 27 Exostratum blumei (Hampe) L.T. Ellis 4 Exostratum sullivantii (Dozy & Molk.) L.T. Ellis 4 Heliconema (Mitt.) L.T. Ellis & A. Eddy 20 Heliconema peguense (Besch.) L.T. Ellis & A. Eddy 21 Leucophanella borneense (Hampe) M. Fleisch. 37 Leucophanella revoluta (Dozy & Molk.) M. Fleisch. 37 Leucophanella rufescens (Hook. & Grev.) M. Fleisch. 40 Mitthyridium H. Rob. 2, 21 Mitthyridium cardotii (M. Fleisch.) H. Rob. 25 Mitthyridium constrictum (Sull.) H. Rob. 4, 22, 23 Mitthyridium fasciculatum (Hook. & Grev.) H. Rob. 21,24 subsp. cardotii (M. Fleisch.) B.C. Tan & L.T. Ellis 4, 24,25 var. cardotii (M. Fleisch.) A. Eddy 25 subsp. fasciculatum 4, 25 subsp. obtusifolium (Lindb.) M. Menzel 4, 24, 25 Mitthyridium flavum (Mull. Hal.) H. Rob. 4, 23, 25 Mitthyridium iwatsukianum B.C. Tan 4, 23, 26 Mitthyridium junquilianum (Mitt.) H. Rob. 2, 4, 23, 27,44 Mitthyridium leucoloma (Mull. Hal.) H. Rob. 24 Mitthyridium luteum (Mitt.) H. Rob. 26, 44 Mitthyridium obtusifolium (Lindb.) H. Rob. 25 Mitthyridium papuanum (Broth.) H. Rob. 5, 26, 28 Mitthyridium repens (Harv.) H. Rob. 4, 27, 28 Mitthyridium retusum (Besch.) W.D. Reese 2 Mitthyridium subluteum (Mull. Hal.) H. Nowak 4, 23, 27 Mitthyridium undulatum (Dozy & Molk.) H. Rob. 44 Mitthyridium wallisii (Mull. Hal.) H. Rob. 4, 27, 29 var. wallisii 28 var. crassum (Broth.) M. Menzel 28 Syrrhopodon Schwagr. 2, 28 Syrrhopodon albovaginatus Schwagr. 4, 29, 30 Syrrhopodon amoenus Broth. 32 Syrrhopodon aristifolius Mitt. 2, 30, 3 1 Syrrhopodon armatus Mitt. 4, 31, 35 Syrrhopodon bartlettii E.B. Bartram 42 Syrrhopodon borneensis (Hampe) A. Jaeger 37 Syrrhopodon ciliatus (Hook.) Schwagr. 2, 30, 32 Syrrhopodon codonoblepharoides Mull. Hal. ex M. Fleisch. 24 Syrrhopodon confertus Sande Lac. 2, 32, 33 Syrrhopodon croceus Mitt. 2, 33 Syrrhopodon curranii Broth. 34 Syrrhopodon fallax Sande Lac. 30 Syrrhopodon fasciculatus Hook. & Grev. 24 Syrrhopodon fimbriatulus Mull. Hal. 31 Syrrhopodon fimbriatus Mull. Hal. 31 Syrrhopodon flammeonervis Mull. Hal. 4, 34, 35 var. robustus Dixon 34 Syrrhopodon flavus Mull. Hal. 25 Syrrhopodon gardneri (Hook.) Schwagr. 4, 28, 34, 36 Syrrhopodon hasagawae Tak. & Iwats. 7 Syrrhopodon hispidissimus Dixon 4, 35, 36 Syrrhopodon hispido-ciliatus E.B. Bartram 35 Syrrhopodon involutus Schwagr. 2, 37, 38 Syrrhopodon japonicus (Besch.) Broth. 4, 37, 38 Syrrhopodon larminatii Broth. & Paris 3 1 Syrrhopodon leucoloma Mull. Hal. 24 Syrrhopodon loreus (Sande Lac.) W.D. Reese 4, 38, 39 Syrrhopodon luzonensis R.S. Williams 25 Syrrhopodon macrotristichus Broth. 42 Syrrhopodon micholitzii Mull. Hal. 29 Syrrhopodon muelleri (Dozy & Molk.) Sande Lac. 3, 39,40 Syrrhopodon obtusifolium Lindb. 25 Syrrhopodon papuanus Broth. 26 Syrrhopodon parasiticus (Brid.) Besch. 4, 39, 40 Syrrhopodon peguense (Besch.) W.D. Reese 21 Syrrhopodon perarmatus Broth. 35 Syrrhopodon perarmatus E.B. Bartram 35 Syrrhopodon philippinensis E.B Bartram 32 Syrrhopodon prolifer Schwagr. 44, 45 var. albidus (Thwait. & Mitt.) S. Orban & W.D. Reese 2, 4, 44, 45 var. papillosus (Mull. Hal.) W.D. Reese 44, 45 var. tosaensis (Cardot) S. Orban & W.D. Reese 2, 4, 44,45 Syrrhopodon repens Harv. 27 Syrrhopodon revolutus Dozy & Molk. 37 Syrrhopodon rufescens Hook. & Grev. 2, 40, 41 Syrrhopodon semperi Mull. Hal. 42 Syrrhopodon spiculosus Hook. & Grev. 4, 41 Syrrhopodon strictifolius Mitt. 17 Syrrhopodon subulatus Sande Lac. 30 Syrrhopodon taitense Sull. 20 Syrrhopodon tenellum Dozy & Molk. 26 Syrrhopodon tjibodensis M. Fleisch. 4, 42, 43 Syrrhopodon trachyphyllus Mont. 4, 42, 43 Syrrhopodon tristichus Nees ex Schwagr. 3, 42, 43 Syrrhopodon tsushimae Cardot 3 1 Syrrhopodon tuberculosus Ther. & Dixon 17 Syrrhopodon victorianus E.B. Bartram 42 Syrrhopodon wallisii Mull. Hal. 27 Syrrhopodon wiemansii M. Fleisch. 39 Thyridium Mitt. 21 Thyridium cardotii M. Fleisch. 25 Thyridium fasciculatum (Hook. & Grev.) Mitt. 24 Thyridium flavum (Mull. Hal.) M. Fleisch. 25 Thyridium luzonensis (R.S. Williams) Broth. 25 Thyridium papuanum (Broth.) M. Fleisch. 26 Thyridium obtusifolium (Lindb.) M. Fleisch. 25 Thyridium repens (Harv.) Mitt. 27 Trachymitrium borneense Hampe 37 Weissia ciliata Hook. 32 Bull. nat. Hist. Mus. Land. (Bot.) 29( 1 ): 47-79 Issued 24 June 1 999 Revision of Hibiscus section Furcaria (Malvaceae) in Africa and Asia F. DOUGLAS WILSON c/o USDA-ARS, Western Cotton Research Laboratory, 4135 E. Broadway, Rd., Phoenix, AZ 85040-8830, U.S.A. CONTENTS Introduction 48 Materials and methods 48 Genome relationships 48 Natural hybrids and introgressant forms 49 The Indian species of Hibiscus section Furcaria 49 Taxonomic treatment 49 Description of Hibiscus section Furcaria 49 Key to African and selected Asian taxa of Hibiscus section Furcaria 50 1. Hibiscus surattensis L 51 2. Hibiscus altissimus Hornby 53 3. Hibiscus furcellatoides Hochr. 53 4. Hibiscus rostellatus Guill. & Perr. 53 5. Hibiscus hispidissimus Griff 55 6. Hibiscus goossensii (Hauman) F.D. Wilson 56 7. Hibiscus furcatus Willd 56 8. Hibiscus sudanensis Hochr. 57 9. Hibiscus flavo-roseus Baker f. 57 10. Hibiscus radiatus Cav 59 11. Hibiscus torrei Baker f. 59 12. Hibiscus noldeae Baker f 59 13. Hibiscus nigricaulis Baker f. 60 14. Hibiscus mechowii Garcke 61 15. Hibiscus subdiversifolius Hochr. 61 16. Hibiscus berberidifolius A. Rich 61 17. Hibiscus sparseaculeatus Baker f 62 18. Hibiscus greenwayi Baker f 62 19. Hibiscus partitus (Hochr.) F.D. Wilson 63 20. Hibiscus diversifolius Jacq 63 20a. Hibiscus diversifolius subsp. diversifolius 63 20b. Hibiscus diversifolius subsp. rivularis (Bremek. & Oberm.) Exell 64 21. Hibiscus reekmansii F.D. Wilson 64 22. Hibiscus sabdariffaL 65 23. Hibiscus cannabinus L 66 24. Hibiscus parvilobus F.D. Wilson 67 25. Hibiscus asperHook.f 68 26. Hibiscus acetosella Welw. ex Hiem 68 27. Hibiscus gilletii de Wild 69 27a. Hibiscus gilletii subsp. gilletii 6< 27b. Hibiscus gilletii subsp. hiernianus (Exell & Mendonc.a) F.D. Wilson 71 27c. Hibiscus gilletii subsp. lundaensis (Baker f.) F.D. Wilson 71 28. Hibiscus moxicoensis Baker f 71 29. Hibiscus cuanzensis Exell & Mendonsa 71 30. Hibiscus mastersianus Hiern '' 31. Hibiscus elongatifolius Hochr. 7^ 32. Hibiscus sineaculeatus F.D. Wilson 72 33. Hibiscus scotellii Baker f. 7- Poorly known and excluded species '- References Index... 79 © The Natural History Museum, 1999 48 F.D. WILSON SYNOPSIS. Hibiscus section Furcaria (Malvaceae) in Africa and Asia is revised. Three new species are described: Hibiscus parvilobus F.D. Wilson from Kenya, Hibiscus reekmansii F.D. Wilson from eastern Democratic Republic of Congo, Rwanda, Burundi, and western Tanzania, and Hibiscus sineaculeatus F.D. Wilson from Nigeria. Four taxa represent new combinations: Hibiscus goossensii (Hauman) F.D. Wilson, Hibiscus partitas (Hochr.) F.D. Wilson, Hibiscus gilletii subsp. hiernianus (Exell & Mendon^a) F.D. Wilson and Hibiscus gilletii subsp. lundaensis (Baker f.) F.D. Wilson. Hibiscus rostellatus Guill. & Perr. and Hibiscus persicifolius Eckl. & Zeyh. are lectotypified. INTRODUCTION This treatment of Hibiscus section Furcaria DC. emend. Hochr. is intended to cover all of the known African taxa. The key also includes the Asian species which I have examined but does not include the two Indian species H. hoshiarpurensis T.K. Paul & M.P. Nayar and the (previously undescribed?) species of Bahadur et al. (1970) because herbarium material was not available for study. Hibiscus section Furcaria is a natural group of over 100 known species, set apart from species in other sections of the large genus Hibiscus by having a calyx with a thickened, rigid midrib and two thickened marginal (or near-marginal) ribs (Wilson, 1994). The sectional name refers to the bifurcate apex of the involucellar bracteoles (epicalyx) of some species. Species are distributed mostly in tropical and subtropical areas of the world but a few penetrate also into the temperate zone (to c. 35°N in North America and 35°S in Australia). Species occur in subsaharan Africa and associated is- lands, on the Indian subcontinent, southeastern Asia, Australia, islands of the Pacific Basin (including New Zealand), the West Indies, and in North, Central and South America. They occur in a variety of habitats, from grasslands to savannas, forests, and marshes. Some are attuned to a wet-dry precipitation cycle, avoiding the dry cycle by either having an annual habit or going dormant. Others are adapted to a fire climax, in which new shoots arise from a protected crown after a fire. Still others are adapted to a more-or-less uniform rainfall regime; some prefer marshy environments (e.g. H. diversifolius). Most are frost-sensitive but at least one species, H. aculeatus Walter, is adapted to the warm-cold seasonal cycle of the Gulf and Atlantic coastal plains of the southeastern United States. Growth habits vary from small, herbaceous annuals to subshrubs, shrubs, vines, and trees. Species have showy, delicate flowers which vary considerably in size; some have potential as ornamentals. Flower colours vary from pure white to various shades of cream, yellow, pink, red, lavender, and purple. Flowers of all species have a red to purple centre (basal petal spot), although mutants have been found which lack this feature. Flowers are adapted to a variety of pollinators; some have rotate, fully opened flowers, others have tubular, pendent flowers. Some have included stigmas and are obviously well adapted for self pollination. Others have exserted stigmas which require cross polli- nation. The red salverform flower of one Mexican species, Hibiscus uncinellus DC., presumably is adapted to hummingbird pollination. The two most important species of commerce are Hibiscus cannabinus, 'kenaf, and//, sabdariffa, 'roselle' (Wilson & Menzel, 1964). Kenaf is a rapidly growing, tall annual fibre plant. The bast (bark) fibres are used mainly for cordage, including twine, rope, and burlap. The bast and woody fibres together are used for paper-pulp, and the woody fibres are used for poultry litter, packing materials, and mulch (Taylor, 1992). Other potential uses include animal bedding, an oil and water sorbent, a horticultural medium, an enhancer of bioremediation in breaking down pollutants, construc- tion boards, textile fibres for blending with other fibres, a component of moldable fibre/resin composites, and as mats to protect forest seedlings (Goforth & Fuller, 1994; Werber, 1993). Young kenaf plants afford a nutritious livestock forage (Perry et al., 1993). Leaves, flowers, and seeds are used for food, the seed oil has various food uses and various parts of the plant are used in folk medicines (Dalziel, 1948; Duke & duCellier, 1993). The commercial cultivars of roselle fall into one of two categories, edible or fibre. The edible cultivars are small, branched annuals which possess fleshy calyces used for making sauces and drinks and for a number of other food uses. Various parts of the plant are used in folk medicines. The fibre cultivars are tall, unbranched annuals from which the bast fibre is extracted for cordage (Crane, 1949; Dalziel, 1948; Duke & duCellier, 1993). A number of species other than kenaf and roselle are also used as fibre, food or medicines by native peoples. The leaves, particularly when young, have an agree- able acid taste (e.g. Hibiscus acetoselld) and are used as salad or cooked as spinach (Dalziel, 1948). MATERIALS AND METHODS Herbarium specimens and living plants in nature, experimental gardens, and glasshouses formed the basis for this study. The late Margaret Y. Menzel and I, at various times, borrowed herbarium material or solicited assistance from the curators of the following herbaria (Holmgren et al., 1990): ASU, B, BM, BOL, BR, BSD, CAL, EA, FSU, G, K, LISC, LISU, MEL, NA, NBG, P, PAL, PRE, S, SRGH, TCD, TEX. I studied living plants in their native habitats in Kenya and Tanzania (Wilson, 1978) and deposited vouchers of 63 accessions in one or more of the following herbaria: ASU, EA, NA. Plants grown from seeds were studied in experimental gardens and glasshouses at USDA-ARS Research Laboratories at Lake Worth, Palm Beach Co., Florida, and at Phoenix, Maricopa Co., Arizona and also at Florida State University, Tallahassee, Leon Co. Over 60000 interspecific pollinations were made and more than 140 different interspecific hybrids were produced (Menzel, 1986). Herbarium specimens examined are listed, insofar as possible, from west to east and from north to south under each taxon. Distribution maps were prepared for all of the African taxa except for those that are mainly cultivated (Hibiscus acetosella, H. radiatus, H. sabdariffa). GENOME RELATIONSHIPS The basic chromosome number for the section is x = 18. Diploid, tetraploid, hexaploid, octoploid, and decaploid species exist in nature (Menzel & Wilson, 1969; Wilson, 1994).Allofthepolyploid species studied thus far are allopolyploid. Thirteen genomes have been confirmed or postulated (A, B, C, D, E, G, H, J, P, R, V, X, Y), either through the study of chromosome pairing in interspecific hybrids or through electrophoresis (Menzel, 1986; Wilson, 1994). The greatest genomic diversity is found in subsaharan Africa; nine of the 13 genomes are represented (A, B, C, D, E, G, H, X, Y) and nine of the 10 confirmed diploid species occur there (all except Hibiscus costatus A. Rich, from Mexico, Central America, and the Caribbean). Tetraploid and octoploid species occur also in Africa. REVISION OF HIBISCUS SECTION FURCARIA Diploid, tetraploid, and octoploid species occur in India and south- eastern Asia. The eight confirmed diploid species carry genomes designated as follows: 'A' (H. asper, H. cannabinus, and H. greenwayi); 'B' (H. surattensis); 'G' (H. sudanensis); 'X' (H. gilletii subsp. hiernianus, H. mastersianus); 'Y' (H. mechowii). The ninth diploid, H. berberidifolius, carries an unknown genome, presum- ably different from the other African diploid genomes. Analysed African tetraploids carry 'AB' (H. acetosella), 'AX' (H. nigricaulis), 'AY' (H. sabdariffa), and 'GH' (H. rostellatus). Analysed or partially analysed African octoploid species carry 'CDE' and a modified 'G' (H. diversifolius), or 'BG' and two unidentified genomes (H. altissimus). Hibiscus surattensis, the African 'B' genome diploid, is found also in India and southeastern Asia. Hibiscus diversifolius, the 'CDEG' octoploid, is found in some parts of southeastern Asia. The only two other analysed Indian species are the 'AB' tetraploid H. radiatus and the octoploid, H. hispidissimus, which carries 'BG' and two unidentified genomes. The diploid African and Asian species display characteristic morphological patterns based upon the four major diagnostic char- acters used in the key and correlated with their genome structure. The four major diagnostic characters are as follows: 1) stems aculeate (+) or not aculeate (-); 2) peduncles articulating nearer the summit (+) or nearer the base (-); 3) involucellar bracteoles forked (bifurcate) (+) or simple (-); 4) nectary on the calyx midrib present (+) or absent (-). The patterns are: 'A ':(+--+); 'B ':(+ + + -); 'G ': (+- + -); 'X': (-- + -); '¥': (+ ). In Hibiscus berberidifolius, the African diploid with an unknown genome, the most common pattern is the same as that of the ' Y' genome H. mechowii, but some plants, having a calyx nectary, show the 'A' pattern. NATURAL HYBRIDS AND INTROGRESSANT FORMS I examined a number of sheets which I and/or others interpreted as natural hybrids or introgressant forms. This interpretation is of course subjective, but my extensive experience with experimental hybrids assisted me materially. The most common hybrid or introgressant forms involved the closely related species Hibiscus asper and H. cannabinus (both 'A genome diploids), interpreted as follows: H. asper x cannabinus: Angola, Welwitsch 5268 (BM); Democratic Republic of Congo, Symoens 7321 (BR), Vanderyst 1833 (BR); Chad,Fotius 1081 (K),Palayer42l (P); Senegal, fler/witf 4551 (P). H. asper introgressed with H. cannabinus: Mali, Chevalier 24919 (P); Guinea, Berhaut 4175 (P); Niger, Boudet 4980 (P); Nigeria, Latilo & Eimuenze FHI 6540 (K). H. cannabinus introgressed with H. asper. Tanzania, Schlieben 2218 (BM); Ethiopia, Welby s.n. (P). Other putative hybrids and introgressive types were interpreted as follows: Hibiscus asper x noldeae(7): Democratic Republic of Congo, DeLoose 203 (BM); Cameroon, deWilde & deWilde-Duyjjes 3751 (BR). H. cannabinus x nigricaulis(l): Mozambique, Barbosa 1086 (BM, LISC); Garcia 490A (LISC). [NOTE. Barbosa 1086 at BR is H. nigricaulis]; [see Gongalves, L.M. in Mem. Junta Invest. Vltram. 2nd sen, 41: 57- 159 (1963)]. 49 H. cannabinus x acetosella: Central African Republic, Chevalier 6914 (G, P); Angola, Welwitsch 5269 (BM); Tanzania, Wilson & Kissie 7562 (NA); Uganda, Jackson 462 (BR). H. diversifolius x nigricaulis(l): Democratic Republic of Congo, Hauman 3 (BR). H. diversifolius x cannabinus(l): Democratic Republic of Congo, Hulstaert 1472 (BR). H. sabdariffa x ?: Democratic Republic of Congo, Dechamps 13088 (BR). H. acetosella introgressed with H. mechowii or H. sabdariffa (?): Zambia, Mutimushi 1949 (K). H. nigricaulis x ?: South Africa, Transvaal, Scheepers 327 (PRE). H. sudanensis x ?: Democratic Republic of Congo, Becquet 108 (BM, BR). THE INDIAN SPECIES OF HIBISCUS SECTION FURCARIA Only one of the species which occur in India, Hibiscus surattensis, is native to both India and Africa. Two others, H. cannabinus and H. sabdariffa, are cultigens developed from wild African ancestors. One, H. mastersianus (including H. beddomei), is native to Africa and apparently adventive in India. Hibiscus hispidissimus is native to India and adventive or cultivated in South Africa Hibiscus radiatus may have originated in India as a hybrid between H. cannabinus and H. surattensis but is now widely cultivated in the Old World and New World tropics (Wilson & Menzel, 1 964; Menzel, 1986). Hibiscus furcatus Willd. occurs in India and Thailand. Hibis- cus hoshiarpurensis and the (previously undescribed?) species called H. furcatus Roxb. (= H. hispidissimus Griff.) by Bahadur et al. (1970) are apparently indigenous in India. Seven of the species which occur in India are included in the key, but Hibiscus hoshiarpurensis and the Bahadur et al. species are not because herbarium material was unavailable to me. The two intro- duced cultigens, H. cannabinus and the fibre form of H. sabdariffa, have aculeate stems, peduncles articulating nearer the base, unforked bracteoles, and calyx nectaries (the 'A' genome pattern). Four native species, H. surattensis, H. hispidissimus, H. furcatus, and//, radiatus, have aculeate stems, peduncles articulating nearer the epicalyx, forked bracteoles, and lack calyx nectaries (the 'B' pattern). The adventive H. mastersianus (including H. beddomei) and the native H. hoshiarpurensis have non-aculeate stems, peduncles articulating nearer the base, forked bracteoles, and lack calyx nectaries (the 'X' pattern). Finally, the species of Bahadur et al. (1970) has aculeate stems, peduncles articulating nearer the base, forked bracteoles, and lacks calyx nectaries (the 'G' pattern). Furthermore, this species occurs on the Upper Gangetic Plain (Dehra Dun andTehri-Gharwal), whereas the true H. hispidissimus occurs in central and southern India and in Sri Lanka. TAXONOMIC TREATMENT Description of Hibiscus section Furcaria Hibiscus section Furcaria DC., Prodr. 1: 449 ( 1 824); emend Hochr. inAnnuaire Conserv. Jard. Bot. Geneve 4: 101 (1900). Lectotype 50 F.D. WILSON species: Hibiscus surattensis L. (Borssum Waalkes, 1966: 57). 3 Calyx nectary absent 4 Fryxell (1988: 195) noted that Kearney had chosen H.furcellatus Calyx nectary present 14 Desr. as lectotype but that this name was inadmissable. 4 Stipules ovate, auriculate, amplexicaul 1. H. surattensis Hibiscus section Sabdariffa DC, Prodr. 1: 453 (1824). A n »* j BL. r; c ie*AMC7A\ Stipules linear to subulate, not amplexicaul 5 Furcaria Kostel., Allg. Med. Pharm. Fl. 5: 1856 (1836). Sabdariffa Kostel., Allg. Med. Pharm. Fl. 5: 1 856 ( 1 836). 5 Peduncle articulated closer to epicalyx than to base 6 Hibiscus section Furcaria subsection Furcaria simplicia Hochr. in ^^ ^^^ at base near base or apparently unjointed .... l , Annuaire Conserv. Jard. Bot. Geneve 4: 41 (1900), pro parte. Hibiscus section Furcaria subsection Furcaria typica Hochr. in 6 Calyx lobes with reticulate venation and simple to 4-fid bristles on Annuaire Conserv. Jard.Bot. Geneve 4: 41 (1900), pro parte. enlarged bases on ribs, otherwise almost glabrous ... 2. H. altissimus Hibiscus section Furcaria series Friesia Ulbr. in Engl., Pflanzenw. Calyx iobe venation not as above 7 Ost-Afrikas 3(2): 402 (1921), pro parte. „ . . r . ,. TT,, 7 Lower and mid-plant leaves entire or shallowly lobed 8 Hibiscus section Furcaria series Furcaria cannabma Ulbr. m Engl., Pflanzenw. Ost-Afrikas 3(2): 400 (1921), pro parte. Lower and mid-plant leaves deeply palmately lobed 9 Hibiscus section Furcaria series Furcaria furcellata Ulbr. in Engl., g ^^ ^ § Qf ^ pubescence; stems steilate.pubescent . Pflanzenw. Ost-Afrikas 3(2): 400 (1921), pro parte. 3 H. furcellatoides Hibiscus section Furcaria series Furcaria diversifolia Ulbr. in Engl., Pflanzenw. Ost-Afrikas 3(2): 402 (1921), pro parte. Leaves without lanate pubescence; stems hirsute with simple hairs to Hibiscus section Furcaria series Furcaria sabdariffa Ulbr. in Engl., 3 mm long 4- H' rostellatus Pflanzenw. Ost-Afrikas 3(2): 402 (1921), pro parte. 9 Peduncles to 10(-13) cm long in fruit; stems with stout, retrorse aculei Hibiscus section Furcaria series Saxicolae Ulbr. in Engl., Pflanzenw. on large bases 5. H. hispidissimus Ost-Afrikas 3(2) : 403 ( 1 92 1 ), pro parte. Peduncles <5 cm long in fruit; stems more finely aculeate or aculeolate Hibiscus [sect.] Cannabini Small, Man. s.e.fl.: 854 (1933). 10 Hibiscus [sect.] Furcellati Small, Man. s.e.fl.: 854 (1933). 10 Peduncles to 5 cm long, articulating 8 mm below epicalyx; ribs of Annual herbs, perennial subshrubs, shrubs, trees, or vines. Stems adaxial leaf surface finely and coarsely stellate-pubescent and other plant parts often aculeate. Stems, leaves, peduncles, and 6. H. goossensii floral parts glabrous or variously pubescent. Leaves alternate, the p^des to 2 cm long, articulating <5 mm below epicalyx; ribs of basal leaves usually petiolate, apical ones sometimes sessile and adaxia] leaf surface with mostly simple (some to 4.fld) hairs reduced to linear bracts; leaf shape variable, from entire to variously 7 H. furcatus palmately lobed, even on the same plant; often with conspicuous foliar nectaries borne basally on the adaxial surface of the midrib ! l Foliar nectary Present L (and sometimes also on secondary ribs). Flowers axillary, solitary, Foliar nectary absent 13 borne in racemes by the reduction or abortion of upper leaves, in ",. 12 Mid-plant leaves not lobed; peduncle articulating near base to 1/3 the clusters at the tips of branches, or in few- to many-flowered sympodia. ^^ from fease to epica,yx 8 H sudanensis Peduncles articulated at or near the base, or above the base, often accrescent. Involucellar bracteoles 5-22, free or connate at the base, Mid-plant leaves deeply 3-lobed, midlobe markedly longer than lateral linear to subulate, simple, channelled, or variously bifurcate at the lobes; peduncle articulating at base or appearing imjoioted^.......... apex. Calyx gamosepalous, 5-lobed, persistent, leathery or woody in fruit (fleshy in edible forms of//, sabdariffa), glabrous or variously 13 Stems above glabrate to finely aculeate; leaves glabrate pubescent, each lobe with a more or less thickened midrib and two 10. H. radiatus marginal (or near marginal) ribs, the midrib often with a conspicu- stems above with stout retrorse aculei and a line of stellate pubescence; ous nectary. Corolla usually delicate, often large and showy; petals leaves scabrous 11. H. torrei varying in colour from white to various shades of yellow, pink, red, lavender and purple, and usually with a reddish to purple petal spot 14 Involucellar bracteoles clearly bifurcate, outer fork 3-6 mm long, not at the base. Staminal column shorter or longer than the petals, reflexed inner fork 1-6 mm long, linear to subulate peduncles articu- ,. latmg above the base, aculeolate and sparsely puberulent above the vanouslycoloured.Styleexceedmgthestaminalcolumn,5-branched ^densely fine-pubescent below the joint 12. H. noldeae above, each branch surmounted by a capitate stigma. Capsule 5- celled, loculicidally dehiscent, glabrous to densely Involucellar bracteoles obscurely bifurcate, outer fork 2 mm long, appressed-pubescent (some South American species have glandular reflexed, inner fork vestigial or absent; peduncles articulating at base or hairs on the capsule). Seeds reniform to angular, often scaly or Appearing ™Jointed> aculeate and Stellate- variously marked, but rarely pubescent. Basic chromosome number, x= 18. 15 Calyx nectary absent 16 Calyx nectary present 18 Key tO African and Selected Asian taxa Of HibisCUS 1 6 Upper 6_ 1 1 mm of involucellar bracteoles lanceolate; leaves through- Section Furcaria out the plant deeply palmately 3-5-lobed 14. H. mechowii Upper part of involucellar bracteoles not lanceolate; leaves entire to shallowly lobed 17 Stems not aculeate j 7 pecjuncie articulating midway between base and epicalyx; leaves densely 2 Involucellar bracteoles clearly or obscurely bifurcate 3 stellate-pubescent throughout 15. H. subdiversifolius Involucellar bracteoles not bifurcate ... ... 15 Peduncle articulating at base or apparently unjointed; leaves with REVISION OF HIBISCUS SECTION FURCARIA 51 simple and stellate hairs on veins, otherwise almost glabrous 16. H. berberidifolius 18 Stems with large stout aculei 19 Stems more finely aculeate or aculeolate 23 19 Aculei borne at the base of stem nodes singly, in pairs, or in threes; leaves to 2 x 4 cm, shed early 17. H. sparseaculeatus Aculei borne ± randomly on the stem; leaves larger, persistent 20 20 Calyx densely and finely stellate-pubescent, sparsely aculeate on ribs; midlobes and lateral lobes of lower leaves rotund to spathulate 18. H. greenwayi Calyx with rigid bristles; leaf lobes not rotund or spathulate 21 21 Calyx aculeate or aculeolate on the ribs; lobes of lower leaves mani- festly secondarily lobed 19. H. partitus Calyx not aculeate or aculeolate on the ribs; leaf lobes not manifestly secondarily lobed 22 22 Stems usually with one or more longitudinal lines of pubescence; flowers yellow 20a. H. diversifolius subsp. diversifolius Stems more densely and uniformly hairy; flowers reddish to purple . 20b. H. diversifolius subsp. rivularis 23 Some flowers subtended by a linear, flattened, sometimes bifid bract resembling the stipules; leaves deeply 5(-7)-lobed, midlobes and lateral lobes of lower leaves often secondarily lobed 21. H. reekmansii Flowers without a subtending bract; leaves variously lobed, but not secondarily lobed 24 24 Involucellar bracteoles subulate to triangular, lengthibase width ratio 3:1 or less, upper 3-6 mm channelled on the inner (ventral) surface . 22. H. sabdariffa Involucellar bracteoles generally narrower, length:base width ratio >3:1, inner surface not channelled 25 25 Vegetative branches in leaf axils well-developed, with up to 7 persistent leaves; newly opened leaves with a gland-like structure at apex of each leaf margin serration; leaves entire to shallowly 3-lobed 16. H. berberidifolius Vegetative branches not as well-developed; gland-like structure not present; leaves variously lobed 26 26 Calyx with white, woolly tomentum basally, or more uniformly distrib- uted; capsule with long, dense appressed pubescence 23. H. cannabinus Calyx without white woolly tomentum; capsule pubescence various . 27 27 Calyx with bristly simple hairs on ribs, almost glabrous between ribs; capsule with long, dense appressed pubescence .... 24. H. parvilobus Calyx with antrorse aculei or pigmented short, coarse bristles on enlarged bases; capsule with short, sparse appressed pubescence 25. H. asper 28 Involucellar bracteoles manifestly to obscurely bifurcate; calyx nectary absent or present 29 Involucellar bracteoles not bifurcate; calyx nectary present 36 29 Calyx and foliar nectaries present; leaves glabrate, sometimes red ... 26. H. acetosella Calyx nectary absent, foliar nectary absent or present; leaves glabrate to pubescent, green 3( 30 Foliar nectary absent; leaves glabrate 10. H. radiatus Foliar nectary present; leaves pubescent 31 3 1 Inner fork of involucellar bracteoles ± twice as long as outer fork . 32 Inner and outer forks of involucellar bracteoles ± of equal length . 34 32 Calyx lobes acute to slightly acuminate in fruit; length:width ratio of midlobe of mid-plant leaves 1:1 to 3:1 33 Calyx lobes long-acuminate in fruit; lengurwidth ratio of midlobe of mid-plant leaves c. 4:1 27c. H. gilletii subsp. lundaensis 33 Lengurwidth ratio of midlobe c. 2:1 to 3: 1 ; outer fork of involucellar bracteoles 2-4 mm long, inner fork 6-9 mm long 27a. H. gilletii subsp. gilletii Length: width ratio of midlobe c. 1 : 1 ; outer fork of involucellar bracteoles 1.5-3 mm long, inner fork 4-5 mm long 27b. H. gilletii subsp. hiernianus 34 Stems lanate-pubescent, patchy below, continuous above; lower leaves deeply 3- to 5-lobed (ratio of base to apex: base to sinus c. 5:1) 28. H. moxicoensis Stems not lanate-pubescent; lower leaves shallowly to moderately 3- to 5-lobed (ratio of base to apex: base to sinus c. 1.3:1 to 2:1) 35 35 Stems finely stellate-pubescent and with coarse stellate hairs; lower leaves moderately 5-lobed (ratio of base to apex: base to sinus c. 2:1), without a foliar nectary 29. H. cuanzensis Stems with stiff, simple to 3-fid or stellate pigmented bristles on enlarged bases; lower leaves shallowly 3-lobed (ratio of base to apex: base to sinus c. 1.3:1), with a foliar nectary 30. H. mastersianus 36 Calyx fleshy in fruit; involucellar bracteoles broadly subulate to trian- gular, upper 3-6 mm channelled on the inner (ventral) surface 22. H. sabdariffa Calyx not fleshy in fruit; bracteoles narrower, not channelled 37 37 Stipules divided into 2 or 3 filiform or linear segments; involucellar bracteoles filiform to narrowly linear; all leaves linear; foliar nectary absent 31. H. elongatifolius Stipules simple; involucellar bracteoles linear to subulate; at least lower leaves broader or palmately lobed; foliar nectary present 38 38 Involucellar bracteoles subulate, c. 1 mm wide at the base, apex pointed; calyx finely stellate-pubescent and with simple to stellate stiff bristles on enlarged red bases 32. H. sineaculeatus Involucellar bracteoles linear, c. 1 mm wide at the base, apex narrowly lanceolate; calyx densely and finely stellate-pubescent, not bristly .... ... 33. H. scotellii 1 . Hibiscus surattensis L., Sp. pi. 2: 696 ( 1 753). Type: India, Surat, Herb. Linn. 875.29 (LINN-holotype). Furcaria surattensis Kostel., Allg. Med. Pharm. Fl. 5: 1856 (1836). Hibiscus surattensis var. genuinus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 1 1 (1900), nom. illeg. (Art. 26.2). Hochreutiner (1900) lists synonymy for//, surattensis sens, strict. (var. genuinus), which will not be repeated here. Hibiscus surattensis var. villosus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 112(1900). Hibiscus surattensis var. villosus Backer, Fl. Batavia 1: 128 (1907), non Hochr. (1900). Hibiscus surattensis var. villosusformaconcolor Backer, Fl. Batavia 1: 129 (1907). Type: Indonesia, Java, Tandjungpriok, Backer 32783 (BO-isotype). Hibiscus furcatus Craib, Fl. siam. 1: 157 (1925), non Roxb. (1814), pro parte quoad specim. Marcan 1517. Hibiscus surattensis forma immaculata Kurz ex Rakshit & Kundu in Sci. & Cult. 27: 194 (1961). Type: Burma, Pegu Yomah, Kurz 1248 (CAL-holotype). 52 Illustrations: Exell & Gonc.alves, Malvaceae in Fl Moqamb.: 25, fig. 8, 4 (1979); Gibson, Wild fl. natal: pi. 64, fig. 6 (1975); Hochreutiner, Fl. Madagasc., Malvac.: 37, fig. X, 1-3 (1955); Paul & Nayar, Fasc.fl. India 19: 154, fig. 32 (1988); Pradeep & Sivarajan, Taxon 40: 636, figs 7, 8 (1991); Icones Roxburghianae no. 1504 (K); Wight, Icon. pi. Ind. orient. 1: pi. 197 (1839). Annual herb, up to 3 m high, prostrate or climbing on shrubs or trees; stems reddish green, with soft simple hairs to 2 mm long, sparsely to densely retrorse-aculeate, the aculei often red-tipped. Stipules up to 15x10 mm, ovate, auriculate, amplexicaul. Leaves: petioles up to 1 1 cm long, hairy and aculeate like the stems, the blades up to 10 x 10 cm, shallowly to densely palmately 3-5-lobed, abaxial surface aculeate and with simple hairs on main and secondary veins and sparsely stellate-pubescent on smaller veins, adaxial surface with mostly simple hairs, the base obtuse to subcordate, the margin serrate, the apex acute to acuminate, the nectary absent. Flowers axillary, solitary; peduncles up to 8 cm long, articulating c. 10 mm below the epicalyx, sparsely aculeate and hirsute below, and densely hispid above the articulation; involucellar bracteoles 8-9, up to 15 mm long, spreading, with simple hairs on margins, the apex bifur- cate, the outer fork up to 5 x 4 mm, spathulate, the inner fork up to 10 mm long, linear, ascending; calyx up to 2.5 cm long, becoming scarious in fruit, the lobes acuminate, aculeate and setose on the ribs, the nectary absent; corolla bright yellow with red-purple centre and a pink stripe along the edge of each petal, the petals up to 6 x 4 cm, obovate, glabrous; staminal column up to 20 mm long, red, the filaments up to 3 mm long, purple; pollen yellow; style branches up to 5 mm long, cream. Capsules up to 15 x 12 mm, ovoid to globose, with dense appressed pubescence, the beak inconspicuous; seeds 3 x 2 mm, subreniform, faveolate and with minute pectinate scales. Chromosome number, n = 1 8. Ruderal and wild, sea level to 1700 m, in cultivated and aban- doned farm fields and plantations, waste ground near habitations, and in a wide variety of natural habitats, many in low-lying areas such as river beds and banks, swamps, and other damp situations, but also in savannas, in grassland and bushland, forest clearings and edges, as well as in coastal habitats such as coastal evergreen bushland and sand dunes. DISTRIBUTION. Fig. 4. Widespread in subsaharan Africa, also in the Indian subcontinent and southeastern Asia, not in Australia, as reported by Masters (1868, 1872) and other subsequent authors. Guinea-Bissau, Susana, 1 December 1960, Raimundo & Guerra 274 (G-2 sheets); entre Nova Lamego e Canjufa, 29 October 1950, Esperito Santo 3545 (BR). Sierra Leone, Kenema (Nongova), 25 November 1952, Deighton 5888 (K). Ivory Coast, banks of Comoe River, opposite Grand Bassam, 15 July 1963, de Wilde 468 (BR); reserve de LAMTO, S/P de Tiassale', 15 December 1987, Gautier- Beguin 728 (G-2 sheets). Ghana, Atwabo, 1931, Fishlock 21 (BR); Kumasi, September 1928, Vigne 1381 (BM, K-2 sheets). Togo, prope Lome, 19007 1902, Warnecke 243 (BM, BR, K). Benin, Ocoja Ouere, 1 1 December 1901, LeTestu 188 (BM-2 sheets); Benin Div., Okomu Forest Reserve, 27 February 1948, Brenan9l66 (BM, K). Nigeria, Western Province, Massambolahun, 3 miles S. of Bolahun, 6 January 1953, Konneh 614 (BR); Ibadan, 1935, Newberry 131 (K). Cameroon, pres Djoum, 21 November 1966, Letouzey 8437 (BR); Bipinde, October 1913, Zenker 423 (G-2 sheets). Central African Republic, 5 km SW of Bambari on the Alindao road, 8 November 1981, Fay 1906(K);Haute-Kotto,31 October 1922, LeTestu 421 \ (BM,BR). Sudan, Equatorial Prov., Zande Land, 24 November 1937, Wyld 332 (BM). Principe, Esperanc.a, 27 December 1932, Exell 678 (BM, BR). Sao Tome, Piedada, 1 July 1885, deWildeman 411 (BR). Equatorial Guinea, Bioco- Riaba, cerca de Bao Basuala, 1 2 February 1 989, Fernandez Casas 11511 (K). Gabon, Cap Esterias, 24 February 1968, Halle & Villiers 5507 (P). Demo- cratic Republic of Congo, Leopoldville, 1 1 June \9\5,Bequaertl516(BR); Yangambi, 22 September 1936, Louis 2642 (K); Kisangani, Zone Lubunga, km 4, route Ubundu, 9 January 1982, Szafranski 1032 (BR). Uganda, F.D. WILSON Kipayo, September 1914, Diimmer 1059 (BM, BOL); W. Mengo Dist., Bat Valley, Kyadondo, 19 November 1985, Rwaburindore 2142 (BR). Kenya, K7, Kilifi Creek, 13 August 1975, Greenway & Kanuri 15556 (K); Kilifi Dist., Mida, Arabuku-Sokoke Forest Reserve, 15 miles S. of Malindi, 2 December \96\,Polhill & Paulo 881 (B, BR). Burundi, Bujumbara, 1 June 1967, Lewalle 1990 (BR); Bururi Prov., Kigwena, 4 June 1980, Reekmans 9253 (BR). Tanzania, Western Prov., Kigoma Dist., Lumbye River mouth, 30 July 1959, Newbould & Jefford 1158 (K); Tanga region, 21 km W. of Pongwa, Hway A 14, 5 August 1975, Wilson & Kissie 75128 (ASU, EA, NA); Bezirk Mahenge, Umgebung der Station Mahenge, 30 June 1932, Schlieben 2148 (BM, G); Lutamba-See, 40 km W. Lindi, 2 September 1934, Schlieben 52 1 9 (BR); Insel Zanzibar, September 1 873, Hildebrandt 9 1 5 (BM ). Angola, Cujenjo (?), July 1909, Gosswe Her 4159 (BM); Pungo Adongo, May 1857, Welwitsch 5253, 5254 (BM); in palmatis ad R. Cuango, May 1 856, Welwitsch 4921 (BM). Zambia, Luapula Prov., Fort Roseberry Dist., 23 August 1952, Angua 301 (BM); Central Province, lolanda, Kafue, 4 June 1965, Robinson 6683 (BR); Victoria Falls, Long Island, April 1918, Eyles 1264 (G). Malawi, Southern Reg., Mulanje, Chisambo Forest Plantation, 1 August 1979, Bandan, Salubeni & Masiya 1538 (BR). Mozambique, Cabo Delgado, 53 km de Diaca para Mocimboa de Praia, 14 April 1964, Torre & Paiva 1 1904 (BR); Maringua, Sabi River, 19 June 1950, Chase 2560 (BM-2 sheets); Maputo (Lourenc.o Marques) Dist., Marracuene, 10 April 1 946, Gomes & Sousa 34 1 7 (K); Delagoa Bay, Inhaca Island, July 1934, Weintraub 20321 (BOL). Comoros, Grande Comore, N. O. T'Sidje-Moroni, 19 August 1981, Doutrelepont 1085 (BR). Seychelles, Mahe . . . road from Victoria to Exile, 18 July 1936, Day 73 (BM). Zimbabwe, Sebwingwe Dist., confluence of Sanyati and Zambesi Rivers, 1 September 1955, Davies 1510 (BR); Umtali Dist., on stream bank beyond Cross Hill, November 1948, Chase 1665 (BM- 2 sheets). Madagascar, Majunga (Mahajanga) Prov., Station forestiere d' Ampijoroa, c. 30 km N. d'Andranofasika Foret, 8 April 1984, Dorr & Koenders 2958 (K); Ambovomba, 2 June 1931, Decary 8963 (G). South Africa, Transvaal, Zoutpanasberg, 4 km NW of Punda Maria, 18 May 1949, Codd & deWinter 5552 (PRE); Transvaal, Lekgalameetse Nat. Res.: Cyprus, near last bridge over Wolfspruit, 3 May 1 985, Stalmans 58 1 (PRE); Natal, 2.5 km from Arboretum turnoff on old Richardsbay road, 12 December 1985, Pienaar 828 (PRE); Natal, Alexandra Dist., Dumisa Station, Friedenau Farm, 26 March 1908, Rudatis 333 (G). Swaziland, Pigg's Peak Dist., Wyldesdale, 25 March \959,Compton 28741 (NBG); MankaianaDist., Ntondozi, 27 June 1963, Compton 31650 (NBG). India, Kerala, Trichur Dist., Chiklai, 29 September 1982, Ramamurthy (Bot. Surv. Ind. 74953) (CAL); Tamil Nadu, Kanniyakumari Dist., Kughithurai, 10 December 1980, Swaminathan (Bot. Surv. Ind. 70302) (CAL); Assam, Garo Hills, Rewar, 16 November 1930, Parry 891 (K). Bangladesh, Chittagong, 3 January 1851(7), Hooker & Thompson 353 (K). China, Hainan, K'iung-shan Dist., Kiung-chow City and vicinity, 30 July 1932, Fung 20256 (B, BM, K); ins. Hong Kong, Hoihan, 1 877, Bullock 559 (BM). Sri Lanka, without specific locality or date, Fraser 68 (BM, BR). Thailand, Pak Chong, 29 December 1923, Marcan 1517 (BM); Phitsanulok Prov., Thoung Saleng Louang, 25 November 1965, Vidal 4559 (P). Laos, Vientiane Prov., env. de Ban Na Hai, 22 October 1965, Vidal 4121 (P). Vietnam, Tonkin, bords de la route conduisant de Tu-Phap aux roches de Notre-Dame, 20 December 1887, Balansa 3715 (P); Onnam, Courane, 18 February 1939, Poilane 28900 (P). Philippines, Luzon, Union Prov., Bauang, February 1904, Elmer 5666 (G); Culio Island, December 1927, Ramos & Edamo s.n. (BR); Occidental Mindoro Prov., Abra de Hog, January 1951,Sw//rs.n. (BR). Singapore, 1 893, Ridley s.n. (BM). Malaysia, c. mile 1 .2 from Kota Belud to Kabaiau, 1 1 March 1954, Darnton 509 (BM). Indonesia, Sumatra, Hoeta Pradang, Asahan, east coast, October-December 1932, Krukoff440\ (BR); Java, sables maritimes, Batavia, 13-15 August 1878, Savinierre 1 170 (BR); Moluccas, Ambon (Island of Amboyna), 3-15 September 1840, Barclay 4136 (BM); Timor, without date, Forbes 3817 (BM). Hibiscus surattensis forma immaculata Kurz ex Rakshit & Kundu is a later name for H. surattensis var. villosus forma concolor Backer, a form with completely yellow flowers, lacking the basal petal spot. Hochreutiner (1900: 112-113) mistakenly lists synonyms for H. surattensis sens, lat., which apply to other taxa included in this paper (see H.furcatus Willd., H. hispidissimus Griff., H. acetosella Welw. ex Hiern, H. mastersianus Hiern, and H. rostellatus Guill. & Perr.). REVISION OF HIBISCUS SECTION FURCARIA 53 Pradeep & Sivarajan (1991: 635) point out that several authors (Borssum Waalkes (1966), among them), wrongly listed Narinam poulli Rheede as a synonym of H. surattensis, while in fact it is synonymous with H. hispidissimus Griff. I examined a number of variant forms of Hibiscus surattensis. The most common is one that has narrow stipules and involucellar bracteoles, occurring throughout much of the range of this species. It may be interpreted as a mutant or as an introgressant form. Examples are: Liberia, Adam 16511 (BM), 20527, 20702 (K); Tanzania, Renvoize 2010 (K); Mozambique, Goodier 1005 (BM); India, Hamilton s.n. (BM); Thailand, Hosseus 468 (BM); Mada- gascar, Herb. Martii 68 (F.V. Thompson s.n.) (BR);Laos, VWa/4121 (P); Vietnam, Pierre s.n. (P). Other variant forms seen are as follows: narrow, acuminate calyx lobes, China, Hainan, Henry s.n. (G); an apparently purple-flowered form, Madagascar, Hildebrandt 2869 (G); atypical leaves, narrow stipules [possibly H. surattensis x H. rostellatus hybrid], 'French West Africa', Roberty 6401, 15860 (G); villous calyx, Cameroon, Zenker423 (G), Tanzania, Schlieben 3690 (G); miniature but typical-shaped stipules, Thailand, Marcan 1517 (BM). 2. Hibiscus altissimus Hornby in Proc. & Trans. Rhodesia Sci. Assoc. 41: 55 (1946). Type: Mozambique, Lusa River Valley, W. of Gurue, 7 May 1942, Hornby 4563 (K!-isotype). Fig. ID. A woody scrambler over bushes; stems to 10 m long, retrorse- aculeate, otherwise almost glabrous or with a line of pubescence, or sometimes more densely pubescent above. Stipules 5-9 mm long, linear, almost glabrous. Leaves: petioles 0.2-10 cm long, retrorse- aculeolate plus a line of pubescence adaxially, the blades 3-1 1 x 2-13 cm, broadly lanceolate to ovate to 3(-incipiently 5)-lobed, the midlobe broadly lanceolate to ovate, the lateral lobes lanceolate to triangular, sometimes unequal in length, the blades abaxially retrorse- aculeolate on the ribs and with sparse coarse 2-fid to stellate hairs on the veins, adaxially with simple to stellate sparse hairs on veins, the base cuneate, truncate, or cordate, the margin serrate or dentate, the apex acute, the nectary 1—4 mm long, slit-like. Flowers axillary, solitary or clustered near the ends of branches; peduncles 1-5 cm long in fruit, articulating 1-3 mm below the epicalyx, or sometimes the joint not evident, accrescent, retrorse-aculeolate and densely and finely stellate-pubescent; involucellar bracteoles 10-12, 9-12 mm long, linear below the apex, with coarse, mainly simple bristles on enlarged bases on the margins, the apex bifurcate, the outer fork 3 mm long, lanceolate, the inner fork 5-9 mm long, linear, ascending, the base free; calyx 2-3 cm long in fruit, the lobes lanceolate, acute to acuminate, with reticulate venation and simple to 4-fid bristles on enlarged bases on the ribs, otherwise almost glabrous or sparse, finely stellate-pubescent, the nectary absent; corolla yellow with a purple centre, the petals 3.5-5 x 2-3 cm, obovate, sparsely and finely stellate-pubescent dorsally, almost glabrous ventrally ; staminal column 22-25 mm long, the filaments 1 mm long; style branches 4- 5 mm long. Capsules 17-27 x 14-21 mm, densely appressed-pubescent, the beak 2 mm long, glabrous; seeds (accord- ing to Exell, 1961) 4 x 3 mm, subreniform, with whitish, irregularly discoid scales. Chromosome number, n = 72. Undergrowth in forest glades and plantations and in serai montane scrub. DISTRIBUTION. Fig. S.Southeastem Africa. Mozambique, Manicae Sofola, Cheringoma, Durundi, 17 May 1948, Barbosa 1660 (BM); Cheringoma, Inhaminga, 24 May 1948, Mendonca 4363 (BM); Sul do Save, entre Bilene eViladeJoaoBelo, 1 May 1 957, de Carvalho 131 (BM); Bilene, Praia deS. Martinho do Bilene, 14 May 1957, de Carvalho 213 (BM); Laurenso Marques, Rikatla, April 1918, Junod 238 (BM, G). South Africa, Transvaal, 20 km along Witvlag Road from Louis Trichardt, 4 April 1971, Oakes 1585 (PRE); Louis Trichardt, Entabeni, 16 August 1989, van HeerdenA33 (PRE); Letaba, Duiwelskloof Rosendal, 26 July 1958, Scheepers 438 (G); Natal, Hlabisa, Dukuduku forest glade, 29 January 1967, Strey 7365 (PRE). Exell (1961) cites, as a synonym of Hibiscus altissimus, H.furcatus 'sensu Harv.' in Harv. & Sond., Fl. Cap. 1: 176 ( 1 860), but Harvey's succinct description and his reference to Drege's specimens labelled 'Hibiscus hamatus E. Mey.' cast some doubt on this conclusion. I examined two of Drege's specimens, collected at Port Natal and labelled H. hamatus (Drege s.n., G; Drege 5289, P); both are referable to the native Indian species that has long been called H. furcatus Roxb. (= H. hispidissimus Griff.). These plants were possi- bly adventive or cultivated in Natal. 3. Hibiscus furcellatoides Hochr. in Annuaire Conserv. Jard. Bot. Geneve 20: 157-158 (1917). Type: Guinea, entre le Konkoure et Timbo, March 1905, Chevalier 12504 (P!-holotype). Fig. 1A. Hibiscus furcatus Hutch. & Dalziel, Fl. W. trop.Afr. 1(2), Isted.: 267 (1928), non Roxb. (1814), pro parte quoad specim. Chevalier 12504. Hibiscus rostellatus Keay, FL W. trop. Afr. 1(2), 2nd ed.: 346 (1958), non Guill. & Perr. (1831), pro parte quoad specim. Chevalier 12504. Habit unknown; stems with stout retrorse aculei on large bases, sparsely stellate-pubescent below, densely and finely stellate-pubes- cent above. Stipules 1 mm long, subulate, almost glabrous. Leaves: petioles 1 .2-2.5 cm long, aculeolate and densely stellate-pubescent, the blades 4.5-6.5 x 4—6 cm, ovate-triangular, the blades of middle leaves and upper leaves entire (lower leaves not seen), abaxially and adaxially with lanate patches, otherwise finely stellate-pubescent, the base truncate, the margin finely serrate, the apex slightly acumi- nate, the nectary to 6 mm long, prominent, slit-like. Flowers axillary, solitary; peduncles 2-3.3 cm long in fruit, articulating 4-9 mm below the epicalyx, with a fine dense stellate tomentum, not ac- uleate; involucellar bracteoles 9-10, 7 mm long below the apex, linear, flattened, densely and finely stellate-pubescent, the apex bifurcate, the outer fork 3-5 mm long, lanceolate to sagittate, the inner fork 7-8 mm long, linear, ascending, the base free; calyx 1 .3- 1.8 cm long, the lobes up to 1 cm wide, ovate, acuminate, densely and finely stellate-pubescent, not aculeate, the nectary absent; corolla yellow(?) with a purple(?) centre, the petals up to 4.5 x 3.5 cm, obovate, sparsely stellate dorsally, glabrous ventrally; staminal col- umn 20 mm long, purple near the summit, the filaments 1.5 mm long, anthers brown(?); style branches 5 mm long. Capsules 17x15 mm, densely appressed-pubescent, the beak 1 .5 mm long, glabrous; seeds unknown. DISTRIBUTION . Fig. 5. Guinea. Known to me only from the type collection. Bahadur et al. (1970) wrongly placed Hibiscus furcellatoides in synonymy with the (previously undescribed?) Indian species which they called H. furcatus Roxb. 4. Hibiscus rostellatus Guill. & Perr. in Guill., Perr. & A. Rich., Fl. Seneg. tent. 1: 55-56 (1831). Type: Apparently not designated. I examined three of Perottet's specimens, one each from BM, G, and P. I designate the specimen from P as the lectotype because it is the most complete and well preserved of the three specimens. The label on the sheet from P reads as follows: 'Herbarium Richard [red ink] Hibiscus rostellatus Nob. afigurer [black ink] 54 F.D. WILSON 10 cm Fig. 1 Leaves and fruits (or flowers) of species of Hibiscus section Furcaria. A: H.furcellatoides, Chevalier 12504 (P); B: H. rostellatus, leaf: Exell & Mendonqa 3171 (BM), fruit: Scott Elliot 7244 (BM); C: H. goossensii, Goossens 1202 (BR); D: H. altissimus, Mendonqa 4363 (BM); E: H. nigricaulis, Exell & Mendonqa 1759 (BM); F: H. flavo-roseus, Young 646 (BM); G: H. torrei, Torre 435 (BM); H: H. noldeae, Leeuwenberg 10288 (BR). REVISION OF HIBISCUS SECTION FURCARIA 55 Senegal, Perottef [red ink]. The sheet from BM has two labels, as follows: 1 ) 'Hibiscus rostellatus Nob., n. 88 [in pencil], ...du Cap Vert, Kounoun 13 Mars 1829, Perottef; 2) 'Perottet, no. 53, 'B' [in pencil] Senegal'. The label on the sheet from G reads as follows: 'Hibiscus rostellatus, fl. Seneg., Kounoun (Capvert), M. Perottet 1831'. The black ink entries on the BM and P sheets were made in the same hand, presumably Perottet's (especially since they say 'Nob.'), while the writing on the G label was in a different hand. Fig. IB. Hibiscus surattensis var. rostellatus (Guill. & Perr.) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 10 (1900). Hibiscus furcatus Hutch. & Dalziel, Fl W. Trop. Afr. 1(2): 267 (1928), non Roxb. (1814), excl. Chevalier 12504 (see Keay, 1958: 346). Hibiscus furcatus Berhaut, Fl. Seneg. : 1 59 ( 1 954), non Roxb. (1814). Coarse prostrate or climbing perennial, much-branched vine up to 3 m tall; stems more or less hirsute (hairs to 3 mm long) [Zenker 4957 almost glabrous], with many conspicuous retrorse aculei on red bases. Stipules up to 1 1 mm long, filiform to subulate, with fine hairs on margins. Leaves: petioles 0.5-9 cm long, with pubescence like that of the stems, the blades 3-13 x 3-15 cm, cordate to very shallowly 3-5-lobed, suborbicular in outline, the lobes triangular, abaxially densely stellate-pubescent and retrorse-aculeolate and with coarse, 2-several-fid bristles on ribs, adaxially sometimes aculeolate on the ribs, usually less densely stellate-pubescent, the base truncate to shallowly or deeply cordate, the margin finely serrate to dentate, the apex acute to long-acuminate, the nectary to 4 mm long, slit-like, on the midrib and sometimes also on two lateral ribs. Flowers axillary, solitary, or sometimes clustered on short flowering branches to 8 cm long, or near the plant apex; peduncles 2-8 cm long, articulating 5-10 mm below the epicalyx, with long simple hairs and retrorse aculeoli below the articulation, dense long straight bristles above the articulation; involucellar bracteoles 8-12, 6 mm long below the apex, linear, bristly, the apex bifurcate, the outer fork 3-7 mm long, foliaceous, more or less enlarged, oval or elliptic, sometimes flattened or reflexed, the inner fork 4-10 mm long, subfiliform to subulate-acuminate, the base free; calyx 1.5-3.5 cm long, the lobes ovate-lanceolate, acute to long-acuminate, very finely stellate-tomentose and also with 2-fid to stellate bristles on the ribs, the nectary absent; corolla lemon yellow with a large red-purple centre, the petals 5-7.5 x 3-6.5 cm, showy, obovate, finely stellate- pubescent dorsally, glabrous ventrally; staminal column 22-30 mm long, white to dark pink, the filaments 2 mm long, light to dark pink, the anthers yellow or pink, the pollen orange; style branches 4-8 mm long, light to dark pink; stigmas pink to red-purple. Capsules 22-30 x 15-20 mm, ovoid, appressed-pubescent, the beak 2 mm long, glabrous; seeds 5x3 mm, subreniform, brown, with parallel striations and fine pectinate scales, the funiculus small, dark brown, glabrous. Chromosome number, n = 36. Ruderal and wild, in farm clearings, plantations, fallow land, savannas, creek and river banks, temporary ponds, swamps, second- ary scrub, gallery and rain forests; apparently never cultivated, but leaves are sometimes eaten [e.g. Democratic Republic of Congo, Staner 1482 (BR)]. DISTRIBUTION. Fig. 4. Western to eastern central Africa from Senegal to Sudan, Angola, Zambia, and Tanzania. Senegal, Laugalkaur, October 1950, Berhaut 678 (BR); Bidahay, 25 January 1931, Trochain 1307 (P). Sierra Leone, Savane Piedmont, est Loma, 16 November 1965, Jaeger 8150 (G); Kambia, Scarcies, 30 December (year?), Scott Elliott 4349 (BM). Guinea, Kindua (Kindia?), March 1905, Chevalier 13031 (P); Koba, 15 November 1956, Jacques-Felix 7262 (P). Liberia, Cape Mount, 18 November 1908, Dinklage 2257 (B); Peatah, 1 6 October 1 926, Under 1 098 (K). Mali, Sikasso- Koutiala, 30 December 1 950, Roberty 1 3337 (G);Ivory Coast, entre Fuenoula etBouafle", 28 December 1963,A/teAra'7292(G);Bingerville, 17-20 February 1907, Chevalier 17355 (P); near Tiassale", right bank of Bandama R., 10 December 1958, Leeuwenberg 2150 (BR). Ghana, road to Amedzofe, 31 January 1 97 1 , Ents GC42474 (K).Benin, Cercle de Zagnanado, Pays de Hollis, Entre Abbo et Masse, 6 February 1910, Chevalier 22969 (P). Nigeria, Bende road, 4.5 miles from Umuahia, 9 November 1 966, Ariwaodo 1 436 (K);Western State, Egbado Dist., Alasinde, 14 November 1973, Eimunjeze & Ekwuno FHI 67996 (K); North East State, Gombe Dist., Malam Sidi, 30 November 1971, Latilo FHI 646447 (K); Ilorin Prov., Omu-Aran Dist., on Ilorin-Kabba motor road, 16 November 1968, Latilo FHI 62279 (K); Ibadan Div., near Ibadan, 5 January 1950, Meikle 959 (BR); southern Nigeria, Eket Dist., main road from Aron to Eket, January 1913, Talbot s.n. (BM). Cameroon, 10 km S. of Ngaoundere', Station Fouragre de Wakwa, 1 November 1966, Leeuwenberg 7672 (BR); Bipinde, Urwaldgebiet, 1896, Zenker 1303 (G-2 sheets); same locality, 1913, Zenker 4957 (B, BM). Chad, Foret galerie du Bakoua, 1 November 1968, Gaston 2481 (P). Central African Republic, 4 km SE of Bambari on the Alindao road, 27 January 1982, Fay 2138 (K); right bank of Mambere R., 2 km S. of Carnot, 6 December 1965, Leeuwenberg 7230 (K); La Haute-Kotto, Oubangui-Chari, 5 October 1921, LeTestu 3325 (BR); Sudan, Mongalla Prov., Valley of R. Yei near source, 28 February 1934, Dandy 543 (BM); Equatorial Prov., Kagelu Stat., Yei, 2 October 1937, Myers 7824 (K). Congo, Impfondo, 24 September 1966, Bouguet 2049 (P). Democratic Republic of Congo, Leopoldville Prov., Thysville Terr., 22 April 1960, Compere 1979 (K); Pare Nat. de 1'Upemba, June 1949, deWitte 6692 (BR, K); Eala, November 1 930, Staner \ 482 (BM); Lac Kivu, He Wahu, 15 March 1953, Van derBen 208 (BR). Uganda, Kyagwe, Mukono U4, 1 km E. of Lugazi town, 25 September 1987, Rwaburindore 2546 (BR); without locality (Ruwenzori Expedition 1 893-94), Scott Elliot 7244 (BM);Albert-Edward-Nyanza, 1893- 94, Scott Elliot 8046 (BM). Rwanda, Kirambo, 6 June 1981, Troupin 16280 (BR). Angola, Cuanza Sul, between Gabela andVil Novade Seles, R. Cuvo, 1 2 July 1937, Exell & Mendonqa 3171 (BM-2 sheets); Pungo Adongo, without date, Welwitsch 5243 (BM). Zambia, Solwezi, 14 May 1969, Mutimushi 3309 (K). Tanzania, Msimbazi stream, 82 km WSW Dar Es Salaam, 7 May 1972, Wingfield 1967 (EA); Mahali Mts, Kasoge, 21 November 1973, Uehara 192 (EA). Bahadur et al. (1970) wrongly placed Hibiscus rostellatus in syn- onymy with the (previously undescribed?) Indian species which they called H. furcatus Roxb. Variant forms of Hibiscus rostellatus were examined, as follows: short bracteole forks and leaves that resemble those of//, goossensii: Tanzania, Faulkner 3667 (BR); fine, short pilose indumentum: Congo, Descoings 5880 (P); reduced pubescence on plant parts: Cameroon, Letouzey 95 1 8 (P). 5. Hibiscus hispidissimus Griff., Not. pi. asiat. 4: 521 (1854); Pradeep & Sivarajan in Taxon 40: 634-637 (1991). Type: India, Kerala, Thanneerpandal, near town of Badagara, Pradeep 5008 (K-neotype, Pradeep & Sivarajan, 1991; CALI-isoneotype). Narinam poulli Rheede, Hort. malab. 6: 75, t. 44 (1686) (see Pradeep & Sivarajan, 1991: 635). Hibiscus furcatus Roxb. [Hort. beng. 51 (1814), nom. nud., nom. illeg. (Art. 53.1)] ex DC., Prodr. 1: 448 (1824), non Willd. (1809) nee Moritzi ( 1 846) nee Mast. ( 1 868) nee Bahadur, Dayal & Raturi (1970). Type: Icones Roxburghianae no. 1582 (K!-holotype). Hibiscus aculeatus Roxb., Fl. ind. 3rd ed.: 206 (1832), non Walter (1788) nee Bahadur, Dayal & Raturi (1970), nom. illeg. (Art. 53.1). Type: Icones Roxburghianae no. 356 (K!-holotype). Furcaria roxburghii Kostel. mAllg. Med. Pharm. Fl. 5: 1 856 ( 1 836). Hibiscus aculeatus T.K. Paul & M.P. Nayar, Bull. Bot. Surv. India 22: 195 (1980), non Walter (1788) nee Roxb. (1832). Illustration: Pradeep & Sivarajan, Taxon 40: 636, figs 1-6 (1991). A trailing or rambling shrub to 2 m tall; stems green tinged with 56 F.D.WILSON pink, with stout retrorse aculei on enlarged bases, otherwise vari- ously hairy from almost glabrous to densely tomentose. Stipules 4-1 1(-15) mm long, linear to foliate, minutely ciliate to tomentose. Leaves: petioles 0.6-10 cm long, aculeate and pubescent like the stems, the blades 3-12 x 2.5-16 cm, blades on lower leaves deeply palmately 3-5(-7)-lobed, the midlobe lanceolate to ovate, the lateral lobes ovate to triangular, the outer lobes triangular, the blades on upper leaves lanceolate, entire or moderately 3-lobed, blades of all leaves abaxially retrorse-aculeolate on the ribs, simple to stellate- pubescent on the ribs and veins, adaxially sparsely pubescent, with mostly simple hairs on ribs and veins, or tomentose, the base truncate to shallowly cordate, the margin crenate to coarsely serrate, the apex acute to acuminate, the nectary 1-2 mm long, slit-like, on the midrib and sometimes also on two lateral ribs. Flowers axillary, solitary and sometimes clustered on the ends of flowering branches; peduncles 2-10(-13) cm long, articulating 5-14 mm below the epicalyx, below the articulation with retrorse aculeoli and sparse to very dense simple hairs, above the articulation with straight simple pigmented bristles, simple to stellate hairs, and sometimes with some retrorse aculeoli; involucellar bracteoles 8-12, 5-9 mm long below the apex, linear, with fine hairs or tomentose and with straight, coarse pigmented bristles, the apex bifurcate, the outer fork 2-A mm long, lanceolate to ovate, sometimes reflexed, the inner fork 6-12 mm long, linear to subulate, ascending, the base free; calyx 1.4-2.2 cm long, the lobes triangular to lanceolate, acute to acuminate, almost glabrous or finely stellate-pubescent to stellate-tomentose and with coarse, mostly simple bristles on bases to 4 mm long on ribs, the nectary absent; corolla yellow with a purple centre, the petals 4-6.5 x 3—4.5 cm, obovate, glabrous; staminal column 1 8 mm long, dark pink, the filaments 2 mm long, the anthers dark pink, the pollen yellow to violet; style branches 6-12 mm long, the styles and stigmas red-purple. Capsules up to 15-20 x 15-17 mm, ovoid, densely appressed-pubescent, the beak 2-4 mm long, glabrous; seeds 4x3 mm, subreniform, dark brown, faveolate and with parallel striations and minute pectinate scales, the funiculus light tan, glabrous. Chromosome number, n = 72. In scrub and evergreen or semi-evergreen forest, often among rocks. DISTRIBUTION. Central and southern India and Sri Lanka; possibly adventive in South Africa. India, Bombay Presidency, southern Marathra Country and North Canara, 23 March 1881, Young s.n. (BM); Goa, Colem, 9 May 1963, Kanodia 88497 (BM); Karnataka, Coorge Dist., Mercara Hill Tops, 30 January 1976, Banner -jee 11542 (CAL); Coorg Region, c. 50 miles from coast, 3 December 1979, Pickett (ASU); Karnataka, 19 km W. of Saklespur, 19 December 1977, van derMaesen 3037 (CAL); Canara, Mangalore, 1854, Metz 1634 (P); Madras, 1885, Beddome 565, 566, 567, 568 (BM); Kerala, Idukki Dist., near Meenmutti, 18 February 1982, Raju 71251 (CAL); Ma- dras, Coimbatore, Vellingiri Hills, 1 8 February 1937, Sebastine 2377 (CAL); Madras, Nilgin Dist., near Woodbriar Estate, 18 November 1958, Sebastine 7396 (CAL); Tamil Nadu, Kanyakumani Dist., 23 January 1978, Henry 53232 (CAL); Sri Lanka, Summo Wadinagala monte, 2 November 1975, Bernardi 15617 (G-2 sheets); Sri Lanka, without specific locality, 1863, Thwaites 727 (BM, P). 6. Hibiscus goossensii (Hauman) F.D. Wilson, comb. & stat. nov. Type: Democratic Republic of Congo (Zaire), Dist. du Bas- Congo: without specific locality, 1920, Goossens 1202 (BR!-holotype). Fig. 1C. Hibiscus rostellatus var. goossensii Hauman in Bull. Jard. Bot. Etat. 31:88(1961). Shrub; stems retrorse-aculeolate, glabrous below, very sparsely hairy above. Stipules 5 mm long, linear, with simple long (>1 mm) hairs on margins. Leaves: petioles 2-7 cm long, retrorse-aculeolate plus a line of pubescence on the adaxial surface, the blades 4-10 x 4- 10 cm, hastately 3-lobed or deeply palmately 5-lobed, the midlobe and lateral lobes lanceolate to narrowly lanceolate, the outer lobes triangular, the sinuses obtuse, the blades abaxially retrorse-aculeolate and finely stellate-pubescent on ribs, sparsely stellate-pubescent on veins, adaxially finely and coarsely stellate-pubescent on ribs, with simple to stellate pubescence on veins, the base truncate to cordate, the margin serrate, the apex acute to acuminate, the nectary 2 mm long, slit-like. Flowers axillary, solitary; peduncles 3-5 cm long in fruit, articulating c. 8 mm below the epicalyx, retrorse-aculeolate and finely and sparsely pubescent below the articulation, more densely and finely pubescent and with bristles to 2 mm long above the articulation; involucellar bracteoles 10, 8-10 mm long below the apex, linear, with fine bristles on margins, the apex bifurcate, the outer fork 3-4 mm long, lanceolate, reflexed, the inner fork 8-10 mm long, linear, ascending, the base free; calyx 2.2 cm long in fruit, the lobes broadly lanceolate, slightly acuminate, the ribs not con- spicuous, with very fine stellate hairs plus a few simple to 3-fid bristles, the nectary absent; corolla yellow with a purple centre, the petals 3.5 x 2.5 cm, obovate, sparsely stellate-pubescent dorsally, glabrous ventrally. Capsules 15 x 14 mm, densely appressed- pubescent; seeds unknown. DISTRIBUTION. Fig. 5. Democratic Republic of Congo. Known to me only from the type collection. 7. Hibiscus furcatus Willd., Enum. pi: 736 (1809). Type: Southern India(?), (B-W no. 12880-holotype; ASU! -photograph). Hibiscus surattensis var. furcatus (Willd.) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 12 (1900). Hibiscus furcatus Craib, Fl. siam. enum. 1: 157 (1925), non Roxb. (1814), pro parte quoad specims. Kerr 2768, Marcan 1078. Herb or shrub to 1.5 m tall; stems with retrorse aculei plus sparse, simple to stellate pubescence below, more dense above. Stipules 1- 10 mm long, linear to foliate, with fine hairs on margins. Leaves: petioles 3-5 cm long on leaves at mid-plant (lower leaves not seen), 0.6-5 cm long on upper leaves, retrorse-aculeate and with fine and dense simple hairs, the blades on leaves at mid-plant 5-7 x 6-7 cm, moderately to deeply palmately 3-5-lobed, the midlobe broadly lanceolate to ovate or obovate, the lateral lobes broadly lanceolate to triangular or ovate, the outer lobes vestigial to broadly lanceolate, the blades on upper leaves 2-6.5 x 2-5.2 cm, moderately to deeply 3-lobed, midlobe and lateral lobes ovate, blades of all leaves abaxially with or without retrorse aculei and with simple to stellate hairs on ribs and veins, adaxially with mostly simple hairs (some to 4-fid) on ribs and veins, the base cuneate to truncate, the margin finely serrate to dentate, the apex obtuse to acute or acuminate, the nectary 1-2 mm long, slit-like, inconspicuous. Flowers axillary, solitary, and sometimes clustered near the ends of flowering branches; peduncles 0.6-1 cm long in flower, up to 2 cm long in fruit, articulating 1-5 mm below the epicalyx, below the articulation almost glabrous, above the articulation retrorse-aculeolate and with a few, fine 1-2-fid hairs; involucellar bracteoles 8-10, 7-9 mm long below the apex, linear, 3- nerved, with fine, sparse simple to 3-fid hairs and coarse simple bristles borne on enlarged bases, the apex bifurcate, the outer fork 5- 8 x 2-3 mm, lanceolate to ovate or foliate, 3-nerved, the inner fork 4-7 mm long, linear, flattened, the base free; calyx 1 .2-1 .5 cm long in flower, up to 2. 1 cm long in fruit, the lobes lanceolate to ovate, acuminate, with or without aculei, and with sparse simple bristles on ribs, otherwise almost glabrous, the nectary absent; corolla yellow with a maroon(?) centre. Capsules 13-16 x 11-14 mm, densely REVISION OF HIBISCUS SECTION FURCARIA 57 appressed-pubescent, the beak 2-5 mm long, conspicuous and glabrous, or inconspicuous and covered with appressed hairs; seeds unknown. Open jungle and dense forest, locally abundant. DISTRIBUTION. India and Thailand. India, without locality or date, Dalzell s.n. (K); without locality or date, Hooker & Thomson s.n. (BM); Waghai, forest, Daugs, 22 October 1955, Santapau 19958 (BR).Thailand, Doi Sutep, 27 October 1912, Kerr 2768 (BM); Ban TaklT, 10 November 1922, Marcan 1078 (BM). Much confusion exists in the literature about the name Hibiscus furcatus. Willdenow published the name in 1 809 for an Indian plant, as did Roxburgh in 1814. Many authors have presumed that these names refer to the same species. However, both Paul & Nayar ( 1 980, 1988: 124) and Pradeep & Sivarajan (1991), after having examined a photograph of the original type material of H. furcatus Willd. (which I have done also), pointed out that the epithet refers to two different species and that the name H. furcatus Roxb. is an illegiti- mate later homonym of Willdenow's name. Paul & Nayar (1980) took up the name H. aculeatus Roxb. as the correct epithet for the Roxburgh plant. However, Pradeep & Sivarajan (1991) pointed out that H. aculeatus Roxb. is also an illegitimate later homonym of H. aculeatus Walter. Pradeep & Sivarajan (1991) took up the name H. hispidissimus Griff, as the correct epithet, and published an excel- lent illustration for the common Indian species that is usually called H. furcatus Roxb. The true H. furcatus Willd. is genuinely different from H. hispidissimus, but it seems to occur much less commonly. 8. Hibiscus sudanensis Hochr. in Annuaire Consent. Jard. Bot. Geneve 10: 18 (1906). Type: Central African Republic, territoire de 1'Oubangi entre la porte de la Noaca et le fort Sibut, 10 December 1903, Chevalier 10757 (P!-holotype; G!-isotype). Fig. 2E. Hibiscus sudanensis var. genuinus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 10: 19 (1900), nom. illeg. (Art. 26.2). Hibiscus sudanensis var. glabrescens Hochr. in Annuaire Conserv. Jard. Bot. Geneve 10: 19 (1900). Hibiscus sudanensis var. glabrescens forma grandiflorus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 10: 20 (1900). Type: 'Cen- tral Africa' , collector and locality unknown, identified by 'XZ063' (P!-holotype). Hibiscus sudanensis var. glabrescens forma minoriflorus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 10: 20 (1900). Type: Central African Republic, territoire du Chari: vallee du Boro, 2 January 1903, Chevalier 7104 (P!-holotype; G!-isotype). Hibiscus rostellatus var. congolanus Hauman in Bull. Jard. Bot. Etat 31: 86-87 (1961). Type: Democratic Republic of Congo (Zaire), District Forestiere Centrale, Yangambi, December 1937, Louis 7032 (BR!-holotype & isotype). Hibiscus rostellatus var. sudanensis Hauman, Fl. Congo beige 10: 119(1963). Variously described as a shrub, shrubby climber, or a trailing woody vine with long internodes, scarcely branched; stems green tinged with pink, or pink, with sparse, fine retrorse aculei, and with or without a line of pubescence. Stipules 3-6(-9) mm long, filiform to linear, usually with fine hairs on margins. Leaves: petioles 2-9 cm long, with pubescence like that of the stems, the blades 6-10 x 3-8 cm, unlobed or incipiently 3-lobed, deltoid to lanceolate, ovate, or cordate, abaxially with or without retrorse aculeoli on the ribs, abaxially and adaxially with sparse, simple to stellate hairs on ribs and veins, or densely stellate-tomentose, the base truncate to cor- date, the margin shallowly dentate (almost entire) or serrate, the apex short- to long-acuminate, the nectary 1-5 mm long, slit-like. Flowers axillary, solitary or clustered near the ends of short flower- ing branches; peduncles 0.3-0.9(-1 .5) cm long, articulating near the base to about one-third the distance from the base to the epicalyx, with or without retrorse aculeoli and simple to several-fid bristles; involucellar bracteoles 8-12, 9-1 1 mm long, linear, with fine simple or stellate hairs and with or without simple to 3-fid bristles on enlarged bases, the apex bifurcate, the outer fork 1 .5-5 mm long, narrowly to broadly lanceolate, reflexed, the inner fork 2-9 mm long, linear to subulate, ascending; calyx 1 .3-1 .9 cm long, the lobes lanceolate to ovate, acute to acuminate, with simple to 4-fid bristles on the ribs, glabrous or finely stellate-pubescent between the ribs, the nectary absent; corolla lemon yellow with a large conspicuous purple centre, the petals 4-5 x 2-3 cm, obovate, finely stellate- pubescent dorsally, glabrous ventrally; staminal column 19 mm long, streaked dark pink and white, the filaments 3 mm long, the staminal column and filaments covered with very small red-purple clavate hairs, the anthers and pollen yellow; style branches 9 mm long, dark pink; stigmas dark red-purple. Capsules up to 17 x 15 mm, ovoid, densely appressed-pubescent, the beak 1-2 mm long (sometimes absent?), glabrous; seeds 4x3 mm, reniform, dark brown, faveolate and with pectinate scales, the funiculus light tan, glabrous. Chromosome number, n = 18. Riverbanks, marshy prairies, swamps, swampy forests. DISTRIBUTION. Fig. 5. Central Africa, Cameroon to Democratic Republic of Congo. Cameroon, sur la riviere Ziege pres de Djemiong (50 km SW de Batouri), 18 April 1962, Letouzey 4794 (P). Central African Republic, Ruisseau pres Riv. Eowe, 40 km N. Alindao, 24 November 1927, Tisserant 2327 (BM-2 sheets; BR-2 sheets; P); Haut Oubangui, Region de Yalinga, 25 December 1922, LeTestu 4247 (BM, P). Democratic Republic of Congo, Leopoldville Prov., Inburbain, Kimuenza, 3 December 1956, Carlier 325 (BR, K); Uele, Station Domest. Elephants, 16 November 1942, Offerman 625 (BR); Ango, November 1945, Germain 4322 (BR); Gandajika, 29 April 1957, Liben 2813 (BR). 9. Hibiscus flavo-roseus Baker f. in /. Bot. 77: 20 (1939). Type: Angola, Lunda: Saurimo, without date, Young 646 (BM!- holotype). Fig. IF. Shrub up to 1 .2 m tall; stems retrorse-aculeolate, sparsely and finely stellate-pubescent below, more densely stellate-pubescent above. Stipules 2 mm long, linear, flattened, a few hairs on margins. Leaves: petioles 0.3-2 cm long, with pubescence like that of the stem, the blades of leaves at mid-plant and upper leaves (lower leaves not seen) 1-6.5 x 0.8-6.5 cm, the blades of leaves at mid-plant deeply 3-5-lobed, the midlobe longer (sometimes markedly so) than the lateral lobes, the blades of upper leaves narrowly ovate to 3-lobed, all blades densely and finely cinereo-stellate-pubescent abaxially, sparsely stellate-pubescent adaxially, the base truncate to cordate, the margin serrate, the apex obtuse to acute, the nectary up to 4 mm long, slit-like. Flowers clustered at the summit of long flowering branches or borne singly or in small clusters on short secondary branches; peduncles 0.3-0.6 cm long in flower, apparently articulat- ing at the base, with pubescence like that of the stem; involucellar bracteoles 10, 6-7 mm long below the apex, linear, long-bristly above, the apex bifurcate, the outer fork 2 mm long, broadened, recurved, the inner fork 5 mm long, linear, ascending; calyx 1.4-1.6 cm long in flower, the lobes triangular, acute, with very fine, dense stellate pubescence and a few coarse 2-fid to stellate hairs on the marginal ribs, the nectary absent; corolla yellowish pink (according to the collector) with a purple centre, the petals c. 3 cm long. Capsules and seeds unknown. DISTRIBUTION. Fig. 5. Angola. Known to me only from the type collection. 58 F.D. WILSON V. ,.-•'•••*. Vv \ •.»••] I - •' • ••• . Sf ^kjN \..\l\\ U/-..4. i •. •-••• '•-'' • ' 10 cm Fig. 2 Leaves and fruits of species of Hibiscus section Furcaria. A: H. parvilobus, Spjut & Ensor 3 1 84 (ASU); B: H. acetosella, leaf: Gossweiler s.n. (BM), fruit: Welwitsch 5271 (BM); C: H. asper, leaf: Dalziel 426 (K), fruit: De Saeger 1038 (K); D: H. reekmansii, Troupin 16264 (BR); E: H. sudanensis, Tisserant 2327 (BM); F: H. greenwayi, leaf: Polhill & Paulo 835 (BR), fruit: Hooper & Townsend 1094 (K); G: H. mechowii, Exell & Mendonqa 1538 (BM); H: H. sparseaculeatus, leaf: Gillett 13198 (K), fruit: Ash 2783 (K). REVISION OF HIBISCUS SECTION FURCARIA 59 10. Hibiscus radiatus Cav., Diss. 3: 150, t. 54, f. 2 (1787). Type: Paris, cult, from seeds collected by Banks, Cavanilles s.n. (P- holotype). Hibiscus unidens Lindl. mBot. Reg. 9: t. 878 ( 1 823). Type: Bot. Reg. 9: t. 878 (holotype). Hibiscus lindleyi Wall., PI. asiat. rar. 1: 4, t. 4 ( 1 830). Type: Burma, Segaing on Mt. Taung Dong, Wallich 1895-1 (K-lectotype, ac- cording to Borssum-Waalkes, 1961; G!-isolectotype-2 sheets). Hibiscus furcatus Moritzi, Syst. Verz.: 29 (1846), non Roxb. (1814). Hibiscus cannabinus Mast, in Oliver, Fl. trap. Afr. 1: 204 (1868), non L. ( 1 759), pro parte quoad syn. H. radiatus. Hibiscus radiatus var. lindleyi (Wall.) Kurz in J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43: 1 10 (1874). Hibiscus cannabinus Hochr. inAnnuaire Conserv. Jard. Bot. Geneve 4: 114 (1900), non L. (1759), pro parte syn. H. radiatus, H. unidens, H. lindleyi. Hibiscus cannabinus var. unidens (Lindl.) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 115 (1900). Hibiscus cannabinus Merr. in Philipp. J. Sci. 3: 244 (1908), non L. (1759). Hibiscus cannabinus var. radiatus (Cav.) Chiov. in Atti 1st. Bot. Univ. Pavia ser. 4, 7: 125-126 (1936). Illustrations: Sims, Bot. Mag.: t. 5098 (1859); Paul & Nayar, Fasc. Fl. India 19: 145, fig. 30 (1988); Icones Roxburghianae no. 1584 (K). Erect, branched herb or subshrub to 2 m tall; stems tinged red, glabrous or sometimes sparsely aculeate. Stipules 8-12 mm long, linear to lanceolate. Leaves: petioles 2-ll(-15) cm long, sparsely aculeate or glabrous, the blades 2- 1 2 x 1.5-12 cm, deeply palmately 3-5(-7)-lobed, glabrate, the base cuneate, truncate, or subcordate, the margin coarsely serrate, the apex acuminate, the nectary absent. Flowers axillary, solitary; peduncles 0.2-0.5(-1.3) cm long, articu- lating near the base to about halfway between the base and the epicalyx, glabrous to pubescent below the articulation, bristly above it; involucellar bracteoles 8-10, 10-15 mm long, spreading, linear, sparsely bristly, the apex bifurcate (or rarely entire or obscurely bifurcate), the outer fork 4-5 mm long, narrowly lanceolate, the inner fork usually shorter, linear, the base free; calyx 1 .5-3 cm long, the lobes lanceolate, long-acuminate, sparse-bristly on the ribs, essentially glabrous otherwise, or minutely pubescent below, the nectary absent; corolla red-purple, or more rarely yellow, with a deep purple centre, the petals 6-7 x 3-4 cm, obovate, finely and sparsely pubescent dorsally, glabrous ventrally; staminal column 25-30 mm long, the filaments 2 mm long, the anthers purple, the pollen yellow-brown; style branches 2-3 mm long, red; stigmas red- purple. Capsules 20-25 x 15-22 mm, ovoid, densely appressed-pubescent, the beak 3 mm long, glabrous; seeds 4x3 mm, subreniform, dark brown, faveolate and beset with minute pectinate scales, the funiculus small, inconspicuous, glabrous. Chro- mosome number, n = 36. DISTRIBUTION. Apparently Indian in origin (an amphidiploid species, pos- sibly from hybridization of Hibiscus cannabinus with//, surattensis: Menzel, 1986: 452), but now widely cultivated. Central African Republic, Chari, 21 November 1902, Chevalier 6347 (G); Oubangi-Chari, 10 November 1924, LeTestu 1687 (BM). India, Assam, November 1890, Dr King's Collectors.n. (CAL); Coromandel, without date, Mace s.n. (P); India, without precise locality or date, Roxburgh s.n. (BM). Burma, Minlen, 1902, Shaik Mokim 548 (G); Wallich 1895c (BM). Vietnam, 'Cochinchine', without precise locality or date, Pierre 3735 (P-2 sheets). Indonesia, 'North Borneo', without precise locality, 1 877-78, Burbidge s.n. (BM); Java, without precise locality or date, Zollinger 1248 (BM). Widely cultivated in both the Old and New Worlds. Whether it ever appears as a truly wild plant is unknown. Saldanha (1985) described it as being 'frequent in deciduous forests' in Karnataka, India. Borssum Waalkes (1966) noted that it was both cultivated and ruderal in Malesia. Fryxell (1988, 1992) states that it is sometimes naturalized in Mexico and Ecuador. 11. Hibiscus torrei Baker f. in / Bot. 75: 101 (1937). Type: Mozambique: Niassa, Vila Cabral, July 1934, Torre 435 (COI- holotype; BM!-isotype, 2 sheets; G! -photograph). Fig. 1G. Shrub or much-branched perennial herb; stems with many stout retrorse aculei and lines of dense stellate pubescence and soft simple hairs. Stipules 4-5 mm long, subulate to lanceolate, with a few small fine hairs. Leaves: petioles 0.5—4.5 cm long, with pubescence like that of the stems, the blades 4. 5-1 Ox 5. 5- 10 cm, palmately shallowly 3-5-lobed, scabrous, abaxially retrorse-aculeolate on the midrib and lateral ribs and veins, adaxially without the aculeoli, the base truncate to cordate, the margin coarsely dentate, the apex acute to obtuse, the nectary apparently absent. Flowers solitary, axillary or sometimes clustered near the apex of flowering branches; peduncles 0.4-0.8 cm long, articulating at the base, with long rigid hairs on enlarged bases and some finer simple or stellate hairs; involucellar bracteoles 8-10, 6-7 mm long below the apex, linear, flattened, finely stellate-pubescent and with a few longer hairs on bases, the apex bifurcate, the outer fork 3-4 mm long, broadly lanceolate to ovate, with a prominent midvein, the inner fork 4—6 mm long, linear to subulate, the base free; calyx 1.7-2.2 cm long, the lobes triangu- lar, acute, finely stellate-pubescent on the ribs and with matted long hairs on enlarged bases and a few aculei on the margins, the nectary absent; corolla yellow with a purple centre, the petals 3-5 cm long, obovate, sparse, simple and stellate-pubescent dorsally, glabrous(?) ventrally. Capsules 19-20 x 15-17 mm, densely appressed-pubes- cent, the beak 1 mm long, glabrous; seeds subreniform, the surface faveolate (according to Exell, 1961). Damp places (Exell, 1961). DISTRIBUTION. Fig. 5. Southern Tanzania to northern Mozambique. Mo- zambique, (type collection). Tanzania, Iringa: Mufindi Dist., Uhafiwa, 3 August 1989, Kayombo 795 A (TEX); Iringa: Mufindi Dist., Luhega forest near Uhafiwa, 10 June 1989, Lovett et al. 3287 (TEX). 12. Hibiscus noldeae Baker f. in / Bot. 77: 20 (1939). Type: Angola, Malange, Quela, April 1 938, Nolde 713 (BM!-holotype). Fig. 1H. Hibiscus eetveldeanus var. asperatus de Wild, in Bull. Jard. Bot. Etat 3: 279 ( 1 9 1 1 ). Type: Democratic Republic of Congo (Zaire), Kasai, Katola, April 1908, Sapin s.n. (BR!-holotype). Hibiscus mechowii Exell & Mendonc,a, Cons. fl. angol. 1(2): 376 (1951), non Garcke (1881), pro parte quoad specim. Milne- Redhead 4100. Hibiscus furcatus Mullend. in Publ. Inst. Natl. Etude Agron. Congo Beige, Ser. Sci. 61: 79 (1954), non Roxb. (1814). Illustration: Maquet, FL rwand. 2: 383, fig. 121, 6A, 6B (1983). Variously described as a herb to 1.5m tall, a shrub to 3 m tall, or a climber with stems to 4 m long; stems retrorse-aculeate and with simple hairs or sometimes stellate-pubescent, more dense near the apex. Stipules 5-1 1 mm long, lanceolate to narrowly ovate, with fine short hairs on the margins. Leaves: petioles 1-1 1 cm long, retrorse- aculeolate and with long, mostly simple hairs, more dense on the adaxial surface, the blades 1-9 x 0.2-10 cm, on lower leaves deeply 60 palmately 3-5-lobed, the midlobe and lateral lobes lanceolate or oblanceolate, the outer lobes vestigial to lanceolate or ovate, the blades on uppermost leaves bract-like, all blades abaxially retrorse- aculeolate and with simple to 4-fid hairs on ribs, mostly simple hairs on veins, adaxially with simple to 4-fid hairs on ribs and veins, the nectary 2 mm long, conspicuous. Flowers axillary, solitary; pedun- cles 0.2-1.2 cm long, articulating from near the base to c. halfway between the base and epicalyx, densely fine-pubescent below the articulation, aculeolate and sparsely puberulent above the articula- tion; involucellar bracteoles 8-10, 7 mm long below the apex, linear to subulate, with long simple hairs or bristles on enlarged bases, mostly on margins, the apex bifurcate, the outer fork 3-6 mm long, lanceolate, not reflexed, the inner fork 1-6 mm long, linear to subulate, the base free; calyx 1.5-2.5 cm long, the lobes triangular, long-acuminate, with simple hairs or bristles on purple enlarged bases plus finer simple to 2-fid hairs, mostly on ribs, more dense below, the nectary < 1 mm long, inconspicuous; corolla yellow with a dark red to purple centre, the petals 2.5-5 x 2-2.5 cm, obovate, sparsely stellate-pubescent dorsally, glabrous ventrally. Capsules 14-20 x 1 1-15 mm, densely appressed-pubescent, the beak 1-2 mm long, glabrous; seeds 3x2 mm, reniform, dark brown, faveolate and with pectinate scales, the funiculus small, inconspicuous, glabrous. Chromosome number, n - 36 (Kachecheba, 1972). Ruderal and wild, in plantations and fallow land, in savannas, edges and clearings in forests, river banks and swamps; apparently sometimes cultivated for fibre and young leaves. DISTRIBUTION. Fig. 4. Western to eastern central Africa, from Sierra Leone to Ethiopia and south to Angola and Zambia. Sierra Leone, c. 5 miles from Kamerea on Kurubonla road, 1 4 November 1 965, Morton SL2486 (K). Ivory Coast, Mt. Nimba, 14 December 1967, Geerling & Bokdam 1713 (BR). Nigeria, NE State, Sardauna Prov., Mambilla Plateau, 9 October 1973, Chapman 3249 (K). Cameroon, cult., 6 km W. of Bandjoum, 26 August 1972, Leeuwenberg 10288 (BR); 36 km N. de Bafia, 22 November 1969, Letouzey 9600 (P); Bamenda, 2 February 1928, Migeod 463 (BM). Central African Republic, Oubangui, Region de la Waka, 10 November 1924, Tisserant 1687 (P). Sudan, Mongalla Prov., S. of Juba-Yei road between Libogo and Yei, 23 February 1934, Dandy 485 (BM). Ethiopia, c. 40 km N. of Lekemti, ...tributary of Angar River, 13 November 1965, deWilde & deWilde-Duyfjes 8866 (B, BR, P). Democratic Republic of Congo, Nioka, 1932, Jurion 128 (BM-2 sheets); Kivu Prov., Kabare Terr., Birava, 30 April 1 960, Meurillon 8 1 9 (BR); Kaniama-Haut Lomani, 1 0 June 1 947, Mullenders 290 (BR). Uganda, SE of Kampala, N. of Nioka, 30 April 1952, Sperry 553 (BR). Rwanda, Butare pref., Gihindansuyagu, 8 June 1978, Raynal 20345 (P). Burundi, Ruyigi: Muzire, 2 May 1980, Reekmans 8981 (K). Angola, Moxico, E. of Lusavo Falls, 13 January 1938, Milne-Redhead 4100 (BM, BR). Zambia, Kawambwa Dist., Mbereshi River swamp, 20 April 1957, Richards 931 '\ (BR). One collection from Cameroon, Letouzey 9600 (P), has unusually long involucellar bracteoles which equal the calyx in length at anthesis. 13. Hibiscus nigricaulis Baker f. in 7. Bot. 77: 19-20 (1939). Type: Angola, Bie: between Coemba and R. Cuanza, 7 May 1937, Ere// & Mendonca 1759 (BM!-holotype). Fig. IE. Hibiscus cannabinus Ulbr. & R.E. Fries in R.E. Fries, Wiss. Ergebn. Schwed. Rhod.-Kongo-Exped. 1: 145 (1914), non L. (1759), pro parte quoad specim. Fries 954. Hibiscus cf. diversifolius Verdoorn & Collett in Farming South Africa 34: 2, fig. 2 (1947), non Jacq. (1789). Hibiscus sabdariffa Mendonc.a &Torre, Contr. Conhec. Fl. Mocamb. F.D. WILSON 1: 1 1 (1950), non L. (1753), pro parte quoad specim. Mendonqa 332 (LISC). Hibiscus meeusei Exell in Bol. Sac. Brot., sen 2, 33: 165 (1959). Type: SouthAfrica, Transvaal, BronkhorstspruitDist., Donkerhoek (Donkerpoort), between Pretoria and Bronkhorstspruit, 19 March 1959, Meeuse 10646 (BM!-holotype; G!, P!-isotypes). Illustration: (asHibiscus cf. diversifolius) Verdoorn & Collett, Farm- ing South Africa 34: 2, fig. 2 (1947). Annual (sometimes biennial?) herb to 1.5 m tall; stems aculeolate and stellate-pubescent. Stipules 4-10 mm long, filiform to narrowly subulate, with short, fine pubescence. Leaves: petioles 1-12 cm long, with pubescence like that of the stems, the blades 2-12 x 0.2- 15 cm, those of lower leaves deeply palmately 3-7(-9)-lobed, the lobes oblanceolate, obovate, or triangular, sometimes secondarily lobed, the blades of upper leaves less deeply lobed or even bract-like at the apex, all blades with fine simple and stellate hairs abaxially and adaxially, the base cuneate, truncate, or very shallowly cordate, the margin regularly to irregularly serrate, the apex obtuse to acute, the nectary 1^4 mm long, slit-like. Flowers axillary, solitary, or clustered on short-to-long flowering branches, the short flowering branches with a subtending, bract-like leaf with a nectary borne above the axillary leaf; peduncles 0.4-0.7 cm long, articulating at the base, with pubescence like that of the stems; involucellar bracteoles 8-9(-12?), 8-10 mm long, linear, flattened, finely stellate- pubescent and with coarse bristles on enlarged bases mainly on the margins, the apex obscurely bifurcate, the outer fork 2 mm long, reflexed, the inner fork <0.5 mm long, vestigial, or sometimes absent, the base free or slightly united; calyx 1 .4-2.0 cm long, the lobes lanceolate to ovate, more or less acuminate, finely stellate- pubescent and with coarse bristles on bases, mostly on the ribs, the nectary 1 mm long, usually conspicuous; corolla cream to light yellow, with a pinkish red to dark red-purple centre, the petals 2-4 x 1 .5-3 cm, obovate, glabrous to stellate-pubescent dorsally, glabrous ventrally; staminal column 10-14 mm long, pink or red, the fila- ments 0.5 mm long, the anthers pink or purple, the pollen yellow or tan; style branches included in staminal column or exserted up to 1 mm long; stigmas red-purple, usually not exserted beyond the anthers. Capsules 14-18 x 11-16 mm ovoid, densely appressed- pubescent, the beak 1-2 mm long, glabrous; seeds 3^4 x 2-3 mm, subreniform, dark brown, finely faveolate and with sparse pectinate scales, the funiculus small, dark brown, glabrous. Chromosome number, n = 36. Ruderal and wild, in old cultivated lands and waste places, disturbed native vegetation, moist places and marshland. DISTRIBUTION. Fig. 4. Southern Africa from Angola, Zambia, and Mozam- bique to South Africa. Hauman (1963) also cites several specimens (as H. meeusei) from southeastern Democratic Republic of Congo (Zaire), none of which I have seen. Angola, Huila, Humpata, 24 April 1964, de Menzies 1061 (BM); Okavongo Nat. Terr., Okavongo River at Dujona Camp, 2 miles E. of Nyangana Mission Station, 18 February 1956, deWinter & Marais 4757 (BM). Zambia, Kasama Dist., 80 km S. of Kasama, 29 April 1962, Robinson 5121 (B); NW Dist., near Kitwe Railway Station, 8 April 1961, Linley 133 (BM); Ndola [North], 20 March 1954, Fanshawe 979 (K); Lusaka Dist., April 1957, Noak 193 (BM); S. Prov., Kalomo Dist., 18 March 1962, Astle 1516 (K). Mozambique, Manica E Sofola, Chimoio, Garuso, 3 March 1948, Barbosa 1086 (BR). Zimbabwe, Lomagundi, UmboiValley, 28 March 1950, Colville 108 (BM); Odzani Banks, March 1935, Gilliland 1721 (BM); Victoria Falls, May 1915, Rogers 13157 (BM). Namibia, Tsumeb, April 1934,D/nter7588 (BM); Grootfontein Nord, zwischen Numkaub und Bumbi im Omuramba, 5 March 1958, Merxmuller & Giess 1827 (BM). South Africa, Transvaal, Waterberg, 4 km Kienaars River Station, 18 March 1947, Codd 2749 (BM, PRE); Transvaal, Kwandebele, farm Gemsbokfontein, 17 REVISION OF HIBISCUS SECTION FURCARIA 61 March 1 98 1 , Hoepen 1 627 (PRE); Transvaal, Zoutpansberg Dist., c. 2 mi W. of Hangklip along road to Buckworth, 4 April 1957, Meeuse 10234 (BM); Natal, 14 July 1929, Salter 382/12 (BM); Natal, Port Shepstone, Izotaha ravine, 8 July 1967, Strey 755 1 (PRE). 14. Hibiscus mechowii Garcke in Linnaea 43: 121 (1881). Type: Angola, Cuanza Norte, without date, Mechow 105 (B-holotype; BM!-drawing). Fig. 2G. Hibiscus lancibracteatus de Wild. & T. Durand in Bull. Soc. Roy. Bot. Belgique 38(2): 25 (1899). Type: Democratic Republic of Congo (Zaire), Coquilhatville, 17 February 1896, Dewevre 752 (BR!-holotype). Hibiscus cannabinus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 14 (1900), non L. (1759), pro parte quoad syn. H. mechowii (4: 115). Illustration: (as//, lancibracteatus) deWild. &T. Durand, Ann. Mus. Congo, ser. 1, Bot. 1: 147, t. 84 (1901). Herb or shrub to 2 m tall; stems aculeolate and with long fine simple hairs and lines of pubescence. Stipules 10-20 mm long, filiform to linear, almost glabrous. Leaves: petioles 2-7 cm long, aculeolate and with long simple hairs, the blades 4-10 x 3-8 cm, very deeply palmately 3-5-lobed, the lobes linear, or rarely, shallowly lobed (e.g. Nannan 24) both surfaces covered with fine simple hairs plus a few stout hairs on enlarged bases on ribs, the base cuneate, the margin serrate or with minute antrorse barbs, the apex acute, the nectary 3 mm long, slit-like. Flowers axillary, solitary; peduncles 0.5-1.2 cm long, articulating at the base, with dense short woolly pubescence and long simple hairs; involucellar bracteoles 9-10, 1 1- 22 mm long, with long simple hairs and pigmented stout bristles, linear below the apex, the upper 6-1 1 mm channelled on the ventral surface, several-nerved on the dorsal surface, becoming lance- shaped upon drying, the apex entire, pointed, the base united; calyx 2-3 cm long, the lobes broadly lanceolate, acuminate, with long simple matted hairs, mostly on the ribs, the nectary absent; corolla lemon yellow with a large red-brown centre, the petals 4-6 x 2-3.5 cm, obovate, glabrous dorsally and with a few simple hairs ventrally; staminal column 15 mm long, light pink, the filaments 1 mm long, the anthers purple, the pollen brownish red; style branches included in the staminal column or to 3 mm long; stigmas white to red. Capsule 20-26 x 15-18 mm, ovoid, densely appressed-pubescent, the beak 5 mm long, pubescent; seeds 4x3 mm, subreniform, with sparse fine matted hairs. Chromosome number, n = 18. Ruderal and wild (rarely cultivated?), in abandoned plantations or cultivated fields, wooded steppes, savannas, sandy, shady river banks, swampy alluvial plains. In Central African Republic, the stem fibres are reportedly used for making fish nets and wrapping steamed manioc (Fay 5295 (K)). DISTRIBUTION. Fig. 4. Central and southern Africa, Central African Repub- lic to Namibia, Botswana, Zimbabwe, and Mozambique; apparently disjunct in extreme western Africa. Guinea-Bissau, Bissau, Quinhamel, 17 Novem- ber 1960, Raimundo & Guerra 48 (K). Senegal, Region de Bignona, Tendouk, 13 December 1963, Berhaut 6773 (P). Central African Republic, Territoire du Chari, 2 December 1902, Chevalier 6546 (P); Bossangoa, 10 October 198 1 , 600 m, Fay 5295 (K); Monova-Gounda-St. Floris Nat. Park, 1 Novem- ber 1983, Fay 6102 (K); Haute-Kotto,Yalinga, 9 October \92\,LeTestu 3339 (BM); Oubangui, Region de Bambari, 10 October 1923, Tisserant 1246 (P). Democratic Republic of Congo, Kivu Prov., UviraTerr., Uvira, 8 June 1 956, Kinet 148 (BR); Terr. Popokabaka, Kinata, 25 April 1959, Pauwels 2661 (BR); cultivated, pres village Bonananga, 22 October 1913, Nannan 24 (BR); Democratic Republic of Congo, without locality, August 1914, Vanderyst 3972 (BM). Burundi, Bubanze, Randa, 5 June 1980, Reekmans 9267 (K). Tanzania, T4, Lukoma, Kigoma Dist., L Tanganyika, 29 May 1975, Kahurananga, Kibuwa & Mungai 2688 (BR); Bezirk Lindi, 50 km W. Lindi, 24 April 1935, Schlieben 6485 (BM). Angola, Mexico, River Luena, near Vila Luso, 3 May \937,Exell& Mendon^a 1 538 (BM);PungoAdongo, April 1857, Welwitsch 5262 (BM). Zambia, Bulaya, June 1950, Bullock 2910 (BR);Abercorn Dist., Kawi Village, 19May 1 936, Burn 61 47 (BM); Zambesi, Gonye", 28 April \925,Pocock 170(BOL);NamwalaDist., 1 934-35, Read 54 (BM); Abercorn Dist., Chilongowelo, 10 May 1952, Richards 1697 (K); Bulaya, NEof Mweru-Wantipa, 1 1 August 1962, Tyrer40l (BM). Namibia, Okavango Native Territory, 4 March 1956, de Winter & Marais 4973 (PRE); Grootfontein-Nord, Siidlich Mavanze, im Omuramba, 5 March 1958, Merxmuller & Giess 1871 (BM). Zimbabwe, Victoria Falls, May 1915, Rogers 13155(BOL). 15. Hibiscus subdiversifolius Hochr. in Annuaire Conserv. Jard. Bot. Geneve 20: 83-85 (191 6). Type: Madagascar, loco collectore haud indicate, n. 22 (G!-holotype). Hibiscus diversifolius var. subdiversifolius Hochr., Fl. Madagasc., Malvac.: 40-41 (1955). Illustration: Hochreutiner, Fl. Madagasc., Malvac.: 39, fig. XI, 1-2 (1955). Shrub 3-4 m tall; stems aculeate and very densely tomentose. Stipules 4 mm long, filiform, with dense, short pubescence. Leaves: petioles 0.5-10 cm long, very densely tomentose, the blades 1.5- 12.5 x 0.5-14.5 cm, lanceolate to ovate or very shallowly 7-lobed, abaxially and adaxially densely stellate-pubescent, the base cuneate, truncate, or deeply cordate, the margin serrate or dentate, the apex acute to acuminate, the nectary 1.5 mm long, slit-like. Flowers apparently clustered near the apex of the stem; peduncles 0.6 cm long, articulating midway between the base and the epicalyx, densely short-tomentose below and above the articulation, and with long simple hairs above it; involucellar bracteoles 6(?), 10 mm long, linear, flattened, channelled on the ventral surface, densely tomentose, the apex entire, the base free; calyx 1.5-1.7 cm long, the lobes triangular, acuminate, densely and finely stellate-pubescent and with long, simple to stellate(?) hairs, mainly on the ribs, the nectary apparently absent; corolla colour unknown, but with dark-coloured centre, the petals 5.5 x 2 cm, obovate, finely stellate-pubescent dorsally, glabrous ventrally; staminal column tan, filaments 1 mm long, anthers purple. Capsules and seeds unknown. Riverbanks and vicinity (Hochreutiner, 1955). DISTRIBUTION. Fig. 5. Madagascar. I have seen only the type collection which cites Madagascar without specific locality, but Hochreutiner (1955) lists two Perrier collections (5343, 5464) from western Madagascar. The specimen mounted on Dale 2479 (G!), collected at the Nairobi, Kenya arboretum in September 1930, is Hibiscus subdiversifolius even though the label states that the plant was collected originally near Embu, Kenya, and was identified as Hibiscus sp. near H. panduriformis Burm.; a later label identifies the plant as Hibiscus diversifolius Jacq. 16. Hibiscus berberidifolius A. Rich., Tent.fl. abyss. 1: 56 (1847). Type: Ethiopia (Abyssynia), Quartin Dillon et Petit s.n. (P!- lectotype, Wilson, 1983; BR!-isolectotype). Hibiscus diversifolius var. witteanus Hochr. in Robyns, Bull. Jard. Bot. Etat 10: 276 (1947). Type: Democratic Republic of Congo (Zaire), Kilisti, route de Kibumba, foret, 17 January 1934, de Wine 1320 (G!-holotype; BR!-isotype); Tshamugussa, 9 August 1934, de Witte 1817 (BR!, G!-paratypes). Illustrations: Maquet, Fl. rwand. 2: 383, fig. 121, 2A, 2B (1983); Wilson, Brittonia 35: 176, fig. 1 (1983). 62 F.D. WILSON Robust, few to many-branched perennial shrub, 1-4 m tall; stems sparsely to moderately aculeate, lower and mid-stem with a longitu- dinal line of dense fine pubescence to 2 mm long, alternating position at each node, also with sparse, mostly simple hairs, upper stems densely and uniformly pubescent. Stipules 2-4 mm long, linear, inconspicuous, very finely pubescent, sometimes lacking (caducous?). Leaves: petioles 0.3-3.5 cm long, aculeate, villous, the blades 0.9-8.5 x 0.4-6 cm, the blades of lower leaves ovate or shallowly 3-lobed, the blades of upper leaves obovate, narrowly obovate or bract-like, vegetative branchlets in leaf axils well-devel- oped, with up to 7 persistent leaves, the blades of leaves on vegetative branchlets ovate to obovate, blades of all leaves abaxially and adaxially with simple and stellate hairs on the veins, the base cuneate to truncate, the margin serrate to coarsely serrate (newly opened leaves exhibit a gland-like, thickened structure at the apex of each serration), the apex acute, the nectary present or absent. Flowers axillary, solitary, disposed in the upper 7-23 cm of the branch; peduncles 0.1-2 cm long, accrescent, articulating at the base or apparently unjointed, densely and finely stellate-pubescent and with dense, simple hairs to 2 mm long; involucellar bracteoles 6-7, 9-13 mm long, subulate, flattened, obscurely several-nerved, apex entire, with long simple hairs on margins; calyx 1.4-2.2 cm long, the lobes short- to long-acuminate, villous below, finely pubescent above, the nectary usually absent; corolla varying from white to several shades of off-white, yellow, and purple, with a small, purple centre, the petals 3-5.5 x 3-5.5 cm, obovate to broadly obovate, sparsely simple- or stellate-pubescent dorsally, glabrous ventrally; staminal column 20-40 mm long. Capsule 1 3-17 x 1 1-17 cm, broadly ovoid, densely appressed-pubescent, the beak absent or up to 1 mm long, glabrous; seeds 4x3 mm, subreniform, dark brown, faveolate and with short, white, many-branched hairs. Chromosome number, 2n = 36. Savanna and grassland, upper elevation forests, bamboo and Hagenia zones. DISTRIBUTION. Mountains of East Africa ( 1 300 to 2700 m) associated with the Rift Valley in Democratic Republic of Congo, Uganda, Rwanda, Tanza- nia, Kenya, and Ethiopia (distributional map and specimens cited in Wilson, 1983). 17. Hibiscus sparseaculeatus Baker f. in J. Bot. 76: 22 (1938). Type: Somalia, Sheik Pass, Freemantle s.n. (BM!-holotype). Fig. 2H. Cienfuegosia sp. Cufod. in Chiovenda, Miss. biol. borana 4: 345, fig. 117(1939). Hibiscus greenwayi Cufod. in Bull. Jard. Bot. Etat. 29: Suppl.: 562 (1959), non Baker f. (1937), pro parte quoad specim. Bally 9128. Hibiscus greenwayi var. megensis J.P. Lebrun inAdansonia 15: 379 (1976). Type: Kenya, Nyiro, 4000-5000 ft, stony hills, Haylett 12 (K-holotype). Illustration: Cheek, Fl. pi. afric. 52: pi. 2058 (1992). Rigid, woody shrub to 3 m tall; stems with conspicuous, long (to 6 mm), red to brown, stout aculei, borne singly, in pairs, or in threes at the base of each node, stems otherwise glabrous and sometimes glaucous below. Stipules not seen. Leaves apparently shed early (some flowering and fruiting specimens are leafless); petioles 0.5-3 cm long, densely stellate-pubescent, the blades 0.6-2 x 2-4 cm, oval or rotund, densely stellate-pubescent abaxially and adaxially, the base truncate to cordate, the margin and apex dentate, the nectary 0.2-0.5 mm long. Flowers semi-pendulous, axillary, solitary; pe- duncles 0.5-3 cm long, articulating at or near the base, densely stellate-pubescent and sometimes with antrorse aculeoli; involucellar bracteoles 7-9, 5-1 1 mm long, narrowly subulate, densely stellate- pubescent, the apex entire, pointed; calyx 1 .2-2.0 cm long, the lobes lanceolate, short- to long-acuminate, densely stellate-pubescent and sparsely aculeolate, the nectary small but conspicuous; corolla yellow with a crimson-purple centre, the petals 4—6 x 3-5 cm, obovate, simple to stellate-pubescent dorsally, glabrous ventrally; staminal column 25-37 mm long, crimson, the filaments 0.5 mm long, the anthers red-brown, the mature pollen purple(?); style branches 9-12 mm long, red-brown below, purple(?) above, the stigmas purple. Capsules 10-20 x 9-15 mm, long appressed-pubes- cent, the beak 1-2 mm long, glabrous; seeds 4x3 mm, with fine pectinate scales, the funiculus with a dense fringe of short hairs. Sandy or red soils, gravelly or rocky slopes, dry bushland domi- nated by Acacia-Commiphora and Sansevieria (Cheek, 1992), 750-2000 m elevation. DISTRIBUTION. Fig. 4. Eastern Africa, from southern Ethiopia and Somalia to northern Tanzania. Ethiopia, Sidamo Prov., 8 miles S. of Mega on International Hwy., 25 December 1974, Ash 2783 (K); Km 10 SE Mega, 4 February 1973, Boudet 8166 (P); Borama, October 1945, LePelley in Bally 4663 (K). Kenya, Northern Province, Dandu, 14 June 1952, Gillett 13198 (BR, K-2 sheets); Northern Province, Furroli Mt., 15 September 1952, Gillett 13874 (B); Kl, N. side of Lolockwe Mt., 28 km NNW of Archers post, 20 April 1973, Gillett20l99 (K). Tanzania, Northern Prov., Mbulu Dist., N. end Lake Eyasi, 23 July 1957, Bally 1 1588 (BR, G-2 sheets). 18. Hibiscus greenwayi Baker f. in /. Bot. 75: 99 (1937). Type: Tanzania, NW Usambara, Mnazi, 12 January 1930, Greenway 2034 (EA-holotype; BM!-isotype). Fig. 2F. Large, perennial, much-branched shrub to 3 m tall; stems almost glabrous below to densely and finely stellate-pubescent above and with few to many stout conical aculei. Stipules 4-6 mm long, filiform, with dense, fine, short hairs. Leaves: petioles 0.4-13 cm long, stellate-tomentose, the blades 2.5-8 x 0.9-1 1 cm, the blades of lower leaves palmately 5-lobed, the midlobe and lateral lobes rotund to spathulate, the outer lobes ovate to oval, the blades of upper leaves entire and oblanceolate or 3-lobed, all blades densely and finely stellate-pubescent abaxially and adaxially, the base cuneate, trun- cate, or cordate, the margin coarsely dentate or repand, the apex obtuse or acute, the nectary 0.5-1 mm long. Flowers pendulous, axillary, solitary and clustered near the apex; peduncles 0.3-0.9 cm long, articulating at the base, stellate-tomentose and sometimes with a few antrorse aculeoli; involucellar bracteoles 6-10, 8-14 mm long, subulate, sometimes slightly acuminate, with fine and dense simple hairs to stellate-tomentose, the apex entire, pointed, the base free; calyx 1.7-2.2 cm long, the lobes lanceolate, acuminate, densely and finely stellate-pubescent and with a few antrorse aculeoli on the ribs, nectary small but conspicuous; corolla funnelform, lemon yellow to primrose yellow, with a red-purple centre, the petals 4-8.5 x 3-6.5 cm, obovate to oval, sparsely and finely to coarsely stellate-pubes- cent dorsally, glabrous ventrally; staminal column 35-60 mm long, dark pink to red-purple, the filaments 0.5-1 mm long, the anthers red to purple, the pollen yellow, orange-tan, or purple; style branches included in the staminal column or up to 9 mm long, red or purple, the stigmas dark red-purple. Capsules 16-18 x 1 1-14 mm, densely appressed-pubescent, the beak 1-2 mm long, glabrous; seeds 5-6 x 3-4 mm, subreniform, dark brown, faveolate and with long (0.5 mm) simple, flattened tan/brown hairs, the funiculus densely covered with simple, flattened straw-coloured hairs. Chromosome number, n = 18. Ditch banks with grasses and shrubs, dry savannas, dense Acacia- Commiphora thombush, edge of Brachystegia woodlands. REVISION OF HIBISCUS SECTION FURCARIA 63 DISTRIBUTION. Fig. 4. Uganda, Kenya, and Tanzania. Uganda, about the Friffen Falls (Mabira Forest), 18 February 1918, Dummer 3908 (BM). Kenya, Kwale Dist., between Samburu and MacKinnon Road, 31 August 1953, Drummond & Hemsley 4078 (K); Taita Dist., Tsavo West, hill c. 4 km SE of Manda Hill, 24 July 1969, Gilbert 4073 (EA); K7, Kilifi Dist., outside SalaGate, 59 kmW. of Malindi, 28 February 1977, Hooper & Townsend 1094 (K-2 sheets); Voi Dist., foot of Taita Hills, Bura, 14 April 1969, Jones 69159 (NA); K4, Kitui Dist., Yakaseva, Voo location, 24 August 1968, Kimani 78 (EA); K4, Kitui Dist., Mutomo, 24 April 1969, Napper & Kanuri 2070 (B, EA); Kilifi Dist., Marafa, 25 miles NW of Malindi, 22 November 1961, Polhill & Paulo 835 (B, BR); Kwale Dist., MacKinnon Rd., c. 54 miles from Mombasa, 12 June 1964, Verdcourt 3904 (EA, K); Kwale Dist., 8 km E. of MacKinnon Rd on Nairobi-Mombasa Hway (A-109), 12 August 1975, Wilson 75140 (ASU, EA, NA); Kwale Dist., 30 km W. of Mombasa on Nairobi-Mombasa Hway (A- 109), 13August 1975, W/5on75143(ASU,EA, NA). Tanzania, Umba Steppe, plains at base of Usambara Mts, 3 February 1954, Faulkner 1339 (B, K);T3, N. of Mombo, 8 September 1972, Flock43\ (EA). 19. Hibiscus partitus (Hochr.) F.D. Wilson, comb, et stat. nov. Hibiscus diversifolius var. partitus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 120 (1900). Type: Madagascar, environ de Tananarive, Goudot s.n. (G). Illustration: Hochreutiner, Fl. Madagasc., Malvac.: 37, fig. X, 9, 10 (1955). Habit unknown; stems large, stout-aculeate and with lines of stellate pubescence below, more densely stellate-pubescent above. Stipules 4-5 mm long, filiform to linear, finely short-pubescent. Leaves: petioles 0.3-5 cm long, aculeolate abaxially and with a line of dense pubescence adaxially, the blades 1.3-5 x 0.2-6 cm, on lower leaves palmately 5-lobed and manifestly secondarily lobed, on upper leaves bract-like or 5-lobed and secondarily lobed, all blades abaxially aculeolate on the ribs, veins, and margins, plus some dense, fine simple to stellate hairs, adaxially with simple to stellate hairs, usually not as dense, the base cuneate to truncate, the margin entire, serrate, or coarsely serrate and secondarily lobed, the apex acute to acuminate, the nectary minute (< 0.5 mm long) but conspicuous. Flowers axillary, solitary; peduncles 0.3-0.9 cm long, articulating at the base, antrorse-aculeolate and finely stellate-tomentose; involucellar bracteoles 7-8, 7-11 mm long, linear to narrowly subulate, with 2-fid or stellate hairs or short bristles, the apex entire, the base free or scarcely united; calyx 1 .2-2. 1 cm long, the lobes lanceolate to narrowly ovate, slightly to long-acuminate, aculeolate on the ribs and with short, rigid bristles, more dense near the base, the nectary small (0.2 mm long) but conspicuous; corolla yellow(?), with a large purple(?) centre, the petals 4-5.5 x 2.5-3.5 cm, obovate, with sparse 2-4-fid hairs dorsally, glabrous(?) ventrally. Capsules 14-20 x 12-15 mm, densely appressed-pubescent, the beak 1 mm long, glabrous; seeds unknown. Rocky habitats and dried-out ponds (Hochreutiner, 1955). DISTRIBUTION. Fig. 5. Central and northern Madagascar. Madagascar, Central, without precise locality and date, Baron 542, 3286, 3945 (K); North- west, without precise locality and date, Baron 5347 (K). Hochreutiner (1955) cites several specimens from west-central Madagascar. 20. Hibiscus diversifolius Jacq., Collectanea 2: 307 (1789). Type: Icon, pi rar.: t. 551 (1792) (neotype, Fryxell, 1988). This was recognized as the holotype by Borssum Waalkes (1966), but as Fryxell (1988) points out, the plate was published several years after the name was published. Perennial subshrub or shrub to 3 m tall (or rarely a tree to 10 m tall), the stems below with many stout conical aculei and with one or more lines of pubescence, or more densely stellate-pubescent, the stems above aculeolate, densely and finely simple- and stellate-pubescent. Stipules 3-7 mm long, linear, pubescent. Leaves: petioles 0.6-22 cm long, with pubescence like that of the upper stems, the blades 3-16 x 0.7-14 cm, the blades of lower leaves ovate to shallowly or very deeply palmately 3-7-lobed, the blades of uppermost leaves bract- like, linear or narrowly lanceolate, all blades abaxially with simple and stellate hairs on the ribs and veins and adaxially with simple and stellate hairs on the ribs, and simple and 2-fid hairs on the margins, the base cuneate, truncate, or cordate, the margin serrate, dentate, or crenate, the apex acute or obtuse, the nectary 1-2 mm long, con- spicuous, on the midrib and sometimes on two lateral ribs. Flowers axillary, solitary, or with 1-3 flowers at each node, or borne in racemes formed by the reduction of the upper leaves and shortening of the internodes; peduncles 0.2-0.8 cm long, articulating at the base, densely stellate-pubescent and with or without long, simple bristles; involucellar bracteoles 5-10, 8-15 mm long, linear to subulate, slightly concave ventrally, bristly, the apex entire, the base free; calyx 2-3 cm long, the lobes lanceolate, acute to short- acuminate, covered with stiff bristles, the nectary conspicuous; corolla light yellow, reddish, or purple, with a dark red-purple centre, the petals 3-6 x 2-5 cm, obovate, sparsely simple- and stellate-pubescent dorsally, glabrous ventrally; staminal column 15- 32 mm long, white, pink, red, or purple, the filaments 1-2 mm long, dark red-purple, the anthers light pink, red, or purple, the pollen orange; style branches included in the staminal column or to 7 mm long, dark pink to purple; stigmas dark red-purple. Capsules 20-24 x 15-19 mm, densely appressed-pubescent, the beak 2 mm long, glabrous; seeds 4x3 mm, subreniform, dark brown, glabrous, with parallel striations, the funiculus tan, conspicuous, glabrous. Chro- mosome number, n = 72. A form from Uganda, Synge S.1042 (BM), is closely allied to Hibiscus diversifolius, but has foliaceous involucellar bracteoles and may be an undescribed species. 20a. Hibiscus diversifolius Jacq. subsp. diversifolius Hibiscus ficulneus Cav., Diss. 3: 148, t. 51, fig. 2 (1787), non L. (1753), nom. illeg. (Art. 53.1). Hibiscus scaber Lam., Encycl. 3: 350 (1792). Type: Mauritius (lie de France), Commerson s.n. (P-LA-holotype). ?Hibiscus biflorusA. Spreng., Tent, suppl.: 19 (1828). Type: South Africa, Zeyher 241 (BM!-sketch of type). Hibiscus macularis E. Mey. ex Harv. in Harv. & Sond., Fl. cap. 1: 171 (1860), nom. nud. Hibiscus decaisneanus Schimp. ex Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4:119 (1900), nom. nud. Type: Ethiopia, Tigre v.Begemder, lONovember \863,Schimper 1479(BM!-isotype?). Hibiscus diversifolius subsp. genuinus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 19 (1900), nom. illeg. (Art. 26.2). Hibiscus paludosus Merr. inPhilipp. J. Sci. 3: 151 (1908).Type: The Philippines, Mindanao, Lake Lanao, Clemens & Clemens s.n. (BO, G, L-isotypes). Hibiscus berberidifolius Cufod. in Bull. Jard. Bot. Etat. 29, Suppl.: 560 (1959), non A. Rich. (1847). Hibiscus diversifolius var. witteanus Cufod. in Bull. Jard. Bot. Etat. 29, Suppl.: 560 (1959), non Hochr. (1947). Hibiscus diversifolius var. angustilobus Hauman in Bull. Jard. Bot. Etat. 31 : 86 ( 1 96 1 ) excl. specims. Laurent 1 1 , de Witte 2435 . Type: Democratic Republic of Congo (Zaire), Haut-Katanga: Sakala, Marungu, April 1944, Dubois 1 157 (BR!-holotype). Illustrations: Maquet, Fl. rwand. 2: 383, fig. 121, 4 (1983); An- 64 drews, Fl. pi. anglo-egypt. sud. 2: 24, fig. 10 (1952); Hochreutiner, Fl. Madagasc., Malvac.: 39, fig. X, 6-8 (1955). Stems usually with one or more longitudinal lines of pubescence, flowers yellow (Exell, 1961). In generally moist habitats, riverbeds, river banks, swamps, in papyrus and other vegetation at lake edges, bays; also reported as ruderal plants on road shoulders and in grass and bushland. DISTRIBUTION. Fig. 4. Circumtropical. Widely distributed in subsaharan Africa from Cameroon and Ethiopia to Madagascar and South Africa, except in the Flora Zambesiaca area (Zambia, Malawi, Zimbabwe, Mozambique, Botswana), where it is replaced by subsp. rivularis (Exell, 1961). Wilson & Kissie 753 1 (subsp. rivularis) and 7532 (subsp. diversifolius) were growing together in a papyrus-7vp//a swamp in the Musoma Region, Tanzania (Wilson, 1978); disjunct in Senegal? Senegal, Niombato, 1950-51, Berhaut 1061 (BR, P). Cameroon, pres village de Pecheurs de Mouhengue (=Malimba), 6 January \914,Letouzey 1 2629 (P);Ndop Plain, 6 March 1962, Bran/ 146 (K- 2 sheets). Ethiopia, Shoa Prov., Lake Shala, 14 November 1971, Ash 1357 (K); Agaro, 24 January 1969, Jones 6908 (K). Congo, Region de Brazzaville, lie M'Bamou, 13 October 1967, Sita 1818 (P); Democratic Republic of Congo, Plaine de la Ruzizi, lac Ranyinia, January 1950, Germain 5650 (G); Kivu Prov., Kabare Territ., Lulonge, 9 January 1967, Vermylen 35 (BR). Uganda, Kyaka, Toro, 1914, Snowden 31 (BM, K). Kenya, 13 miles S. of ThikaonNairobi-ThikaRoad(HwayA-2), 17 July 1975, Wilsonl5\\ (ASU, EA, NA); c. 9 km from Nairobi on road to Nakuru, 24 June 1977, Gachathi 287 (B); Nairobi Prov., Kiririchusa Ndogo Valley, 10 July 1944, Bally 3407 (G).St. Helena, above Lufkins, 23 February 1955, Kerr 108 (BM). Rwanda, 22 August 1958, Michel 5562 (BR). Tanzania, Musoma Region, 8 km N. of Nyakanga on Hwy B-6, 23 July 1975, Wilson & Kissie 7532 (ASU, EA, NA); Laliondo Dist., 6 July 1956, Williams 702 (BR); Kirumba, Ilemera, Mwanza, 24 June 1953, Tanner 1608 (K); Ndingo, Songea Dist., 29 September 1956, Semsei 2490 (K). Malawi, Station Kyimbila, 29 September 1910, Stolz 307 (G); N. Prov., Mzimba Dist., Vipya Plateau, 14 July 1975, Pawek 9887 (K). Madagascar, Nord Betsileo, Linabe, August 1 880, Hildebrandt 3567 (BM, G); central Madagascar, 1855, Baron 4169 (BM). South Africa, Natal, Alexandra Dist., Dumisa, 12 May 1908, Rudatis 380 (BM); Cape Peninsula, Retreat, April 1 95 1 , Pillans 10565 (G); Cape Prov., Voeklip camping area, 29 August 1985, Williams 1087 (PRE); Cape Prov., Humansdorp, November 1921, Fourcade 1781 (BOL); Cape Peninsula, Granger Kloof swamp below farm, 9 May 1897, Wolley 2496 (BM, BOL). Mauritius, Riviere Citron near Balaclava, 21 November 1961, Duljeet 10244 (BM). Hulstaert 1472 (BR!), listed under Hibiscus diversifolius var. angustilobus by Hauman (1963) is probably a hybrid involving H. diversifolius (H. cannabinus x diversifolius?), as Hauman's annota- tion of the specimen indicated. 20b. Hibiscus diversifolius subsp. rivularis (Bremek. & Oberm.) Exell, Fl. zambes. 1: 444 (1961). Hibiscus rivularis Bremek. & Oberm. in Ann. Transvaal Mus. 16(3): 424 (1935). Type: Botswana, Chobe R., Kabulabula, July 1930, van Son s.n., in Herb. Transv. Mus. 28936 (PRE-holotype; BM!- isotype). Stems more densely and uniformly hairy, flowers reddish to purple (Exell, 1961). Generally in habitats in or close to standing water, riverbanks, swamps, lake shores, forest edges. DISTRIBUTION. Fig. 4. Eastern, southeastern and southwestern Africa from Uganda to Botswana. Angola, Cuanza Sul, banks of River Longo, Quissama Country, 7 June 1921 , Gossweiler 8300 (BM, P) [Exell & Mendonc.a (1937) cite Gossweiler 8300 as subsp. diversifolius - note on BM specimen says 'flower a beautiful dusky purple']- Democratic Republic of Congo, Lac Mokoto,Terr. Masisi, 22 January 1959, Leonard 2651 (BR). Rwanda, Territ. Biumba, Reg. du Mutara, env. de Mimuli, 2 April 1957, Troupin 3079 (BR). F.D. WILSON Burundi, Kigamba (preT. Ruyigi), 3 October 1974, Auquier 4367 (BR). Uganda, Koki, 9 August 1903, Bagshawe 380 (BM); Igara, Ankole, March 1939, Purseglove 612 (BR, K-2 sheets). Tanzania, West Lake Province, Keza, Bushubi, Ngara, 30 May 1960, Tanner 4950 (K); Bezirk Lindi, Lutamba-See, 40 km W. Lindi, 9 November 1934, Schlieben 5287 (BM, G); Musoma Reg., 8 km N. of Nyakanga on Hwy B-6, 23 July 1975, Wilson & Kissie 7531 (ASU, EA, NA); Zanzibar, June 1894, Sacleux 2148 (P). Zam- bia, Western Prov., Fort Rosebery Dist., 20 August 1952, Angus 353 (BM, BR); near Nabwalya's, Luangwa Valley, 16 August 1966, Astle 4932 (K). Malawi, Benga by Lake Nyasa, 28 August 1950, Foster 1 1 (K); Kota-Kota Dist., Benga, W. shore of Lake Nyasa, 2 September 1946, Brass 17480 (BR); Chipoka, Fort Johnston, 7 August 1954, Banda 41 (BM). Mozambique, Gaza, Bilene, Praia de S. Martinho, 7 November 1969, Correia & Marques 1450 (K); Sul do Save, Louren?o Marques, Catembe, 10 June 1957, de Carvalho 211 (BM); Delagoa Bay, Rikatla, 1890, Junod 38 (G-2 sheets); Louren90 Marques, Marracuene, 30 June 1971, Marques 2313 (NBG); Kurumadgi R. Jihu, 12 August 1906, Swynnerton2\09 (BM). Botswana, N. District, Linyanti R. at Shalie, 28 October 1972, Gibbs Russell 2413 (PRE); banks of Chobe R., 8 miles N. of Kachikau, 9 July 1937, Erens 366 (BR). 21. Hibiscus reekmansii F.D. Wilson, sp. nov. Type: Burundi, Muramwya Prov., bois sacre de Mptosa jachere bord route, 17 May 1979, Reekmans 7993 (BR!-holotype). Fig. 2D. Hibiscus diversifolius var. angustilobus Hauman in Bull. Jard. Bot. Etat 31: 86 ( 1961 ), pro parte quoad specims. Laurent 1 1 , deWitte 2435. Hibiscus diversifolius var. angustilobus P. Maquet in G. Troupin (Ed.), Fl. rwanda 2: 382 (1983), non Hauman (1961), pro parte quoad specims. Auquier 3937, Troupin 16264. Caules aculeati; folia profunde 5-7 lobata; pedunculi fructificantes, <1 cm longi; bracteolae involucellorum non bifurcatae, apicibus auctis; calyces pilis argenteis vel cinereis vel stramineis, nectariis in quoque costa. Shrub to 2 m tall; stems with antrorse to retrorse aculeoli and with lines of stellate pubescence below, densely stellate-pubescent above. Stipules 2-6 mm long, filiform or linear, with fine, short hairs. Leaves: petioles 0.4-5 cm long, with pubescence like that of the stem, the blades 0.6-6 x 0.2-7 cm, the blades of lower leaves deeply 5(-7)-lobed, the midlobe linear to ovate or obovate, the lateral lobes linear to ovate, the outer lobes lanceolate or triangular, the midlobe and lateral lobes often secondarily lobed, the blades of upper leaves bract-like or 3-lobed, all blades abaxially aculeolate and with simple and stellate hairs on the ribs, and simple to 4-fid hairs on the veins, adaxially with dense simple to stellate hairs on the ribs and fine, short, mostly stellate hairs on the veins, the base cuneate, truncate, or cordate, the margin serrate to irregularly serrate, the apex acute or acuminate, the nectary 0.2-0.5 mm long. Flowers axillary, solitary, or with 2-3 nodes clustered near the apex, some flowers with a subtending bract resembling the stipules, linear, flattened, pink, sometimes with a bifid apex; peduncles 0.2-0.6 cm long, articulat- ing at the base, with simple, appressed grey-to straw-coloured, dense, fine hairs to 2 mm long; involucellar bracteoles 6-8, 6-10 mm long, linear, flattened, with short bristles or with hairs like those of the peduncles, the apex entire but slightly enlarged, the base free; calyx 1.0-1.5 cm long, the lobes triangular-acute to lanceolate- acuminate, with bristles to 3 mm long or with hairs as on the peduncles, the nectary small but conspicuous; corolla bright yellow to primrose yellow, with a small purple centre, the petals 3-4 x 2- 2.5 cm, obovate, finely stellate-pubescent dorsally, glabrous ventrally ; staminal column 16 mm long, yellow(?), the filaments 0.5 mm long, the anthers yellow-brown(?), the pollen yellow; style branches 5 mm long, yellow-brown(?). Capsule 12-15 x 10-13 mm, densely REVISION OF HIBISCUS SECTION FURCARIA 65 appressed-pubescent, the beak 1-2 mm long, glabrous; seeds 3x2 mm, subreniform, dark brown, with a faveolate background and minute protuberances, the funiculus small, dark-brown, glabrous. Mountainous habitats, 1 800-2400 m elevation, in savannas, rock talus, fields, and pastures. DISTRIBUTION. Fig. 5. Eastern Democratic Republic of Congo, Rwanda, Burundi, and western Tanzania. Democratic Republic of Congo, Kivu, environs de Kabare, 25 July 1942, Laurent 1 1 (BR); Pare Nat. de L'Upemba, Mukana, 24 March 1942, de Witte 2435 (BR). Rwanda, route Butare- Cyangugu, vers km 30, Pre"f. Gikongoro, 28 May 1981, Troupin 16264 (BR); Colline Gisego, village Rugera, au sud de lat Pref. Gikongoro, 6 September 1974, Auquier 3937 (BR). Burundi, Muramvya Prov., Teza, 19 June 1980, Reekmans 9395 (BR).Tanzania, Iringa Prov., Mbosi, May 1935, Horsbrugh- Porter s.n. (BM). 22. Hibiscus sabdariffa L., Sp. pi. 2: 695 (1753). Type: resolved but not yet published (P. A. Fryxell, pers. comm., 1996). Hibiscus cannabinus Hiern, Cat. afr. pi. 1: 72 (1896), non L. (1759), pro parte quoad specims. Welwitsch 5263 (BM ! , LISU !), Welwitsch 5265 (BM!, LISU!). Hibiscus masuianus de Wild. & T. Durand in Bull. Soc. Roy. Bot. Belgique 38(2): 20 (1899). Type: Democratic Republic of Congo (Zaire), Boma, 1 August 1895, Dewevre 71 (BR!-holotype & isotype). Hibiscus sabdariffa var. albusA. & G. Howard in Mem. Dept.Agric. India, Bot. Ser. 4:32(1911). Hibiscus sabdariffa var. intermedius A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 32 (1911). Hibiscus sabdariffa var. ruberA. & G. Howard in Mem. Dept.Agric. India, Bot. Ser. 4: 32 (1911). Hibiscus sabdariffa var. bhaghalpuriensis A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 33 (1911). Hibiscus sabdariffa var. altissima Wester in Philipp. Agric. Rev. 7: 266(1914). For additional synonymy, see Hochreutiner, 1900: 116, 117;Hauman, 1963: 114; Borssum Waalkes, 1966: 64; Fryxell, 1988: 225. Illustrations: Andrews, Fl. pi. anglo-egypt. sud. 2: 24, fig. 12 (1952); Hochreutiner, Fl. Madagasc., Malvac.: 39, fig. XI, 3 (1955); Ochse, Veg. Dutch E. Ind.: 476, fig. 296 (1960). Annual herb, branched or unbranched, up to 4.5 m tall; stems and foliage green, green-red, or red, glabrous to sparsely pubescent, sometimes sparsely aculeate. Stipules 5-13 mm long, linear to narrowly subulate. Leaves: petioles 0.3-12 cm long, pubescence like that of the stems, the blades 2-15 x 2-15 cm, blades on lower leaves almost entire to shallowly or deeply 3-5(-7)-lobed, on upper leaves almost entire to shallowly to deeply 3-5-lobed, the middle lobe longest, at the plant apex narrowly lanceolate to bract-like, all blades almost glabrous or abaxially and adaxially with mostly simple hairs, and abaxially with a few aculei on the ribs, the base cuneate to truncate, the margin finely to coarsely serrate, the apex obtuse to acute or acuminate, the nectary 1-3 mm long, slit-like. Flowers axillary, solitary; peduncles 0.6-2.0 cm long, articulating near the base to halfway between the base and the epicalyx, glabrous to finely pubescent below the articulation, and almost glabrous to bristly or sometimes aculeolate above the articulation; involucellar bracteoles 8-12, united at the base, the free part 6-18 mm long, the upper 3-6 mm channelled on the ventral surface, flattened, subulate to triangu- lar, short-aculeolate, the apex entire, pointed; calyx l.l-5(-5.5) cm long, green, green-red, or red, fleshy and edible, or leathery and inedible, the lobes triangular to ovate, acute to acuminate, glabrous, sparsely pubescent or bristly, the nectary inconspicuous; corolla pale yellow, with or without a dark red-purple centre, the petals 2-5 x 1.2-3.5 cm, obovate, glabrous to simple or stellate-pubescent dorsally, glabrous ventrally; staminal column 10-20 mm long, pink, filaments 1 mm long, pollen tan; style branches included in the staminal column; stigmas dark red-purple. Capsules 13-22 x 1 1-20 mm, ovoid, nearly glabrous to sparsely or densely appressed-pubes- cent; seeds 3-5 x 2-4 mm, subreniform, dark brown, faveolate, with or without minute pectinate or stellate scales, the funiculus small, dark brown, glabrous. Chromosome number, n = 36. Widely cultivated in tropical and subtropical areas of the world, including northern Africa; sometimes escapes from cultivation (e.g. Karnataka, India, Saldanha, 1985). The most common form of Hibiscus sabdariffa in Africa is a culinary form with fleshy edible calyces (segregated as//, sabdariffa var. sabdariffa by Rakshit & Kundu, 1970: 163). The calyces have various uses, including drinks and jams. The leaves also are eaten sometimes as a vegetable (Exell, 1961; Ochse, 1960). On a collec- tion from Darfur Province, Sudan, Lynes 538 (BM), it was noted that the plant was used to preserve and water-proof skins, but it did not specify which part of the plant was used, or how it was used. Several cultivars have been released (e.g. Abdallah et al., 1976). In fact, the varieties of Howard & Howard (1911) and of Wester (1914) repre- sent horticultural cultivars rather than botanical varieties. The culinary form is found mostly in cultivation but sometimes becomes natural- ized. A short description follows: Branched, branches long, plant up to 1 .5 m tall; stems almost glabrous; involucellar bracteoles up to 1 8 mm long; calyx 3(-5) cm long in fruit, fleshy and edible, glabrous or sparsely pubescent; petals up to 5 x 3 cm. DISTRIBUTION (in subsaharan Africa and Asia). Senegal, Pare Nat. de lat Basse Casamance, 26 November 1984, Bamps 7785 (BR). Mali, Madina Diassa, 1 9 November 1974, Audru 5928 (P). Upper Volta, W. Dinderesso, 14 November 1980, Lejoly 80/129 (BR). Sierra Leone, Njala, 19 November 1930, Deighton 1849 (K). Guinea-Bissau, Bedanda-Cadique, 5 January 1962, Pereira 2613 (G-2 sheets). Togo, Lomi (Nord), 5 November 1985, Schiller 112 (G). Nigeria, North-east State, Gombe Dist., near Gombe Railway Station, 19 October 1971, Latilo FHI 63844 (K). Central African Republic, 6 km SE of Bambari on the Alindao road, 1 4 November 1 98 1 , Fay 1955 (K). Cameroon, Mogode, Gomtemale Mt., 9 October 1972, Leeuwenberg & v. Beek 10474 (BR). Sudan, Tombouctou, 18 July 1899, Chevalier 1250 (BR). Democratic Republic of Congo, Leopoldville, 17 April 1915, Bequaert 7410 (BR). Uganda, without specific locality, June 1917, Dummer 3194 (BM). Burundi, Bujumbura Prov., Katumba Com- mune, 15 April 1977, Reekmans 6021 (BR). Tanzania, West Lake Prov., Ruganzo, Bugufi, Ngara, 10 July 1960, Tanner 5033 (K). Angola, Lunda, 1 1 May 1903, Gossweiler 1477 (BM). Zambia, Balovale Dist., July 1933, Trapnell 1262 (K). Malawi, Rifu Village, 13 July 1936, Burn 6088 (BM). Mozambique, Montepuez, Aldeamento de Nairo, 27 August 1972, Mafumo 19 (BR). Madagascar, Vallee de la Betsiboka, 5-6 July 1928, Humbert & Swingle 4412 (B). Namibia, Okahamoja, September 1934, Vedderl%26 (B); Righini Terr., Lovanium, 7 April 1961, Evrard 6290 (BR). India, Tilaikani, Gangpur State, Orissa, 14 November 1947, Mooney 2970 (K); Gdukki Dist., 15 December 1982, Mohanan (Bot. Surv. Ind. 76099) (CAL). Nepal, East Nepal, Nissim-Dunham, 30 October 1 963, Hara etal. (Univ. Tokyo 6300743) (BM). Burma, Taungdut, Chindwin River, 28 March 1935, Kingdon-Ward 1 1277 (BM). Thailand, Bangkok, 16 November 1924, Marcan 1899 (BM). The Philippines, Manila, 17 December 1903, Merrill 3801 (BM). A short description follows of the fibre type of Hibiscus sabdariffa cv. 'altissima' (segregated as H. sabdariffa var. altissima Wester by Rakshit & Kundu, 1970): Unbranched or branches short, plant up to 4.5 m tall; stems aculeolate or pubescent; involucellar bracteoles up to 7 mm long; calyx up to 2.2 cm long in fruit, leathery, inedible, bristly; petals up to 3 x 2 cm. This form is not found commonly in Africa. Morton & Jarr 3250 from Sierra Leone is a tall, erect and unbranched cultivated plant, but 66 F.D. WILSON it is not clear from the label whether it is cultivated for fibre (could be cultivated for leaves or seed). According to Dempsey (1975: 306) the fibre type is grown as a fibre crop on a large scale in the Central African Republic and in seven countries in south-central and south- eastern Asia. DISTRIBUTION. Sierra Leone, Kain Kordu, near Lagbwema, Lono Dist., 1 2 December 1 965, Morton & Jarr SL 3250 (K). India, Carnatic, 1 0 km from Ulundurpet on the Salem road, 1 3 December 1 978, Matthew & Perumal RHT 20145 (CAL). Indonesia (Java), Zollinger 2998 (BM). Forms occur in Africa that are neither the edible, culinary form nor the fibre form; they are usually branched and have bristly and/or aculeate plant parts. Some may be cultivated for seeds (Wilson & Menzel, 1964) but some appear to be truly wild or at least ruderal. For example, the seeds which gave rise to Menzel & Wilson HV220 (FSU) were collected originally in the wild near Maradi, Niger by D.W. Fishier. This plant is apparently truly wild because it is branched and woody, has aculeate stems, bristly calyces, and small seeds. DISTRIBUTION. Niger, provenance Maradi, grown in cultivation in U.S.A., Florida, Palm Beach Co., Everglades Exp. Stn., 12 December 1964, Menzel & Wilson HV220 (FSU). Ghana, mile 73, Arebubi-Yeji road, grass savannah, 13 December 1955, Afruah (WACRI Herb. 5097) (K); Danongo, Gonja Dev. Corp. farm, ruderal, 30 September 1956, Innes GC 30194 (K); N. of Pong- Tamale, in cultivated land, 12 December 1953, Morton GC 9870 (K). Nigeria, Plateau Prov., Jemaa Emirate, about 8 miles E. of Jagindi, Madaki Dist., open savannah woodland just W. of Kurmin Damiss, 19 November 1946, Keay & Onochie (FHI 212724) (K); Angola, GolungoAlto, May 1855, Welwitsch 5263 (BM); Quicuxe(?), July 1854, Welwitsch 5265 (BM). 23. Hibiscus cannabinus L., Syst. nat. 10th ed.: 1 149 (1759). Type: resolved but not yet published (P. A. Fryxell, pers. comm., 1996). Hibiscus sabdariffa var. 5 L., Sp. pi. 2: 695 (1753). ?Hibiscus verrucosus Guill. & Perr., Fl. Seneg. tent. 1: 57 (1831). ^Hibiscus cannabinus var. verrucosus (Guill. & Perr.) Garcke in Linnaea 43: 56 (1880). Hibiscus cannabinus var. genuinus Hochr. in Annuaire Conserv. lard. Bot. Geneve 4: 1 15 (1900), nom illeg. (Art. 26.2). Hibiscus cannabinus var. simplex A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 16 (191 1). Hibiscus cannabinus var. purpureus A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 17 (1911). Hibiscus cannabinus var. ruber A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 17 (1911). Hibiscus cannabinus var. viridis A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 17 (1911). Hibiscus cannabinus var. vulgaris A. & G. Howard in Mem. Dept. Agric. India, Bot. Ser. 4: 17 (191 1). Hibiscus sabdariffa subsp. cannabinus (L.) G. Panigrahi & S.K. Murti, Fl. Bilaspur dist. 1: 127 (1989). For additional synonymy, see Exell, 1961. Hochreutiner (1900: 114-115) lists an extensive synonymy for Hibiscus cannabinus, most of which I have not been able to check. Hochreutiner and other authors generally have treated H. verrucosus as a synonym of H. cannabinus, but it may well be synonymous with H. asper (in which case, the correct name for this entity is H. verrucosus). In the absence of an authentic type specimen of H. verrucosus, this ques- tion must go unanswered. Some of Hochreutiner's synonyms obviously refer to other species, as follows: H. radiatus Cav., //. unidens Lindl. (= H. radiatus Cav.), H. lindleyi Wall. (=H. radiatus Cav.), H. asper Hook, f., H.. mechowii Garcke, H. acetosella Welw. ex Hiern. Illustrations: Icones Roxburghianae no. 355 (K);Verdoorn & Collett, Farming South Africa 34: 1, fig. 1 (1947); Andrews, Fl. pi. anglo- egypt. sud. 2: 24, fig. 11 (1952); Hochreutiner, Fl. Madagasc., Malvac.: 37, fig. X, 4, 5 (1955); Gibson, Wildfl. natal: pi. 65, fig. 3 (1975). Annual herb up to 2 m tall (up to 5 m tall in cultivars), stems pale green to pink or red-purple, sparsely aculeate, otherwise nearly glabrous. Stipules 4-7 mm long, filiform. Leaves: petioles 0.4-30 cm long, aculeate like the stems, the blades 1-19 x 0.1-20 cm, very shallowly to very deeply palmately 3-7-lobed below, sometimes unlobed above, or even bract-like near the apex, nearly glabrous, sparsely aculeolate on the ribs, the base cuneate to shallowly cor- date, the margin serrate or dentate, the apex acute, the nectary 3 mm long, prominent. Flowers axillary, solitary, or sometimes clustered near the apex; peduncles 0.3-0.6 cm long, articulating at the base, aculeolate or bristly; involucellar bracteoles 7-8, 7-18 mm long, linear to subulate, aculeolate or bristly, the apex entire, the base free; calyx 1.1-2.5 cm long (up to 3.4 cm in cultivars), the lobes acumi- nate to subcaudate, aculeate, or sparsely to densely bristly, with a characteristic white, woolly tomentum, especially near the base and margins, the nectary conspicuous; corolla off-white, cream, or various shades of purple, from almost grey to deep purple, and with a deep purple centre, the petals 4-6 x 3-5 cm, obovate, sparsely stellate-pubescent dorsally, glabrous ventrally ; staminal column 1 7- 23 mm long, dark red, the filaments 1-2 mm long, the anthers tan, the pollen yellow; style branches 2-4 mm long, dark red; stigmas dark red. Capsule 12-20 x 11-15 mm, ovoid-acuminate, densely- appressed pubescent, the beak 1 mm long, glabrous; seeds 3-^ x 2-3 mm, subreniform, dark brown, faveolate, and sometimes with minute pectinate scales, the funiculus small, tan, glabrous. Chromosome number, n = 18. Cultivated, ruderal, and wild, in cultivated land, old gardens, dikes between irrigated fields, ridge tops in shallow soil, rocky fissures, talus, open grassland plains, savannas, flood plains, sea- sonal swamps. DISTRIBUTION. Fig. 4. Widely distributed as a cultivated fibre, food and medicinal plant and as a ruderal and native plant in subsarahan Africa; grown by Makrakra Tribe in Sudan for game nets, Cartwright 2 (K); seeds eaten by natives in Sudan, Imperial Institute s.n. (K); used medicinally in Nigeria for stomach complaint, Wickens 3546 (K). It is cultivated as a fibre crop and sometimes naturalized in India (Saldhana, 1 985), and cultivated as a fibre and paper-pulp plant in a number of other tropical and subtropical countries (Dempsey, 1975; Taylor, 1992). The following listing includes only collec- tions from native habitats. Senegal, 1952, Berhaut 263 (P). Mali, near Kidal, 25 October 1 968, Popov 1 1 (BM). Ghana, along Comtomenlo-Achimota Rd, Accra E. P., 18 August 1955, Lovi (WACRI Herb. 4200) (K). Nigeria, NE State, Kirinava, 9 December 1975, Wickens 3546 (K). Cameroon, Waza, 21 November 1 969, Hepper 3957 (K); presYafounou (50 km ENE de Meiganga), 17 October 1963, Letouzey 6200 (P); Mogode", Gomtemale Mt., 9 October 1972, Leeuwenberg 10473 (BR). Chad, c. 8 km S. of Ft. Lamy, along road to Bongor, 3 January 1965, deWilde & deWilde-Duyfies 5175 (BR). Central African Republic, Manova-Gounda, St. Floris Nat. Park, 2 October 1982, Fay 3683 (K). Sudan, Equatorial Prov., 6 miles S. of Gogrial, without date, Myers 7692 (K); Yei River District, without date, Cartwright 2 (K); Equato- rial Prov., Zande Land, 1 2 September 1 940, Wyld 84 1 (BM). Ethiopia, 70 km SW of Jimma on Bonga Road, 25 January 1969, Jones 6913 (K); Kaffa Province, Omech Village, 2 January 1962, Meyer 7887 (K); road from Soddo to Arba Mintch, 55 km from Sotto, 2 December 1967, Westphal & Westphal- Stevels 2927 (BR); Democratic Republic of Congo, Gimbi, July 1949, Brynaert 134 (BR); plaine du lac Edoard et de la riviere Rutshuru, 28 July 1937, Louis 4788 (BR-2 sheets); Ruzizi, 22 January 1950, Germain 5770 (G); Kivu Dist., Beni Prov., Kangali, 8 December 1954, deWitte 1 1387 (K). Uganda, elephant grassland, W. Kipayo, September-October 1914, Dummer 1055 (BM); U4, 1 mile E. of Kakoge, Mengo, 14 December 1955, Langdale- Brown 1711 (K); Ul, Mt. Debasien, 1936, Eggeling E2820 (K). Kenya, 70 REVISION OF HIBISCUS SECTION FURCARIA 67 miles from Isiolo, 23 May 1945, Adamson 79 (G); Kajiado Dist., Athi Plains 30 miles from Nairobi on the Namanga Road, 1 June 1963, Verdcourt 3647 (BR); K7, Lamu Dist., Kanwe-Mayi pool between Witu and Kipini, 4 March 1977, Hooper & Townsend 1182 (K); Kl, Northern Frontier Prov., S. Turkana, c. 12 m NNE of Kangetet, 25 June 1970, Matthew 6383 (K); Kwale Dist., 38 kmW. of Mombasa, 1 3 August 1975, Wilson 75 141 (ASU,EA,NA). Somalia, Webi Schabeli, 1891, Keller 53 (K); Sheik husin, 25 September 1 894, Donaldson-Smith 200 (BM). Rwanda, Territ. Buinda, Reg. du Mutaro, environ de Karukwangi, 15 April 1958, Troupin 7097 (BR); Rwabiega, Pare Nat. de la Kagera, January 1938, Lebrun 9806 (B). Burundi, Bujumbura, 12 May 1971,L€H'a//e5719(BR);routedeBugarama, km 10,territ. Bujumbura, 24 May 1966, Lewalle 844 (K). Tanzania, Bahi, on the Dodoma to Manyoni Rd, mile 38, 23 April 1962, Polhill & Paulo 2129 (BR); Shinyanga Reg., 5 km S. of Shinyanga on Hwy B-6, 28 July 1975, Wilson & Kissie 7579 (NA); Tabora Region, 1 2 km W. of Tabora on road to Urambo, 29 July 1975, Wilson & Kissie 7580 (EA, NA); T7, Njombe Dist., 19 m E. of Rujewa on road from Mbeya to Iringa, 6 April 1969, Jones 69 1 30 (K); T6, Masada, 28 June 1973, Greenway & Kanurl 15280 (K); Kondoa Dist., Mapapu, Mbuga, 15 May 1929, Bum 2067 (BM);TangaProv., Handeni Dist., HandeniTown, 5 August 1989, Mhoro 6301 (TEX); Cheju Plain, Zanzibar, 6 August 1933, Vaughan 2158 (BM). Angola, Huila, Cassinga, Chamutete, 2 April 1960, Barbosa & Correia 8965 (BM); Mossamedes Dist., 1 August 1859, Welwitsch 4931 (BM); Huila, Quipungo, Handa, 1 May 1967, Henriques 1079 (BM); Pungo Adongo, April 1857, Welwitsch 5274 (BM); Huila, Chicungo, 9 May 1962, Texeira &Almeida 5625 (BR). Zambia, Mkushi Dist., Tiwila Congo Zambesi Basin, 1932, Hewitt 37 (BM); Marumba (Livingstone), June 1909, Rogers 7142 (BOL). Zimbabwe, Umtali, 22 February 1960, Head22 (BM); Nartley Dist., Poole Farm, 15 April 1950, Nornby 3171 (BM); Manica Dist., Umtali Div., Odranis River Valley, 1914, league 130 (BOL). Malawi, Mafina Hills, Chisenga, 24 August 1962, Tyrer 535 (BM); S. Region, Thyolo Dist., Boumbure, 7 May 1983, la Croix 2272 (BM). Mozambique, Lourenso Marques Dist., Sul do Save, Costa do Sul, 3 March 1949, Barbosa 2637 (BM); Sul do Save, Guija, 4 May 1957, de Carvalho 151 (BM); Sul do Save, estrada Chissano-Chibuto, 8 July 1948, Myre 46 (BM); Cabo Delgado, Macondes, lOApril 1964, Torre & Paiva 1 1849 (K); Maputo, Marracuene, ao km 34 da Estrada Nacional no. 1 , 25 May 1 977, Nuvunga, Zunguze & Doane 1 (NBG). Namibia, Grootfontein Dist., Farm Bergen, 25 April 1963, Giess, Volk & Bleissner 6453 (PRE); Tsumeb, 22 April 1934, Dinter 7549 (B-2 sheets). Botswana, floodplain grassland of the Sibuyu River, 12 April 1983, Smith 4250 (PRE); Savuti, Chobe Nat. Park, 25 March 1948, Jacobsen 3053 (PRE); Northern Dist., Thamalakane River, Maun, 24 March 1976, Smith 1 666 (B). South Africa, Transvaal, Grobiersdal, Mossiesdal, 11 March 1958, Meeuse 10321 (PRE); Transvaal, Waterburg, 6 December 1931, Galpin 11569 (BOL); Transvaal, Shilouvane, 1904-1905, Junod 2265 (G); Natal, Uvongo Beach, April 1968, Liebenberg 8099 (PRE); Natal, sandy flat near Durban, 27 April 1893, Wood 4848 (BM); Natal, Port Shepstone, 3 April 1966, Strez 6540 (BR, G). Swaziland, Lubombo Dist., 7 km W. of Siteki, 31 March 1977, Kemp 825 (PRE). I also examined cultivated specimens from Afghanistan, Yemen, Eritrea, Saudi-Arabia, Mauritania, India, Vietnam, Pakistan, Egypt, Thailand, and Nepal. A number of variants in the Hibiscus cannabinus-asper group occur in nature, some of them representing hybrids or introgressive types. Others possibly represent undescribed taxa, but material is insuffi- cient to describe them adequately, as follows: a form with very long bristly bracteoles (longer than the calyx) and bristly calyces: Ethio- pia, Schimper 1484 (BM, K); Tanzania(?), Schweinfurth 1626 (B, BM, BR, G), Uganda, Wilson s.n. (K); a form with dense, stiff bristles mainly on the calyx midrib and marginal ribs: Zambia, Fanshawe 8786 (BR). The varieties of Howard and Howard represent horticultural cultivars rather than botanical varieties. The key characters used to differentiate their varieties are generally simply inherited and are found in various combinations in cultivars, land races, and in wild forms of H. cannabinus. Hauman (1963) recognized var. simplex, but it is merely an entire-leaved form of this polymorphic species (deWitte 10527, 10617 (BR!)). 24. Hibiscus parvilobus F.D. Wilson, sp. nov. Type: Kenya, Rift Valley Prov., Nakuru Dist., 35°39'E, 0°9'S, c. 9 km NE of Londiani, 2743 m, 8 February 1973, Spjut & Ensor 3184 (EA!- holotype; BR!, K!-isotypes; sheets are also lodged at ASU which were prepared from plants grown at Phoenix from seeds of the original collection). Fig. 2A. Hibiscus berberidifolius F.D. Wilson in Brittonia 35: 178 (1983), non A. Rich. ( 1 847), pro parte quoad specim. Spjut & Ensor 3 1 84. Caules aculeati; folia 5-lobata, apicibus et sinibus obtusissimis; pedunculi fructificantes <1 cm longi; bracteolae involucellorum non bifurcatae, lineares, subulatae vel triangulares, apicibus auctis; caly- ces in costas pilis densis, glabrati inter costas, nectariis in quoque costa prominentibus. Many-branched perennial shrub, 1-4 m tall; stems with antrorse aculei and with fine, simple hairs up to 1 .5 mm long, aculei and hairs sparse below, more dense above, and with a line of crisped pubes- cence changing position at each node. Stipules 6 mm long, linear, with sparse, fine, simple hairs. Leaves: petioles 1.6-6.5 cm long, with dense, woolly pubescence adaxially, sparse simple hairs abaxially, the blades 3.4-5.5 x 1-9 cm, the blades of all leaves on vegetative branches moderately to deeply 5-lobed, not differing much in size from the base to the apex of the plant, abaxially and adaxially with simple antrorse hairs to 1 mm long on midrib and principal veins, less dense or lacking on smaller veins and absent from the lamina, the base cordate, the margin finely serrate, the apex and sinus obtuse; the blades of mid-plant leaves on flowering branches 3-lobed, otherwise similar to those on vegetative branches, blades of upper leaves on flowering branches obscurely 3-lobed to entire, the base truncate, the margin finely serrate, the apex acute; nectary present on all leaves, 2 mm long; vegetative branchlets in the leaf axils to 2.5 cm long, blades up to 3 x 3 cm, ovate, the base truncate to very slightly cordate, the margin finely serrate, the apex obtuse. Flowers borne singly in the leaf axils of flowering branches; peduncles 0.3-0.5 cm long, articulating at the base or joint not evident, with dense, simple bristly hairs 2-3 mm long, borne at right angles to the peduncle; involucellar bracteoles 6-8, 5-1 1 mm long x 1-3 mm wide at the base, linear to subulate or triangular, flattened, several nerved, below with pubescence like that of the peduncle, above almost glabrous, apex entire; calyx 1 .2-1 .4 cm x 0.6 cm wide at the base in flower, 1.5-1.8 cm x 0.7-0.8 cm wide at the base in fruit, the lobes slightly acuminate, with pubescence like that of the peduncle on the ribs below, sometimes sparsely aculeolate on the ribs above, almost glabrous between the ribs, the nectary prominent, enlarged, 2 mm long; corolla light yellow with a prominent deep red- purple centre, the petals 3.5 x 3.5 cm, obovate, finely and densely pubescent dorsally, glabrous or pubescent ventrally; staminal col- umn on only flower seen 23 mm long, deep red-purple; filaments 1 mm long, anthers and pollen tan; style branches 8 mm long, deep red-purple; stigmas deep purple. Capsule 16-17 x 15-16 mm, ovoid, densely appressed-pubescent, the beak 3 mm long, glabrous; seeds 4x3 mm, subreniform to triangular, dark brown, with minute pectinate scales, funiculus very dark brown, with a fringe of colour- less hairs on the large end. Collected in a more or less dry, scrubby forest with Tarchonanthus camporatus L., Trichocladus ellipticus Eckl. & Zeyh., Euclea divinorum Hiern, Rhus natalensis Bernh. ex Krauss, R. vulgaris Meikle, Maytenus heterophylla (Eckl. & Zeyh.) N. Robson, and Scutia myrtina Kurz. DISTRIBUTION. Known to me only from the type collection. 68 F.D. WILSON 25. Hibiscus asper Hook, f., Niger fl.: 228 (1849). Type: Sierra Leone, Miss Turner s.n. (K-holotype). Fig. 2C. Hibiscus verrucosus var. punctatus A. Rich., Tent. fl. abyss. 1: 59 (1847). Type: Ethiopia, crescit in provincia Chire', Petit s.n. Hibiscus cordofanus Turcz. in Bull. Soc. Nat. Mosc. 31: 193 (1858). Type: Ethiopia, Cordofan, 1 837- 1838, Kotschy 65 (KW-holotype). Hibiscus cannabinus Mast, in Oliver, Fl. trap. Afr. 1: 204 (1868), non L. (1759), pro parte quoad syn. H. asper. Hibiscus cannabinus Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 15 (1900), non L. (1759), pro parte quoad syn. H. asper. Hibiscus cannabinus var. chevalieri Hochr. in Annuaire Conserv. Jard. Bot. Geneve 5: 125 (1901). Type: Hochreutiner (1901) did not designate a holotype for H. cannabinus var. chevalieri but lists two syntype specimens (neither of which I have seen), as follows: Mali, Koulikoro, moyen Niger, 6-14 October 1899, Chevalier s.n.; Mali, Sindou, terres cultive"es parmi les planta- tions de cotonniers, 10 May 1899, Chevalier 856. Chevalier (1920) cites these two and a later collection from Guinea, Cheva- lier 14909 (G!, P!) under H. cannabinus var. chevalieri. Hochreutiner annotated a sheet at G as a '2nd form of Hibiscus cannabinus var. chevalieri, leg. Chevalier, Sindou et Niger moyen ex Herb. Chevalier' which is referable to Hibiscus scotellii Baker f. Hibiscus vanderystii de Wild, in Bull. Jard. Bot. Etat. 5: 35 (1915). Type: Angola, Kwango, July 1913, Vanderyst 1377 (BR!). Hibiscus cannabinus var. punctatus (A. Rich.) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 20: 82 (1916). Hibiscus malangensis Baker f. in J. Bot. 77: 22 (1939). Type: Angola, Malange, River Cuango, near Xa-Sengue, c. 1075 m, 5 April 1937, Exell & Mendon§a 274 (BM!). Hibiscus asper var. punctatus (A. Rich.) Berhaut, Fl. Senegal: 160 (1954). Herb to 2 m tall; stems with antrorse aculeoli (aculeoli sometimes 2- fid) and with fine, simple to stellate hairs, and sometimes with a line of pubescence changing at each node. Stipules 3-6 mm long, filiform, with short, fine pubescence. Leaves: petioles 0.3-18 cm long, with pubescence like that of the stems or sometimes less or more pubescent, the blades extremely variable in size and shape, the blades of lower leaves 4-18 x 1.5-14 cm, strap-like, lanceolate, ovate, oval, or shallowly to deeply palmately 3-5(-7)-lobed (some blades even 2- or 4-lobed), the lobes sometimes secondarily lobed, the blades of upper leaves 2-7 x 0.2-6.5 cm, bract-like, lanceolate, oblanceolate, ovate, or 3-lobed, the blades of all leaves with 2-fid to stellate hairs on ribs and veins, more sparse on the adaxial surface, the base cuneate, truncate, or cordate, the margin regularly or irregularly finely to coarsely serrate or repand, the apex acute, the nectary 0.5-2 mm long, slit-like. Flowers axillary, solitary, or often with more than one flower at the same node; peduncles 0.2-0.8 cm long, articulating at or near the base, with or without antrorse aculeoli and straight bristles and with finer, simple to stellate hairs; involucellar bracteoles 6-7, 9-18 mm long, subulate to narrowly lanceolate-acuminate, with or without antrorse aculeoli or short, coarse pigmented bristles on enlarged bases plus some finer simple to stellate hairs, the apex entire, pointed, the base free or scarcely united; calyx 0.8-2.6 cm long, the lobes lanceolate to broadly lanceolate, short- to long-acuminate, with antrorse aculei or pigmented short, coarse bristles on enlarged red bases on the ribs, otherwise almost glabrous or with some fine hairs, the nectary small but conspicuous; corolla light yellow with a red-purple centre, the petals 2.8-4.5 x 2-3 cm, obovate, glabrous to sparsely stellate- pubescent dorsally, glabrous ventrally; staminal column 10-12 mm long, dark pink, the filaments 0.5-1 mm long, the anthers tan-orange or red-purple, the pollen yellow; style branches included in the staminal column; stigmas red-purple. Capsule 12-19 x 12-16 mm, sparsely and finely short appressed-pubescent, the beak 1-2 mm long, pubescent, inconspicuous; seeds 3-4 x 2-3 mm, subreniform, dark brown, with parallel striations and minute protuberances, the funiculus with simple, coarse flattened hairs. Chromosome number, n=18. Ruderal and wild, in abandoned farm fields, savannas, grassland, river alluvium, moist rock basins, and edges of gallery forest. DISTRIBUTION. Fig. 4. Widely distributed in subsaharan Africa, from Sen- egal to Ethiopia, Angola, Zambia, Mozambique, and Madagascar (Sprague, 1913). Senegal, Pare Nat. de la Basse Casamance, 26 November 1984, Bamps 7773 (BR). Guinea-Bissau, Bafata, entre Cambasse et Geba, 19 October 1952, Esperito-Santo 3102 (BR). Guinea, Kouria et environs, 20 October 1905, Chevalier 14909 (G, P). Upper Volta, Duarkoye Station, 30 October 1973, Audru 5443 (P). Sierra Leone, Waterloo, 1 December 1936, Brighton 3346 (K). Ivory Coast, Bouake, 6 February 1981, Ake Assi 15791 (BR); reserve de LAMTO, 11 November 1986, Gautier-Beguin 438 (G). Ghana, Northern, Mole Nat. Park, 25 km N. of Damango, 27 November 1995, Schmidt, Amponsah & Welsing 1790 (TEX); Dukwesein, N. of Agogo, Ashanti, 23 December 1913, Clupp 599 (K); Weijaa, near Accra, October 1961, Irvine 4807 (K). Mali, Domaine soudanien vallee Si Kasso-Koutiala, I January l95\,Roberty 1 3324 (G). Niger, Pare Nat. duW,du Niger, presde la Mekrou, 2 December 1986, Robbrecht & Leman 3460 (BR). Nigeria, Zunjeru, 10 October 1905, Dalziel 128 (K); Sokoto, October 1910, Dalziel 426 (K-2 sheets); nearToupe' on grassy plain along Comoe R., 28 November 1967, Geerling & Bokdam 1521 (K). Chad, Sarh Station, 7 November 1974, Palayer 420 (P). Cameroon, Waza, 2 1 November 1 969, Hepper 3956 (B, K). Central African Republic, Bambari Reg., 6 km SE of Bambari on the Alindao Road, 29 November 1984, Fay 1947 (K). Sudan, Sugura, Gedaref Dist., 28 September 1951, Babiku Beshir 145 (K). Ethiopia, in montibus prope Dscheladscheranne, 3 November 1839, Schimper 717 (G). Congo, bords du Congo, 14 April 1966, Farron 5100 (P). Democratic Republic of Congo, Orientale, Pare Nat. de la Garamba, 5 March 1951, De Saeger 1038 (BR, K). Uganda, Namaraprure(?), June 1915, Dummer 2565 (BM). Tanza- nia, Mbeya Dist., base of Punguluma Hills, Parinari woodland, 21 April 1 990, Lovett & Kayombo 4562 (TEX); Iringa Region, Mufindi Dist., Ngwazi, 15 April 1989, Kayombo 441 (TEX); 50 km W. of Lindi, 24 April 1935, Schlieben 6486 (BM). Angola, Dist. Ambaca, inter Mandiocae plantationes prope Ngombe, October 1856, Welwitsch 5267 (BM, LISU); Dist. Cuanza Norte, Cassualala, Cuanza, 10 July 1923, Gossweiler 8927 (BM). Zambia, Kapiri Mposhi, 13 July 1932, Young 38 (BM). Malawi, Central Prov., Dowa Dist., 31 July 195 1 , Chase 391 1 (BM); without precise locality, 24 May 1937, Lawrence 411 (BR). Mozambique, Nemagoa, Dist. Mocuba, November 1947, Faulkner 159 (K); Montepuez, proximidades do aldeamento de Nairo, I 1 August 1972, Mafumo 12 (NBG). Madagascar, Beravi Interior, Gebinge, July 1879, Hildebrandt 3086 (BM). Berhaut (1954) lists four collections of his H. asper var. punctatus (none of which are involved in the typification of//, asper): Senegal, Berhaut 1800, 4175, 4453, 4551; I have seen Berhaut 4175 (P), which I regard as an H. asper x cannabinus hybrid, and Berhaut 455 1 (P), which perhaps also is a hybrid derivative but closer to the true //. asper. 26. Hibiscus acetosella Welw. [ex Ficalho in Bol. Soc. Geogr. Lisbon, se"r. 2: 608 (1881), nom. nud.] ex Hiern, Cat. Afric. PI. 1: 73 (1896). Type: Angola, Cuanza Norte, Golungo Alto, Decem- ber 1854, Welwitsch521l (BM!-holotype; LISUI-isotype). Exell & Mendonga (1937) listed the specimen at LISU as 'tipo' (pre- sumably the holotype), but it is more appropriate to list the BM specimen as holotype because the best set of Welwitsch's speci- mens were retained at BM (Hiern, 1896). Fig. 2B. REVISION OF HIBISCUS SECTION FURCARIA 69 Hibiscus eetveldeanus de Wild. & T. Durand in Bull. Soc. Bot. Belgique 38(2): 24 (1899). Type: Democratic Republic of Congo (Zaire): Monbanga, Dewevre s.n. (BR-holotype). Hibiscus surattensis var. furcatus (Willd.) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4:112(1 900), pro parte quoad syn. H. eetveldeanus. Hibiscus cannabinus Hochr. inAnnuaire Conserv. Jard. Bot. Geneve 4: 1 14 (1900), non L. (1759), pro parte quoad syn. H. acetosella. Hibiscus surattensis var. eetveldeanus (de Wild. & T. Durand) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 6: 49-51 (1902). Illustration: Ochse, Veg. Dutch E. Ind.: 474, fig. 295 (1960). Annual or perennial subshrub to 2 m tall; stems and leaves green or red; stems glabrous to sparsely pubescent, not aculeate. Stipules 12- 15 mm long, linear to very narrowly lanceolate, with a few simple hairs on the margins. Leaves: petioles 3-1 1 cm long, glabrous or finely pubescent, the blades 2-10 x 2-10 cm, shallowly to moder- ately 3-5-lobed below, sometimes entire above, glabrous or sparsely pubescent, the base cuneate, truncate, or subcordate, the margin crenate, the apex obtuse, the nectary 2 mm long, slit-like. Flowers axillary, solitary; peduncles 0.4-1.0 cm long, articulating near the base, glabrous to hirsute; involucellar bracteoles 8-10, 7-12 mm long below the apex, linear, with simple hairs on margins, the apex bifurcate, the outer fork 3 mm long, lanceolate, ascending or recurved, the inner fork 3 mm long, linear, ascending; calyx 1.5-2.5 mm long, the lobes ovate to ovate-lanceolate, acute to acuminate, nearly glabrous or sometimes with long simple hairs on the ribs, particu- larly near the base, the nectary present; corolla dusky pink (red-foliaged plants) or lemon yellow (green-foliaged plants), with a red-purple centre, the petals 2-5.5 x 1.5-3.5 cm, obovate, gla- brous; staminal column 10-20 mm long, the filaments 1-2 mm long, the anthers purple, the pollen orange or tan; style branches included in the staminal column or up to 5 mm long; stigmas pink to red- purple. Capsule 19-25 x 12-18 mm, ovoid, almost glabrous to appressed-pubescent, the beak 1-2 mm long, glabrous; seeds 3 x 2.5 mm, subreniform, dark brown, the surface finely faveolate and with pectinate scales, the funiculus small, dark brown, glabrous. Chro- mosome number, n = 36. Cultivated, ruderal, and naturalized; abandoned plantations, marshy habitats, forest clearings. DISTRIBUTION. Probably African in origin (an amphidiploid species, possi- bly from hybridization of Hibiscus asper with//, surattensis: Menzel, 1986: 451). Now widely distributed in tropical and subtropical areas of both hemispheres as a cultigen used for ornamental and food purposes. Westphal & Westphal-Stevels 8978 (PRE) noted that the leaves are used for making tea in Cameroon. Ivory Coast, Adiopodonne, December 1963, Ake Assi 7376 (G). Cameroon, Ipono, near Ntem River, 3 December 1975, Westphal & Westphal-Stevels 8978 (PRE). Sudan, Kagelu, 1938, Myers 11897 (K). Congo, Kinsuka, 6 April 1967, Pauwels 4987 (BR). Democratic Republic of Congo, Bambesa, 23 October 1961, Gerard 5041 (BR); Yangambi, 13 January 1953, Germain 1880 (G). Uganda, Entebbe Region, 13 May 1925, Maitland s.n. (K). Tanzania, Mwanza Region, Sengerema, 26 July 1975, Wilson & Kissie 7561 (NA); Tanga Region, Muheza, 5 August 1975, Wilson & Kissie 75129 (ASU, EA, NA). Principe, S. Antonio, 31 August 1956, Khonua 1 2 1 87 (BM). Sao Tome, S. Vicente, 7 January 1 949, Espirito Santo 77 (BR). Angola, CuanzaNorte, Golungo Alto, without date, Welwitsch 5270 (BM, LISU). Lunda, Dala, 29 June \937,Exell & Mendon^a 1456 (BM); Bie, Chitembo, 2 November 1966, Teixeira 10911 (BR); Moxico, Vila Cuzo- Cuchipoque(?), without date, Gossweiler s.n. (BM). Zambia, Mivinilunga, 1 1 June 1963, Edwards 739 (BR). Zimbabwe, Harare (Salisbury), 23 June 1953, Bertram 43476 (BM, SRGH). Mozambique, Niassa, Vila Cabral, 1 March 1964, Torre & Paiva 10932 (BOL). Mauritius, garden at Phoenix, 19 April 1963, Vaughan 10752 (BM). 27. Hibiscus gilletii deWild. in Ann. Mus. Congo, Ser. Bot. 5(1): 166 ( 1904). Type: Democratic Republic of Congo (Zaire), Bas-Congo, Kimuenze, March 1901, Gillet2Q57 (BR!-holotype). Annual (or sometimes perennial?) herb, stems branching near the base, sparsely fine stellate-pubescent, with or without coarse stellate hairs on enlarged bases below, coarse stellate hairs becoming more dense above. Stipules 2-6 mm long, filiform to linear, with fine, short pubescence. Leaves: petioles 0.2-7 cm long, with pubescence like that of the stems, the blades 1-5 x 0. 1-5 cm, the blades of lower leaves shallowly to deeply palmately 3-5-lobed, midlobe lanceo- late, ovate, obovate, or rhombic, length-width ratio from 1:1 to 4:1, the blades of upper leaves from bract-like near the apex to deeply 3- lobed, all blades finely stellate-pubescent and sometimes with a few coarse stellate hairs on bases on the veins abaxially and adaxially, the base cuneate, truncate, or cordate, the margin serrate to repand, the apex acute or obtuse, the nectary 1-3 mm long, prominent, slit- like. Flowers clustered at the apex of flowering branches, the branches up to 54 cm long; peduncles 0.2-0.8 cm long, articulating at the base, densely and finely stellate-pubescent and with or without coarse stellate hairs; involucellar bracteoles 10-14, 5-8 mm long below the apex, filiform to linear, densely and finely stellate- pubescent throughout and long-bristly on the margins, the apex bifurcate, the outer fork 1 .5-4 mm long, linear, reflexed, the inner fork 4-9 mm long, filiform, ascending, the base free; calyx 1.0-1.6 cm long, the lobes lanceolate, acute to long-acuminate, finely stellate- pubescent with coarse, 2-fid to stellate hairs on the ribs, the ribs conspicuous, the nectary absent; corolla yellow with a red or reddish brown centre, the petals up to 3.5 cm long; staminal column 12 mm long, filaments 0.5 mm long; style branches 2 mm long. Capsule up to 12 x 11 mm, ovoid, densely appressed-pubescent, the beak inconspicuous; seeds 3x2 mm, subreniform, brownish, with paral- lel striations and a few pectinate scales. 27a. Hibiscus gilletii subsp. gilletii Fig. 3C. Hibiscus gilletii de Wild, in Ann. Mus. Congo, Ser. Bot. 5(1): 166 (1904). Hibiscus poggei Giirke ex Engl. in Engl., Pflanzenw. Ost-Afrikas 3(2): 400 (1921 ). Type: Angola, 'Lunda-Kasai Bezirk', Pogge s.n. (B). Stems finely stellate-pubescent and with sparsely and coarsely stellate hairs throughout; leaves at mid-plant usually 5-lobed, length- width ratio of midlobe c. 2:1 to 3:1; outer fork of involucellar bracteole 2-4 mm long, the inner fork 6-9 mm long; calyx lobes acute to slightly acuminate in fruit. Ruderal and wild, abandoned cultivated fields, savannas, and gallery forests. DISTRIBUTION. Fig. 5. Democratic Republic of Congo, Angola and Zam- bia. Democratic Republic of Congo, Kisai Occidentale, Dibaya, 26 March 1957, Liben 2725 (BR); Miao, 4 June 1913, Sparano 18 (BR); Difuma (Kinshe), 9 March 1934, Rossignol 99 (BR); Katanga, Kafubu, 28 March 1927, Quarre 186 (BR-2 sheets); Wombali, without date, Vanderyst 1297 (BM); Nioki, Dist. du Lac Leopold, Goossens 6070 (BR). Angola, Lunda, Xa-Sangue, 4 April 1937, Exell & Mendonqa 251 (BM); Lunda, near Vila Henrique de Carvalho, 15 April 1 937, Exell & Mendonqa 618 (BM); Moxico, Teixeira de Sousa, 4 July 1940, Gossweiler 125 10 (BM); Nordeste da Lunda, Luachimo, 8 November 1946, Gossweiler 13.82K (BM). Zambia, Kabompo Dist., 6 kmW. of Kabompo, 24 March 1961, Drummond & Rutherford-Smith 7267 (K). 70 F.D. WILSON 10 cm Fig. 3 Leaves and fruits of taxa of Hibiscus section Furcaria and //. trichospermoides. A: H. cuanzensis, Welwitsch 5241 (BM); B: H. mastersianus, leaf: Exell, Mendonga & Wild 1458 (BM), fruit: Beddome 581 (BM); C: H. gilletii subsp. gilletii, Quarre 186 (BR); D: H. sineaculeatus, leaf: Olorunfeni FHI 24354 (K), fruit: fery FHI 22587 (K); E: H. moxicoensis, Exell & Mendonqa 1599 (BM); F: H. scotellii, Glanville 1 1 1 (K); G: H. elongatifolius, Zenker 1440 (P); H: H. trichospermoides, Humbert & Capuron 19291 (G). REVISION OF HIBISCUS SECTION FURCARIA 71 E. Morith at B wrote to E.G. Baker, 5 December 1938. Morith had compared Exell & Mendonga 618 with the type of Hibiscus poggei at B and said that they were the same species. Exell & Mendonga 6 1 8 (BM!) is H. gilletii subsp. gilletii. 27b. Hibiscus gilletii subsp. hiernianus (Exell & Mendonga) F.D. Wilson, comb, et stat. nov. Hibiscus mastersianus Hiern, Cat. Afric. PL 1: 71 (1896), pro parte quoad specims. Welwitsch 4927 et 4928. H. surattensis var. mastersianus (Hiern) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 112 (1900), pro parte quoad specims. Welwitsch 4927 et 4928. Hibiscus hiernianus Exell & Mendonga in J. Bot. 74: 136-137 (1936).Type: Angola, Huila, January 1860, Welwitsch 4927 (BM!- holotype). Stems finely stellate-pubescent and with occasional coarse stellate hairs below, increasing in density above; leaves at mid-plant mainly 3-lobed, the midlobe often conspicuously obovate or rhombic, the length-width ratio approaching 1:1; outer fork of involucellar bracteole 1.5-3 mm long, the inner fork 4-5 mm long; calyx lobes acute to slightly acuminate in fruit. Chromosome number, n = 18. River banks, river marshes, grasslands, and woodlands. DISTRIBUTION. Fig. 5. Angola and Zambia. Angola, River Tiengo Cuito, 5 March 1 906, Gossweiler3665 (BM) [See Exell & Mendonga, 1 95 1 ]; Huambo, 22 February 1927, Lynes s.n. (BM); Huila, May 1860, Welwitsch 4928 (BM); S. Angola, without date, Pearson 2102 (BOL). Zambia, Kasama Dist., Chibutubutu, 24 February 1960, Richards 12562 (K); Mwinilunga, 16 June 1963, Edwards 796 (K). 27c. Hibiscus gilletii subsp. lundaensis (Baker f.) F.D. Wilson, comb, et stat. nov. Hibiscus lundaensis Baker f. in/. Bot. 77: 21 (1939). Type: Angola, Lunda: Cacolo, River Cuilo, 11 April 1937, Gossweiler 11821 (BM!-holotype). Stems finely and coarsely stellate-pubescent throughout; leaves at mid-plant deeply 5-lobed, length- width ratio of the midlobe c. 4:1; outer fork of involucellar bracteole c. 3 mm long, the inner fork 7- 8 mm long; calyx lobes long-acuminate in fruit. Annual herb among grasses. DISTRIBUTION. Fig. 5. Angola. Known to me only from the type collection. 28. Hibiscus moxicoensis Baker f. in /. Bot. 77: 21 (1939). Type: Angola, Moxico: River Luena, near Vila Luzo, c. 1240 m, 4 May 1937, Exell & Mendonga 1599 (BM!-holotype). Fig. 3E. Perennial (annual?), prostrate herb; stems patchily lanate-pubescent below, continuous above; petioles, leaf blades, peduncles, involucellar bracteoles, and calyx also lanate or patchily lanate-pubescent (indi- vidual hairs stellate), interspersed with fine to coarse, simple to stellate hairs. Stipules 6 mm long, filiform, with short, fine hairs. Leaves: petioles 0.4-2.0 cm long, the blades 1-3.5 x 0.3-3.8 cm, deeply 3- to 5-lobed below to bract-like at the apex, the midlobe of lower leaves oblanceolate, the lateral lobes narrowly lanceolate, the outer lobes lanceolate, the midlobe and lateral lobes also secondarily lobed, the base cuneate, truncate, or subcordate, the margin regu- larly to irregularly serrate, the apex acute, the nectary 2 mm long, slit-like. Flowers sometimes axillary, solitary but mostly clustered at the apex of the main shoot or on flowering branches up to 8.5 cm long; peduncles 0.4 cm long in fruit, articulating at the base; involucellar bracteoles 10-12, 4 mm long below the apex, filiform to linear, the apex bifurcate, the outer fork 3 mm long, linear, reflexed, the inner fork 4 mm long, linear, ascending, the base free; calyx 1.1- 1 .4 cm long in fruit, the lobes lanceolate, slightly acuminate, the ribs conspicuous, the nectary absent; corolla yellow with a red-brown centre. Capsules 11x10 mm, ovoid, densely appressed-pubescent, the beak 1 .5 mm long, glabrous; seeds 2x2 mm, subreniform, very dark brown, with parallel striations, the funiculus inconspicuous, glabrous. Habitat unknown. DISTRIBUTION. Fig. 5. Angola. Known to me only from the type collection. 29. Hibiscus cuanzensis Exell & Mendonga in /. Bot. 77: 136 (1936). Type: Angola, CuanzaNorte, Pungo Adongo, April 1857, Welwitsch 5241 (BM!-holotype; G!, LISU!, P!-isotypes). Fig. 3A. Hibiscus furcatus Garcke in Linnaea 43: 121 (1881), non Roxb. (1814). Hibiscus mastersianus Hiern, Cat. Afr. PL 1:1 \ (1896), pro parte quoad specim. Welwitsch 5241. Perennial herb to 1 .5 m tall; the stems finely stellate-pubescent and with coarse stellate hairs, increasing in density towards the apex, not aculeate. Stipules 5-7 mm long, linear, densely and finely pubes- cent. Leaves: petioles 0.5-7 cm long, densely and finely stellate-pubescent, the blades 2-7.5 x 0.6-9 cm, the blades on lower leaves moderately 5-lobed, the midlobe and lateral lobes ovate, the outer lobes triangular, the blades of upper leaves lanceolate, ovate, or shallowly 3-lobed, the blades of all leaves densely stellate- pubescent and with a few short simple hairs abaxially, densely stellate-pubescent and with a few longer (to 2 mm) simple hairs adaxially, the base cuneate to truncate, the margin finely serrate to irregularly dentate, the apex acute, the nectary 2 mm long, slit-like, on upper leaves and on mid-plant leaves, apparently absent on lower leaves. Flowers axillary, solitary, or sometimes with more than one flower at a node or on a short flowering branch; peduncles 0.3-0.6 cm long, articulating at the base, densely and finely stellate-pubes- cent; involucellar bracteoles 9-10, 5-6 mm long below the apex, linear, finely pubescent and with longer bristles, the apex bifurcate, the outer fork 4 mm long, reflexed, the inner fork 5 mm long, ascending, the base free; calyx 1 .4-1 .6 cm long, the lobes triangular, acute to slightly acuminate, purple-punctate, densely and finely stellate-pubescent and with a few coarse 2-fid to stellate hairs on bases, the nectary absent; corolla yellow. Capsules 14 x 10 mm, ovoid, densely appressed-pubescent, the beak < 1 mm long, obscure; seeds unknown. 'Ad dumetria junta lafera rivulorum', etc. (Welwitsch 5241); 'erva vivaz do mato xerofilo' (Exell & Mendonga, 1937: 167). DISTRIBUTION. Fig. 5. Angola. Narrow endemic, collected only from 'Cuanza Norte, Pungo Adongo'. Exell & Mendonga (1937) cite, besides the type specimen, Mechow 103 (B), from the same general location. Exell (1936) and Exell & Mendonga (1937) also cited Welwitsch 5242 under Hibiscus cuanzensis, but it is in fact H. mastersianus Hiern. 30. Hibiscus mastersianus Hiern, Cat Afric. PL 1: 71 (1896) emend, excl. specim. angol. Welwitsch 4927, 4928, 5241. Type: Mozambique, Lupata, Kirk s.n. (K-lectotype, designated by Milne-Redhead, Bull. Misc. Inform. 1935: 273 (1935)). Fig. 3B. Hibiscus furcatus Mast, in Oliver, Fl. Trop. Afr. 1: 201 (1868), non Roxb. (1814), pro parte excl. specim. ex Gambia. 72 F.D. WILSON Hibiscus surattensis var. mastersianus (Hiern) Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 12(1900),excl. specims. Welwitsch 4927, 4928. Hibiscus pachmarhicus Haines in Bull. Misc. Inform. 1914: 24 (1914). Type: India, Central Provinces; common in the middle Gondwana sandstones about Pachmarhi in the Satpura range, 900 m, October 1911, Haines 197P (K-holotype). IHibiscus beddomei Rakshit & Kundu in Sci. & Cult. 27: 192 ( 196 1 ).Type: India, South India, without precise locality, Beddome 91 & 92 (CAL-holotype). Illustrations: Haines, Bull. Misc. Inform. 1914: 25, figs 1-7 (1914) (as//, pachmarhicus); Rakshit & Kundu, Sci. & Cult. 27: 192, fig. 1 (1961) (as//, beddomei). Annual erect herb to 2 m tall; stems with sparse to dense, stiff simple to 3-fid or stellate pigmented bristles on enlarged bases and one or more lines of dense pubescence, not aculeate. Stipules 5-8 mm long, filiform. Leaves: petioles <1-18 cm long, hairy like the stems, the blades 2-16 x 0.3-15 cm, very shallowly palmately 3-lobed below to entire above, or bract-like near the apex, sparsely to densely stellate-pubescent on both surfaces, the base cuneate to truncate or cordate, the margin serrate to dentate, the apex acute to slightly acuminate, the nectary 1-3 mm long, slit-like, near base of midrib, and sometimes on the two adjacent ribs. Flowers axillary, solitary; peduncles 0.4-0.8 cm long, articulating at the base, hispid and with a few long bristles; involucellar bracteoles 9-10, 9-11 mm long, linear, ascending, hispid and with 1-2-fid bristles on the margins, the apex bifurcate, the outer fork 3—4 mm long, lanceolate, the inner fork 2-5 mm long, linear, the base free; calyx 1.1-2.0 cm long, the lobes lanceolate, slightly to long-acuminate, bristly mainly on the ribs, the nectary absent; corolla yellow with a purple centre, the petals 3-5 x 2-3 cm, obovate, sparsely stellate-pubescent dorsally, glabrous ventrally; staminal column 15 mm long, the filaments 1 mm long; style branches 1 mm long. Capsules 12-15 x 9-11 mm, ovoid-acuminate, densely appressed-pubescent, beak 2 mm long, glabrous; seeds 4x3 mm, angular to subreniform, faveolate, beset with numerous pectinate scales. Chromosome number, n = 18. Ruderal and wild, in old cultivation, in open and mixed grassland, savannas, dense bush and woodland. DISTRIBUTION. Fig. 5. Eastern, southern and southwestern Africa, Tanza- nia to South Africa; possibly adventive in India. An anonymous reviewer reported it as having been recorded from Mutomo Hill, Kitui Dist., Kenya, but I saw no specimens from Kenya Tanzania, Kondoa Dist., Sambala, 28 March 1929, Burtt 2152 (BM); Iramga Plateau, April 1959, Hammond 263 (K); Nbulu Island, Lake Tanganyika, 1 April 1955, Richards 5389 (BR); Western Province, Tabora-Itigi, 27 April 1962, Tallantire T628 (K); Mpulungu...Pier L'ile de Niamkolo, 27 March 1947, Van Meel 1352 (BR); Mwanza Region, 27 km E. of Geita on Hway B-163, 26 July 1975, Wilson & Kissie 7559 (ASU, EA, NA). Angola, PungoAdongo, 1 May 1857, Welwitsch 5242 (BM); Okavongo Native Territory, 25 February 1956, deWinter & Marais 4865 (BM). Zambia, Wankie Dist., Victoria Falls Village, 14 March 1 974, Gonde 69/74 (K); Livingstone, lOApril \930, Jenkins 44 (BM); 1 mile S. of Kafue on road to Livingstone, 2 April 1969, Jones 69126 (NA); Livingstone and common all along the line, April 1909, Rogers 7007 (BOL, K). Mozambique, Tete, Cabora Bassa, 19 April 1972, Pereira & Correia 2143 (BM). Namibia, Tsumeb, 20 March 1934, Dinter 7462 (BM); Grootfontein-Nord, zwischen Karakowisa und Numkaub im Omuramba, 5 March 1958, Merxmiiller & Giess 1819 (BM); Botswana, Northern Div., Chobe-Zambesi confluence, 11 April 1955, Exell, Mendonca & Wild 1458 (BM); Savuti, Chobe Nat. Park, 25 March 1984, Jacobsen 3030 (PRE); Northern Div., 51 km S. of Shakawe on Sepopa Rd, 16 March 1965, Wild & Drummond 7 102 (BR). Zimbabwe, Harley Dist., 1 3 January 1952, Hornsby 3256 (BM); Victoria Falls, 1 1 April 1925, Pocock 9 (BOL). South Africa, Transvaal, 14 January 1894, Schlecter 4160 (BOL). India, Madras, Jubbulpore, 1885, Beddome 581 (BM). Hiern (1896) cited Welwitsch 4927, 4928, 5241, and 5242 from Angola as specimens of Hibiscus mastersianus. Welwitsch 4927 (BM!) and Welwitsch 4928 (BM!, LISU!) are referable to//, gilletii subsp. hiernianus; Welwitsch 5241 (BM!, G!, LISU!, P!) is referable to H. cuanzensis; Welwitsch 5242 (BM!, G!, LISU!) is typical //. mastersianus, not a form of H. cuanzensis, as Exell & Mendonsa (1937) surmised. Sacleux 2141 (P-2 sheets, one collected July 1894, the other collected February 1898), from Kenya, is near //. mastersianus, but may be an undescribed species; it has very distinc- tive deeply 7-lobed leaves; the lobes are obovate and are themselves secondarily lobed. 3 1 . Hibiscus elongatifolius Hochr. in Annuaire Conserv. Jard. Bot. Geneve 4: 1 17 (1900). Type: Cameroon, Sanaga, Urwaldgebiet, 1897, Zenker 1440 (P!-holotype; BM!, G! [2 sheets]-isotypes). Fig. 3G. Habit unknown; stems pilose, with matted grey hairs. Stipules 8-1 1 mm long, filiform or linear, each stipule divided into 2 or 3 filiform or linear segments, finely appressed-pubescent. Leaves', petioles 0.3-0.8 cm long, with pubescence like that of the stem, the blades 2.5-12.5 x 0.3-0.8 cm, linear, abaxially densely grey stellate- pubescent, adaxially with less dense, simple to 3-fid grey hairs, the base cuneate, the margin serrate, the apex acute, the nectary absent. Flowers borne on a leafless shoot, at first clustered near the apex, later axillary, solitary, as the shoot elongates; peduncles 1 .7-2.2 cm long, articulating at the base, with pubescence like that of the stem; involucellar bracteoles 11-13, 9-12 mm long, filiform-linear, as- cending, with dense, soft, appressed pubescence, the apex entire, the base free; calyx 1.4-1.5 cm long, the lobes lanceolate, acuminate, densely and softly appressed-pilose, the nectary 1 mm long, slit-like; corolla orange-red(?), petals 2 cm long, glabrous(?), the petal spot not seen. Capsules 10x10 mm, oval, densely appressed-pubescent, the beak inconspicuous; seeds unknown. DISTRIBUTION. Fig. 5. Cameroon. Known to me only from the type collec- tion. 32. Hibiscus sineaculeatus F.D. Wilson, sp. nov. Type: Nigeria, Zaria Province, Kan Gimi Veg. mapping area, Dutsen Gwagwa, 20 October 1947, Keay FHI 201 17 (K!-holotype). Fig. 3D. Hibiscus scotellii Keay, Fl. W. trop. Afr. 2nd ed. 1(2): 347 (1958), non Baker f. (1894), pro parte quoad specims. Keay FHI 20017, Lely 609, P686. Caules non aculeati; folia ovata ad 3-lobata; pedunculi fructificantes <1 cm longi; bracteolae involucellorum 2 mm latae ad basim, non bifurcatae, subulatae, apicibus acutis; calyces pilis stellatis tenuis atque setis rigidis ad basim rubris, nectariis in quoquo costa. Erect shrubby herb to 2 m tall; stems, petioles, leaf blades, pedun- cles, and involucellar bracteoles densely and finely stellate-pubescent and sometimes with some coarse stellate hairs, not aculeate. Stipules 2-4 mm long, filiform to linear, with short, fine hairs. Leaves: petioles 0.8-6 cm long, the blades 2-8 x 0.7-5 cm, on lower leaves ovate to palmately shallowly or deeply 3-lobed, on upper leaves bract-like to narrowly lanceolate, the base cuneate, truncate, or cordate, the margin serrate, dentate or repand, the apex acute to obtuse, the nectary 0.5-1.5 mm long, slit-like. Flowers axillary, solitary, or with an additional smaller flower on a short branch; peduncles 0.2-1.0 cm long, articulating at the base; involucellar bracteoles 6-7, 6-10 mm long, 2 mm wide at the base, subulate, several nerved, the apex entire, pointed, the base free; calyx 1.1- 1 .8 cm long, the lobes triangular to broadly lanceolate, short- to REVISION OF HIBISCUS SECTION FURCARIA 73 long-acuminate, finely stellate-pubescent and with simple to stellate stiff bristles on enlarged red bases on the ribs, the nectary small but conspicuous; corolla pale yellow to yellow with a deep crimson to purple centre, the petals 3—4- x 1-2 cm, obovate, with sparse simple to stellate pubescence dorsally, glabrous(?) ventrally. Capsules 10- 14 x 12-14 mm, with long, dense appressed pubescence, the beak 1-1.5 mm long, glabrous; seeds unknown. Ruderal and wild, in farm fields, in rocks in grasslands and in open savanna woodlands. DISTRIBUTION. Fig. 5. Nigeria. Nigeria, Panshanu, N. Nigeria, 20 Septem- ber 1921 , Lely 609 (K); N. Nigeria, without precise locality, September 1930, Lely P686 (K); Zari Prov., Zari Dist., Anara F. R., Kangimi, 23 September 1948, Olorunfeni FHI 24354 (K); Oyo Prov., Oyo Dist., about 3 miles to Oke- iho on the Iseyin road, 1 7 October 1 959, Onochie FHI 40873 (K); Ondo Prov., Idanre Hills, Carter Peak, 24 October 1948, Keay FHI 22587 (K). 33. Hibiscus scotellii Baker f. in /. Linn. Soc., Bot. 30: 74—75 ( 1 894). Type: Sierra Leone, on gneissose rocks by Scarcies River, 2 miles N. of Sasseni, January, Scott-Elliott 4535 (K-holotype; BM!-isotype). Fig. 3F. Hibiscus cannabinus var. chevalieri Hochr. (1901), pro parte quoad specim. Chevalier s.n. (G!). Erect, little-branched herb to 2 m tall; stems, stipules, petioles, leaf blades, peduncles, bracteoles, and calyx densely covered with very fine stellate pubescence. Stipules 2-^ mm long, linear to subulate. Leaves: petioles 0.2-5 cm long, the blades 1-7 x 0.3-7 cm, the blades of lower leaves shallowly to deeply 3-5-(7)-lobed, the lobes usually oblanceolate to secondarily lobed, the blades of upper leaves strap-like or bract-like, the base cuneate, the margin distantly and finely serrate, the apex acute or obtuse, the nectary 0.5 mm long, usually on the midrib and two lateral ribs. Flowers axillary, solitary or sometimes clustered on short branches; peduncles 0.2-1 cm long, articulating at the base; involucellar bracteoles 6-7, 4-10 mm long, 1 mm wide at the base, linear, flattened, several-nerved, the apex entire, slightly broadened to lanceolate, the base free; calyx 0.7-1.4 cm long, the lobes triangular to broadly lanceolate and acuminate, the nectary present but sometimes inconspicuous; corolla yellow to primrose yellow, with a dark red to purple centre. Capsules 6-1 1 x 6-10 mm, ovoid, densely appressed-pubescent, the beak <1 mm long, inconspicuous; seeds 3x2 mm, subreniform, shiny, dark brown, sparsely covered with minute, 2-fid to stellate hairs, the funiculus small, with very fine hairs. In open, sunny locations in grasslands and among rocks. DISTRIBUTION. Fig. 5. Sierra Leone, Ivory Coast, and Mali. Sierra Leone, Tisana, Bonthe Island 14 November 1931, Deighton 2318 (K); Lungi, 28 November 1928, Glanville 111 (K-2 sheets); E. of York on road, 2 April 1958, Melville & Hooker 381 (K); Falaba Rest House, 26 November 1965, Morton SL 2852 (K); Sugarloaf, 1 892, Scott Elliott 403 1 (BM). Mali, Sindou et Niger moyen, Chevalier s.n. (G). Ivory Coast, near Brafoudi, 75 km NW of Abidjan, 30 December 1958, Leeuwenberg 2293 (B). POORLY KNOWN AND EXCLUDED SPECIES Hibiscus baumanii Ulbr. inNotizbl. Bot. Gart. Mus. Berlin-Dahlem 8: 167 (1922). Type: Togo, tree prairies and savannahs N. to 7° N, Baumann 400 (B-holotype). Ulbrich stated that this taxon is related to Hibiscus elongatifolius Hochr. and placed it in his series Furcaria cannabina. No specimen was found at B. Hutchinson & Dalziel (1928) treated H. baumannii as a synonym of//, congestiflorus Hochr. (section Ketmia) [Annuaire. Conserv. Jard. Bot. Geneve 10: 21 (1906)] and cited Baumann 400 under this species. Hibiscus beddomei Rakshit & Kundu in Sci. & Cult. 27: 192 (1961 ).Type: India, South India without precise locality, Beddome 91 & 92 (CAL-holotype). This plant is probably conspecific v/ithHibiscus mastersianus Hiern. Both have stiff stem pubescence, ovate, shallowly 3-5-lobed hairy leaves with cordate bases, 9-10 stiffly pubescent involucellar bracteoles, acuminate calyx lobes with pubescence on the ribs (otherwise almost glabrous), and beaked, densely pubescent cap- sules. The plant in the photograph accompanying the description of H. beddomei (Rakshit & Kundu, 1961), even though poorly pre- sented, strongly resembles H. mastersianus. Unfortunately, I have been unable to examine any authentic specimens of H. beddomei. Hibiscus campanulifolius Ulbr. in Notizbl. Bot. Gart. Mus. Berlin- Dahlem 8: 166 (1922). Type: Namibia, Tsumeb, Dinter 1692 (B-holotype). Ulbrich places this species in series Furcaria cannabina but pencil- led notes on two sheets at BM of sketches of the type specimens of Hibiscus campanulifolius suggest that it is referable to//, lunarifolius Willd. or H dongolensis Del., in section Ketmia. Hibiscus comoensis A. Chev., Expl. hot. Afrique occ. franc. 1: 65 (1920), nom. nud. Hutch. & Dalziel, Fl. W. trop. Afr. 1: 265, in clavi, 267 (1928); Hutch. & Dalziel, Kew Bull. 1928: 298 (1928) descr. Type: Ivory Coast, Middle Comoe: between Akabossue and Ebrinahone, 19 December 1909, Chevalier 22613 (P!- holotype; BR!, P!-isotypes). Hibiscus comoensis, a plant of southeastern Ivory Coast (Chevalier 22613) and southwestern Ghana [between Kwapon and River Tano, c. 90 km W. of Kumasi, 23 December 1963, Oldeman 807 (BR)] (Figs 5, 6), was associated with other species of Hibiscus section Furcaria by Hutchinson & Dalziel (1928) in their key because the involucellar bracteole has a foliaceous appendage. This species does not fit into any of the currently recognized sections of Hibiscus because of its umbellate inflorescence, much reduced calyx lobes (0.4-0.5 cm long) with poorly developed ribs, and alternating involucellar bracteoles that are linear below and ovate above and much longer than the calyx lobes (6-7 mm long below and 3-6 mm long and 2-4 mm wide in the ovate portion). Hibiscus friesii Ulbr. in Feddes Repert. 13: 521 (1915). Type: Zambia, Chiruktu bei Broken Hill, Fries 239 (UPS-holotype). Ulbrich placed this species in section Furcaria in his series Friesia. I did not see a specimen from B. Baker (1937) moved series (subsection) Friesia to the genus Kosteletskya. Exell (1961) placed Hibiscus friesii in synonymy with Hibiscus pan duriformis Burm.f., but suggested the possiblity that it is a separate taxon. At any rate, //. friesii does not belong in section Furcaria. Hibiscus furcatus Bahadur, Dayal & Raturi in J. Bombay Nat. Hist. Soc. 70: 495^96, pi. II (1970), non Roxb. (1814). This plant, collected in northwestern India, is certainly not conspecific with Hibiscus furcatus Roxb. (H. hispidissimus Griff.). In fact, it may be a previously undescribed species of Hibiscus section Furcaria. 74 F.D. WILSON H mgncaulis H. green way i H. sparseaculeatus H. ttiversifolius: • subsp diversifolius • subsp. rivularis H.asper Fig. 4 African distributions of taxa of Hibiscus section Furcaria. REVISION OF HIBISCUS SECTION FURCAR1A 75 H gilletii. • subsp gilletii m subsp. hiemianus A subsp lundaensis H. mastersianus • H. scotellu • H smeaculeatus A H comoensis T H. cuanzensis H. sudanensis H. reekmansii H. furcellatoMes H. elongatifolius H. goossensii H. flavo-mseus H. moxicoonsis H.tomi Fig. 5 African distributions of taxa of Hibiscus section Furcaria. Bahadur et al. (1970) wrongly placed the following taxa in syn- onymy with the species that they called H. furcatus Roxb.: H. aculeatus Roxb., H. furcellatoides Hochr., H. hispidissimus Griff., and H. rostellatus Guill. & Perr. Hibiscus furcatus var. microcarpus Mattei in Boll Reale Orto Bot. Giardino Colon. Palermo!: 103 (1908). Type: Somalia, Giumbo in reg. Goscia, Macaluso 55 (PAL-holotype). Mattei differentiates this taxon from the Indian species Hibiscus furcatus [referring to//, furcatus Roxb., not loH. furcatus Willd.] by its short peduncles, small, deep yellow corolla, included capsule, and very short calyx. He also states that it has an affinity with H. surattensis. No specimen was found at PAL. Hibiscus hispidissimus A. Chev., Rev. Bot. Appliq. 20: 326 (1940), non Griff. (1854), nom. illeg. (Art. 53.1) [based on Hibiscus abelmoschus var. hispidissimus A. Chev., Explor. Bot. I'Afrique occ. franc,. 1: 63 (1920), nom. nud.] = Abelmoschus esculentus (L.) Moench (see Keay, 1958: 348). Type: Ivory Coast, Cercle du Baoule-Nord Vallee du Nzi moyen; entre Korakissikro et Manikro Casut, 1909, Chevalier 22307 (P!). My opinion is that Chevalier 22307 is a typical specimen of Abelmoschus esculentus. Hibiscus hoshiarpurensis T.K. Paul & M.P. Nayar in Bull. Bot. Surv. India 25: 188-189, figs 1-6 (1983). Type: India, Punjab, Dholbah, Hoshiarpur Dist., 22 September 1970, Misra 41888 (BSD-holotype). This plant is closely related to Hibiscus mastersianus (including H. beddomei) but seems to be a genuine species, according to the description and accompanying illustration. It has lanceolate, unlobed leaves, and shorter petioles and longer calyx lobes in relation to the capsule than does H. mastersianus. It occurs in extreme northwest- ern India (Punjab), whereas H. mastersianus has a more southerly distribution in India. Hibiscus imerinensis Ulbr. in Notizbl. Bot. Gart. Mus. Berlin- Dahlem 8: 168 (1922). Type: Madagascar, Imerina, near Tananarive, Hildebrandt 4033 (BM!, G!-isotypes). Hochreutiner [Fl. Madagasc., Malvac.: 46 (1955)] lists Hibiscus imerinensis as a synonym of H. stenophyllus var. solandroketmia (Hochr.) Hochr. [section Solandra] . At any rate, it does not belong in section Furcaria. Hibiscus keilii Ulbr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 8: 681 (1924). Type: Tanzania (Ostafrika), Usumbura, Luwironsa- Ufer bei Mugera, 4 August 1905, Keil 183 (B-holotype). 76 F.D.WILSON 10 cm Fig. 6 Hibiscus comoensis, Oldeman 807 (BR). REVISION OF HIBISCUS SECTION FURCARIA 77 Ulbrich placed this species in series Furcaria furcellata and noted that it is closely related to Hibiscus rostellatus. Hibiscus paolii Mattei in Chiovenda, Result. Sci. Somalia ital. 1: 32 (1916). Type: Somalia, Bur-Meldac, Paoli 699 (PAL-holotype). A sheet at BM (012890) has a poor sketch of this species. An abbreviated description on this sheet shows that this species belongs to section Furcaria and suggests a close affinity with Hibiscus rostellatus (rhomboid leaves and bifurcate involucellar bracteoles). Hibiscus persicifolius Eckl. & Zeyh., Enum. pi. afric. austral.: 38 (1835); Harvey & Sonder, Fl Cap. 1: 171-172 (1860). Type: South Africa, in the Great Karroo, between Graaf Reynet and Uitenhage, Ecklon & Zeyher 305 (Herb. Sond.). I made inquiries at six herbaria (BM, K, MEL, NBG, S, and TCD) known to have Sonder specimens. A fragmentary sterile specimen of Ecklon & Zeyher 305 was found at NBG (photocopy seen), and two fragmentary specimens, one sterile and one flowering, were found at S (not seen). I designate the flowering specimen of Ecklon & Zeyher 305 at S as lectotype and the other specimens of this same collection as isolectotypes. In addition, a fragmentary sterile specimen of apparently the same species (Ecklon & Zeyher s.n.) was found at MEL (photocopy seen). No specimens were found at the other three herbaria The appearance of the plant fragments and the sketchy description of Hibiscus persicifolius suggests strongly that this is the Australian species, H. heterophyllus Vent. (In particular, the pedun- cles and calyx clothed with golden-yellow echinate tubercles). If so, the specimen must have been collected from a cultivated plant or one that had escaped cultivation. Ecklon & Zeyher's description of the flower as purplish is a little disturbing because H. heterophyllus has white or yellow flowers. However, the corollas of the white-flow- ered form are pink-tinged, particularly on the dorsal surface, and dry to a dark pink color. Hibiscus procerus Roxb. ex Wall., Cat. no. 2692. Roxburgh's illustration [Icones Roxburghianae 1506 (K!)] is of the white-flowered form of Hibiscus heterophyllus from Australia. Masters in Hooker, Fl. Brit. India 1: 339 (1872) gives as type: Burma, Wallich. If this plant was collected in Burma, it was obvi- ously cultivated, not native there. Hibiscus saxicola Ulbr. in Notizbl. Bot. Gart. Mus. Berlin-Dahlem 7: 179-1 80 (1917).Type: Cameroon, Ebolowa region, Mildbraed 5515 (B-holotype). In his original description, Ulbrich placed this species in section Azanza, but later erected series Saxicolae under section Furcaria to accommodate it. Hibiscus trichospermoides Hochr. inAnnuaire Conserv. Jard. Bot. Geneve 20: 82 (1916). Type: Madagascar, Gres liasiques, bois sees, Mt Ampohipirika (Herbier Perrier de la Bathie n. 62) (G!- holotype). Fig. 3H. Hibiscus bernieri van trichospermoides (Hochr.) Hochr., Fl. Madagas., Malvac.: 65, fig. 17 (1955). Illustration: Hochreutiner, Fl. Madagasc., Malvac.: fig. XII, 3-6 (1955). The sectional placement of Hibiscus trichospermoides is in doubt. My opinion is that it does not belong in section Furcaria. The calyx nervation resembles more closely other species of section Solandra rather than section Furcaria, but the capsule and seed characteristics are not like other species of either of these sections. No other African or Asian species of section Furcaria shows the same combination of diagnostic characters: stem not aculeate, peduncles long, articulat- ing above the base, involucellar bracteoles not bifurcate, calyx nectary absent. Furthermore, the capsule has a dense stellate, unappressed tomentum, and the seed surface is covered with silky hairs to 3 mm long, characters that appear in no other species of Furcaria with which I am familiar. Hochreutiner, in his description of H. trichospermoides, thought that its closest affinity was with H. heterophyllus and related Australian species of section Furcaria, but later regarded it merely as a variety of H. bernieri Baill., which he inexplicably placed in section Lilibiscus. I have seen no authentic specimens of Hibiscus bernieri var. bernieri but Hochreutiner's illustration and description suggest that these two entities, both endemic in Madagascar, are indeed closely related. Figure 3H is Basse Mahahavy du Nord, au Sud d' Ambilobe, April 1951, Humbert & Capuron 19291 (G). ACKNOWLEDGMENTS. I thank O.J. Blanchard, P.A. Fryxell, and two anony- mous reviewers for their helpful comments, the curators of various herbaria from which I borrowed specimens or solicited assistance, and Donald J. Pinkava and Leslie R. Landrum, Arizona State University, for their cheerful assistance and for providing storage and working space. I also thank Thomas J. Henneberry, Laboratory Director, USDA-ARS, Western Cotton Research Laboratory, for providing office, field, and glasshouse space and technical help. Finally, I thank Benny R. Stapp and Patrick Alexander for maintaining the plants, and Marsha Minniss for preparing the illustrations. REFERENCES Abdallah, M.S., Sa'ad, F.M. & El-Kholy, S.A. 1976. Taxonomical studies in the flora of Egypt VI. Culinary cultivars of Hibiscus sabdariffa L. in Egypt. Notes Agricul- tural Research Centre Herbarium Egypt 3: 1-20. Bahadur, K.N., Dayal, R. & Raturi, D.P. 1970. New plant records for the upper Gangetic Plain. Journal of the Bombay Natural History Society 70: 495^4-96, pi. II. Baker, E.G. 1937. Notes on tropical African species of Hibiscus. Journal of Botany 75: 98-102. Berhaut, J. 1954. Flore du Senegal, Cle Pratique. Dakar. Borssum Waalkes, J. van 1966. Malesian Malvaceae revised. Blumea 14: 1-213. Cheek, M. 1 992. Hibiscus sparseaculeatus. Flowering Plants of Africa 52( 1 ): pi. 2058. Chevalier, A. 1920. Malvaceae. Exploration Botanique de L'Afrique Occidental Franfaise 1: 56-75. Paris. Crane, J.C. 1949. Roselle-a potentially important plant fiber. Economic Botany 3: 89- 103. Dalziel, J.M. 1948. The useful plants of West Tropical Africa. London. Dempsey, J.M. 1975. Fiber crops. Gainesville. Duke, J.A. & duCellier, J.L. 1993. CRC Handbook of alternative cash crops. Boca Raton. Exell, A.W. 1936. Notes on the flora of Angola-Ill. Journal of Botany 74: 132-140. 196 1 . Hibiscus. In A.W. Exell & H. Wild (Eds), Flora Zambesiaca 1(2): 434-472. London. & Mendonca, F.A. 1937. Hibiscus. In L.W. Carrisso (Ed.), Conspectus Florae Angolensis 1(1): 158-176. Lisbon. 1951. Hibiscus. In L.W. Carrisso (Ed.), Conspectus Florae Angolensis 1(2): 177-180, 374-376. Lisbon. Fryxell, P.A. 1988. Hibiscus. In Systematic Botany Monographs 25: 192-232. Ann Arbor. 1992. Malvaceae. In G. Marling & L. Anderson (Eds), Flora of Ecuador. Copenhagen. Goforth, C.E. & Fuller, MJ. 1994. A summary of kenaf production and product development research 1989-1993. Mississippi Agriculture and Forestry Experimen- tal Station Bulletin 101 1. Mississippi State. 11 ;iu man. L. 1 963. Hibiscus. In W. Robyns (Ed.), Flore du Congo et du Ruanda-Urundi 10: 94-136. Brussels. 78 F.D. WILSON Hiern, W.P. 1 896. Catalogue of the African plants collected by Dr. Friederich Welwitsch in 1853-61. Dicotyledons. London. Hochreutiner, B.P.G. 1 900. Revision du genre Hibiscus. Annuaire Conservatoire & du Jardin Botanique Geneve 4: 23-191. 1901. Malvaceae Chevalieranae. Annuaire Conservatoire & du Jardin Botanique Geneve 5: 120-126. 1955. Hibiscus. In H. Humbert (Ed.), Flore de Madagascar et des Comores, Malvaceae: 8-83. Paris. Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. 1990. Index herbamrium Pan I: The herbaria of the world, 8th ed. Bronx. Howard, A. & Howard, G. 1911. Studies in Indian fibre plants 2. On some new varieties of Hibiscus cannabinus L. and Hibiscus sabdariffa L. Memoirs of the Department of Agriculture in India, Botanical Series 4: 1-36. lliitdiinM.ii. J. & Dalziel, J.M. 1928. Flora of West Tropical Africa 1st ed. 1(2). London. Kachecheba, J.L. 1972. The cytotaxonomy of some species of Hibiscus. Kew Bulletin 27: 425-433. Keay, R.WJ. 1958. Flora of West Tropical Africa 1(2), 2nd ed. London. Masters, M.T. 1 868. Hibiscus. In D. Oliver (Ed.), Flora of Tropical Africa 1: 194-208. London. 1872. Hibiscus. In J.D. Hooker (Ed.), The flora of British India 1(1): 334-344. London. Menzel, M.Y. 1986. Genetic relationships among the relatives of Hibiscus cannabinus and H. sabdariffa. In K.A. Siddiqui & A.M. Faruqui (Eds), New genetical ap- proaches to crop improvement'. 445—456. Karachi. & Wilson, F.D. 1969. Genetic relationships in Hibiscus sect. Furcaria. Brittonia 21: 91-125. Ochse, JJ. 1960. Vegetables of the Dutch East Indies. Buitenzorg. Paul, T.K. & Nayar, M.P. 1980. The status of Hibiscus furcatus Roxburgh. Bulletin of the Botanical Survey of India 22: 194-195. 1988. Hibiscus. In M.P. Nayar, K. Thothathri & M. Sanjappa (Eds), Fascicles of Flora of India: 1 19-159. Calcutta. Perry, R.C., Jones, D.E. & Bhangoo, M.S. 1993. A preliminary study on kenaf as a feed for livestock. In M.S. Bhangoo (Ed.), Proceedings 1993 International Kenaf Conference'. 45-48. Fresno. Pradeep, A.K. & Sivarajan, V.V. 1 99 1 . Hibiscus hispidissimus, the correct name for H. furcatus Roxb. ex DC. non Willd. and H. aculeatus Roxb. non Walter (Malvaceae). Taxon 40: 634-637. Rakshit, S.C. & Kundu, B.C. 1961. New species and varieties of Hibiscus. Science & Culture 27: 192-194. 1970. Revision of the Indian species of Hibiscus. Bulletin of the Botancial Survey of India 12: 151-175. Saldanha, CJ. 1985. Hibiscus. In Flora of Karnataka 1: 248-253. Rotterdam. Sprague, T.A. 1913. Hibiscus asper. Kew Bulletin of Miscellaneous Information 1913: 418^19. Taylor, C.S. 1992. Kenaf: annual fiber crop products generate a growing response from industry. In U.S. Department of Agriculture Yearbook, New Crops, New Uses, New Markets: 92-98. Washington, D.C. Werber, F.X. 1993. Kenaf product and process research in ARS. In M.S. Banghoo (Ed.), Proceedings 1993 International Kenaf Conference: 45—48. Fresno. Wester, P.J. 1914. New varieties of roselle. Philippine Agricultural Review 7: 266-269. Wilson, F.D. 1978. Wild kenaf, Hibiscus cannabinus L. (Malvaceae) and related species in Kenya and Tanzania. Economic Botany 32: 199-204. 1983. The taxonomic status of Hibiscus (sect. Furcaria) berberidifolius A. Rich. Brittonia 35: 175-179. 1994. The genome biogeography of Hibiscus L. section Furcaria DC. Genetic Research & Crop Evolution 41: 13-25. & Menzel, M.Y. 1964. Kenaf (Hibiscus cannabinus L.) and roselle (H. sabdariffa L.). Economic Botany 18: 80-91. REVISION OF HIBISCUS SECTION FURCARIA 79 INDEX Accepted names are in roman, synonyms in italic. Abelmoschus esculentus (L.) Moench 75 Cienfuegosia sp. Cufod. 62 Furcaria Kostel. 50 Furcaria mxburghii Kostel. 55 Furcaria surattensis Kostel. 5 1 Hibiscus abelmoschus var. hispidissimus A. Chev. 75 H. acetosella Welw. ex Hiern 68 H. aculeatus Bahadur, Dayal & Raturi 75 H. aculeatus T.K. Paul & MR Nayar 55 H. aculeatus Roxb. 55 H. aculeatus Walter 48, 57 H. altissimus Hornby 53 H. asper Hook.f. 68 H. asper van punctatus (A. Rich.) Berhaut 68 H. baumanii Ulbr. 73 H. beddomei Rakshit & Kundu 72, 73 H. berberidifolius A. Rich. 61 H. berberidifolius Cufod. 63 H. berberidifolius F.D. Wilson 67 H. bernieri Baill. 77 H. bernieri van bernieri 77 H. bernieri var. trichospermoides (Hochr.) Hochr. 77 H. biflorus A. Spreng. 63 H. campanulifolius Ulbr. 73 H. cannabinus Hiern 65 H. cannabinus Hochr. 59, 61, 68, 69 H. cannabinus L. 66 H. cannabinus Mast. 59, 68 H. cannabinus Merr. 59 H. cannabinus Ulbr. & R.E. Fries 60 H. cannabinus var. chevalieri Hochr. 68, 73 H. cannabinus var. genuinus Hochr. 66 H. cannabinus var. punctatus (A. Rich.) Hochr. 68 H. cannabinus var. purpureus A. & G. Howard 66 H. cannabinus var. radiatus (Cav.) Chiov. 59 H. cannabinus var. ruber A. & G. Howard 66 H. cannabinus var. simplex A. & G. Howard 66 H. cannabinus var. unidens (Lindl.) Hochr. 59 H. cannabinus var. verrucosus (Guill. et Perr.) Garcke 66 H. cannabinus var. viridis A. & G. Howard 66 H. cannabinus var. vulgaris A. & G. Howard 66 H. comoensis A. Chev. 73 H. congestiflorus Hochr. 73 H. cordofanus Turcz. 68 H. costatus A. Rich. 48 H. cuanzensis Exell & Mendonsa 71 H. decaisneanus Schimp. 63 H. diversifolius Jacq. 63 H. diversifolius Verdoorn & Collett 60 H. diversifolius var. angustilobus Hauman 63, 64 H. diversifolius var. angustilobus P. Maquet 64 H. diversifolius subsp. diversifolius 63 H. diversifolius subsp. genuinus Hochr. 63 H. diversifolius var. partitus Hochr. 63 H. diversifolius subsp. rivularis (Bremek. & Oberm.) Exell 64 H. diversifolius var. subdiversifolius Hochr. 61 H. diversifolius var. witteanus Cufod. 63 H. diversifolius var. witteanus Hochr. 61 H. dongolensis Del. 73 H. eetveldeanus de Wild. & T. Durand 69 H. eetveldeanus var. asperatus de Wild. 59 H. elongatifolius Hochr. 72 H. ficulneus Cav. 63 H. flavo-roseus Baker f. 57 W./ri«iiUlbr. 73 H. furcatus Bahadur, Dayal & Raturi 73 H. furcatus Berhaut 55 H. furcatus Craib 51, 56 H. furcatus Garcke 7 1 H. furcatus Harv. 53 H. furcatus Hutch. & Dalziel 53, 55 H. furcatus Mast. 71 H. furcatus Moritzi 59 H. furcatus Mullend. 59 H. furcatus Roxb. 55 H. furcatus Willd. 56 H. furcatus var. microcarpus Mattei 75 H. furcellatoides Bahadur, Dayal & Raturi 53, 75 H. furcellatoides Hochr. 53 H. furcellatus Desr. 50 H. gilletii de Wild. 69 H. gilletii subsp. gilletii 69 H. gilletii subsp. hiernianus (Exell & Mendonc.a) F.D. Wilson 71 H. gilletii subsp. lundaensis (Baker f.) F.D. Wilson 71 H. goossensii (Hauman) F.D. Wilson 56 H. greenwayi Baker f. 62 H. greenwayi Cufod. 62 H. greenwayi var. megensis J.P. Lebrun 62 H. hamatus E. Mey. 53 H. heterophyllus Vent. 77 H. hiernianus Exell & Mendonc,a 7 1 H. hispidissimus Bahadur, Dayal & Raturi 75 H. hispidissimus A. Chev. 75 H. hispidissimus Griff. 55 H. hoshiarpurensis T.K. Paul & M.P. Nayar 75 H. imerinensis Ulbr. 75 H. keilii Ulbr. 75 H. lancibracteatus de Wild. & T. Durand 61 H. lindleyi Wall. 59 H. lunarifolius Willd. 73 H. lundaensis Baker f. 7 1 H. macularis E. Mey. ex Harv. 63 H. malangensis Baker f. 68 H. mastersianus Hiern 71 H. masuianus de Wild. & T. Durand 65 H. mechowii Exell & Mendonc.a 59 H. mechowii Garcke 6 1 H. meeusei Exell 60 H. moxicoensis Baker f. 7 1 H. nigricaulis Baker f. 60 H. noldeae Baker f. 59 H. pachmarhicus Haines 72 H. paludosus Merrill 63 H. panduriformis Burm.f. 61, 73 H. paolii Mattei 77 H. partitus (Hochr.) F.D. Wilson 63 H. parvilobus F.D. Wilson 67 H. persificolius Eckl. & Zeyh. 77 H. poggei Giirke ex Engl. 69 H. procerus Roxb. ex Wall. 77 H. radiatus Cav. 59 H. radiatus var. lindleyi (Wall.) Kurz 59 H. reekmansii F.D. Wilson 64 H. rivularis Bremek. & Oberm. 64 H. rostellatus Bahadur, Dayal & Raturi 55, 75 H. rostellatus Guill. & Perr. 53 H. rostellatus Keay 53 H. rostellatus var. congolanus Hauman 57 H. rostellatus var. goossensii Hauman 56 H. rostellatus var. sudanensis Hauman 57 H. sabdariffa L. 65 H. sabdariffa Mendonc,a & Torre 60 H. sabdariffa subsp. cannabinus (L.) G. Punigrahi & S.K. Murti 66 H. sabdariffa var. albus A. & G. Howard 65 H. sabdariffa var. altissima Wester 65 H. sabdariffa var. bhaghalpuriensis A. & G. Howard 65 H. sabdariffa var. 8 L. 66 H. sabdariffa var. intermedius A. & G. Howard 65 H. sabdariffa var. ruber A. & G. Howard 65 H. saxicola Ulbr. 77 H. scaber Lam. 63 H. scotellii Baker f. 73 H. scotellii Keay 72 H. sineaculeatus F.D. Wilson 72 H. sparseaculeatus Baker f. 62 H. stenophyllus var. solandroketmia (Hochr.) Hochr. 75 H. subdiversifolius Hochr. 61 H. sudanensis Hochr. 57 H. sudanensis var. genuinus Hochr. 57 H. sudanensis var. glabrescens Hochr. 57 H. sudanensis var. glabrescens forma grandiflorus Hochr. 57 H. sudanensis var. glabrescens forma minoriflorus Hochr. 57 H. surattensis L. 51 H. surattensis forma immaculata Kurz ex Rakshit & Kundu 51 H. surattensis var. eetveldeanus (de Wild. & T. Durand) Hochr. 69 H. surattensis var. furcatus (Willd.) Hochr. 56, 69 H. surattensis var. genuinus Hochr. 5 1 H. surattensis var. mastersianus (Hiem) Hochr. 71, 72 H. surattensis var. rostellatus (Guill. & Perr.) Hochr. 55 H. surattensis var. villosus Backer 5 1 H. surattensis var. villosus Hochr. 5 1 H. surattensis var. villosus forma concolor Backer 5 1 H. torrei Baker f. 59 H. trichospermoides Hochr. 77 H. uncinellus DC. 48 H. unidens Lindl. 59 H. vanderystii de Wild. 68 H. verrucosus Guill. & Perr. 66 H. verrucosus var. punctatus A. Rich. 68 Narinam poulli Rheede 55 Sabdariffa Kostel. 50 [section] Cannabini Small 50 section Furcaria DC. emend. Hochr. 49 [section] Furcellati Small 50 section Sabdariffa DC. 50 series Friesia Ulbr. 50 series Furcaria cannabina Ulbr. 50 series Furcaria diversifolia Ulbr. 50 series Furcaria furcellata Ulbr. 50 series Furcaria sabdariffa Ulbr. 50 series Saxicolae Ulbr. 50 subsection Furcaria simplicia Hochr. 50 subsection Furcaria typica Hochr. 50 Bulletin of The Natural History Museum Botany Series Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Volume 22 No. 1 An account of southern Australian species of Lithophyllum (Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. Campbell. 1992. Pp. 1-107, 63 figs. £41.25 No. 2 Palynological evidence for the generic delimitation of Sechium (Cucurbitaceae) and its allies. J.L. 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Mounting techniques for the preservation and analysis of diatoms. S.J. Russell. 1993. Pp. 51-54. 1 fig. £43.25 No. 2 New taxa of Gentiana (Gentianaceae) from Western China and the Himalayan region. T.-N. Ho and S.-W. Liu. 1993. Pp. 55- 60. 2 figs. New combinations, names and taxonomic notes on Gentianella (Gentianaceae) from South America and New Zealand. T.-N. Ho and S.-W. Liu. 1993. Pp. 61-66. Studies in Hypericum: validation of new names. N.K.B. Robson. 1993. Pp. 67-70. Generic monograph of the Asteraceae-Anthemideae. K. Bremer and C.J. Humphries. 1993. Pp. 71-177. 12 figs. £43.25 Volume 24 No. 1 Pre-Linnaean references for the Macaronesian flora found in Leonard Plukenet's works and collections. J. Francisco-Ortega, A. Santos-Guerra and C.E. Jarvis. Pp. 1-34. Studies on the lichen genus Sticta (Schreber) Ach.: II. Typification of taxa from Swartz's Prodromus of 1788. D.J. Galloway. Pp. 35^8. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 4. Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F. Prud'homme van Reine and W.J. Woelkerling. Pp. 49-90. Studies on the Cretan flora 3. Additions to the flora of Karpathos. N.J. Turland and L. Chilton. Pp. 91-100. £43.25 No. 2 Observations on the benthic marine algal flora of South Georgia: a floristic and ecological analysis. D.M. John, P.J.A. Pugh and I. Tittley. Pp. 101-1 14. Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical species (excluding Australia). D.J. Galloway. Pp. 1 15-160. Morphology and ecology of seedlings, fruits and seeds of Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp. 161-172. A study of Bixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character. R.E. Dempsey and N.C. Garwood. Pp. 1 73- 1 80. £43.40 Volume 25 No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. 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Rhodophyta (Florideae) 6. Genera [Q] R-Z, and an update of current names for non- geniculate Corallinales. W.J. Woelkerling, G.W. Lawson, J.H. Price, D.M. John and W.F. Prud'homme van Reine. Pp. 115- 150. INTENTS ••T^ 1 The moss family Calymperaceae (Musci) in the Philippines LT. Ellis 47 Revision of Hibiscus section Furcaria (Malvaceae) in Africa and Asia F.D. Wilson jral History BOTANY SERIES Vol.29, No. 1, June 1999 luseur