ISSN 0968-044 Bulletin of The Natural History Museum THE NATURAL HISTORY 22 KOV 2000 Q6NEKAI LIBRARY THE NATURAL HISTORY MUSEUM VOLUME 30 NUMBER 2 30 NOVEMBER 2000 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 R. Bateman 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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The Natural History Museum, 2000 Botany Series ISSN 0968-0446 Vol. 30, No. 2, pp. 33-130 The Natural History Museum Cromwell Road London SW7 5BD Issued 30 November 2000 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.) 30(2): 33-79 Issued 30 November 2000 The genus Polystichum (Dryopteridaceae) in Africa . ./ " , \") JACOBUS P. ROUX National Botanical Institute, Compton Herbarium, Private Bag X7, Claremont 7735, South Africa 22 NOV CONTENTS igfgKAL LJ6RARV Introduction 33 Materials and methods 34 Terminology 34 Taxonomic treatment 35 Key to the African species of Polystichum (including Marion and Prince Edward Islands) 36 References 78 Systematic index 79 SYNOPSIS. Polystichum Roth is a fern genus of 1 60 to 200 species occurring throughout the temperate parts of the world and the montane tropics, but absent from the arid regions as well as the lowland tropics. Although floristic accounts of Polystichum exist for many parts of the world, the genus remains poorly understood and a taxonomic account for the group as a whole has never been undertaken. Polystichum is poorly represented in Africa (including the Marion Island group) with merely 16 species and one known hybrid having been recorded from the region. Most of the species are confined to sub-Saharan Africa, occurring mainly along the eastern mountain ranges and Mt. Cameroon in the west. In this review of the African Polystichum species detailed observations are presented especially on the paleae, as these structures were found to provide the best characters on which a subgeneric classification can be based. Diagnostic features and relationships, variation, and the distribution and ecology of each species are included. INTRODUCTION The genus Polystichum Roth consists of between 160 (Tryon & Tryon, 1982) and 200 (Daigobo, 1972) species. It occurs throughout the temperate parts of the world as well as the montane tropics, but is mostly absent from the lowland tropics. Within the range two distinct centres of diversity can be identified, namely a larger Asiatic centre with approximately 70 species and a tropical American centre with approximately 55 species. Both these regions fall within the mountainous tropics characterized by mild, moist climates that are hardly seasonal. They also correspond with the areas of highest species diversity for homosporous ferns (Tryon, 1985). Within the Dryopterideae Polystichum is most closely related to Arachniodes Blume, Cyrtomium C. Presl, Dryopteris Adans. and Phanerophlebia C. Presl. Cyrtomium and Phanerophlebia have often been included in Polystichum (Kramer, 1990), but it has subsequently been shown that both these genera have a closer affinity to Polystichum than to each other (Yatskievych, 1996). Polystichum in a strict sense is a natural and relatively homogene- ous group of plants characterized by 1 -pinnate to 3-pinnate laminae with acroscopically developed ultimate segments, anadromous free venation, uni- or biseriate circular sori positioned medially, termi- nally or near terminally on abbreviated or unabbreviated vein branches, and peltate indusia (a number of species are exindusiate). The receptacle appears to be nude in most species. Although Polystichum as a genus is easily recognized, many species are superficially very similar. The delimitation of species is further hampered by the fact that many, mostly common species are allopolyploids (Vida & Reichstein, 1975; Wagner, 1979). Also the frequent occurrence of Fl -hybrids in some groups obscures species limits. A formal subgeneric classification for the genus has thus far only been provided for the east Asian species by Tagawa (1940) and Daigobo (1972). Some of these sections have since been subdivided further by Zhang & Kung (1995, 1996a, b) and Kung & Zhang (1998) to make provision for some of the Chinese species. Since most species remain poorly known a phylogeny for the genus cannot be proposed. Although floristic accounts of the genus have been published for many parts of the world, some being very old and outdated, no single monographic treatment exists and most species remain poorly known. Some of the modern-day regional treatments provide no detailed observations that may suggest affinities. Within the study area floristic accounts for Polystichum are available for North Africa (Maire, 1952), West tropical Africa (Alston, 1959), Cameroon (Tardieu-Blot, 1964), Mozambique, Malawi, Zambia and Zimba- bwe (Schelpe, 1970), Rwanda, Burundi and Kivu (Democratic Republic of Congo) (Pichi Sermolli, 1985), southern Africa (Schelpe & Anthony, 1986) and Bioko (Benl, 1991). It is a well-known fact that Africa, when compared with other tropical parts of the world, supports a floristically impoverished vascular flora. This phenomenon is also reflected in the pteridophyte flora of the continent. The cause of this floristic poverty is ascribed to the isolation of Africa from the other continents since the mid- Cretaceous and the subsequent significant changes in the climate as a result of uplift, continental drift and aridification caused by extra- tropical glaciation. All these changes may well have resulted in a The Natural History Museum, 2000 34 J.P. ROUX progressive elimination of the once rich tropical and subtropical forests that existed towards the late Jurassic and the establishment of extensive deserts and semi-deserts by the early Pliocene (Coetzee, 1993). Within the study area two regions can be identified: an African and a sub-Antarctic region. The origin, composition and floristic affinities of these regions differ markedly. The African region is the largest and today there are three fundamentally different floras or biogeographical subregions which can be identified: a southern African flora, a tropical African flora and a North African flora. The flora of the southern African subregion is believed to have evolved gradually since the mid-Tertiary, derived partly from an ancient southern African temperate flora and partly from a tropical African forest flora (Goldblatt, 1978). Elements of the southern African flora currently extend into tropical Africa along the eastern escarpment. The southern Cape forests are believed to be impover- ished remnants of the tropical African forest flora (Coetzee & Muller, 1984). Also the tropical African flora is believed to be an impoverished remnant of a once much richer tropical rainforest flora that extended over a far greater area than it currently occupies. North Africa has also experienced significant changes in its climate and vegetation. During the Palaeocene the present Sahara desert was clothed by a rich tropical lowland rainforest that also covered part of Europe (Greenway, 1973; Raven & Axelrod, 1974), but by the Oligo-Miocene it was replaced by a subtropical woodland savanna (Axelrod & Raven, 1978). From the Pliocene a desert climate established itself in the major part of the Sahara (Quezel, 1978), serving as an effective barrier to migration from the south. The formation of glaciers on the high mountains during the Pleistocene permitted the establishment of circumboreal elements. Many of these elements are present in the North African flora since it is composed of relict elements of African origin as well as elements from Eurasia, not frequent in the present sub-Saharan flora. The mediterranean influence on the flora of North Africa justifies it being considered as a biogeographical subregion of its own. The Marion and Prince Edward Island group forms part of the sub- Antarctic region, a phytogeographical area completely different from the foregoing. This island group is of volcanic origin and is estimated to be 0.5 million years old (Verwoerd, 1971). Situated in the Southern Ocean some 1 800 km from Africa, its biota consists of taxa capable of long-distance dispersal and the ability to establish themselves in habitats not always favourable for plant growth. About two-thirds of the African pteridophy tes are limited in their occurrence to the continent (Kornas, 1993). The majority of these, however, are closely related to taxa in either tropical America and/ or southeast Asia. Pteridophytes of the sub-Sahara biogeographical region exhibit three discontinuous distribution patterns: an Ameri- can-African disjunction, an African-Madagascan disjunction and an African-Asian disjunction. Polystichum in Africa is largely confined to the Afromontane Phytochorion. White (1978) divided this montane archipelago into seven regional mountain systems. The North African Atlas moun- tain ranges are here added as an eighth. Although the sub-Saharan mountain ranges are sufficiently distinct, the systems are connected by a complex series of intermediate floras (Fig. 1 ). The Drakensberg system, with six Polystichum endemics, is the richest. This is also true for the angiosperms (White, 1978). The only other mountain systems with true endemics are the Imatongs-Usambara system with two endemics (P. kilimanjaricum & P. volkensii) and the Ethiopian system with one endemic (P. magnificum). Other African Polystichum species have wider distributions. Polystichum zambesiacum occurs in the Chimanimani, Uluguru-Mulanje, and Imatongs-Usambara mountain systems, whilst P. transvaalense and P. wilsonii are dis- tributed throughout seven mountain systems. The distribution of P. wilsonii, however, also extends along the Himalaya mountains to Bhutan, Japan and Taiwan. Polystichum luctuosum has an almost similar eastern distribution but is confined to the Drakensberg and Chimanimani mountain systems in Africa. Polystichum luctuosum and P. wilsonii also show a disjunct distribution with the Madagascan region. Based on observations taken from the systematic treatment, and judging from wide-ranging species, southern and tropical African Polystichum has a closer affinity with taxa from Asia than with those from the Americas. Polystichum in the North African phyto- geographical subregion shows a closer affinity with Polystichum from Europe than from Africa as P. aculeatum and P. setiferum are widespread in that region. Only P. marionense, endemic to Marion, Prince Edward and Crozet Islands, occurs in the sub-Antarctic phytogeographical region. MATERIALS AND METHODS This review is based on observations made during extensive field- work in southern Africa and on cultivated plants collected during these travels. The collections of several herbaria were also studied. These include: B, BM, BOL, BR, ETH, GRA, K, L, M, MAL, NBG, NH, NU, P, PRE, RAB, SAM, SRGH, WAG (abbreviations follow Holmgren et al., 1990) and the private herbarium of Prof. R.E.G. Pichi Sermolli (PIC.SERM.). Palea and indusium observations were made by removing a small number of these structures from selected specimens. These were cleaned and cleared in diluted household bleach, after which they were semi-permanently mounted in glycerine and the cover slips sealed with Entellan. Observations were made with an Olympus CH-2 light-microscope fitted with a drawing tube. The collections studied are all listed under 'Material examined'. These are arranged alphabetically according to country of origin. South African (including Lesotho and Swaziland) collections are further arranged according to the quarter-degree square-grid system (Edwards & Leistner, 1971). In this system each one-degree square is known by a standardized name, derived from a town or other feature of importance in the square. Each one-degree square is divided into four half-degree squares (30' x 30'), numbered A, B, C and D from left to right and top to bottom. Each half-degree square is again subdivided into quarter-degree squares (15' x 15'), again numbered A, B, C and D. By using these co-ordinates a geographical area can immediately be identified. Unless cited otherwise, the chromosome numbers provided here are based on the author's own observations and will be published elsewhere. Terminology This study is principally based on a detailed comparative morpho- logical analysis of the sporophyte, where palea structure proved to be most informative in suggesting species groups. The terms used to describe the apex or the apical cell of the paleae are defined as: - apex flagelliform: the apex of the palea terminates in a uniseriate series of slender cells. - subulate cell: the apical cell is less than 0.4 mm long and the apex is usually blunt. - acicular cell: the apical cell is slender, straight, more than 0.4 mm long, and the apex is usually sharp. POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 1 35 r V VIII. NORTH AFRICAN U-y IV. IMATONGS-USAMBARA \ P. kilimanjaricum VII. ORAKENSBERG '. drakensbergense . incongruum . macleae . monticola . pungens . transkeiense Fig. 1 Distribution of Polystichum in Africa. filiform cell: the apical cell is slender, twisted, and more than 0.6 mm long. thin-walled cell: these cells can vary considerably in size and the cell wall is conspicuously thinner than that of the surrounding cells, and is therefore often lost. In dry material the cellular contents are usually yellowish in colour and appear crystalline. TAXONOMIC TREATMENT Polystichum Roth, Tent.fl. Germ. 3: 31, 69 (1799). Type species: Polystichum lonchitis (L.) Roth (= Polypodium lonchitis L.). Hypopeltis Michx., Fl. bor.-amer. 2: 266 (1803). Type species: Hypopeltis lobulata Bory (= Polystichum aculeatum (L.) Roth). Plecosorus Fee, Mem.foug. 5: 1 50 ( 1 852). Type species: Plecosorus mexicanus Fee, nom. superfl. for Cheilanthes speciosissima Kunze (= Polystichum speciosissimum (Kunze) R.M. Tryon & A.F. Tryon). Sorolepidium H. Christ in Bot. Gaz. 51: 350 (1911). Type species: Sorolepidium glaciale (H. Christ) H. Christ (= Polystichum glaciate H. Christ). Hemesteum H. Lev., Fl. Kouy-Tcheou: 450, 496 (1915), non Newm. (1851). Type species: several Polystichum species are listed. Aetopteron House inAmer. Fern J. 10: 88 (1920), nom. nud. Papuapteris C. Chr. in Brittonia 2: 300 (1937). Type species: Papuapteris linearis C. Chr. (= Polystichum lineare (C. Chr.) Copel.). Acropelta Nakai in Bull. Natl. Sci. Mus. Tokyo 33: 5 (1953). Type species: Acropelta omeiensis (C. Chr.) Nakai (= Polystichum omeiense C. Chr.). Plants terrestrial, or epilithic, rarely low-level epiphytes. Rhizome erect to suberect and mostly unbranched, or creeping, or decumbent and branched; rarely stoloniferous; dictyostelic; set with roots, closely to widely spaced persistent stipe bases, and paleae. Fronds monomorphic, caespitose or closely to widely spaced, to 1 .8 m long: stipe proximally convex adaxially, becoming slightly to deeply sulcate distally; with two larger near-circular vascular bundles dorso-laterally, ventrally with three to five smaller circular vascular bundles; initially moderately to densely paleated, becoming near glabrous later, the paleae often appearing heteromorphous, variable: lamina 1 -pinnate to 3-pinnate, anadromous, sometimes bearing 1 to several paleated proliferous buds adaxially along the rachis near the 36 J.P. ROUX lamina apex: rachis adaxially shallowly to deeply sulcate, the sulcus Distal pinnae never folded ventrally along the rachis; spores aborted proximally not open to sulci of lower order axes, moderately to 9. P. x saltum densely paleated; paleae variable: pinnae short-stalked, opposite to 10 Rhizome to 10 mm in diameter; sori exindusiate 1 1 alternate, closely to widely spaced, often imbricate, simple to 2- pinnate, acroscopically auricled: pinna-rachis adaxially sulcate, Rhizome more than 10 mm in diameter; sori indusiate 12 open to sulci of costae, sparsely to densely paleated; paleae variable: u Rhizome to 5 mm in diameter; stipe and rachis paleae with long pinnules proximally mostly short-stalked, opposite to alternate, flagelliform outgrowths along the margin 10. P. marionense closely to widely spaced, often imbricate, the proximal acroscopic _ Rhizome to , Q mm - n diameter; stjpe ^ fachis pa , eae ^ Qr without pinnule mostly longer than the next in 2-pmnate or more dissected thin-walled cells along the margin 1 1 . P. transkeiense species, herbaceous to firmly coriaceous, inaequilateral, ovate to ovate-rhomboid or trullate, often somewhat falcate, mostly 12 Lamina with a proliferous bud along the rachis near the apex acroscopically auricled in 2-pinnate or more dissected species, 12. P. magm icum lobate, dentate or serrate, sharp-tipped or aristate; variously paleated. Lamina without proliferous buds along the rachis 13 Venation free, pinnately branched, anadromous, terminating near or ._..,. , . . 13 Rhizome paleae conspicuously rugose, often with a few long filiform at the margin when sterile, immersed or raised. Son circular, essen- outgrowths along the margin; smaller stipe, rachis and pinna-rachis tially unisenate, borne medially on unabbreviated vein branches, or paleae basally with short and/or , ong flliform outgrowths along the near or at a vein ending of mostly anadromous vein branches: margin 13. P. zambesiacum sporangium with 8-(13)-30 indurated annulus cells; stalk with glandular cells or eglandular, 3-seriate below capsule: indusium Rhizome paleae not conspicuously rugose, mostly with short straight or curved marginal outgrowths; smaller stipe, rachis and pinna-rachis absent or present, peltate, mostly persistent, the margin variously ... f, , ., J h J paleae basally without short and/or long unisenate outgrowths along the sculptured, with or without gland-like cells. Spores monolete, the margin 14 laesura 2 / 3 to 3 / 4 of the spore length, the perispore irregularly folded, mostly somewhat spinulose, often perforate. Chromosome number 14 Proximal acroscopic pinnule to 22 mm long; larger stipe base paleae n=41, 82, 164; 2n=82, 164, 328; apogamous 123, 246. often bicolorous r Proximal acroscopic pinnule usually more than 22 mm long; larger stipe Key tO the African Species Of PolystictlUm (includ- base paleae never bicolorous 16 ing Marion and Prince Edward Islands) 1 5 Rhizome short-decumbent with crowded stipe bases, closely branched; apogamous (32 spores per sporangium) .... 14. P. monticola 1 Lamina 1 -pinnate (rarely 1 -pinnate-pmnatifid) l.P. macleae - , - Rhizome decumbent, stoloniferous; sexual (64 spores per sporangium) - Lamina 2-pinnate to 3-pmnate 2 15. P. dracomontanum 2 Rhizome short, erect to suberect, mostly unbranched 3 16 Pinnules maequilaterally ovate to narrowly trullate, to 60 x 13 mm; Rhizome short-decumbent to widely creeping, mostly branched .... 10 sporangium stalk glandular or eglandular; indusium with or without unicellular thin-walled cells along the margin 16. P. incongruum 3 Larger rhizome and stipe base paleae with long unisenate hairs along the margin and superficially 2. P. luctuosum Pinnules inaequilaterally ovate, ovate-oblong, ovate-rhomboid or trullate, to 50 x 1 9 mm; sporangium stalk and indusium always eglandular Larger rhizome and stipe base paleae without long unisenate hairs along ,7 p Dun o ens the margin and superficially 4 4 Lamina with 1-3 paleated proliferous buds along the rachis near the 1. Polystichum macleae (Baker) Diels in Engl. & Prantl, Nat. lamina apex 5 Pflanzenfam. 1(4): 190(1902), as macleanii. Type: South Africa, - Lamina without proliferous buds along the rachis .... ... 6 in convallibus humidis - Drakensbergen prope 'Pilgrim's Rest Gold Fields', McLea 34 sub Bolus 3030 (Kl-lectotype, desig- 5 Pinnule margins obtusely serrate to crenate, never aristate natedby Schelpe & Anthony (1986); BOL!, SAM!-isolectotypes). 3. P. volkensii p- ^ Pinnule auricle and apex aristate 4. P. kilimanjaricum ... . \ti i/i /ioo/:\ Aspidium macleae Baker in Hook.f., Icon, pi.: t. 1654 (1886), as 6 Apices of paleae terminating in a short subulate cell or a small thin- macleaii. walled cell, the margins set with short straight and/or angular outgrowths 7 Plants terrestrial, epilithic, or rarely epiphytic. Rhizome decumbent, to 200 mm long x 20 mm in diameter, densely set with roots, Apices of paleae always terminating in an acicular cell, the margins set per sistent stipe bases, and paleae; paleae ferrugineous, membranous with long straight and/or long twisted emarginate to forked outgrowths tQ chartaceouSi narrowly ovate or lanceolate , to 7 x 2 mm. Fronds caespitose, to 7 per plant, arcuate, to 1.47 m long: stipe proximally 7 Stipe and rachis moderately paleated; paleae mostly flat or irregularly castaneous, stramineous distally, adaxially sulcate, to 670 mm long folded 5. P. aculeatum x g mm j n diameter, proximally densely paleated; larger paleae Stipe and rachis densely paleated; paleae mostly helically twisted broadly attached, concolorous or bicolorous, the concolorous paleae 6. P. setiferum ferrugineous, the bicolorous paleae centrally dark brown or black, 8 Conspicuously larger paleae mostly confined to the stipe, rugose ... ovate ' narrowl y ovate ' or narrowl y oblon g' cordate ' entire or with 7 P. transvaalense short and/or long marginal outgrowths proximally, the apex flagelliform, terminating in a long filiform cell or an oblong thin- :onspicuously larger stipe paleae extending to the rachis, never rugose walled cell to 37 x 8 mm; smaller paleae concolorous, ferrugineous, membranous, narrowly ovate, lanceolate, narrowly triangular, nar- 9 Distal pinnae folded ventrally along the rachis (conduplicate); spores rowly oblong to acicular, cordate, cordate-imbricate, or short-stalked, not aborted 8. P. wilsonii proximally erose and/or with long twisted, filiform outgrowths, the POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 37 apex flagelliform, terminating in a filiform cell or an oblong thin- walled cell: lamina 1 -pinnate (rarely 1-pinnate-pinnatifid), oblong to narrowly elliptic, with up to 37 free pinna pairs, to 840 mm long, the proximal pinnae slightly reduced, usually deflexed: rachis stramineous, adaxially sulcate, moderately paleated; paleae short- stalked, ferrugineous, membranous to chartaceous, narrowly ovate, narrowly lanceolate, or oblong to subulate, cordate, cordate-imbri- cate, proximally usually erose and/or with a few short or long twisted marginal outgrowths, the apex long-attenuate to flagelliform, terminating in an acicular cell or an oblong thin-walled cell, to 3 mm long: pinnae firmly herbaceous, olive-green adaxially, paler abaxially, generally not overlapping, short-stalked, narrowly ob- long-attenuate, straight, auriculate acroscopically, the base unequally broad-cuneate to truncate, doubly serrate, to 168 mm long x 16 mm wide, the acroscopic auricle on proximal pinnae often free, ovate to trullate, to 24 x 22 mm; costa adaxially sulcate, sparsely paleated, the paleae taeniform, sessile or short-stalked, entire, the apex ter- minating in an acicular cell or an oblong thin-walled cell, to 3 mm long, abaxially moderately to densely paleated, the paleae ferrugin- eous, membranous, narrowly lanceolate to narrowly trullate, often bullate, cordate to cordate-imbricate, the margin proximally with short and/or long irregular outgrowths, entire distally, the apex terminating in an acicular cell or an oblong thin-walled cell, to 2.4 mm long. Venation raised. Sori circular, to 1.5 mm in diameter, variable in size, those closest to the costa largest, discrete at matu- rity, medial to inframedial on unabbreviated vein branches: sporangium with 12-(16)-28 indurated annulus cells; stalk eglandular: indusium brown, persistent, peltate, circular to irregular, repand to erose, often with flabellate central processes, the maxi- mum radius 0.29-(0.49)-0.7 mm. Spores 64 per sporangium, brown, the perispore folded to form a sparse reticulum of low compressed ridges, variously granulate, verruculate to echinulate, closely perfo- rated, the exospore 40-(51.31)-66 x 28-(37.89)-48 urn. Chromosome number 2n=164. MATERIAL EXAMINED SOUTH AFRICA. 2330 (Tzaneen): Tzaneen, Woodbush Forest Reserve (CC), Balsinhas 2166 (PRE); Wolkberg, Agatha Forest Reserve, 1500 m, Mutter 264 (PRE); Woodbush, Van Jaarsveld6\ 10 (BOL). 2430 (Pilgrim's Rest): Haffenden Heights, Zoutpansberg (AA), Junod 4069 (P, PRE); Mariepskop (DB), Van der Schijf 4305, (B, NU, PRE), 5597 (PRE); Mariepskop summit, 1800m, Van der Schijf '4861 (PRE); Mariepskop, below radar station, Krynauw 786 (PRE); Graskop, Erasmus Kop, Hardcastle 59 (PRE); Mariepskop, Schweickerdt 4305 (BOL); Ohrigstad Nature Reserve, 6000 ft (DC), Jacobsen 1556 (PRE); Pilgrim's Rest, Mount Sheba Nature Reserve, Roux 2555 (NBG); Mount Sheba Nature Reserve, Jacobsen 4436 (PRE); Mount Sheba Nature Reserve, Crouch 633 (NU); Graskop, Cigar Rock (DD), Rauh & Schlieben 9744 (PRE); Graskop, Kowyn's Pass, Rauh & Schlieben 9725 (PRE); Graskop, Driekop Gorge, Wager 173 (PRE); Pil- grim's Rest, Rogers 14925, 14927 (PRE); Blyde Bosboustasie, Bredenkamp s.n. (PRE); Pilgrim's Rest, MacLea 170 (PRE); Kowyn's Pass, Schelpe 1641 (BOL, NH, NU), 6092 (BOL); Graskop, Fairyland, Roux 2548, 2549 (NBG); Graskop, The Pinnacle, 4500 ft, Braithwaite 207 (BOL). 2530 (Lydenburg): Lydenburg, Hartbeesvlakte (BA), Kluge 2039, 2333 (PRE); Lydenburg, Hartbeesvlakte, 1 960 m, Mohle 288 (PRE); Pilgrim's Rest, Mount Anderson, Smuts 38 (PRE); Sabie, forest at Tweefontein (BB), Wager 53 (PRE); Sabie/ Lydenburg road, Roux 2242, 2561 (NBG); Witklip Staatsbos (BD), Kluge 806 (PRE); Belfast (CA), Wager s.n. (PRE); Kaapse Hoop (DB), Van Jaarsveld2Q8%a (NBG, PRE), 3376 (BOL); Kaapse Hoop, Wager 73 (BOL), 1496c (PRE). 2531 (Komatipoort): Barberton, Tiger Creek, 4500 ft (CC), Thorncroft96 (BR, P, Herb. PIC.SERM., PRE); Barberton, Maid of the Mist, Thorncroft 50 (P, PRE), 68 (NBG, P, PRE); 17 miles SE of Barberton towards Havelock, 5000 ft, Schelpe 41 15 (BOL, PRE); W. of Havelock, Songimvelo Game Reserve, on farm Josefsdal, 1640 m, Kunitz & Otto 15 (J, PRE). SWAZILAND. 2531 (Komatipoort): New Havelock, 12 miles from Havelock (CC), Schutte4 (BOL); Havelock Mine, Dyer51 (NU). 2632 (Bela Vista): Mbabane, Ngwenya Mountain (AA), Compton 31405 (NBG) WITHOUT EXACT LOCALITY: loco incerto, Bolus s.n. (PRE); South Africa, Wood s.n. (NU). The change of the specific epithet macleaii to macleae is in concord- ance with Article 60.11 (Recommendation 60C.l.a) of the International Code of Botanical Nomenclature (Greuter et al., 1994). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum macleae is the only 1 -pinnate (rarely 1-pinnate-pinnatifid) species in Africa and is quite similar to P. kalambatitrense Tardieu from Madagascar. Baker (1886) considered P. macleae to be related to P. munitum (Kaulf.) C. Presl from North America and P. falcinellum (Sw.) C. Presl from Madeira. This assumption was probably based on the 1- pinnate lamina morphology in these taxa. Both these species, however, have ciliated indusia. Also, the palea morphology of both these species differs from that in P. macleae. In P. macleae the marginal outgrowths of the lamina paleae are pluricellular whilst those of P. munitum and P. falcinellum are unicellular. VARIATION. Polystichum macleae shows considerable variation in the number of indurated annulus cells per sporangium and in indu- sium and pinna morphology. The number of indurated annulus cells per sporangium ranges from 1 2 to 28. The mean number of indurated annulus cells per sporangium is 16.67 (n=650, SD=0. 14) taken from 13 populations throughout the species' distribution. Some popul- ations have a larger number of indurated annulus cells than others. Although no definite correlation could be made between habitat and the number of indurated annulus cells, plants collected from an exposed streambank on the Hartbeesvlakte near Lydenburg [Kluge 2039 (PRE)] have a number of indurated annulus cells that ranges from 19 to 28 (x=23.86, SD=1.91, n=50), which is significantly higher than for a plant growing in a forest habitat [Thorncroft 68 (NBG)], where the number of cells ranges between 12 and 16 (x=13.28, SD=0.75, n=50). Intermediates between these extremes do occur. Indusia are mostly simple, but on some plants they may bear one or more small wings, whilst on others they may bear numerous flabellate central processes. The margins vary from repand to erose. A 1-pinnate-pinnatifid form of Polystichum macleae has been recorded from Mpumalanga with the central pinnae bearing up to 1 1 nearly free pinnule pairs. Pinnules are inaequilaterally narrowly trullate to oblong-attenuate in outline with the margins obtusely serrate. In the distal pinnae the margins are merely lobed midway to the costa. Sori are uniseriate on either side of the costa and are borne inframedially. The size of the acroscopic auricle varies considerably, and on the proximal pinnae it is often detached from the rest of the pinna. The auricle sometimes overlaps with the pinna directly above. Pinna margins are generally obtusely serrated or doubly serrated but rarely the margins are also deeply lobed and serrated. DISTRIBUTION AND ECOLOGY. Polystichum macleae is confined to the Drakensberg Escarpment and Wolkberg in the Mpumalanga province of South Africa and the northern parts of Swaziland, occurring at elevations ranging between 1350 and 1960 m. The species is largely confined to forests where it grows on banks above streams, in forest margins, among rocks and often as a low-level epiphyte. Plants often form large stands in deep shade, but rarely also occur in exposed habitats. 2. Polystichum luctuosum (Kunze) T. Moore, Ind.fil.: 95 (1858). Type as for Aspidium luctuosum Kunze. Fig. 3. 38 J.P. ROUX 10 mm Fig. 2 Polystichum macleae. A, proximal part of lamina; B, rhizome; C, section of abaxial surface of fertile pinna. A & B, drawn from Roux 2548 (NBG); C, drawn from Roux 2242 (NBG). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 39 Aspidium luctuosum Kunze in Linnaea 10: 548 (1836). Type: In monte Katriviersberg in sylvis, Ecklon s.n. (LZf-syntype); ad fontes fl. Katrivier prope Philipstown, in sylvis montium, Ecklon s.n. (LZt-syntype). Aspidium tsus-simense Hook., Sp. fil. 4: 16, t. 220 (1862). Type: Island of Tsus Sima, in the Straits of Korea, Wilford s.n. (K- holotype, 2 sheets; NBG! -photograph). Polystichum tsus-simense (Hook.) J. Sm., Hist, fil.: 219 (1875). Polystichum lobatum var. luctuosum (Kunze) H. Christ in Ber. Schweiz. Bot. Ges. 3: 34 (1893). Plants terrestrial, epilithic, or rarely low-level epiphytes. Rhizome short, erect to suberect, to 10 mm in diameter, densely set with roots, persistent stipe bases, and paleae; larger paleae broadly attached, castaneous, chartaceous, ovate, narrowly ovate, or lanceolate, cor- date, with long twisted uniseriate, gland-tipped hairs on the apical margin and surface, the apex flagelliform, terminating in an oblong thin-walled cell, to 10.5 x 3.3 mm; smaller paleae short-stalked, narrowly triangular to subulate, cordate, the margins proximally with numerous long and twisted uniseriate hairs, distally with widely spaced apically and basally directed marginal outgrowths that become smaller apically, the apex flagelliform, terminating in a small thin-walled cell. Fronds crowded, caespitose, 7-16 per plant, suberect to arching, to 0.93 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 450 mm long x 5 mm in diameter, densely paleated; proximal paleae broadly attached, castaneous, chartaceous, ovate, cordate, proximally entire or with a few short and/or long uniseriate hairs, distally with numerous multicellular hairs as for rhizome paleae; distal paleae short-stalked, narrowly oblong, narrowly triangular or subulate, cordate to hastate, the margins bearing a few long and/or short multicellular hairs proximally, distally with widely and irregularly spaced outgrowths reduced in size and number towards apex, the apex flagelliform, terminating in a small thin-walled cell, to 15 x 1.5 mm: lamina 2- pinnate to 2-pinnate-pinnatifid, with up to 25 free pinna pairs, to 480 mm long, firmly herbaceous to coriaceous, olive-green adaxially, paler abaxially, narrowly ovate to ovate, the proximal pinnae slightly reduced, often somewhat deflexed: rachis stramineous, adaxially sulcate, densely paleated; paleae short-stalked, dark brown to black, glossy, chartaceous to crustaceous, narrowly triangular to subulate, cordate to hastate, the auricles usually bearing long and twisted multicellular and uniseriate hairs some of which terminate in a thin- walled cell, the margins either distally with short, widely and irregularly spaced outgrowths that reduce in size and number to- wards the apex, or more or less entire in smaller paleae, to 7 mm long: pinnae short-stalked, 1 -pinnate to 1-pinnate-pinnatifid, with up to 12 free pinnule pairs, narrowly lanceolate, proximally widely spaced, distally often somewhat overlapping, to 173 mm long: pinna-rachis stramineous, adaxially sulcate, densely set with paleae similar to but less complex than those on the rachis: pinnules widely spaced to overlapping, the proximal acroscopic pinnule the largest, often significantly longer than the next, up to 40 mm long and 1 2 mm wide, inaequilateral, narrowly trullate to rhomboid, basiscopically cuneate, acroscopically truncate and auricled, often somewhat falcate, lobate-serrate, aristate; proximal pinnules short-stalked, often acroscopically incised to or nearly to the costa; costa adaxially sulcate, glabrous, abaxially sparsely paleated, the paleae castaneous, chartaceous, narrowly triangular-hastate to subulate-hastate, cor- date to cordate-imbricate, proximally with long and/or short filiform outgrowths often terminating in a thin-walled cell, the apex always terminating in a small thin-walled cell, to 0.3 mm long. Venation immersed. Sori circular, c. 1.2 mm in diameter, terminal or nearly terminal on abbreviated vein branches, essentially uniseriate: spor- angium with 10-(13)-19 indurated annulus cells; stalk eglandular: indusium peltate, circular, entire, repand or crenulate, persistent, brown, pale brown and often dark centred before drying, cupulate when dry, the maximum radius 0.5-(0.73)-0.95 mm. Spores 32 per sporangium, brown, the perispore unevenly folded to form narrow and broad reticulate ridges, the ridges and areas between ridges echinulate, spiculate or verruculate, the exospore 30-(38.84)-50 x 22-(28.2)-36 urn. Chromosome number 2n=123, apogamous. MATERIAL EXAMINED LESOTHO. 2828 (Bethlehem): Leribe (CC), Phillips s.n. (SAM). 2927 (Maseru): Roma Valley (BC), Schmitz 6963 (PRE); Roma, Ruch 1909 A- only (PRE). SOUTH AFRICA. 2430 (Pilgrim's Rest): The Downs (AA), Junod s.n. & 4044 (BR, P, PRE); Mt Sheba (DC), Kluge 2320 (NBG, PRE); Blyde Forest Reserve (DD), Jacobsen 4365, 4376 (PRE); Driekop Gorge, Graskop, Wager 178 (PRE); Pilgrim's Rest, Ponies Krantz, Braithwaite 229 (BOL); Sabie, just outside Ceylon Forest Reserve, Braithwaite 135 (BOL). 2530 (Lydenburg): Lydenburg District, Spitzkop, Wilms 1781 (B, BM); Sabie Gorge (BB), Wager s.n. (PRE); Lone Creek Falls, Sabie, Burrows 1342 (BOL); Sabie, Rogers 20379 (PRE); Sudwala Caves, 1500 m (BC), Kluge 2463 (NBG, PRE); Lydenburg, Buffelskloof Nature Reserve, Burrows 3860 (GRA); between Machadodorp and Badplaas (CD), Steel 242 (PRE). 2531 (Komatipoort): Rimers Creek, Barberton (CC), Thorncroft 35 (P); Baberton, Pott-Leendertz 5574 (PRE); Lomati Falls behind Barberton, Wager 154 (PRE); Barberton, Thorncroft 36, 104c (PRE); creeks near Barberton, Thorncroft 2475 (L). 2729 (Volksrust): Newcastle, Nkandu Reserve, 4900 ft (DD), Smith 64 (NU). 2730 (Vryburg): road to Luneburg (AD), Roux 2268 (NBG); Pongola Bush Nature Reserve, 1500 m (BC), Glen 2390 (PRE); Utrecht, Donkerhoek, 5500 ft, Devenish 1 144 (PRE); Hlobane, Mtola Forest, Johnstone 296 (NU). 2828 (Bethlehem): Farm Boschkloof (DB), Roux 1228 (NBG); Witsieshoek, Junod s.n. (P); Royal Natal National Park, Okell 60 (NU). 2829 (Harrismith): Van Reenen, 5000 ft (AD), Schlechter 6718 (B, BM, GRA, PRE, SAM); Van Reenens Pass, Rehmann 7204 (P); Van Reenen, 5000 ft, Lidey 42 (NU); Robinson's Bush, Oliviershoek Pass (CA), Schelpe 7967 (BOL); near Cathedral Peak, Box 3371 (BM). 2929 (Underberg): Giants Castle Nature Reserve (AB), Roux 2503 (NBG); Injasuti Nature Reserve, below Cataract Valley, Roux 2718 (NBG); Champagne Castle, Bayer 1444 (NU); Cathedral Peak, bank of Kweliquala River, 4700 ft, Schelpe P4 (NU); Cathedral Peak, 5700 ft, Killick 1155 (NU, PRE); Cathe- dral Peak, Rainbow Gorge, 5500 ft, Cowan 96 (NU); Cathedral Peak, 1550 m, Goetghebeur4552 (BR, PRE); Cathedral Peak, c. 5000 ft, MacGregor43 (NU); Cathkin Park, Howlett 53 (NH); Estcourt, Nolema Forest, 4200 ft, Edwards 2685 (NU, PRE); Cathkin Peak, Ndema Forest, 4400 ft, Hillary 106 & 107 (NU); Cathkin Park, Howlett 53 (NH); Injasuti area, 5000 ft, Esterhuysen 26034 (BOL, NBG, PRE); above Dalton Bridge, above Bushmans River, c. 4500 ft, Wright et al. 27 (NH, PRE); Mooi River, The Hoek, 4700 ft (BC), Bourquin 320 (NU); Polela District, Ndumduma, Glengariff (CB), Rennie 913, 940 (NU); Cobham Forest Station, Whale Rock, Hill 48 (GRA); Bulwer Mountain (DB), Van Jaarsveld 6468 (NBG, PRE); Bulwer (DD), Clarkson 111 (NH, NU); Bulwer, Sunset, 5200 ft, Rennie 546 (NU); Bulwer, Allsopp 839 (NU); near Bulwer, Schelpe P52 (NU). 2930 (Pietermaritzburg): Balgowan, farm Boschfontein, 4000 ft (AC), Fisher 638 (NH, NU), 642 (NH, NU); Lions River District, Dargle, Smook 624 (NU); Balgowan, Thomas 71 (NU); Balgowan, Devlin 62 (NU); Balgowan, 4000 ft, Lindahl 107 (NU); Balgowan District, Thienel 109 (NU); Balgowan, 3500 ft, Bernele 1 13 (NU); Balgowan, 3500 ft, Crookes 105 (NU); Nottingham Road, McClean 899 (NH, PRE); Nottingham Road, sine coll. NH-26790 (NH); Dargle, Griffin's Farm, 1500 m, Jones 20 (NH); Lions River, Dargle, Esterhuysen 26202 (BOL); Balgowan, Bosch Hoek, 1400 m, Moll 905 (BOL, NU, PRE); Lions River, Lions Bush Forest, Moll 829 (BOL, NU); Pietermaritzburg (CB), Tyson s.n. (PRE); Pietermaritzburg, c. 2200 ft, Carnegie 692 (NU); Pietermaritzburg, Blackridge, F.G.C. 692 (NU); Inanda, Wood s.n. (B). 3029 (Kokstad): Langewacht Forest Reserve near Kokstad, c. 1200 m (CB), De Joncheere s.n. (PRE); Mt Currie, Kokstad, Stephany 505 (BOL); Glen Hope, Jaconet & Jaconet 539 (BM). 3126 (Queenstown): Woodvale Forest, Gwatyn, 4200 ft (AA), Galpin 8203 (PRE). 3127 (Lady Frere): Engcobo (DB); McLoughlin 1022 (PRE); Engcobo, Flanagan 2781 (PRE). 3128 (Umtata): Maclear, farm Woodcliffs (AB), Roux 2479 (NBG); Engcobo, 40 J.P.ROUX 10 mm Fig. 3 Polystichum luctuosum. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2433 (NBG). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA Ku-Hlophekazi Forest (AC), Cawe 111 (BOL); Tsolo, Gxalibomvu Forest (AD), Cawe 660 (BOL); Tsolo, Bele Forest (BC), Cawe 731 (BOL); Nqadu Forest, Hatchings 39 (BOL); Mount Baziya, Baur644 (B). 3225 (Somerset East): in sylvis ad pedem montis Boschberg (DA), Barkley s.n. (GRA, P, SAM); in sylvis ad pedem montis Boschberg, MacOwan 1884 (B, BOL); Boschberg, Bolus 95 (BOL). 3226 (Fort Beaufort): Katberg Forest (BC), Holland s.n. (NBG); Katberg, Hutton s.n. (B, L); Katberg Forests, c. 3000 ft, Adams 142 (NU); Hogsback, Madonna and Child Falls (DB); Greatheads.n. (SRGH); Hogsback Forest, Auckland Kloof, Griffen x46 (PRE); Brambledene, Menziesberg, Acocks 11112 (PRE); Hogsback, Gibbs-Russel 3832 (PRE). 3227 (Stutterheim): Isidinge Forest (CA), Roux 1986 (NBG); Keiskamma Hoek, Gxulu Mountain, 5500 ft, Story 3509 (PRE); Kalogha Forest Station (CB), Roux 2433 (NBG); Pine Forest, along Amatola trail, Roux 2709 (NBG); Pirie Forest, Flanagan 1758 (PRE); Fort Cunningham, Roux 2427 (NBG); Pirie (CC), sine coll. s.n. (GRA); Amabele (DA), sine coll. s.n. (PRE); Pirie, Sim s.n. 505, 1727c (GRA, PRE); Komgha (DB), Flanagan s.n. (SAM). SWAZILAND. 2631 (Mbabane): Gobolo, c. 3500 ft (AC), Dlamini s.n. (NBG, NH, PRE); Stroma, c. 4000 ft, Campion 25822 (NBG, PRE). ZIMBABWE: Inyanga, Nyangani, 6000 ft, Chase 3813 (NU, SRGH); Gweni, Mt. Cashel, Chase 1083 (SRGH). WITHOUT EXACT LOCALITY: Gold Fields, Ayres s.n. (NH); Cap de Bonne Esperance, Drege s.n. (P); Johannesburg, Westeman s.n. (P); Cap b. Spei, Ecklon s.n. (P); Natal, sine coll. s.n. (P); Mor Bridge, Hill 692 (PRE); Drakensberg, Bottomley s.n. (PRE); Natal, Buchanan s.n. (BOL, M); O.F.S. TM1761c (PRE); Natal, Wood 504 (PRE); in vollbus montium seciis Katrivier, prope Philipstown, 3000-4000 ft, Ecklon & Zeyhers.n. (P); Natal, Buchanan 21 A-only (M); Kaffrarian forests, Sim s.n. (B); Natalia, Buchanan 74, 84 (B); CapTPr. b. sp., Ecklon & Zeyher 38.6 (B); ceded territory, Quellen des Katrivier, 3000-4000 ft, Drege s.n. (B); Prom. b. Spei, Drege s.n. B & C only (B); Natal, Wood s.n. (NU); Xumeri Forest, Rycroft 518 (NU); loco incerto, sine coll. s.n. NH-9785 (NH); below Mwndali, 5000 ft, Anderson s.n. (BM); Himalaya, Ravi Valey, Chanjii, 7000 ft, McDonnell 34 (BM); South Africa, Barckley 95 (GRA); sine coll. s.n. (L); loco incerto, sine coll. s.n. NH-26468 (NH). The African Polystichum luctuosum (Kunze) T. Moore and the Asian P. tsus-simense (Hook.) J. Sm. have been considered either as distinct taxa(Mitui, 1965, 1968;Hirabayashi, 1969;Daigobo, 1973; Nakaike, 1975;Gibby, 1985;Punethaetal., 1988) or as synonymous (Hope, 1902; Hooker in Hooker & Baker, 1868). Plants occurring in the western Indo-Himalayan mountains have been ascribed to either P. tsus-simense (Dixit, 1983) or to P. luctuosum (Khullar, 1987; Punetha et al., 1988). Fraser- Jenkins (in Gibby, 1985) considers the two taxa as vicariants. I have studied material throughout the distribution range of these taxa and find them to be conspecific. DIAGNOSTIC FEATURES AND RELATIONSHIPS. Diagnostic of Polystichum luctuosum is the olive-green colour of the adaxial surface of the lamina and the darker veins seen in living plants. It is also separated from other taxa in the region by the usually very dark and narrow paleae occurring along the stipe and rachis. The larger rhizome and stipe base paleae bear long filiform outgrowths along the margin and palea surface. Indusia are large, persistent and entire, and take on a cupulate form when mature. Polystichum luctuosum is furthermore a triploid apomict with 32 spores per sporangium and has a somatic chromosome number of 2n=123. Within the study area Polystichum luctuosum is the only member belonging to section Xiphopolystichum Daigobo. VARIATION. Polystichum luctuosum shows little variation in stipe, lamina and basal pinna length within the study area. A comparison of these parts with Asian material shows that African (including Madagascar) plants are slightly larger than the plants from Asia. Guard-cell length in African material shows little variation, but in Asian plants the variation is pronounced. Asian plants also have larger guard cells than African plants. Sori may be uniseriate or 41 biseriate, variation that appears to be environmentally induced. Indusia show a large degree of variation in both African and Asian plants, with African plants having larger indusia than Asian plants. Also the number of indurated annulus cells per sporangium shows some variation. In African plants the number ranges from 10 to 19, whereas in Asian plants the number ranges between 10 and 21. Spores too show variation, with Asian plants having larger spores than African plants (Table 1). Although variations in palea colour occur, their morphology remains fairly stable throughout the distri- bution of the species. DISTRIBUTION AND ECOLOGY. In South Africa Polystichum luctuosum occurs from the Eastern Cape through KwaZulu-Natal to the northeastern parts of the Free State, Mpumalanga, and the Northern Province. It also extends to the lower elevations in the western parts of Lesotho, the higher-lying part of Swaziland, and with isolated populations occurring along the eastern escarpment in Zimbabwe. Outside of Africa the species occurs on Reunion and the central parts of Madagascar, extending to the Indian subcontinent, Pakistan, China, Vietnam, South Korea and Japan (Honsyu, Sikoku and Kyusyu). Polystichum luctuosum occurs in the eastern parts of the summer rainfall regions of southern Africa where it appears to be restricted to the drier forest types such as Dohne Sourveld in the Eastern Cape, Highland Sourveld along the Drakensberg foothills, 'Ngongoni Veld in the KwaZulu-Natal midlands and Northeastern Mountain Sourveld in Swaziland, and along the lowveld escarpment and Soutpansberg. In South Africa P. luctuosum occurs at elevations ranging from 670 m to 1740 m, whereas in Zimbabwe on Mount Nyangani it occurs at elevations as high as 1825 m. Polystichum luctuosum mostly grows on rocks along streams, but often also as a low-level epiphyte in moist forests. Plants often also grow on rocks away from water and in fairly dry conditions. 3. Polystichum volkensii (Hieron.) C. Chr., Index filic.: 97 (1905). Type as for Aspidium volkensii Hieron. Fig. 4. Aspidium volkensii Hieron. in H.G. A. Engler, Pflanzenw. Ost-Afrikas: 86 ( 1 895). Type: Tanzania, an der oberen Grenze des Waldes iiber Kiboscho, 3000 m, Volkens 1520 (B!-holotype). Polystichum barbatum C. Chr. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 178 (1924). Type: Kenya, Mt. Aberdare, pr. Kinangop, regio Hagenia abyssinica, c. 3300 m, Rob. E. & Th. C.E. Fries 2735 (K!-holotype; B!-isotype). Plants terrestrial. Rhizome short, erect, to 10 mm in diameter, densely set with roots, persistent stipe bases, and paleae; paleae sessile or short- stalked, ferrugineous, chartaceous, narrowly lanceo- late, cordate, entire, the apex terminating in an acicular cell, to 15 mm long. Fronds caespitose, to 14 per plant, erect, to 1.2 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 520 mm long x 10 mm in diameter, densely paleated; larger paleae short-stalked, ferrugineous, membranous, translucent, shrivelled, elliptic to ovate, cordate to cordate-imbricate, minutely fimbriate, the apex long, shrivelled, filiform, entire, terminating in an acicular cell, to 34 x 10 mm; smaller paleae short-stalked, convolute, ovate to narrowly ovate, cordate to cordate-imbricate, proximally with short straight or angular marginal outgrowths, becoming entire towards the apex, the apex subulate, terminating in an acicular cell: lamina 3-pinnate, herbaceous, narrowly elliptic, to 925 mm long, olive-green adaxially, slightly paler abaxially, with a single paleated proliferous bud on the rachis near the lamina apex, the proximal pinnae decrescent, often somewhat deflexed: rachis stramineous, adaxially sulcate, densely paleated; paleae short-stalked, convolute, 42 J.P. ROUX Table 1 Variation in metric characters for African and Asian Polystichum luctuosum (Kunze) T. Moore. Character X African Range n X Asian Range n Stipe length 197mm 90-450 mm 35 193mm 98^50 mm 21 Lamina length 271 mm 158^48 mm 46 235 mm 162-465 mm 29 Basal pinna length 66.8 mm 31-178 mm 46 51.8mm 30-98 mm 21 Guard cell length 41.16 urn 30-52 urn 580 46.9 (am 36-60 urn 760 Indusium size 0.73 mm 0.56-0.95 mm 90 0.58 mm 0.31-0.80 mm 115 Number of indurated annulus cells 13.52 10-19 600 14.44 10-21 537 Spore length 38.84 (am 30-50 urn 275 40.75 Mm 30-52 Mm 405 Spore width 28.2 urn 22-36 urn 275 28.83 Mm 22-40 Mm 680 stramineous to ferrugineous, narrowly ovate, narrowly lanceolate, or transversely elliptic, cordate to cordate-imbricate, proximally erose or with short straight or angular outgrowths, becoming entire towards the apex, the apex subulate, terminating in an acicular cell, the smaller paleae to 18x6 mm: pinnae generally not overlapping at the lamina base, overlapping towards middle of the lamina, oblong-attenuate, somewhat falcate, basal pinnae to 54 mm long, the middle pinnae to 190 x 40 mm, proximal acroscopic pinnule slightly enlarged; pinna-rachis stramineous, adaxially sulcate, densely paleated; paleae similar to but smaller than those on the rachis: pinnules opposite to alternate, asymmetric, acroscopically auricu- late, ovate, to 23 x 11 mm, deeply lobed, the acroscopic auricle ovate, cuneate, lobes oblong to narrowly oblong, serrate to crenate, adaxially moderately paleated; paleae castaneous to ferrugineous, chartaceous, convolute, filiform, to 15 mm long, abaxially moder- ately to densely paleated; paleae short-stalked, castaneous to ferrugineous, chartaceous, convolute, filiform, narrowly linear or subulate, cordate to cordate-imbricate, proximally with short angu- lar outgrowths, entire towards apex, the apex terminating in an acicular cell, to 16.5 mm long. Venation immersed. Sori circular, <1 mm in diameter, essentially uniseriate, discrete at maturity, terminal or near-terminal on abbreviated vein branches, or dorsally on unabbreviated vein branches: sporangium with 1 2-( 1 4)- 1 9 indurated annulus cells; stalk eglandular: indusium ferrugineous to castaneous, peltate, circular, elliptic or irregular, coarsely erose, the maximum radius 0.48-(0.66)-0.92 mm. Spores 64 per sporangium, brown, the perispore folded to form a close reticulum of compressed ridges, the ridges and areas between granulate, verruculate or echinulate, vari- ously perforated, the exospore 34-(42.64)-52 x 24-(30.32)-38 (im. Chromosome number unknown. MATERIAL EXAMINED. KENYA: Mt. Nyandarua, forest belt, 10800-1 1000 ft, Rabb & Nightingale 1 (K). TANZANIA: Kilimanjaro, highest forest above Kibosho, Uhlig 186 (B), Uhlig 242 (B, K); Kilimanjaro, cave above Moschi, Uhlig 76 (B, K); Kilimanjaro, forested area just below 1 st hut and also above Machame route, 1820 m, Schippers T1452 (WAG); Kilimanjaro, Machame route, 3450 m, Pocs s.n. (WAG); Kilimanjaro, B-only, Brenner s.n. (P); Kilimanjaro, S. slope along the Mweka route, near Mweka base hut, 2850 m, Pocs 6718/A (K). WITHOUT EXACT LOCALITY: loco incerto, sine coll. BOL-5726 (BOL). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum volkensii is unique among African Polystichum species in the narrowly elliptic lamina outline and the long-decrescent, deflexed or arcuate pinnae. The densely paleated stipe and rachis, and the single proliferous bud borne near the frond apex are also characteristic. The finely divided pinnules and the morphology of especially the smaller paleae ensure that it cannot be mistaken for any other species in the region. The affinity of Polystichum volkensii is yet to be determined. VARIATION. Polystichum volkensii shows little infraspecific mor- phological variation. Variation is largely restricted to pinnule size and pinna orientation and this may be ascribed to environmental influences. DISTRIBUTION AND ECOLOGY. Polystichum volkensii appears to be confined to Mount Kilimanjaro in Tanzania and the Aberdare Moun- tain Range in Kenya. At lower elevations (1820 m) on Mount Kilimanjaro it occurs in Undifferentiated Afromontane forests but higher up, at 3450 m, it occurs in the Ericaceous belt with Erica arborea and Podocarpus milanjianus. On the Aberdare Mountain Range the species occurs in Undifferentiated Afromontane forests but also in Single-dominant Afromontane forests such as Hagenia abyssinica-forests at elevations ranging between 3300 and 3610 m (White, 1983). 4. Polystichum kilimanjaricum Pic.Serm. in Webbia 27: 445 (1972). Type: Tanzania, Kilimanjaro, presso la Bismarck's Hut, terrestre, nel sottobosco rado nella parte piu alta della foresta umida montana a Podocarpus milanjianus, Hagenia abyssinica ed Ilex mitis, c. 2850 m, 8 July 1 956, Pichi Sermolli 5171 (Herb. PIC.SERM. 20640-holotype; Herb. PIC.SERM. 25150!, K!- isotypes). Fig. 5. Plants terrestrial. Rhizome erect to suberect, to 1 80 mm long, closely set with roots, persistent stipe bases, and paleae. Fronds 8-12 per plant, caespitose, suberect to arching, to 1.05 m long: stipe proxi- mally castaneous, stramineous distally, adaxially sulcate, to 430 mm long x 8 mm in diameter, proximally densely paleated; proximal paleae broadly attached, ferrugineous, crustaceous, narrowly trian- gular, truncate to cuneate, the margins irregularly set with large recurved outgrowths, the apex terminating in an acicular cell, to 9 x 0.8 mm; distal paleae of two types, the larger broadly attached, bicolorous, with a central ebeneous to castaneous, glossy, crusta- ceous band, and a dull brown, chartaceous margin, narrowly ovate to broadly ovate, truncate to cuneate, the margins minutely fimbriate, the fimbriae straight or twisted, simple or apically forked, the apex terminating in an acicular cell, to 18x7 mm, the smaller short- stalked, concolorous, ferrugineous to stramineous, chartaceous, narrowly ovate to subulate, the margins minutely fimbriate, the subulate paleae always with long and/or short, simple or branched, often apically forked fimbriae at the base and widely spaced, recurved or apically directed outgrowths distally, the apex termin- ating in an acicular cell: lamina 2-pinnate, with up to 35 free pinna pairs, triangular to ovate, to 685 mm long, with 1-3 often widely spaced proliferous buds in pinna axils near the apex: rachis stramineous, adaxially sulcate, moderately to densely set with paleae similar to but smaller and paler than those on the stipe: pinnae 1- POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 43 10 mm 5 mm Fig. 4 Polystichum volkensii. A, middle pinnae of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A, drawn from Volkens 1520 (B); B, drawn from Radd & Nightingale 1 (K); C, drawn from Pocs s.n. (WAG). 44 J.P. ROUX Fig. 5 Polystichum kilimanjaricum. A, proximal pinnae of lamina; B, lamina apex showing proliferous bud; C, abaxial surface of fertile pinnule. All drawn from Pichi Sermolli5\l\ (Herb. PIC.SERM.). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 45 pinnate, with up to 14 free pinnule pairs, not overlapping proxi- mally, the basal pinnae not or slightly reduced in size, often somewhat deflexed, ovate, narrowly ovate or oblong-attenuate, to 140 x 35 mm: pinna-rachis stramineous, adaxially sulcate, set with acicular paleae with numerous long and often twisted outgrowths at the base, the apex terminating in an acicular cell, to 7 mm long: pinnules slightly imbricate, short-stalked proximally, firm-herbaceous to subcoriaceous, adaxially olive-green, somewhat paler abaxially, inaequilateral, ovate-rectangular to ovate-rhomboid, basiscopically cuneate, acroscopically truncate and weakly auriculate, shallowly undulate or serrate, the teeth and aristae bent inwards, the auricle and apex aristate, the proximal acroscopic and basiscopic pinnules on basal pinnae often slightly reduced in size, the proximal acroscopic and basiscopic pinnules on upper half of lamina slightly larger than the next, to 20 x 10 mm; adaxially sparsely set with short-stalked, acicular, somewhat twisted paleae, often with a few long straight or twisted, often branched outgrowths at the base, the apex terminating in an acicular cell, to 2.5 mm; abaxial surface with similar paleae but more densely set. Venation immersed or raised. Sort circular, c. 1 .4 mm in diameter, terminal on abbreviated vein branches, uniseriate or biseriate on acroscopic auricle, discrete: sporangium with 8- (13)-20 indurated annulus cells; stalk eglandular: indusium peltate, subcircular to irregular, the maximum radius 0.73-(0.87)-1 .02 mm, persistent, brown. Spores brown, the perispore folded to form inflated or compressed tubercles, echinulate, verruculate to echinu- late, sparsely to closely perforated, the exospore 34-(43.44)-56 x 26-(32.45)^M \im. Chromosome number unknown. MATERIAL EXAMINED TANZANIA : Kilimanjaro, below 1 st hut, Machame Route, 2950 m, Schippers T1465 (WAG); Kilimanjaro, above Mandate Hut, 1830 m, Schippers7l234A (WAG). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum kilimanjaricum is characterized by the proliferous buds on the lamina and the palea morphology. Paleae on the proximal part of the stipe are narrowly triangular and are either castaneous throughout, or may often have a very narrow paler brown margin. Larger paleae higher up the stipe have an ebeneous to castaneous centre and a broader ferrugineous to stramineous margin. These paleae are mostly oblique in outline. Larger paleae on the upper two-thirds of the stipe are ovate to broadly ovate in outline with only the central part of the apices being ferrugineous to castaneous in colour. Pichi Sermolli (1972) considered this species to closely resemble P. pauciaculeatum Bonap. (as P. coursii Tardieu) and P. tsaratananense Tardieu from Madagascar and went on to describe how P. kilimanjaricum can be distinguished from them. Morpho- logically P. kilimanjaricum is more similar to P. tsaratananense than it is to P. pauciaculeatum. Polystichum kilimanjaricum belongs to section Lasiopolystichum Daigobo. DISTRIBUTION AND ECOLOGY. Polystichum kilimanjaricum appears to be endemic to Mount Kilimanjaro in Tanzania, occurring at elevations ranging between 1830 and 2950 m. The species is evi- dently confined to the Hagenia abyssinica montane forests and thickets associated with the Ericaceous belt where it mostly occurs on the forest floor and on rocky streambanks. 5. Polystichum aculeatum (L.) Roth, Tent. fl. Germ. 3(1): 79 (1799). Type as for Polypodium aculeatum L. Fig. 6. Polypodium aculeatum L., Sp. pi.: 1090 (1753). Type: Habitat in Europa. H.L.B. 908,311.72 (L-lectotype, designated by Alston (1940)). Polypodium lobatum Huds., Fl. angl: 390 (1762). Type: Habitat in umbrosis et ad sepes. Haller, Hist, stirp. Helv. : 1 7 1 2 ( 1 768); Pluk., Phytographia: 180, f. 1 (1691); Ray, Syn. meth. stirp. brit.: 121 (1690)-syntypes. Aspidium aculeatum (L.) Sw. in Jl. Bot. (Schroder) 1800(2): 37 (1801). Aspidium lobatum (Huds.) Sw. in Jl. Bot. (Schroder) 1800(2): 37 (1801). Polystichum lobatum (Huds.) Bastard, Essaifl. Maine et Loire: 367 (1809). Chevall., Fl. Belg., Pterid.: 107 (1950). Dryopteris aculeata (L.) Kuntze, Revis. gen. pi. 2: 812 (1891). Dryopteris setifera subsp. lobata (Huds.) Maire in E. Jahandiez & R.C.J.E. Maire, Cat. pi. Maroc 1: 3 (1931). Plants terrestrial or epilithic. Rhizome short, erect to suberect, to 1 20 mm long, to 15 mm in diameter, set with roots, closely spaced persistent stipe bases, and paleae. Fronds crowded, caespitose, 8-11 per plant, erect to arching, to 935 mm long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 170 mm long x 6 mm in diameter, proximally densely set with conspicuously larger and smaller paleae, moderately paleated distally; larger paleae sessile, castaneous, chartaceous, broadly ovate, cordate, the margins proximally closely set with short curved outgrowths, the apex usually short-flagelliform, terminating in a small thin-walled cell, to 15 x 9 mm; smaller paleae sessile, castaneous to stramineous, chartaceous, lanceolate or narrowly to broadly ovate, cordate to cordate-imbricate, the margins proximally with short curved outgrowths, the apex terminating in a subulate cell or a small thin- walled cell: lamina 2-pinnate, with up to 41 free pinna pairs, coriaceous, adaxially dark green, slightly paler abaxially, narrowly elliptic, to 770 mm long, closely spaced and often imbricate distally, proximally more widely spaced, the proximal pinnae reduced, often slightly deflexed: rachis stramineous, adaxially sulcate, moderately paleated; paleae sessile, ferrugineous, chartaceous, broadly ovate, ovate, narrowly elliptic or hastate, cordate to cordate-imbricate, the margins with short, somewhat curved outgrowths extending nearly to the apex, the apex terminating in a subulate cell, a long acicular cell, or a small thin-walled cell, to 6 x 3 mm: pinnae short-stalked, pinnatifid to 1 -pinnate, with up to 16 free pinnule pairs, narrowly oblong-attenuate, the middle pinnae to 110 mm long, the proximal pinnae to 88 mm long: pinna-rachis stramineous, adaxially sulcate, moderately paleated; paleae sessile or short-stalked, ferrugineous, chartaceous, ovate, narrowly lanceolate to hastate, cordate to cor- date-imbricate, the margins proximally with short or long, usually curved outgrowths, the apex terminating in a subulate cell or a small thin-walled cell, to 2 x 0.8 mm: pinnules opposite to alternate, somewhat imbricate, the proximal acroscopic pinnule usually slightly longer than the next, asymmetric, trullate to narrowly trullate, basiscopically cuneate, acroscopically cuneate to truncate and au- riculate, serrate to long-aristate, to 15 mm long; adaxially with a few membranous, filiform paleae terminating in a subulate or thin- walled cell confined to proximal part of pinnule, to 1 .75 mm long; abaxially moderately set with membranous, narrowly trullate or narrowly lanceolate paleae with a few short and straight marginal outgrowths, or the paleae filiform, short-stalked, with the apex terminating in a subulate cell or a small thin-walled cell, to 2.5 mm long. Venation immersed. Sori circular, to 1.5 mm in diameter, terminal or near terminal on abbreviated vein branches, essentially uniseriate, discrete to confluent at maturity: sporangium with 12- (13)-17 indurated annulus cells; stalk eglandular: indusium chartaceous, peltate, circular, entire to repand, the maximum radius 0.63-(0.92)-1 .26 mm, persistent, brown. Spores brown, the perispore folded to form short echinate ridges or crests, the areas between 46 J.P. ROUX 10 mm Fig. 6 Polystichum aculeatum. A, proximal part of lamina; B, fertile pinnae; C, rhizome. A & B drawn from Lindberg 2793 (B); C, drawn from Cosson s.n. (S). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 47 fenestrate with pores of variable sizes, the exospore 32-(37.46) 48 x 24-(27.93)-34 jam. Chromosome number 2n= 1 64 (Manton, 1 950). MATERIAL EXAMINED ALGERIA: Montagnes du Djurdjura, cercle de Dra el Mizan, Cosson s.n. (B); La Gourraya de la Bougie, Corzeillea Tere s.n. (S). MOROCCO: Haul Atlas, Ourika, 1400 m, Litardiere s.n. (M, P); Great Atlas Mountains, Si Chamharouch, 2280 m, Polunin 2 1 84 (BM); N. face G- bou Orionl, 2950 m, Newbould 109, 1 10 (BM); Arromiel, Balls 2972 (B, BM, S); Taddert, Marrakesh-Quarzazat road, High Atlas, 1600 m, Chatworth- Musters 362 (BM); Atlas Magnum, Amismiz, 1400 m, Lindberg 2793 (B, S); Meknes, Aguelmane Azigza, 1600 m, Casas et al. s.n. (B); Grand Atlas, Ourika, 1 300-1400 m, Maire s.n. (RAB); Haul- Atlas, Ourika, 2600 m, sine coll. s.n. 18633 (RAB); env. de la maison forestier de Khanolak-Anasar, Jovet-Astetal 13313 (RAB). Variation in Polystichum aculeatum and P. setiferum and the occur- rence of intermediate forms and hybrids (P. x bicknellii (H. Christ) Hahne) between these species have resulted in diverse interpreta- tions as to their delimitation. The result has been some nomenclatural confusion (Newman, 1844; Alston, 1940; Elliot, 1950; Meyer, 1960). Hudson ( 1 762), unaware of the existence of the name Polypodium setiferum Forssk., recognized two forms in European P. aculeatum and divided plants into two species. The rigid and less divided form he named Polypodium lobatum, and the lax and more divided form he retained in P. aculeatum. Hudson's interpretation off. aculeatum is therefore synonymous with P. setiferum (Forssk.) T. Moore ex Woyn. and P. lobatum with P. aculeatum as now interpreted. Al- though the name P. lobatum never became well-established, arguments in favour of its retention were made by Meyer (1960). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum aculeatum is a fertile sexual species intermediate in morphology between P. lonchitis (L.) Roth and P. setiferum (Forssk.) T. Moore ex Woyn., although it is closer in appearance to the latter. Manton (1950) demonstrated P. aculeatum to be a tetraploid of hybrid origin between the putative parents P. lonchitis and P. setiferum. Daigobo (1972) placed this species and P. setiferum in different sections, but their palea morphology suggests them to be related. Both belong to section Metapolystichum Daigobo. Polystichum aculeatum is characterized by having a coriaceous lamina and smooth, shiny, dark green pinnules. The longest pinnae occur at or near the middle of the lamina with the most proximal pinna pair distinctly shorter than the middle pinnae. The stipe/ lamina ratio in P. aculeatum is 1:3.27 (n=12). Palea density and morphology are also diagnostic and differ from that of P. setiferum with which it may be confused. In P. aculeatum the stipe, rachis and pinna-rachis are moderately paleated, whereas in P. setiferum they are usually densely set with twisted paleae. Marginal outgrowths in larger paleae are short and curved, and gradually phase out towards the apex. Smaller paleae are mostly broad-based and have short curved marginal outgrowths, but distally they terminate abruptly in an almost simple subulate apex. Paleae occurring abaxially on the pinnules are short and proximally bear a few small, straight or curved, marginal outgrowths, with the apex terminating in a subulate cell or a small thin-walled cell. Polystichum aculeatum also differs from P. setiferum in a number of micromorphological characters, with the mean adaxial epidermal cell length, guard cell length, maximum radial length of the indusium, and the spores being larger than those in P. setiferum. VARIATION. Polystichum aculeatum varies in the degree to which the pinnae reduce in size towards the base of the lamina and in the length of the stipe in relation to the length of the lamina. European plants appear to have shorter stipes than plants from Africa. Pinnules of plants from the study area are remarkably stable and show little variation. DISTRIBUTION AND ECOLOGY. Polystichum aculeatum is widespread in Europe but in North Africa its distribution is more restricted. Hansen & Sunding (1993) and Derrick et al. (1987) considered the species to also occur on Madeira and the Canary Islands, but no material originating from Madeira could be traced by Manton et al. (1986), Gibby & Paul (1994), or myself. I have also not seen any material of this species from the Canary Islands. In Algeria and Morocco Polystichum aculeatum is restricted to the High Atlas Mountains. The lithology of the region consists largely of basement rock and unconsolidated clay marls (White, 1983). The rainfall is low and seasonal at lower elevations but at higher elevations precipitation may occur throughout the year. The species occurs at elevations ranging between 1400 m and 2950 m, where it is confined to moist shaded rock crevices along streams and at waterfalls. 6. Polystichum setiferum (Forssk.) T. Moore ex Woyn. in Mitt. Naturwiss. Vereines Steiermark 49: 181 (1913). Type as for Polypodium setiferum Forssk. Fig. 7. Polypodium setiferum Forssk., Fl. aegypt.-arab.: 185 (1775). Type: Turkey, Dardanelles ('Ad Dardanelles'), Forsskal 814 (C!- lectotype, designated by Hepper & Friis (1994)). Aspidium angulare Kit. ex Willd., Sp. pi 4, 5(1): 257 (1810). Type: Habitat in Hungaria, sine coll. s.n. (B-Willd.-holotype, NBG!- photograph). Aspidium hastulatum Ten., Semina 1830: 15 (1830). Type: In nostri regni nemoribus, et abunde in vallibus circa Neapolim, 5. Rocco, Ponti Rossi & Camaldoli s.n. (not located). Polystichum angulare (Kit. ex Willd.) C. Presl, Tent, pterid.: 83 (1836). Aspidium aculeatum subsp. angulare (Kit. ex Willd.) Asch. in P.F.A. Ascherson & K.O.R.P.P. Graebner, Syn. mitteleur. Fl. 1: 39 (1896). Polystichum aculeatum subsp. angulare (Kit. ex Willd.) Vollm., Fl. Bayern: 9 (1914). Dryopteris aculeata subsp. angularis (Kit. ex Willd.) Schinz & Thell. in H. Schinz & R. Keller, Fl. Schweiz 3rd ed., 2: 3 (1914). Dryopteris setifera (Forssk.) Woyn. ex Schinz & Thell., Vierteljahrsschr. Naturf. Ges. Zurich 60: 340 (1915). Dryopteris setifera subsp. angularis (Kit. ex Willd.) Maire in E. Jahandiez & R.C.J.E. Maire, Cat. pi. Maroc 1: 3 (1931). Plants terrestrial or epilithic. Rhizome erect to suberect, short, to 18 mm in diameter, set with roots, closely spaced stipe bases, and paleae; paleae broadly attached, stramineous to ferrugineous, chartaceous, ovate to broadly ovate, often somewhat bullate, cor- date, the margins minutely fimbriate to erose, the apex generally entire, terminating in a subulate cell or a small thin-walled cell. Fronds 8-22 per plant, suberect to arching, to 1 .2 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 520 mm long x 6 mm in diameter, densely paleated; paleae ferrugi- neous, chartaceous, the larger paleae sessile, narrowly ovate, ovate, or broadly ovate, often somewhat bullate, cordate, the margins finely fimbriate to erose, the apex entire, terminating in a subulate cell or a thin-walled cell, to 20 x 1 1 mm, the smaller paleae narrowly oblong, narrowly ovate, or subulate, mostly helically twisted, short- stalked, cordate-imbricate, the margins proximally with short and/or long outgrowths, the outgrowths straight, narrowly triangular, or angular, reduced in size towards the apex, the apex terminating in a 48 J.P. ROUX Fig. 7 Polystichum setiferum. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinna. A & C drawn from Mandon 291 (S); B, drawn from Tullgren 21 (S). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA subulate cell or a small thin-walled cell: lamina 2-pinnate to 2- pinnate-pinnatifid, with up to 45 free pinna pairs, firmly herbaceous to coriaceous, dark green adaxially, somewhat paler abaxially, ovate to elliptic, to 830 mm long, the proximal pinnae reduced, often somewhat deflexed: rachis stramineous, adaxially sulcate, densely paleated; paleae narrowly oblong, narrowly ovate, narrowly lanceo- late, or subulate-hastate, helically twisted, short-stalked, cordate to cordate-imbricate, the margins proximally with short and/or long straight, curved or angular outgrowths that reduce in size towards a usually entire apex, the apex terminating in a subulate cell or a small thin-walled cell, to 6 x 2 mm: pinnae with up to 26 free pinnule pairs, closely to widely spaced, often overlapping towards the apex, narrowly oblong-attenuate, the middle pinnae to 162 mm long, the proximal pinnae to 1 40 mm long: pinna-rachis stramineous, adaxially sulcate, moderately to densely set with paleae similar to but smaller than those on the rachis: pinnules opposite to alternate, closely spaced, the proximal acroscopic pinnule not or slightly enlarged, inaequilateral, acroscopically auriculate, trullate or oblong-acumi- nate, lobate-dentate, aristate, to 13x5 mm; adaxially with a few twisted filiform paleae confined to the costa on the proximal part of the pinnule, these terminating in a subulate cell or a small thin- walled cell, to 4.5 mm long; abaxially moderately set with narrowly triangular, subulate-hastate, or filiform paleae, the larger paleae proximally usually with a few long marginal outgrowths, the apex entire, terminating in a subulate cell, to 2.9 mm long. Venation immersed. Sori circular, c. 1 mm in diameter, terminal or near terminal on abbreviated vein branches, essentially uniseriate, dis- crete to confluent at maturity: sporangium with 1 1 -( 14)-20 indurated annulus cells; stalk eglandular: indusium pale brown, persistent, peltate, circular, repand, the maximum radius 0.48-(0.85)-1.21 mm. Spores 64 per sporangium, brown, the perispore folded to form inflated tubercles and ridges, echinulate to verruculate, sparsely perforated, the exospore 26-(34.52)-44 x 1 8-(25.92)-36 Jim. Chro- mosome number 2n=82 (Manton et al., 1986). MATERIAL EXAMINED ALGERIA: Mont Magnis, l500m,Reverchon371 (BM, P); Djebel Edough, Cosson s.n. (P); gorge de la Chiffa, Cosson s.n. (P); Djebel Marouf, petite Kabylie, Prov. de Constantine, Cosson s.n. (P); dans la fout du Dirah aux environs d' Aumale, Chaoy 834 (P); montagnes du Djurdjura, cerde de Dra el Mizan, Prov. d'Alger, Cosson s.n. (P); 3 miles W. of the Hotel Lambert, Adekar, c. 900 m, Alston & Simpson 37578 (BM); Djebel, Stephenson s.n. (BM); Remain, Nud el Kebin, Alwah, sine coll. s.n. (BM); Algeria, Eichard s.n. (WAG). MOROCCO: entre les rochers humides et umbrage du mont Amareza, Atlas, Bove s.n. (P); Al Hoceima. cerca de Ketama, subiendo al monte Koudiet Imougras, 1 880 m, Casas 7237 (B); Hafa-es-Sabbaba (Ben-Hosmar), ad 500 m, Quer s.n. (B, S). TUNISIA: NV d'ain-Draham, Cosson s.n. (P); Massif d' El-Fedja, Cosson & Duval s.n. (P); Ain Draham, open cork-oak forest, c. 900 m, Simpson 38370 (BM); foret du Feidja, 20 km W. of Ghardimaou, 800 m, Jansen 462 (WAG). WITHOUT EXACT LOCALITY: Herb. Luerssen 5242, sine loc. (P); loco incerto, sine coll. B-96812 (B); loco incerto, De Buck s.n. (B); Kaap de Goede Hoop [error, not a native of the Cape], sine coll. 9 (L). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Problems are fre- quently experienced in separating Polystichum setiferum from P. aculeatum. The former, however, has larger fronds that are softer in texture. Also the proximal pinnae are not usually markedly reduced in P. setiferum as they are in P. aculeatum. In P. setiferum the paleae are characteristically helically twisted and the apices more fre- quently terminate in a small thin-walled cell than those of P. aculeatum. Micro morphological characters separating the two taxa are reported under P. aculeatum. 49 VARIATION. Considering the wide geographical distribution of the species, it shows remarkably little variation. Dyce (1963) reported that a wide range of minor variations in shape and habit can be expected in any colony of this species. I found variations in the size and shape of the fronds to be most pronounced, but since no obvious geographic pattern was detected, it is here considered to be environ- mentally induced. Plants from drier areas, in particular the North African region, have fronds that are often merely 1 -pinnate or 1- pinnate-pinnatifid. In large specimens from moist areas, however, the lamina may be 2-pinnate-pinnatifid with the proximal acroscopic pinnule being 1 -pinnate and often twice as long as the next pinnule. Irrespective of habitat and environmental conditions, the palea structure shows little variation. DISTRIBUTION AND ECOLOGY. Polystichum setiferum is widespread in Britain, Europe south of 53 N latitude, the Crimea, Macaronesia (Azores, Canary Islands and Madeira), and Africa north of the Sahara. In North Africa P. setiferum occurs at elevations ranging between 500 and 1 880 m in the Saharan Atlas-, High Atlas- and Anti-Atlas Mountain ranges in Tunisia, Algeria and Morocco. In this region of low rainfall plants are restricted to well-protected rock crevices and moist banks. In Tunisia, however, the species also occurs in open cork-oak (Quercus suber L.) forests. 7. Polystichum transvaalense N.C. Anthony in Contr. Bolus Herb. 10: 146 (1982). Type: South Africa, Transvaal (Northern Prov- ince), Pietersburg District, Woodbush Forest Reserve, Bredenkamp & Van Vuuren 450 (BOL!-holotype; PRE!-isotype). Fig. 8. Plants terrestrial or epilithic. Rhizome short, erect, to 8 mm in diameter, densely set with roots, persistent stipe bases, and paleae; paleae broadly attached, castaneous, chartaceous, narrowly linear to narrowly lanceolate, the margins proximally entire, distally with numerous short, apically or basally directed outgrowths, the apex terminating in an acicular cell, to 14 x 1 mm. Fronds caespitose, 5- 17 per plant, suberect to arching, to 1.045 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 535 mm long x 5 mm in diameter, densely paleated, the paleae twisted; larger paleae mostly confined to the stipe, concolorous or bicolorous, castaneous to ferrugineous or with the central part castaneous to black, rugose, narrowly ovate-acuminate to ovate-acuminate, short- stalked, the margins irregularly lacerate-fimbriate, the apex terminating in an acicular cell, to 20 x 6 mm; smaller paleae short- stalked, narrowly ovate to narrowly lanceolate, the margins proximally lacerate, distally irregularly lacerate-fimbriate, the apex terminating in an acicular cell, to 6.5 x 1 .4 mm: lamina 2-pinnate, with up to 26 free pinna pairs, herbaceous, ovate to narrowly ovate, to 670 mm long, pale green adaxially, paler abaxially, the proximal pinnae often slightly reduced, often deflexed: rachis stramineous, adaxially sulcate, often flexuous distally, densely paleated; paleae short-stalked, twisted, castaneous to ferrugineous, narrowly ovate, narrowly triangular, or linear, the margins proximally lacerate, distally irregularly and widely fimbriate, the apex terminating in an acicular cell, to 4.5 mm long: pinnae 1 -pinnate, with up to 20 free pinnule pairs, proximally widely spaced, distally closely spaced and somewhat overlapping, oblong-attenuate, the basal pinnae to 140 mm long x 28 mm wide, proximally often slightly reduced, the basalmost acroscopic pinnules longer towards the middle of the lamina: pinna-rachis stramineous, adaxially sulcate, densely paleated; paleae similar to but smaller than those on the rachis: pinnules opposite to alternate, inaequilateral, acroscopically auricu- late, ovate to obliquely transversely rhomboid, to 15 mm long. 50 J.P.ROUX Fig. 8 Polystichum transvaalense . A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2414 (NBG). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA serrate to lobate-serrate, often short-aristate, the proximal pinnules often pinnatifid; adaxially subglabrous or with a few twisted, fili- form paleae proximally on costa, the apex terminating in an acicular cell, to 2 mm long; abaxially moderately paleated; paleae short- stalked, twisted, narrowly linear to narrowly triangular, the margins proximally long-lacerate or fimbriate, the apex terminating in an acicular cell, to 2.5 mm long. Venation immersed. Sori circular, c. 1 mm in diameter, terminal or near terminal on abbreviated vein branches, discrete at maturity: sporangium with 10-(14)-22 indurated annulus cells; stalk eglandular: indusium stramineous, peltate, circular, simple or often with a few long central processes, repand to erose, persistent, the maximum radius 0.48-(0.76)-0.97 mm. Spores 64 per sporangium, brown, the perispore folded to form tubercles and inflated or compressed reticulate ridges, echinulate, closely perforated, the pores to 1 .5 urn in diameter, the exospore 30- (38.54)-48 x 22-(28.86)-38 urn. Chromosome number 2n= 1 64. MATERIAL EXAMINED BIOKO: Bioko (Fernando Po), 9000 ft, Mann 340 (K); carratera del pico Basile, km 18-19, nacimento del rio Cope, 32NMJ7597, 2470 m, Carvalho 3682 (B, BR). CAMEROON: Pisted'Acha-Abaw au lac Oku, 40 km NE Bamenda, Letouzey 1 3439 (P); montane forest between hut 1 and hut 2, 1 950 m, Breteler et al. 266 (K, P, WAG); Buea, Preuss 719 (B); Mannsquell, Luckhardt 636 (B); Mount Cameroon, Kalbreyer 133 (B); Uauenzuba, 1900 m, Schaeter90 (B). DEMOCRATIC REPUBLIC OF CONGO: Mare de Kikeri, Volcan Hekeno, Jean-Louis 5 1 9 1 (BR, P, PRE); entre le Hikeno et le Heisfumangabo, Jean-Louis 5002 (BR, P, PRE); Kivu Province, Goma, petite mare de Kikeri, au pied du Mikeno Pare National Albert, 2200 m, Lebrun 7225, 7228 (BR, K, P); entre Kibumba et le Ngamuragira, Lebrun 7087 (BR, P); Kivu, Pare des Virunga, Karisimbi, vers le SW Zaire, 3465 m, Van der Veken PV 9130 (B); Kivu, Buhavu-Goma, Gupffert 169 (BR); 748'S, 2944'E, Marungu route Kieluzi-Mwela km 45 Ravin Lukole, 1 890 m, Bodenghiem & Malaisse 1459 (BR); Pare National Albert, versant S. du Mikens, 2400-2600 m, Lebrun 7307 (BR); Omigi, Mickule, Bequaert 6294 (BR); route Goma-Rwindi 30 km, Breyne 1785 (BR); Volcan Niamlagyra, c. 2000 m, Germain 1381 (BR); Kivu, Rumangabo, 1525 m, Germain 3025 (BR); Numbi territory, Kalehe, 2300 m, Leonard 4554 (BR); Mt. Kiniki territory, 1960 m, Gutzwiller 1236 (BR); Mt. Kiniki, Wambalyro, 1960 m, Gutzwiller 1 106 (BR); Pare National Albert, Kalonge, Butahu, vallee du la Nyamwumba, 2010 m, Demaret5\92 (BR). ERITREA: Eritrea- Assaorta: bosco del Caribozza, c. 2700 m, Pappi 2812 (BOL, BR). ETHIOPIA: c. 5 km NW of Addis Ababa, c. 2500 m, De Wilde 598 1 (BR, ETH, WAG); Kaffa Province, 35 km W. of Bonga along the road to Shewa Ghimmira,c. 1 950 m, Fr/w et al. 2171 (ETH, K); near Wush-Wush,c. 20km NW of Bonga, c. 1800 m, De Wilde 7756 (BR, ETH, WAG); Gara Ades, Burger 2580 (K); Mount Wachacha, near Addis Ababa, 2400 m, Mooney 7895 (K); Wofasha Forest, Shoa, Mooney 7003 (K); Bellete State Forest, 40 km SW of Jimma, c. 2000 m, De Wilde 6999 (BR, WAG); Kaffa Province, village c. 2 hours walk NW of Maji, 2200 m, De Wilde 6194 (BR, WAG); W. slope of Mount Uociacia, c. 15 km W. of Addis Ababa, 2700 m, De Wilde 9580 (WAG); Mount Uociacia, c. 15 km W. of Addis Ababa, c. 2600 m, De Wilde 8532 (WAG); Kaffa Province, Limmu, Monti Botor, c. 2250 m, Pichi Sermolli 7069 (SRGH); 717'N, 365'E, Kaffa Province, 35 km W. of Bonga along the road to Shewa Ghimmira, 1950 m, Friis 211 \ (BR). KENYA: Nyambeni Hills, base of Kirima, 6400 ft, Polhill & Verdcourt 295 (BR, K, PRE); Aberdares, Cave Waterfall, Coe 794 (PRE); Molo, Mail Escarpment, 2440 m, Alluaud 55 (BR, P, PRE); Kiambu District, Katamayu River Forest, 2200-2250 m, Faden & Evans 69/236 (BOL); Meru District, Ngambeni Hills, above Kiegoi, 2250 m, Faden et al. 69/678 (K); Aberdares, S. Kinangop, 8600 ft, Molesworth-Allen 3637 (K); Kericho District, crossing at the Kitinges River, c. 8 km ENE of Kericho, 2060 m, Faden 72/302 (K); Kericho District. W. Mau Forest, SW of Mt. Blacket, Faden et al. 72/356 (K); Samburu District, Nyiro Mountain, 2400 m, Bono 23 (K); Taita Hills, Vuria Hill, 1920-2200 m, Faden 72/255 (BOL, K): Kinangop, Brown Trout Inn, 9000 ft, Verdcourt 880 (K); S. Kinangop, near Brown Trout Inn, Molesworth- 51 Allen 3620 (K); Kinangop, above Isanga farm, 8500 ft, Andrews 4461 (K); Chyulu Hills, 6800 ft, Van Someren 7572 (K); Chyulu Hills, 2250 m, Bally 1 163 (K); Mount Meru, 5000-6000 ft, Leighton s.n. (K); Aberdare Range, base of Mount Kenya, Dawson 96a (K); Taita Hills, Vuria Forest, c. 7000 ft, Schippers K27 1 (WAG); Thompson Falls, c. 7600 ft, Schippers K 1 7 (WAG); Kinangop, Brown Trout Inn, Verdcourt & Moggi 2486 (B, SRGH); Elgon Forest, Webster 9055 (K); prope 'West Kenia Forest Station', 2300 m, Friis 594 (B, BR, S); Mount Elgon, 4300 ft, Barrele 92 (NU); Aberdare Range, near W. part of the Nyeri track, 3 1 00 m, Hedberg 1 533 (S); Samburu District, Mt. Nyiru, 8000 ft, Cameron 147 (BR). LESOTHO. 2927 (Maseru): gorge dans la montagne Ma-Khrarane, au dessus de la station missionaire de Morija (DA), Dieterlen 1309 (P, PRE). MALAWI: Nyika, Zovochipolo, 2225 m, La Croix 4634 (PRE); Nyika Plateau, Zovochipolo forest patches, 2200 m, Dowsett-Lemaire 297 (MAL); Kirk Range, Dzonze Forest, 1750-1800 m, Dowsett-Lemaire 1079 (K); Mwanembu Mountain, McClounie 6 (K); S. region, Malosa Mountains (N. of Zomba), 1900 m, Dowsett-Lemaire 973 (K); Mount Mulanje, Tuchila Pla- teau, 6000 ft, Newman & Whitemore 214 (SRGH). MOZAMBIQUE: Penhalonga Waterfall, Chase 3247 (NU, SRGH); Penhalonga Forest, Chase 3219 (SRGH). RWANDA: Plantation Gasiza au N. de Ruhengeri au pied Ngahinga et du Muhavura, 2350 m, Van der Veken PV 10265 (B, BR); Kirunga Vulcan, 2500 m, Poetsen 8 1 (B); Dalinghi, Zappelli 262 (BR); Kissenyi, Sake, Zappelli 1 77 (BR); Chaine des Birunga, pied SE du Gahinga, 2500 m, Lambinon 74/1 534 (BR). SOUTH AFRICA. 2329 (Pietersburg): Louis Trichardt, Hanglip Forest Station (BB), Roux 2572 (NBG); Louis Trichardt, Zoutpansberg Suds, farm Rustfontein, c. 1400 m, Schlieben 7342 (BR); Tzaneen, Dap Naude Dam, Woodbush (DD), Burrows 3269 (BOL, PRE). 2330 (Tzaneen): Duiwelskloof, Westfalia Estate (CA), Scheepers 419 (PRE); De Hoek Forest Station (CC), Roux 2563 (NBG); Woodbush Forest Reserve, Grootbos, Roux 2564, 2570 (NBG); Magoebaskloof near De Hoek Forest Station, Van Jaarsveld 6093 A (BOL, NBG); Woodbush, Jenkins s.n. TM 919c (PRE); Woodbush, Wager s.n. CH7464 (PRE); Woodbush, Reynolds s.n. CHI 0246 (PRE); De Hoek, Schweickerdt s.n. (NBG, PRE); Pietersburg, Woodbush, Schelpe 6050 (BOL). 2430 (Pilgrim's Rest): Mariepskop, base of Klaserie Waterfall (DB); Bur- rows 3113 (BOL, PRE); Mount Sheba Nature Reserve (DC), Roux 2556 (NBG); Mount Sheba, Kluge 2320 (NBG); Mount Sheba Nature Reserve, Jacobsen 4420, 4428 (PRE); Ohrigstad Nature Reserve, Jacobsen 1413 (PRE); Pilgrims Rest (DD), Collins s.n. TM895c (PRE); Graskop, Blyde River Forest Reserve, Jacobsen 4363 (PRE). 2530 (Lydenburg): Lydenburg, Coromandel farm (AD), Roux & Burrows 13 (BOL); Coromandel farm, Burrows 1309 (BOL); Sabie, Tweefontein (BB), Wager 48 (PRE); Sabie Gorge, Wager 25 (PRE); Sudwala Caves, forest 2 km N. of caves, 1500 m (BC), Kluge 2465 (PRE); Sudwala Caves, Burrows 3193 (BOL); Nelspruit, Witklip Staatsbos (BD), Kluge 853 (PRE); Kaapsehoop (DB), Wager s.n. TM149c (PRE); Lydenburg, Clivia Pass (DD), Edwards 1149 (NU). 2531 (Komatipoort): Lomati falls and kloof behind Barberton (CC), Wager 151 (PRE); Barberton, Williams 104 (P); Baberton, Thorncroft 104 (GRA). 2630 (Carolina): Marieriestad (CA), Pott-Leendertz 4848 (BOL, PRE). 2730 (Vryheid): Piet Relief- Wakkerstroom road, 6 km from turnoff to Liineburg (AD), Roux 2269 (NBG); Wakkerstroom, Oshoek, Devenish 2 (PRE). 2828 (Bethlehem): Royal Natal National Park, Goodoo Forest (DB), Doidge s.n. (PRE); Tugela Valley, Hafstrom & Acocks 1970 (PRE); Royal Natal National Park, Devils Hoek, 5000 ft, Schelpe 7973 (BOL). 2829 (Harrismith): Van Reenens Pass (AD), Rehmann 7205 (B, P); Oliviershoek Pass, Begonia Falls (CA), Roux 2514 (NBG); Qualeni Valley, 800 ft (CC), Schelpe 7270 (NU, PRE). 2929 (Underberg): Cathedral Peak Forest Research Station, 6050 ft (AB), Killick 1 134 (PRE); Lions River, Lions Bush (BD), Moll 829 (PRE). 2930 (Pietermaritzburg): Lidgetton (AC), Mogg CHI 764 (PRE); Lions River District, Karkloof, 'Braco', 4300 ft, Schelpe 51 19 (BOL); Lidgetton, Roberts 87 1 (PRE); Zwaartkop (CB), Sim s.n. PRE-9045 (PRE); Zwaartkop, 4500 ft, Sim s.n. (NU); Pietermaritzburg, Ferncliff Nature Reserve, Crouch 593 (NU). 3029 (Kokstad): Kokstad (CB), McLoughlin 753 (BOL). 3127 (Lady Frere): Cala (DA), Young 5 1 1 (PRE); Engcobo (DB), McLoughlin s.n. CH7677 (PRE). 3128 (Umtata): Maclean farm Woodcliffs (AB), Roux 2482 (NBG). 3129 (Port St Johns): Port St Johns (DA), Wager s.n. CH2905 (PRE). 3226 (Fort Beaufort): Katberg Forest Reserve (BC), Roux 2700 (NBG): Hogsback Forest Reserve, Fern Walk, 800 m (DB), Dahltrand 1853 J.P. ROUX (PRE); Hogsback, Zingcuka Forest, Roux 2414 (NBG). 3227 (Stutterheim): Stutterheim, Isidinge Forest (CA), Roux 1982 (NBG); Keiskamma Hoek, Ely 526 (PRE); Cathcart, Fort Cunnyngham Forest Reserve (CB), Roux 243 1 (NBG); Pirie, Sim s.n. TM5 14 (PRE); Kingwilliamstown, Pirie Forest along Amatola trail, Roux 2708 (NBG). 3325 (Port Elizabeth): Johana Kloof (BC), Breutel s.n. (L). 3419 (Caledon): Riviersonderend, farm 'Oubos' (BB), Roux 2585 (NBG). SUDAN: Gilo, Imatong Mountains, Ngairigi River, 5000 ft, McLeay 455 (K). TANZANIA: Station Kyimbila, Fundort Rungwe, 1300 m, Stolz 889 (B, L, P, S, WAG); Kilimanjaro, environs de Kibosho, 2500 m, Daubenberger s.n. PRE-6788 (PRE); Kilimanjaro, Kibosho, 2000-4000 m, Daubenberger 35 (B, BR, P, PRE, S); Mt. Meru, NE end of the caldeira wall, c. 8500 ft, Greenway & Fitzgerald 13613 (K, PRE); Kilimanjaro-Slid, c. 1900 m, Schlieben 4596 (BOL, BR, PRE, SRGH); region de Kilimanjaro, environs de Kibosho, Kilema-Machame, Daubenberger s.n. (BR, P); Kilimanjaro, 2800 m, Alluaud 310 (P); British East- Africa, forets de Lamoru, Le Petit s.n. (P); forets de plateau Kikuyu, 2000 m, Le Petit s.n. (P); Usambara Mountains, Mahali Mountains, 6000 ft, Newbould & Jefford 1731 (K); Morogoro, Glover 268 (K); Marangu, SE Kilimanjaro, 4600 ft, Beesley 14 (K); Mbeya District, Kikondo camp, Poroto Mountains, 1950m, Richards 13972(B,BR, K); Moshi District, Kilimanjaro, c. 1900 m, Schlieben 4596 (K); Arumeru District, banks of Engare Olmotonyi River, c. 4 km N. of Olmotonyi Forestry Institute, Mtui 143 (K); Mount Meru, Engarenanyuki, 7600 ft, Vesey- FitzGerald 3031 (K); Mbeya District, Mount Kikondo, 6500 ft, M.R. 13972 (K); Mount Meru, end of Olmotonyi, Schippers T778 (WAG); Mount Meru, 2090 m, Schippers T729 (WAG); South Pare Mountains, Mugambo Forest Reserve, 1480 m, Schippers T95 1 (WAG); W. Usambara Mountains, on hill above Shume Forest Meteorological Station, 2050 m, Schippers T1506 (WAG); Kilimanjaro, oberhalb Marangu, Volkens 1266 (B); Kilimanjaro, 1900 m, Schlieben 4596 (B); Usambara, Lutindi, Liebush s.n. (B); Kondoa- Frangi, Ndiomeberg, 1800 m, Ledemann s.n. (S); Kilimanjaro, above Marungu, 2000 m, Pedersen 527 (BR); Morogoro Mountains, 2300 m, Chisongela 9 (BR). UGANDA: near Luhiza-Kigezi, 7000 ft, Rose 10311 & 10312 (K); forest near Mt. Debasien, 6000 ft, Eggeling 2683 (K); Ruwenzori Mountains, 7000 ft, Hazel 1 14 (K); Luhiza-Kigezi, 7000 ft, Rose 10299B (K). ZAMBIA: Nyika Plateau, Chowe Forest, 2100 m, Dowsett-Lemaire 220 (K). ZIMBABWE: near Umtali, Holland s.n. (NBG); Vumba Mountains, near Umtali, 6000 ft, Obermeyer 2099 (K, PRE); Melsetter, Bridal Veil Falls, Jacobsen 3087 (PRE); Melsetter, in gulley border of 'Skyline' & 'Thornton' areas, Chase 7482 (BOL, K); Umtali, Banti south, 5800 ft, Jacobsen 3864, 3879 (SRGH); Inyanga, above Pungwe rest hut 2, 5300 ft, Chase 5655 (BOL, PRE, SRGH); Melsetter, Musapa mountain, Grosvenor 264 (BOL, SRGH); Melsetter, Gwendingwe, Muller 2880 (SRGH); Umtali District, Cashel, Black Mountain Inn, Chase 4021 (NU); Melsetter District, Bridal Veil Falls, Chase 4020 (NU); Inyanga, Patterson 24 (GRA); Inyanga, Pungwe Rest Huts, 5300 ft, Schelpe 5679 (BOL). WITHOUT EXACT LOCALITY: Natal, Tyson s.n. CH2168 (PRE); Natal, Gerrard 1931 (P); Zululand, Gerrard & McKen s.n. (P); Zimbabwe, Wild 1470 (K); loco incerto, Mann 2067 - pro parte (K); near Bamenda, 7500 ft, Migeod3&3 (K); Natal, sine coll. s.n. (NBG); Natal, Buchanan 27 (B-only) (M); Natal, Plant 328 (B); Natal, Buchanan 75 (B); loco incerto, Bergius s.n. (B); Rebfall bei Gaffat, Haidner s.n. (B, S); Kissenye, Ninagongo, 2500- 2900 m, Mildbraed 1341 (B); loco incerto, Hoist 3837 (B); loco incerto, Sim s.n. CH4171 (PRE); Gold Fields, Ayres s.n. (NH); Burungo, De Witte 1472 (BR); Natal, Holub s.n. (BR); Kikuku, 1750 m, Ban 367 (BR); Natal, Buchanan s.n. (BOL). Polystichum transvaalense and P. wilsonii are often confused. Pichi Sermolli (1977, 1985) ascribed material of P. transvaalense to P. fuscopaleaceum Alston var.fuscopaleaceum, while Schelpe (1967, 1970, in part) and Jacobsen (1978) ascribed material of this species to P. setiferum var. fuscopaleaceum (Alston) Schelpe. Jacobsen & Jacobsen (1989), however, considered P. fuscopaleaceum and P. transvaalense to be conspecific. Aware that two forms exist, Schelpe (1967) concluded that no clear differentiation at the specific level was possible and considered plants with dark stipe base paleae as P. setiferum var.fuscopaleaceum. This classification was largely followed by Jacobsen (1978), al- though he considered the high elevation collections a Drakensberg form. Although he ascribed several collections to this form he refrained from giving it any formal taxonomic status. Pichi Sermolli (1977), however, considered P. fuscopaleaceum distinct from P. setiferum. He also recognized two 'altitudinal vicariants' with P. fuscopaleaceum var.fuscopaleaceum occurring at lower elevations than P. fuscopaleaceum var. ruwensoriense, a subdivision he re- tained in 1985. I consider the two groups sufficiently distinct to warrant specific status, a conclusion supported by the discovery of a sterile hybrid between these putative parents. DIAGNOSTIC FEATURES AND RELATIONSHIPS. Diagnostic of Polystichum transvaalense is its confinement to moist forests, the presence of up to 17 caespitosely arranged, suberect to arching fronds that may reach a length of up to 1 .045 m on a short erect to suberect rhizome, the stipe, rachis and pinna-rachises bearing mostly ferrugineous, twisted and somewhat shrivelled, proximally lacerate paleae terminating in an acicular cell, and the mostly fimbriated but often erose and rarely repand indusium. The perispore is highly porate. Polystichum transvaalense appears to be most similar morpho- logically to P. wilsonii H. Christ; the two belong to section Lasiopolystichum. An analysis of the differences between these species is provided under P. wilsonii. VARIATION. Polystichum transvaalense shows considerable varia- tion in the length of the frond, stipe, lamina and basal pinna (Table 2). This may be ascribed to the diverse altitudes, climates and vegetation types it occupies throughout its broad range. The species, however, shows little variation in pinnule outline and palea struc- ture, distribution and density. Stipe-base paleae are mostly ferrugineous, but in rare cases the larger paleae are densely impregnated with secondary compounds giving them a dark brown colour. The indusium also shows significant variation in size, shape and the presence or absence of central processes. Basal pinnae may be deflexed or not. Possible causes of these variations in plants occurring in close proximity under similar growing conditions remain unknown. Table 2 Variation in frond, stipe lamina and basal pinna length in Polystichum transvaalense N.C. Anthony. Ramge (mm) x (S.D.) Frond 212-1045 713.9(204.1) 41 Stipe 77-535 283.3(101.6) 42 Lamina 135-670 422(116.1) 52 Pinna 25-140 86.3 (27.2) 52 DISTRIBUTION AND ECOLOGY. Polystichum transvaalense is widely distributed in temperate and tropical Africa. The distribution largely follows the escarpment and mountain ranges on the eastern parts of the continent. In South Africa it occurs from the Drakensberg foothills in the Eastern Cape along the KwaZulu-Natal Drakensberg escarpment, the Eastern Cape and southern KwaZulu-Natal mid- lands, the Free State-KwaZulu-Natal and Mpumalanga-Northern Province escarpments to the Soutpansberg in the Northern Province. A single collection is also known from the foothills of the Riviersonderend Mountains in the southern Cape. In Zimbabwe it is found in the Chimanimani and Vumba Mountains extending to the Zomba Plateau, the Kirk Mountains and the Nyika Plateau in Malawi and Zambia. In Tanzania it occupies the Uluguru- and POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 53 Usambara Mountains, Mt. Meru and Mt. Kilimanjaro. Further north it occurs in the mountainous areas of Kenya, Uganda, Ethiopia, the Imatong Mountains in Sudan and the Kivu Ridge in the Democratic Republic of Congo. A disjunction in this pattern arise in that P. transvaalense is also found on Mt. Cameroon and Bioko in the Gulf of Guinea. The lithology, climate and vegetation associated with Polystichum transvaalense varies considerably throughout its range. In the south- ern Cape it occupies isolated forest patches in Mesic Mountain Fynbos (Moll et al., 1984) at c. 365 m in acidic sandy soils and on rocks of the Table Mountain Sandstone formation. Polystichum transvaalense is an exclusively forest growing species, generally growing as individuals on streambanks or on rocks along streams but rarely also as low-level epiphytes. Polystichum transvaalense occurs in several forest types as defined by Acocks (1988). In the Eastern Cape it is found in Pondoland Coastal Plateau Sourveld forests and rarely in Typical Coast-belt forests. In the Natal mid- lands and Drakensberg escarpment where it may appear at elevations up to 1840 m it occurs in 'Ngongoni Veld forests and Highland Sourveld forests. In Mpumalanga these forests are replaced by Northeastern Mountain Sourveld forests that extend to the Soutpansberg. North of the Limpopo the species is found at eleva- tions ranging between 1300 and 2825 m in largely Undifferentiated Afromontane forests, as defined by White (1983), but often also in single-dominant Afromontane forests such as Juniperus procera forests at 2400 m in Ethiopia and in Afromontane bamboo at 1 828 m on Mt. Malati in Tanzania and at 2745 m at Kinangop in Kenya. 8. Polystichum wilsonii H. Christ in Bot. Gaz. 51: 353 (1911). Type: China, Szechuan Province, Mupin, woodlands, 4000- 6000 ft, Wilson 2614 (BM!-holotype). Fig. 9. Polystichum lobatum var. ruwensoriense Pirotta in L.A. di Savoia, // Ruwensori I: 478 (1909). Type: Ruwenzori, nella foresta scendendo da Kichuchu a Nakitava, Roccati et Cavalli-Molinelli s.n. (TO-holotype). Polystichum aculeatum var. mildbraedii Brause in Bot. Jahrb. Syst. 53: 379 (1915). Type: Fernando Po (Bioko), Nordseite des Pics Sta. Isabel oberhalb Basile, Grasflur-Region des Gipfels mit viel Ericinella, zwischen Gras, c. 2700 m, Mildbraed 7180 (B!- holotype). Polystichum aculeatum var. rubescens Bonap., Notes Pterid. 14: 214 (1923). Type: Tanzania, Kilimanjaro, zone super des forets, 2760 m, Alluaud 48 (P!-holotype). Polystichum aculeatum var. stenophyllon Bonap., Notes Pterid. 14: 215 (1923). Type: Kenya, Mont Kenya, versant ouest, foret interfere, 2400 m, Alluaud 241 (P!-holotype). Polystichum fuscopaleaceum Alston in Bol. Soc. Brot. ser. 2, 30: 22 (1956). Type: Cameroon, Victoria District, Cameroon Mountain, SW of hut 2, in gully woodland, 9100 ft, Keay FHI 28602 (BM!- holotype). Polystichum setiferum var. fuscopaleaceum (Alston) Schelpe in Bol. Soc. Brot. ser. 2, 41: 216 (1967). Polystichum fuscopaleaceum var. ruwensoriense (Pirotta) Pic.Serm. inWebbia32:90(1977). Polystichum alticola Schelpe & N.C. Anthony in Contr. Bolus Herb. 10: 144 (1982). Type: South Africa, Ladismith, Swartberg, Toverkop, 2160 m, Esterhuysen 26699 (BOL!-holotype; NBG!, PRE!, isotypes). Plants terrestrial or epilithic. Rhizome short, to 130 mm long, erect to suberect, to 10 mm in diameter, rarely branched, set with roots, crowded, persistent stipe bases, and paleae; paleae broadly attached, castaneous, chartaceous, narrowly linear, the margins with small, widely spaced cellular outgrowths, the apex mostly terminating in an acicular cell, rarely in a small thin-walled cell, to 12 x 1 mm. Fronds caespitose, 8-12 per plant, suberect to arching, to 1.05 m long; stipe proximally castaneous, stramineous distally, adaxially sulcate, to 450 mm long x 5 mm in diameter, sparsely to densely paleated; larger paleae broadly attached, often slightly bullate, spreading, extending to the rachis, concolorous or bicolorous, chartaceous to crustaceous, broadly ovate-acuminate to ovate-acu- minate, cordate to cordate-imbricate, the margins widely to closely fimbriate, fimbriae generally straight, the apex entire, terminating in an acicular cell, to 23 x 9 mm; smaller paleae apically or basally directed, stramineous, chartaceous, narrowly triangular to subulate, short-stalked, often somewhat auricled, the margins proximally with long straight, angular or curved outgrowths, distally with few widely spaced, short or long marginal outgrowths, the apex entire, terminating in an acicular cell, to 13 x 7 mm: lamina 2-pinnate, with up to 29 free pinna pairs, herbaceous to firmly herbaceous, pale to dark green adaxially, paler abaxially, narrowly elliptic, to 625 mm long, the proximal pinnae reduced, deflexed: rachis stramineous, adaxially sulcate, densely set with paleae similar to but smaller than those on the stipe, paleae restricted to the abaxial surface, to 9 x 3 mm: pinnae 1 -pinnate, with up to 12 free pinnule pairs, proximally widely spaced, distally closely spaced and somewhat overlapping, folded ventrally along the rachis (conduplicate), narrowly triangular to oblong-attenuate, the proximal pinnae to 88 mm long x 20 mm wide: pinna-rachis stramineous, adaxially sulcate, densely paleated; paleae short- stalked, narrowly ovate to narrowly triangular, the margins proximally with long straight or angular outgrowths, apically with few widely spaced short or long outgrowths, the apex entire, terminating in an acicular cell: pinnules asymmetric, acroscopically auriculate, narrowly trullate to trullate, to 12 mm long, serrate, long- aristate; adaxially with straight or slightly twisted filiform paleae, simple or proximally with short straight or curved marginal outgrowths, the apex terminating in an acicular cell; abaxially with straight or proximally somewhat twisted, subulate-hastate paleae, the margins with short straight or angular outgrowths at the base, the apex entire, terminating in an acicular cell. Venation raised. Sori circular, c. 1 mm in diameter, terminal or near terminal on abbrevi- ated vein branches: sporangium with 1 1-(15)-24 indurated annulus cells; stalk eglandular: indusium stramineous, peltate, circular or reniform, repand to erase, often with small central processes, per- sistent, the maximum radius 0.51-(0.75)-1.09 mm. Spores 64 per sporangium, brown, the perispore smooth or tuberculate, spiculate, closely perforated, the exospore 32-(41.74)-52 x 24-{30.16)-40 um. Chromosome number 2n=164. MATERIAL EXAMINED BIOKO: Fernando Po, Mann s.n. (K); cratera del pico Basile, km 23, junto a la cumbre, 3000 m, Carvalho 3652 (B, BR). CAMEROON: Mt. Cameroon, 3700 m, Breteleret al. 69 (K, P, WAG); Mt. Cameroon, 1950 m, Breteleret al. 75 (K, P, WAG); Bambutos, 2600 m, Felix 5430 (P); Mt. Cameroon, 3600 m, Annet 126 (P); Cameroon, mont versant, 3000 ft, Meurillon 1 158 (BR, K, P); piste du village d'Okon au mert Okon, 3008 m (45 km SSO de Nkambe), Letouzey 8940 (K, P); Mt. Cameroon, haut plateau, 3600 m, Annet 128 (P); Bambutos, station mi-ombragee, vers 2300 m, sine coll. 30 (P); Mt. Cameroon, 7-10000 ft, Mann 1376 (K); Cameroon Mountain, above 2nd hut, 12000 ft, Hutchinson & Metcalfe 48 (K); Mt. Cameroon, 11000 ft, Steele 22, 27 (K); Mt. Cameroon, 11000 ft, Migeod 190 (K); Mt. Cameroon, oberhalb Buea, 2800 m, Mildbraed 10883 (B, K); Buea, 3000 m, Preuss 787 (B), 788 (B, S); Kamerun-Berg, standort iiber Buea, unteren Fako Plateau, 2800 m, Mildbraed 3377 (B); Kamerungebirge, Buea, Deistel s.n. (B); Kamerun-pitz, 3500-3600 m, Bornmuller 26 (B); Mt. Cameroon, 3800 m, Hintz 29 (B); Buea, Wonjombia faco, Reder 1026 (B). 54 J.P. ROUX Fig. 9 Polystichum wilsonii. A, proximal part of lamina; B, rhizome; C, adaxial surface of pinnule; D, abaxial surface of fertile pinnule. All drawn from Roux 2529 (NBG). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 55 DEMOCRATIC REPUBLIC OF CONGO: Kivu District, Virunga west, Nyamuragira, Stauffler 178 (BR, PRE); P.N.A. Kabara, flanc N. du volcan Karisimbi, 3000 m, Jean Louis 5301 (BR, K, P); Volcan Karisimbi (au NE du lac Kivu), 3500 m, Humbert 8563 (BR, P); Kivu, Volcan Mikeno, 2500-3400 m, Humbert 8010 (BR, P); Mt. Kinangop, 2800 m, Alluaud 262 (P, S); P.N.A. Nyamlagira, 3000 m, Germain 3476 (BR); Kivu District, SW side of Mt. Mikeno, 10500 ft, Chaplin 373 (BR); 129'S, 2926'E, Goma territory, versant ouest du Karisimbi, 3360 m, Bamps 2984, 2995 (BR); Pare National Albert, ruisseau affl. de la Nososa, (a Test de Mahungu), 3180 m, Fredericq 9152 (BR); Viroenga Park, Karisimbi-massif, 3465 m, Van der Veken 9130, 9135 (BR); Kivu, upper Ruamoli Valley, 1 180 ft, /to 77 8 (BR); Viroenga Park, Karisimbi-massif, 3 km van de gite Rukumi, 3330 m, Van der Veken9\40 (BR); massif du Karisimbi, a 500 m env. du gite de Rukumi, 3500 m,Auquier 2290 (BR); Kivu Province, upper Ruamoli Valley, 12 500 ft, Ross 743 (BR); Pare National Albert, selle de Kabora entre le Karisimbi et le Mikeno, 3000-3100 m, Lebrun 7344 (BR). ETHIOPIA: Bale Region, 45 km N. of Goba, Sannetti Plateau, 3900 m, Tadesse 5545 (ETH); SE of Dinsho on road from Goba to Shashemene near proposed HQ of Bale National Park, 10400 ft, Gilbert 1812 (ETH, K); Bale Province, Bale Mountain National Park, E. of Garba Goracha camp, 4070 m, Hedberg 5649 (ETH); pass just N. of the summit of Cara Mulatta Mountain, c. 10200 ft, Burger 1480 (ETH); Choke Mountain, Gojjam, vicinity of the upper Ghiedeb Valley, Flenley & Evans 327 (ETH, K), Begemdir Province Simien, Buahit, 3870 m, Hedberg & Aweke 5461 (ETH); Bale Mountains, Finchaya Habera, 3510 m, Miehe 266, 335 (ETH); Bale Mountains, above Goba, 3500 m, Miehe 3086 (ETH); Bale Mountains, above Rira, 3530 m, Miehe 2343 (ETH); Shoa Province, Arussi Mountains, 35 km S. of Mount Chillalo 32 km on track to Ticcio via Robie turnoff, 35 km S. of Asella, 3275 m, Ash 2330 (ETH); Bale region, Mendeyou Auraja, c. 5-7 km on Fincha Haberra-Soddota track, 3500-3580 m, Tadesse 7813 (ETH); Shoa, Lake Wonchi, outer rim of caldeira, Gilbert & Tewolde 3279 (ETH, K); Bale region, Mendeyou Awraja Fincha Heberra, 3490-3510 m, Tadesse 7713 (ETH); in rupibus umbrosis Demeski, 10500 ft, Schimper 244 (P); Matssehe Dedschem, 12000 ft, Schimper 1398 (P); Arussi Prov., Chillalo Awraja, Galama Mountains (30 km ESE of Asella) c. 3 km E. of Boraluco, 3750 m, Hedberg 4233 (K, PIC.SERM.); Bale region, Dello Awraja, c. 3.7 km N. of Kecha towards Rira, 2620 m, Tadesse 5143 (ETH); Shewa region, Wonchi Mountains, edge of volcanic crater 20 km to SSE of Ambo, Pavlov & Petelin 138 (ETH); in regio media montis, Schimper 180 (B, K, P, S); Begemder Province, Semian Mountains, De Wilde 175 (BR, WAG); Mt. Borullccu, along road to Ticco c. 30 km SE of Asella, c. 4000 m, De Wilde 9039 (WAG); c. 25 km SE of Asella, W. slope of Mt. Boruluccu, c. 3800 m, De Wilde 8089 (WAG); c. 3 km E. of Asella, c. 175 km SSE of Addis Ababa, W. slope of Mt. Cilalo, c. 2700 m, De Wilde 6623 (BR, WAG); Shoa Province, Menagesha State Forest on the W. slope of Mt. Wuchacha, 2600 m, Friis 1 209 (K); c. 1 75 km from Addis Abeba on Dessie road, 10500 ft, Gilbert 454 (K); Bale region, 10-15 km SE of Goba on road towards Masslo, 3200-3400 m, Thulin 3678 (K); Gara Mullato Mountains, 10800 ft, Burger 1907 (K); Scioa, Monte Wochacha, 3250-3300 m, Pichi Sermolli 6740 (B, BR, K); Mussolini Pass, between Dera Sina and Debra Berhan, c. 3000 m, De Wilde 9654 (WAG); Ethiopia, Schimper 1398 (B); ad rupes locis humidis umbrosis pr. Demerts, 10500 ft, Schimper 244 (B); Arussi, Catena dei Monti Galamo-Sagatu, c. 3100 m, Pichi Sermolli 6828 (BR). KENYA: Mt. Kenya, forest end, 9600 ft, Mclaughlin 676 (BOL, PRE); Mt. Elgon, versant est, Arambourg et al. 134, pro parte (P); W. slopes of Mt. Kenya, along the trail from West Kenya Forest Station to summit, c. 3630 m, Mearns 1421, 1502, (P); W. slopes of Mount Kenya, along trail from West Kenya Forest Station to summit, c, 3000 m, Mearns 1702 (B, P, S); Mt. Kenya, c. 7500 ft, Schippers K164 (WAG); Aberdares National Park, c. 10500 ft, Schippers K110 (WAG); Aberdares National Park, c. 10300 ft, Schippers K78 (WAG); Shira Plateau, W. Kilimanjaro, 3200 m, Schippers T1052 (WAG); Mt. Meru, c. 2000 m, Schippers T777 (WAG); Mt. Kenya, Kinangop, Aberdare, 8800-8900 ft, Chandler 2266 (K); Aberdare Moun- tains, James s.n. (K); Aberdare Mountains, Ramsden s.n. (K); Aberdare range, near W. part of the Nyeri track, 3 1 00 m, Hedberg 1533 (K); near Molo, Mau Forest, 8000 ft, Gardner 975 (K); Narok District, 20 miles from Olokurto on road to Elburgon, c. 9600 ft, Glover et al. 1096 (K); SE Aberdares, Kitikuya, 8500 ft, Gardner s.n. (K); Rift Valley, Nakuru District, E. Mau Forest Reserve, 2750 m, Geesteranus 5908 (BR, K, L, PRE, S); Mt. Elgon, E. slope above Tweedie's saw-mill, 2550 m, Hedberg 68 (K, S); North Forest, 9200 ft, Schippers K364 (WAG); Mt. Aberdare, c. 3200 m, Fries 2644 (B); Mt. Elgon, 2700 m, Gravik s.n. (S); Mt. Elgon, 3800 m, Gravik s.n. (S); Mt. Kenya, Sagana Valley, 10500 ft, Schelpe 2713 (BR); Mt. Aberdare, pr. 'West Kenya Forest Station', 2350 m, Fries 775 (BR); Mt. Kenya, 10000 ft, Meyerscough K2, K3, K16, K18, K22, K26, K30, K31 (BOL). LESOTHO. 2828 (Bethlehem): Butha Buthe District, Khatibe B camp, 9500 ft (DC), Troughton B26 (GRA); Leribe, Dieterlen 167 (BOL, P, pro parte); 1 km from Moteng store, Roux 1 294 (NBG). 2927 (Maseru): between Blue Mountain Pass and Likholaneng, 8700 ft (BD), Schmitz 7266 (PRE); Morija, Dieterlen 1309 B-only (PRE). 2928 (Marakabei): Mamalapi, 8000 ft (AC), Jacot-Guillarmod 690 (PRE); Mamalapi, 9000 ft, Compton 21331, 21334, 21339 (NBG); hill at Bushmen Pass, beyond little Bokong, 9000 ft, Bevis 102 (PRE); Blue Mountain Pass, Roux 2227 (NBG); mountain road, 60 miles from Maseru, 8000 ft, Bowmaker 23 (BOL); mountain road, 38 miles from Maseru, 7500 ft, Bowmaker 25 (BOL); Lehaha-la-Sekhomgana, 9 1 00 ft (AD), Jacot-Guillarmod 206 (PRE); Semonkong, waterfall gorge, c. 7000 ft (CC), Davidson 3023 (PRE); Semonkong, at Le Bihan Waterfall, Roux 1493 (NBG). 2929 (Underberg): Between Mokhotlong and Sani top, 15 km from Mokhotlong, 2200 m (AC), Matthews 887 (NBG, PRE); 15 km past Thaba-Tseka turnoff on Sani road, Roux 1 344 (NBG); Sehlabathebe National Park (CC), Schmitz 7122 (PRE); Sehlabathebe National Park, Matthews 987 (NBG); Sehlabathebe area, on way to Devils Knuckles, 9500 ft, Davis 181 (NU); Sehlabathebe area, Devils Knuckles, c. 9000 ft, Davis 176 (NU); Sehlabathebe National Park, 2250 m, Hoener 1658 (BOL). SOUTH AFRICA. 2730 (Vryheid): Wakkerstroom, Oshoek, 6400 ft, (AD), Devenish 195, 638 (PRE). 2731 (Louwsburg): Nongoma, c. 1000 ft (DC), Tosh s.n. (NU). 2828 (Bethlehem): Clarence (CB), Van Hoepen s.n. TM 18230 (PRE); gully between the Witches and the Sentinel (DB), Roux 1906, 2529 (NBG); versant N. du Mont-aux-Sources, region de Witzies Hoek, c. 1800 m, Junod 14 (P); Royal Natal National Park (DB), Hafstrom & Acocks 1699 (PRE); Royal Natal National Park, Gudu Forest, Roux 2510a, 2511 (NBG); Mont-aux-Sources, 8000 ft (DD), Dyke 5489a (NBG); Royal Natal National Park, Plowmans Kop, Aerck 1966 (S); Mont-aux-Sources, 10000 ft, Sim s.n. TM521c (PRE); Mont-aux-Sources, Mogg 4222 (PRE); Mont-aux-Sources, 3100 ft, Marloth 2862 (BOL). 2829 (Harrismith): Harrismith, Platberg, Zig-Zag Pass, 1800 m (AC), Jacobsz 4715 (PRE); Harrismith, Platberg, Donkie Pass, 1850 m, Jacobsz 4729, 4730 (PRE); Platberg, 6800 ft, Roux 782 (NBG); Harrismith, Platberg, Roux 2521, 2524, 2526 (NBG); Harrismith, farm Bosch Hoek (AD), Roux 892 (NBG); Harrismith District, farm Klavervlei (CA), Roux 876 (NBG); Oliviershoek Pass, S. of Seheletwane,/tou*2516, 2517 (NBG); MnWeni Pass, 8000-9000 ft (CB), Esterhuysen 27838 (BOL); MnWeni area, Pinnacles Gully, 9000 ft, Esterhuysen 29595 (BOL); MnWeni area, Mbunduni scree, c. 6000 ft, Esterhuysen 27816 (BOL); Drakensberg, Injasuti area, 6500-8500 ft (CC), Esterhuysen 26045 (BOL, K, NBG, PRE); along Cathedral Peak path, 1550 ft, Goetghebeur457l (BR, PRE); Cathedral Peak, Ruch 2030, 2300 (PRE); Cathedral Peak area, 5000 ft, Harding 38 (NU); Cathedral Peak Forest, Killick 981 (NU). 2929 (Underberg): Cathedral Peak Forest Research Station, 6100 ft (AB), Killick 981 (PRE); summit of Cathedral Peak, 7700 ft, Schelpe p.30 (NU); Cathedral Peak area, Cleft Peak path, 8000 ft, Schelpe 557 (NU); upper Tsanatalana Valley, near Cleft Peak, 9800 ft, Schelpe 7227 (BOL); Champagne Castle, Bayer 1443 (PRE), 1445 (NU, PRE); Giants Castle (AD), Symons 134 (PRE); Mpendhle Distr., Mulangane Ridge, above Carter's Nek, 7000-7300 ft (BC), Milliard & Burtt 1 695 1 (BOL, NU), 1 6969 (BOL, NU, PRE); Mpendhle District, Highmoor Forest Reserve, ridge SE of Giants Castle, headwaters of Elandshoek River, c. 8100 ft, Milliard & Burtt 1 6 1 92 (BOL, NU); near Rosetta, 5000 ft (BD), Thode s.n. (NBG); Drakensberg Garden State Forest Reserve, 9500 ft (CA), Van Jaarsveld 6531 (NBG); Garden Castle Forest Reserve, Mlambonya Valley, 6200 ft, Hilliard & Burtt 14972 (BOL, NU); upper tributaries S. of Mkomazi River (CB), Hilliard & Burtt 15853 (NU, PRE); Bamboo Mountain, McClean 684 (PRE); Sani Pass, wet slope below waterfall, 6900 ft, Hilliard & Burtt 1 7976 (NU, PRE), 1 7983 (BOL, NU, PRE); Sani escarpment, c. 9000 ft, Marker s.n. (GRA); Underberg District, 5-7 miles NNW of Castle View farm, headwaters of Mlahlangubo River, 8500 ft, Hilliard & Burtt 15331 (BOL, K, NU); headwaters of Mlahlangubo River, c. 7800 ft, Milliard & Burtt 13714 (NU); Underberg District, Gxalingenwa Valley between Sani Pass and Polela Valley, 7400 ft, Milliard & Burtt 1 7 1 99 (BOL, NU); Ndumeni area (CC), Everson s.n. (BOL); 56 Bulwer (DD), Allsopp 850, A-only (NU); Bulwer, Henkel s.n., A-only (NU); Xumeni Forest, Rycroft 519 (NU). 2930 (Pietermaritzburg): Nottingham Road District, 'Drayton', 5400 ft (AC), Smith 147 (NU); York, 'Benuie', c. 4000 ft (AD), Fisher 1040 (NU); Impendhle, Boston, 4500 ft (CA), Beanie 77 (NU); Pietermaritzburg, Zwaartkop (CB), Sim s.n. (NU, PRE). 3027 (Lady Grey): Lady Grey, mountain left of summit of Jouberts Pass on road to Barkley East (CA), Roux 1 136 (NBG); Wittebergen, Ben McDhui, 9550 ft (DB), Galpin 6934 (BOL, GRA. PRE), 6935, 6939 (BOL, PRE); road between Naude's Nek and Ben McDhui, Roux 1180 (NBG); Barkley East District, Ben McDhui, Bell River Gorge, c. 8000 ft, Milliard & Burn 16526 (BOL, K, NU); zwischen Passtrasse Maclear und Naude's Nek, Werdermann & Oberdieck 1118 (B); Barkley East District, Ben McDhui, 9550 ft, Galpin 6939 (B); Ben McDhui, c. 9000 ft, Milliard & Burn 16406 (BOL, NU); Ben McDhui, 8900 ft, Milliard & Burn 16495 (NU). 3028 (Matatiele): near summit of Ongeluks Nek Pass (AD), Roux 1383 (NBG); Rhodes, Naude's Nek Pass (CA), Roux 2475, 2477 (NBG). 3029 (Kokstad): upper slopes of Inungi Range, Matatiele, c. 5500 ft (CA), Acocks 12207 (PRE); Kokstad (CB), McLoughlin S38 (PRE); Mt Currie Nature Reserve, Kokstad, Crouch 511 (NU); Kokstad, McLoughlin 746, 753 (BOL). 3030 (Port Shepstone): Oribi Gorge (CB), Slinger59 (NU). 3127(Lady Frere): Barkley Pass between Elliot and Barkly East (BB), Roux 2469 (NBG); Bastervoetpad, between Ugie and Barkley Pass, Roux 247 1 , 2474 (NBG). 3128 (Umtata): summit of Biziya Mountain, 1250 m (AD), Stever 898 (PRE). 3225 (Somerset East): near Somerset East (DA), MacOwen s.n. (P). 3226 (Fort Beaufort): Upper Zwart Kei, Mount Hope farm, 5300 ft (BC), Galpin 5621 (GRA, PRE); Katberg Pass summit, farm Pleasant View, Roux 2698 (NBG). 3319 (Worces- ter): Hex River mountains, shale band between Buffels Dome and Milner Peak, 5000 ft (AD), Esterhuysen 28708 (BOL, NU, PRE); Roodeberg (Matroosberg group), 6000 ft (BC), Esterhuysen 27695a (BOL); Worcester Division, shale band below Milner Peak, 5000 ft (CB), Esterhuysen 14885 (PRE); Hex River Mountains, Moraine kloof, 4000 ft (DD), Esterhuysen 28075 (BOL). 3321 (Ladismith): Swartberg near Ladismith, Toverkop (AD), Esterhuysen 28241 (BOL). TANZANIA: Mt Meru, Arumeru District, Gereau 1623 (PRE); Kilimandjaro-Siid, Korongo, c. 3000 m, Schlieben 4869 (B, K, PRE, SRGH); Kilimanjaro, tra la Peters Hut a la Bismarks Hut, c. 2900 m, Pichi Sermolli 5 1 36 (BR, K, P); Ob. Urwald iiber Kibosho, c. 2800 m, Uhlig 1 85 (B, K); Mt. Meru, W. slopes above Olkakola Estate, 3300 m, Medberg 2306 (K, S); Kilimanjaro, above Marungu, c. 2 km from Peter's Hut, 3700 m, Hedberg 1288 (BR, K, S); Mbeya, Kilando, 8000 ft, Herb. I.R.L.C.S. 6700 (K); Kilimanjaro, Petershutte, 4100 m, Peter 1212 (B); Kilimanjaro, 2000-3000 m, Meyre s.n. (B); Kilimanjaro, Volkens 1 155 (B); Usambara, Hoist 3824 (B); Kissenye, Ninagongo, 3000 m, Mildbraed 1372 (B); NO Kivu, W. Kalago, c. 2300 m, Mildbraed 1651 (B); Arusha National Park, crater of Mt. Meru, below Njeku Hut, 2560 m, Pocs & Kornas 652 I/A (BR). UGANDA: Mt. Elgon, 9000 ft, Dummer3560 (BOL, K, NBG); Ruwenzori Mountains, Nyamagasani Valley, 12500 ft, Loveridge 197 (K, SRGH); Ruwenzori, Lanuri c. 3500 m, Bequaert 4544 (P); Ruwenzori, le vallee du Mobuku, Val de Kabuamba, 3500 m, Alluaud 274 (P); Ruwenzori, vail, du Mobuku, abri sous roche de Buamba, 3500 m, Alluaud 275, (K, P); Ruwenzori (Est), vallee du Mobuku, rocher de Kichuchu, 3000-3200 m, Alluaud 310 (P); Ruwenzori (Est), vallee du Mobuku, abri sous roche de Buamba, 3500 m, Alluaud 276 (P); Toro District, Ruwenzori, Bigo, 3350 m, Osmaston 3921 (K); Ruwenzori, Nyamudamba, c. 10000 ft, Scott Elliot 8094 (K); NE Elgon, Tweedie 2745 (K); Ruwenzori, Mijusi Valley, 3500 m, Medberg 613 (K, S); Mt. Elgon, c. 1 1 000 ft, Allen 3676 (K); on Elgon at Benet, 9100 ft, Eggeling 2454 (K); Mt. Elgon, Rose 10267 (K); Western Province, Bigo, R. Bujuku Valley, 3550 m, Osmaston 1738 (BR); Ruwenzori, c. 3500 m, Bequaert 4544 (BR). ZIMBABWE: Inyanga District, 6500 ft, Chase 5100 (NU); Inyanga, 7000 ft, Patterson 29 (GRA); Vumba Mountains, Umtali District, Eagle School road, Jackson 29 (GRA). WITHOUT EXACT LOCALITY: Natalia, Buchanan 83 (B); loco incerto, ex Herbario Natalensi, sine coll. s.n. (S); loco incerto, Buchanan s.n. TM522c (PRE); Natal, Medley-Wood s.n. TM520c (PRE); loco incerto, Dinter 575, A-only (B); Muhonora, 3500 m, De Wine 1962 (BR); Karisimbi (versant sud) nr. Biuri, c. 3000 m, De Wine 1246 (BR); South Africa, ?Rivier, Lincke 57 (BR); Basutuland, Koopoeitz s.n. (GRA); loco incerto, sine coll. s.n. NH-9784 (NH). J. P. ROUX Sledge (1973) cited Polystichum fuscopaleaceum as synonymous with P. setiferum var. nigropaleaceum (H. Christ) Sledge [= P. nigropaleaceum (H. Christ) Diels]. Christ (1893) described this variety from a single specimen collected by H.F. Blanford at 4000 ft in the Jumna valley between Mussoorie and Lokwah, western Himalayas. Sledge did not examine the type of this variety as he was unable to locate it. Fraser-Jenkins (Fraser-Jenkins & Khullar, 1985) reported he had studied the type in the Manchester Herbarium (MANCH). A Blanford specimen from the same locality has since been located in the Paris Herbarium (P!) and may serve as an isotype. This plant shows no clear affinity with either R fuscopaleaceum or P. setiferum, but rather to the P. luctuosum group (section Xiphopolystichum) as was suggested by Christ. Fraser- Jenkins & Khullar (1985) consider it synonymous with P. discretion (D. Don) J. Sm. DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum wilsonii and P. transvaalense occur sympatrically and often grow side by side. This has led to a great deal of confusion in separating the two taxa from one another. Pichi Sermolli (1977) referred to them as altitudinal vicariants, with P. wilsonii (as P. fuscopaleaceum var. ruwensoriense) occurring at higher elevations, having narrower blades, and a denser covering of wider, approximately rounded, acuminate paleae with those on the stipe being pale. In 1985 Pichi Sermolli added further observations as to how the two taxa differ. In var. ruwensoriense (= P. wilsonii) he found the apical part of the pinnae to be acute, moderately incised and usually provided with sori, whereas in var. fuscopaleaceum (= P. transvaalense) it is acuminate, deeply incised and devoid of sori. Jacobsen (1978), who refers to P. wilsonii as the 'Drakensberg Form' of P. setiferum var. fuscopaleaceum, provides some charac- teristics of the species and ascribes several collections in the National Herbarium, Pretoria (PRE) to it. Many of these collections, how- ever, belong to other species. Polystichum wilsonii is separated from P. transvaalense by the slightly shorter and narrower fronds, shorter stipe, a more pro- nounced reduction and deflexing of the basal pinnae, and in the paleae. Larger paleae in P. wilsonii extend from the stipe to the rachis, are generally complanate at maturity and somewhat pol- ished. The smaller paleae are more rigid with shorter and less divided marginal outgrowths. In P. transvaalense the larger paleae are mostly restricted to the stipe and proximal part of the rachis and become somewhat shrivelled at maturity. The marginal outgrowths on the proximal part of the smaller stipe paleae are also longer, more divided, and more twisted. Polystichum wilsonii, although often present in forests at lower elevations with P. transvaalense, is predominantly a high altitude species occurring in exposed conditi- ons. A natural hybrid between the putative species was described as P. x saltum (Roux, 1997a). Polystichum wilsonii forms part of the section Lasiopolystichum Daigobo assemblage of species. More recently (Kung & Zhang, 1998) P. wilsonii has been placed as a synonym of P. sinense H. Christ. I choose to maintain the two as distinct species pending critical study. VARIATION. Morphological variation in Polystichum wilsonii is mostly restricted to the larger paleae present on the stipe and abaxially on the rachis. Variation is most apparent in the size, density and colour of the paleae. Larger paleae are broad in plants growing in more exposed habitats; they are more densely set. In plants from deeply shaded forest habitats, however, the larger paleae cannot be readily separated from the smaller paleae, especially on the rachis. Palea size and density thus appear to be environmentally influenced. No correlation could be drawn between habitat and POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 57 palea colour, consistent with Schelpe's (1967) remark that forms with dark paleae intergrade with forms with pale stipe paleae. The large paleae are generally stramineous and concolorous. In some plants, however, the larger stipe paleae are bicolorous with the proximal central part of the paleae being densely impregnated with phenolic substances and castaneous. In some plants these bicolorous paleae are restricted to the proximal part of the stipe, whereas in others they may extend to the basal pinnae. More rarely the larger paleae on the rachis are nitid, densely impregnated throughout, almost black, and extend to the lower half of the rachis. Schelpe & Anthony (1986), in their key to the South African Polystichum species, used the direction and length of the pinnule aristae to distinguish between taxa. In some plants the basal basiscopic aristae of each pinnule may fold over the adaxial surface of the pinnule lamina, but in others they may not. Arista length also varies considerably. In some plants it may be relatively short but in others unusually long. In some plants the basal basiscopic aristae curve away from the pinnule lamina. Indusium size shows some variation with the margins ranging from repand to erose. DISTRIBUTION AND ECOLOGY. Polystichum wilsonii has a wide distribution ranging from Africa to the Uttar Pradesh mountains in northern India, and to Bhutan, China (Szechuan) and Taiwan (Ilan, Taichung, Hsinchu). In the study area P. wilsonii has a disjunct distribution. In South Africa it occurs on the southern Cape moun- tains, along the KwaZulu-Natal Drakensberg and into Lesotho, extending along the Free State-KwaZulu-Natal escarpment as far north as the Vryheid District. To the north it occurs in the mountain- ous areas of Zimbabwe, Tanzania, Kenya, Uganda, Ethiopia and the Kivu Ridge in the Democratic Republic of Congo. The species is also known from Mt. Cameroon and the island of Bioko, 32 km from the mainland in the Gulf of Guinea. It has furthermore been recorded from Grande Comore c. 300 km from the mainland in the Mozam- bique channel. Although the higher ground of the Zambezian Region, which includes Zimbabwe and Malawi, supports Afromontane plant communities (White, 1983), it is rare in this region with only one collection known from Zimbabwe. The lithology, climate and vegetation associated with Polystichum wilsonii vary considerably through its range. In the southern Cape P. wilsonii occurs at 1500-2000 m in acidic sandy soils derived from sediments of the Cape Supergroup. These soils support the unique Mesic Mountain Fynbos (Moll et al., 1984). This area is the only part of the distribution range of the species that experiences winter rainfall (April-September). In the Drakensberg Polystichum wilsonii is associated with the Clarens Sandstone formation, the Drakensberg Basalt Formation and the intrusive Karoo dolerites. At lower elevations in the Drakensberg (1250-1800 m) the species commonly occurs along streambanks or on rocks in Undifferentiated Afromontane forests and scrub forests confined to sheltered ravines and mountain slopes. These forests are largely associated with the Clarens Sandstone Formation. At higher elevations (> 1600- 1800 m) the Drakensberg Basalt formation and the intrusive Karoo dolerites are prevalent. These formations support the Themeda-Festuca Alpine veld (Acocks, 1988). Here P. wilsonii occurs among boulders along streams, in dry exposed rock crevices or in wet and shaded rock overhangs. The Ethiopian and Kenyan highlands, Mt. Elgon, Mt. Meru, Mt. Kilimanjaro, Mt. Cameroon, Bioko and the Comoro Islands are all of volcanic origin or consist in part of volcanic deposits. Many of the isolated mountains are still volcanically active today. Also the Kivu Ridge, which is largely composed of Precambrian rocks, has local exposures of volcanic deposits. On all these mountains the vegeta- tion diminishes in structure from the lower slopes to the summit. Local features such as aspect, exposure incidence of frost, depth of soil and overall patterns of climate contribute to modify the vegeta- tion (White, 1983). At these elevations the plants become smaller and the apical pinnae more pronouncedly conduplicate along the rachis. In tropical Africa Polystichum wilsonii occurs in a wide range of vegetation types. At lower elevations on Mt. Elgon (2550 m) and the Ethiopian highlands (2700 m) it occurs in Undifferentiated Afromontane forests. On the Ethiopian highlands the species also occurs in single-dominant Afromontane forests such as Juniperus procera forests on Mt. Wuchada (2600 m) and Hagenia abyssinica forests in the Bale Mountains. On Mt. Kenya it has been recorded from the Afromontane bamboo zone. On Mt. Kenya (3200 m), Mt. Meru (3300 m), Mt. Elgon (3500 m) and the Bale Mountains (3500 m) it occurs in Afromontane bushland and thicket. Again on the Bale Mountains (3500 m) and on the rim of the caldeira round Lake Wanchi (3650 m) it occurs in Afromontane and Afroalpine shrubland. With an increase in elevation the latter vegetation type is replaced by Afromontane and Afroalpine grassland. Polystichum wilsonii has been recorded from this vegetation type on Mt. Kilimanjaro (3000 m) and the Ethiopian highlands (3900 m). On Mt. Cameroon P. wilsonii has been recorded from 1950 m to 3800 m and on the island of Grande Comore from 1000 m to 1400 m. In both cases the plants were associated with lava flows. Growth in Polystichum wilsonii shows a degree of seasonality. In the Drakensberg several new fronds are produced almost simultane- ously at the onset of the rainy season in November. This pattern is retained in cultivated plants. Several Afroalpine vegetation types are subject to periodic burning. Fires, however, appear to have little or no damaging effect on the rhizomes. 9. Polystichum x saltum J.P. Roux in Bot. J. Linn. Soc. 124: 376, fig. 1 (1997). Type: South Africa, KwaZulu-Natal. 2828 (Bethle- hem): Royal Natal National Park, Gudu Forest, near Gudu Waterfall, c. 1800 m (DB), Roux 2510b (NBG!-holotype). Plants terrestrial or epilithic. Rhizome erect to suberect, to 20 mm in diameter, densely set with roots, closely set persistent stipe bases, and brown to ferrugineous paleae. Fronds caespitose, to 19 per plant, erect to arching, to 400 mm long: stipe proximally stramineous, greenish distally, shallowly sulcate adaxially, to 1 10 mm long, to 4 mm in diameter, densely paleated, the paleae of two types; larger paleae broadly attached, brown to ferrugineous, chartaceous, lan- ceolate to narrowly lanceolate, cordate, often slightly auriculate, the margins closely to widely set with short and long, straight or curved, often forked projections, the apex always terminating in a long acicular cell, to 1 1 x 3.5 mm; smaller paleae brown to ferrugineous, chartaceous, short- or long-stalked, narrowly triangular, cordate to cordate-imbricate, the proximal margins closely set with short and long, straight or angular, simple or branched projections, the number and size of the projections reduced distally, the apex usually simple, terminating in a long acicular cell, to 6 x 1 mm: lamina 2-pinnate, narrowly ovate, to 300 mm long, with up to 17 free pinna pairs: rachis greenish throughout, adaxially shallowly sulcate, densely paleated, the proximal paleae of two types; larger paleae similar to those on the stipe and reduced in size towards the middle of the lamina; smaller paleae short- or long-stalked, ferrugineous, chartaceous, narrowly lanceolate to narrowly triangular, slightly cordate to cordate-imbricate, often slightly auriculate, the margins proximally with short and long, straight or curved, often branched projections that are reduced in size and frequency distally, the apex usually simple, terminating in a long acicular cell, to 6 x 1 mm: pinnae 1 -pinnate, long-stalked, proximally widely spaced, slightly 58 J.P. ROUX reduced, deflexed, with up to 10 free pinnule pairs, slightly overlap- ping distally, narrowly ovate to oblong-attenuate, to 75 x 24 mm: pinna-rachis greenish, adaxially shallowly sulcate, sparsely paleated; paleae ferrugineous, chartaceous, long-stalked, narrowly triangular to narrowly oblong, cordate to cordate-imbricate, the margins proxi- mally with long, straight or twisted, simple or forked projections reduced in size and frequency distally, the apex usually simple, terminating in a long acicular cell, to 3.5 x 0.5 mm: pinnules opposite to alternate, firmly herbaceous, pale- to olive-green adaxially, slightly paler abaxially, the proximal acroscopic pinnule usually slightly longer than the next, asymmetric, ovate to ovate- rhomboid, basiscopically narrowly cuneate, acroscopically broadly cuneate and auriculate, the auricle often incised midway to costa, serrate to doubly serrate, long-aristate, to 18 mm long; adaxially sparsely set with a few twisted paleae chiefly along proximal part of costa, stramineous, chartaceous, filiform or with a few short mar- ginal projections near the base, the apex always terminating in a long acicular cell, to 2 mm long; abaxially sparsely paleated, stramineous, chartaceous, long-stalked, narrowly deltate to filiform, the margins proximally with long, curved or angular, simple or branched projec- tions, the apex simple, always terminating in a long acicular cell, to 3 mm long. Venation raised abaxially. Sori circular, c. 1.2 mm in diameter, terminal or near terminal on abbreviated vein branches, essentially uniseriate, discrete: sporangium with 13-(14)-17 indurated annulus cells; stalk eglandular: indusium brown, chartaceous, persistent, peltate, circular, frequently with long cen- tral processes, fimbriate, the maximum radius 0.8-(0.9)-1.02 mm. Spores aborted, the perispore closely perforated. Chromosome number 2n=164, meiosis yielding univalents and bivalents (Roux, 1997a). DIAGNOSTIC FEATURES. Polystichum x saltum closely resembles P. wilsonii in size, frond and pinnule morphology, and to a certain degree in the characteristics of the paleae. The erose to fimbriate indusium, however, is more characteristic of P. transvaalense. The mean guard cell length, the adaxial epidermal cell length, and the mean maximum radius of the indusium are anomalous - being larger than that of either progenitor (Roux, 1997a). Perhaps the most distinctive diagnostic feature of the taxon is the varying number of aborted spores borne in the sporangia. DISTRIBUTION AND ECOLOGY. Polystichum x saltum is currently known from only one forest in the foothills of the KwaZulu-Natal Drakensberg. This forest fragment forms part of the Highland Sourveld vegetation type (Acocks, 1988) and is nestled in a shel- tered ravine on a steep mountain slope. Forests of this type, situated at 1500 to 1700 m, are mostly cool and moist throughout the year, even though most of the rainfall occurs during the summer (Novem- ber-March). Like its putative parents, P. x saltum also occurs on moist moss-covered boulders along streams or on the forest floor in permanently moist conditions. Polystichum wilsonii, a taxon mostly associated with higher elevations where it occurs in more exposed habitats, frequently grows sympatrically with P. transvaalense in forests along the Drakensberg. lO.Polystichum marionense Alston & Schelpe in J. S. African Bot. 23: 106, fig. la, t. 34 (1957). Type: Marion Island, Moseley s.n. (BM!-holotype). Fig. 10. Plants terrestrial or epilithic. Rhizome short, decumbent, branched, stoloniferous, to 5 mm in diameter, set with roots, closely spaced persistent stipe bases, and paleae; paleae sessile, ferrugineous to castaneous, scarious. Fronds crowded, to 8 per plant, erect, to 940 mm long: stipe proximally castaneous, distally stramineous, adaxially shallowly sulcate, to 290 mm long x 4 mm in diameter, proximally close-set with unicellular pyriform glands, also sparsely to densely paleated; paleae sessile, ferrugineous to castaneous, scarious, lan- ceolate to broadly ovate, cordate to cordate-imbricate, the margins with irregularly spaced, unicellular pyriform glands (which also occur superficially) and short or long flagelliform outgrowths ter- minating in either a long filiform cell, a long filiform thin-walled cell, or rarely in a pyriform glandular cell, the apex terminating in an acicular or small thin- walled cell, to 1 1 x 3 mm: lamina 1-pinnate- pinnatifid to 2-pinnate, with up to 16 free pinna pairs, narrowly ovate to oblong-acute, to 285 mm long, the pinnae proximally wide- spaced, imbricate towards the apex, the most proximal pinna pair slightly to strongly reduced: rachis stramineous, shallowly sulcate adaxially, moderately set with unicellular pyriform glands and sparsely to densely paleated; paleae sessile, ferrugineous to stramineous, scarious, lanceolate to narrowly ovate, cordate to cordate-imbricate, the margins irregularly set with unicellular pyri- form glands (which also occur superficially), short cuneate-emarginate outgrowths often terminating in a unicellular glandular cell, and short or long flagelliform outgrowths (which also occur superficially) terminating in either a long filiform cell, a long filiform thin-walled cell, or rarely in a pyriform glandular cell, the apex teminating in an acicular or thin-walled cell, to 7 x 2 mm: pinnae pinnatifid to 1 -pinnate, with up to 5 free pinnule pairs, short- stalked, triangular, ovate, deltoid or oblong, to 36 x 18 mm: pinnules opposite to alternate, proximally short-stalked and widely spaced, sessile and imbricate towards the apex, firmly herbaceous to coriaceous, dark green adaxially, slightly paler abaxially, broadly ovate to circular, broadly cuneate, the margins shallowly crenate to dentate, revolute in plants from exposed habitats, to 1 1 mm long; adaxially sparsely set with a few twisted, cartilaginous, castaneous paleae chiefly along the pinna-rachis or costa; paleae short-stalked, linear to oblong, the margins subentire, with a few short cuneate- emarginate outgrowths or rarely with a few unicellular pyriform glandular cells and/or flagelliform outgrowths, the apex terminating in an acicular cell, to 3 mm long; abaxially sparsely to moderately set with hairs and scarious, stramineous to ferrugineous paleae, the paleae sessile, narrowly lanceolate to narrowly ovate, cordate, the margins (and often superficially) with unicellular, pyriform glandu- lar cells, short cuneate outgrowths that often terminate in a unicellular glandular cell, and often with a few flagelliform outgrowths termin- ating in an acicular or thin-walled cell, to 5 mm long. Venation raised. Sori circular, to 1 .5 mm in diameter, medial to inframedial, uniseriate, discrete but slightly confluent in depauperate plants; exindusiate: sporangium with 11-(14)-20 indurated annulus cells. Spores castaneous, the perispore folded to form closely set low tubercles, verruculate to echinulate, the exospore 30-(37.78)-78 x 24-(28.62)-36 |jm. Chromosome number unknown. MATERIAL EXAMINED MARION ISLAND (4654'S, 3745'E): Black Hagless River near Kildalkey Bay, Gremmen s.n. (WAG); Macaroni Bay en route to Stony Ridge, Rand 3270 (BOL, PRE); grey lava cliffs near Duikers Point, 10 m, Huntley 466 (NBG-2 sheets, PRE-2 sheets); Marion Island, Mostert 15 (NBG, PRE); valley in cliffs above Prinsloo Lake, 25 m, Huntley 788 (NBG-2 sheets); Nellie humps, 40 m, Huntley 137 (BOL, NBG); stream adjacent Kildalkey hut, 100 m, O'Connor 1003 (BOL, NBG); cliffs at Goodhope Bay, Rand 3653 (BOL); between station and Skua Ridge, Rand 3766 (BOL); Rand 3 1 92 (BM, BOL), 3271 (BM, BOL), 3690 (BM, BOL). PRINCE EDWARD ISLAND (4638'S, 3757'E): cliffs S. of cave on E. coast, 25 m, Huntley 657 (BOL, NBG). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum marionense differs from any other taxon in the study area in having POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 59 1 mm Fig. 10 Polystichum marionense. A, habit; B, abaxial surface of fertile pinna. All drawn from Huntley 466 (NBG). 60 J.P. ROUX a thin, decumbent and branched rhizome, pinnae that are not acroscopically developed, raised veins with few dichotomies, exindusiate sori situated along the veins and not at the vein endings, and paleae with unicellular glandular cells and flagelliform outgrowths along the margins and frequently also superficially. Alston & Schelpe (1957) considered the species to belong to the Polystichum mohrioides (Bory) C. Presl group. In this group they included P. plicatum (Poepp.) Hicken from the Andes and South Georgia, P. elegans J. Remy from the Andes, P. scopulinum (R.J. Eaton) Maxon and P. lemmonii Underw. from the western United States, and P. cystostegia (Hook.) J.B. Armstr. from New Zealand. Polystichum plicatum and P. elegans are sometimes considered to be varieties of P. mohrioides, but I found P. mohrioides to be extremely variable and could not on the grounds of the palea, indusium, sporangium and spore morphology distinguish between these taxa. I furthermore do not consider them related to P. marionense, as in this species the paleae bear long flagelliform outgrowths along the margins which often terminate in a glandular cell and are exindusiate. In P. mohrioides the paleae margins are subentire, but mostly bear a few short angular outgrowths. The palea apex always terminates in a short acicular cell. The indusia are large and mostly bear a variable number of pyriform cells along the margin and often also on the adaxial and abaxial surfaces. Polystichum cystostegia and P. mohrioides are clearly related as both are characterized by similar paleae and indusia. Polystichum scopulinum and P. lemmonii are related as can be judged from the paleae with short angular marginal outgrowths and apices that terminate in either an acicular cell or a small thin-walled cell. This is supported by the findings of Wagner (1979). Polystichum scopulinum and P. lemmonii are not considered to be related to either P. marionense or P. mohrioides. The affinity of P. marionense remains obscure. VARIATION. Variation in Polystichum marionense on Marion and Prince Edward Islands can be ascribed to environmental influences. Plants from well protected sites are large, the pinnae widely spaced, and the stipe sparsely paleated. Plants from more exposed sites are generally depauperate, densely paleated, and the pinnae coriaceous and closely imbricate with the pinnule margins strongly revolute. Pinnae of depauperate forms are often arranged perpendicular to the lamina axes. Minor variations also occur in the paleae. In some collections the flagelliform marginal outgrowths are extremely long whereas in others they are short. Also the occurrence of such outgrowths from the surface of the palea varies from collection to collection. The limitation of unicellular glands to the proximal part of the palea surface seems to be fairly constant. Unicellular pyriform glandular cells on the adaxial and abaxial surfaces of the lamina have only been observed in Huntley 788 (NBG). DISTRIBUTION AND ECOLOGY. Polystichum marionense is known only from Marion Island, Prince Edward Island and Possetion Island of the Crozet group in the Southern Ocean (Alston & Schelpe, 1957; Gremmen, 1982). Since the floras of the subantarctic islands are poorly known the species may have a wider distribution than is currently known. On Marion and Prince Edward Islands the species is only known from low-lying areas, with most collections having been made at elevations between 10 and 100 m above sea-level. The plants form large clumps in basalt rock crevices and at boulder and cliff bases. Huntley ( 1 97 1 ) reported the plant to always occur in sites protected from the predominantly westerly and north-westerly winds. 1 1. Polystichum transkeiense W. Jacobsen in /. 5. African Bot. 44: 169 (1978). Type: South Africa, Transkei, Port St Johns, near road to Second Beach, deep shade in forest, 67 m, W.B.G. Jacobsen 4301 (PRE!-holotype). Fig. 11. Plants terrestrial or epilithic. Rhizome prostrate, widely creeping, branched, to 10 mm in diameter, set with roots, closely to widely spaced persistent stipe bases, and paleae (which are restricted to apical region); paleae broadly attached, stramineous to castaneous, chartaceous, narrowly lanceolate, cordate to cordate-imbricate, the margins repand to erose, generally without thin-walled hair-like cells, the apex often flagelliform, mostly terminating in a thin- walled cell, to 8.5 x 1 .5 mm. Fronds usually widely spaced, 4-6 per plant, arching, to 1.34 m long: stipe firm, adaxially sulcate, proxi- mally castaneous, stramineous distally, to 710 mm long x 4 mm in diameter, proximally densely paleated; paleae broadly attached, castaneous to stramineous, chartaceous, narrowly to broadly ovate, cordate to cordate-imbricate, the margins repand, erose to fimbriate, with or without thin-walled cells, the apex often flagelliform, ter- minating in a thin-walled cell, to 7 x 2.5 mm; distally sparsely paleated, becoming glabrous with age: lamina 2- or 3-pinnate, with up to 22 pairs of free pinnae, firmly herbaceous, adaxially dark green, somewhat paler abaxially, ovate to broadly ovate, to 655 mm long, the proximal pinna pair reduced in size: rachis stramineous to greenish, adaxially sulcate, sparsely paleated; paleae short-stalked, stramineous, chartaceous to membranous, narrowly oblong to nar- rowly ovate, cordate to hastate, the margins proximally repand, erose, or set with short and/or long irregular outgrowths, often with filiform outgrowths terminating in a thin- walled cell, distally repand to entire, flagelliform, terminating in a filiform cell or a thin- walled cell, to 6 x 1 mm: pinnae 1 -pinnate or 2-pinnate, with up to 20 pairs of free pinnules, proximally widely spaced, mostly not overlapping, distally frequently overlapping; proximal pinnae narrowly ovate to narrowly oblong-attenuate, those towards the middle of the lamina ovate, narrowly oblong to oblong-attenuate, to 240 x 75 mm: pinna- rachis stramineous, adaxially sulcate, sparsely to densely paleated; paleae short-stalked, stramineous, chartaceous to membranous, lin- ear, narrowly triangular to narrowly ovate, cordate to hastate, the margins proximally with short and/or long irregular outgrowths often terminating in a thin-walled cell, distally entire, twisted, the apex terminating in a filiform or thin-walled cell: pinnules short- stalked, opposite to alternate, widely spaced to overlapping, the proximal acroscopic pinnule the largest, the proximal basiscopic pinnule on basal pinna pair generally significantly smaller than the next basiscopic pinnule, inaequilateral, ovate, ovate-oblong to ovate- rhomboid, acuminate to obtuse, acroscopically auricled, shallowly to deeply incised, lobate-serrate, the lobes oblong, the proximal acroscopic auricle obovate, sharp-tipped to aristate, the costa adaxially proximally sulcate, sparsely paleated; paleae stramineous, membranous, twisted, simple or proximally with short or long angular outgrowths, the apex terminating in a filiform or a thin- walled cell, to 2.6 mm long, abaxially sparsely to moderately paleated; paleae stramineous, membranous, narrowly triangular to narrowly ovate, short-stalked, cordate to cordate-imbricate, the margins proximally erose or with short and/or long angular outgrowths or with long filiform outgrowths terminating in a thin- walled cell, distally entire, flagelliform, twisted, the apex terminating in a filiform or thin- walled cell, to 1 .5 mm long. Venation raised. Sori circular, c. 1 mm in diameter, near or at the apex of abbreviated veins, discrete at maturity: sporangium with 10-(13)-19 indurated annulus cells; stalk eglandular: indusium absent. Spores 64 per sporangium, brown, the perispore folded to form a reticulum of inflated ridges, the ridges with a high crest, variously but mostly sparsely echinulate, minutely perforated, the exospore 32-(38.8)- 46 x 22-(28.4)-36 urn. Chromosome number 2n=164. MATERIAL EXAMINED SOUTH AFRICA. 2330 (Tzaneen): Woodbush (CC), Jenkins 919 (PRE). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 61 10 mm Fig. 11 Polystichum transkeiense. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A & C, drawn from Roux 2541 (NBG); B, drawn from Roux 2539 (NBG). 62 J.P.ROUX 2430 (Pilgrim's Rest): Mariepskop (DB), Burrows 3150 (PRE). 2531 (Komatipoort): Barberton, Ida Doyer Nature Reserve, 1 100 m (CC), Mutter 2107 (PRE); Barberton, Maid of the Mist, Thorncroft 40 (PRE). 2730 (Vryheid): Pongola Bush Reserve, Stinkwood Falls, 1550 m (BC), Glen 2436 (PRE). 2731 (Louwsburg): Ngome Forest Reserve (CD), Roux 2535, 2536, 2537, 2538, 2539, 2540, 2541 (NBG); Ngome Forest, Reid 68 (PRE); Ngome Forest, 4100 ft, Schelpe 6223 (BOL); Ngome Forest, Strey 8378, 9381 (BOL, NH), 10487 (NH); Ngome Forest, c. 1000 ft, Schelpe 6244 (BOL); Ngome Forest, along waterfall path, 1000 m, Glen 93, 97 (PRE). 2830 (Dundee): Eshowe, Hospital Wood (CD), Lawn 63 (NH); Qudeni Forest, 5000 ft (DB), Fisher 802 (NH, NU, PRE), 877 (NU), Qudeni Forest, Jordaan 698 (NH, PRE); Qudeni Forest, 5500 ft, Schelpe 6267, 6268 (BOL); Qudeni Forest, 5000 ft, Clarkson 87 (NU), 130 (BOL, NU); Qudeni Forest, c. 5000 ft, Allsopp 743 (NU); Qudeni, 5000 ft, Fisher 83 1 (NH, NU); Qudeni, Ekombe Forest, 5000 ft, Fisher 8 1 7 (NU); Qudeni Forest, 1 1 00 m, MacDevette 702 (PRE); Qudeni Forest Reserve, Van Wyk 7306 (NH); Qudeni Forest, 5000 ft, Fisher & Schweickerdt 109 (NH). 2831 (Nkandla): Nkandla Forest (CA), Roux 1932 (NBG); Eshowe to Nkandla, 1080 m, Goetghebeur 4443 (BR, PRE); Nkandla Forest, Schelpe 1701 (BOL); Nkandla, 3000 ft, Meebold 12614(M); Nkandla, Schelpe 1701 (NU); Nkandla, Mjton s.n. (NU); Nkandla, Lawn 2001 (NH). 2930 (Pietermaritzburg): Karkloof, farm Shawswood (AC), Roux 1915 (NBG); Karkloof, Van Jaarsveld 5026 (BOL, NBG); Karkloof, farm Ehlateni, Roux 1006, 1007, 1008, 1009, 1010, (NBG), 1011 (NBG, PRE); Karkloof, 'Braco', 4300 ft, Schelpe 5115 (BOL); Balgowan, 'Boschfontein', 4000 ft, Schelpe 606, 610 (NU); Balgowan, 4000 ft, Lindahl 102 (NU); Balgowan, 'Boschfontein', 4000 ft, Fisher 630 (NH, NU); Karkloof, 'Elderslie', Rycroft s.n. (NU); Balgowan, Graham 107 (NU); Karkloof Forest, Ehlatine, Rycroft 89 (NU); Karkloof Forest, Wirminghaus 610 (NU); Karkloof Forest, bank of Mshwati River, Wirminghaus 902 (NU); Karkloof, Colsbourne farm, Vos & McGregor s.n. (NU); Balgowan, Thomas 67 (NU); Ahrens, 'Mowbray', c. 5000 ft (BB), Fisher 993 (NU); Dargle, Kilgoblin (CA), Smook 579 (NU), 566 (BOL, NU); Dargle, Esterhuysen 26200 (BOL); Lions River District, Lions Bush, Moll 832, 833 (NU); Dargle, Kilgoblin, Smook 661 (NU); Pietermaritzburg, 2500 ft, (CB), Sanderson s.n. (PRE); Pietermaritzburg, Winters Kloof, Doige s.n. (PRE); Pietermaritzburg, Worlds View, Venter 736 (PRE); Pietermaritzburg, Ferncliff, Schelpe s.n. (BOL); Zwaartkop, Sim s.n. (BOL); Pietermaritzburg, Ferncliff Nature Re- serve, c. 2500 ft, Cowan 120 (BOL); Pietermaritzburg, Town Bush Valley, 2500 ft, Tosh et al. (K); Swartkop, Hillary 69 (NU); Pietermaritzburg, Swartkop, Duncan-Vale 19 (NU); Sweetwaters, Stinger 58 (NU); Pietermatitzburg, Swartkop, 4000 ft, Fisher 123 (NH, NU); Swartkop, 4500 ft, Nixon 36 (NU); Cascades, Town Bush, 2900 ft, Sidly 49 (NU); Pietermaritzburg, Claridge, Carnegie s.n. (NU); Hilton Road, Devlin 43 (NU); Town Hill, Carnegie 706 (NU); Sweetwaters, Uhjati 52 (NU); Pietermaritzburg, Town Bush Valley, Fisher 693 (NU); upper Town Bush Valley, 3300 ft, Wand 28 (NU); Town Bush Valley, Tosh Robinson & De Villiers 1 (NU); Town Bush Valley, c. 2250 m, Doni 72 (NU); Pietermaritzburg, Ferncliff Nature Reserve, 1000 m, Crouch 556, 570, 598 (NU); Swartkop, Clarkson 19 (NU); Town Bush Valley, Devlin 34 (NU); Town Bush Valley, 3000 ft, Fisher 667 (NU); Ferncliff Nature Reserve, 2500 ft, Cowan 155 (NU); Town Bush Valley, 3000 ft, Nieuwoudt 56 (NU); Winters Kloof, Doidge P54 (PRE); Cottingham, farm Keerom, 4500 ft (CC), Strey 8429 (BOL, NH); Richmond, Enon Forest (CD), Van Jaarsveld 5044 (PRE); Inanda (DB), Wood s.n. (B, PRE); Camperdown, Nagle Dam, 3000 ft (DD), Wells 1551 (NU). 2931 (Stanger): 10 km from Kwasizabantu towards Mapumulo (AA), Van Jaarsveld & Lang 5096, 5098 (BOL, NBG), Van Jaarsveld & Jacobs 5851 (NBG); Alexandra District, Moyeni, 750 m (BA), Rudatis 1 100 (B, K, NBG, P); Richmond, Enon Forest (CD), Van Jaarsveld 5044 (BOL, NBG). 3029 (Kokstad): 22 miles E. of Kokstad, 4850 ft (CB), Schelpe 4417, 4418 (BOL); Tabankulu Forest Reserve (CD), Wilkins 40 (PRE); Ingeli Bush (DA), Taylor 5227 (NBG, PRE); Mpetsheni Forest, Weza, Roux 1960, 1961 (NBG); Weza Forest, Roux 2493, 2494, 2495, 2497, 2498 (NBG); Mpetsheni Forest, Weza, c. 1000 m, Nicholas & Marais 1675 (PRE); Weza Forest, Roux 623 (BOL); Weza Forest, Bangeni Forest, 1 200 m, MacDevette 1 534 (NH). 3030 (Port Shepstone): Burntwood, Paddock (CC), Strey 5994 (BR, K, NU); Umtamvuna Nature Reserve, Long Kloof, 360 m, Abbott 1818 (NH); Umtamvuna Nature Reserve, Gogosa Kloof, Abbott 2101 (NH); Umtamvuna Nature Reserve, Verassend Kloof, 360 m, Abbott 1821 (NH). 3128 (Umtata): Tsolo, Nqadu Ridge, c. 1 1 00 m, (BC) Keeler & Cloete 449 (NH). 3129 (Port St Johns): Port St Johns, Egossa Forest (BC), Strey 8869 (BOL, NH, NU, PRE); Lusikisiki, Magwa Falls, Strey 67 1 8 (NH, PRE); Egossa Forest above Magwa Falls, 1300 m, Venter & Vorsterll (BR, PRE); Port St Johns, stream at S. end of airstrip (DA), Roux 582 (BOL, NBG); Port St Johns, Hardcastle s.n. (NBG); woods at Port St Johns, Flanagan 2973 (PRE); Port St Johns, Hardcastle 28 1/283 (PRE); Port St Johns, McLoughlin 788 (BOL, PRE); Port St Johns, edge of plateau, 1200 ft, Hardcastle 285 (PRE); Port St Johns, McLoughlin S36 (PRE); Port St Johns, Agate Terrace, McLoughlin 780 (BOL); Port St Johns, Isaac s.n. (BOL); Port St Johns, Moffets Glen, Roux 589 (BOL); Port St Johns, Schelpe 357, 358 (NU); Port St Johns, Flanagan 2473 (PRE). SWAZILAND. 2531 (Komatipoort): Piggs Peak, Kings Forest (CD), Compton 27831 (NBG, PRE); Havelock, Kings Forest, 5000 ft, Schelpe 6163, 6169 (BOL). WITHOUT EXACT LOCALITY: Port Natal, Krauss 258 (BM); loco incerto, Hill 36 (PRE); loco incerto BOL 5771 3 (BOL); Natal, Buchanan s.n. (BOL); Natalia, in sylvis montis humidis, sine coll. s.n. (P); Pondoland, Buchanan 30 (P); Hlokozi, Alexandra City, 2700 ft, Rudatis 2309 (NBG); in sylvis montanis umbrosis humidis, Guienzius 28 (B); Natalia, in sylvis humidis, sine coll. B-97066 (B); Natal, McKen s.n. (B); Natal, Buchanan 75 (B); Pondoland, Buchanan 30 (B); zwischen dem grossen Wasserfall und Omsamcaba, Drege s.n. (B); loco incerto, Drege s.n. B-97092 (B); Natal, Guienzius s.n. B-97064 (B); loco incerto, sine coll. s.n. NH-26388 (NH); Prom. b. spei, Guienzius s.n. (BR); Natal, sine coll. s.n. (BR); Zululand, Haygarth s.n. (NH); loco incerto, sine coll. s.n. NH-26465 & 26466 (NH); Inanda, Great Noodsberg, Town Hill, P.M.B., sine coll. s.n. NH-26791 (NH); loco incerto, sine coll. s.n. NH-26387 A-only (NH). DIAGNOSTIC FEATURES AND RELATIONSHIPS. The thin, widely creep- ing rhizome, the thin-walled cells on the paleae, and the exindusiate sori are the most diagnostic features of the species. The affinity of Polystichum transkeiense is yet to be determined. VARIATION. Polystichum transkeiense shows extreme variation in frond morphology, even within a population. Lamina dissection varies between 2-pinnate and 3-pinnate with the pinnules showing various degrees of dissection. Pinnule lobes may vary between broadly elliptic to obovate and narrowly obovate to narrowly ob- long. Palea variation is not as significant as that of the lamina. Paleae do, however, vary in outline, the degree to which the margins are sculptured, the absence or presence of thin-walled cells along the margins, and in the apex terminating in a thin-walled cell or a filiform cell. Both conditions are usually present in the same plant. DISTRIBUTION AND ECOLOGY. Polystichum transkeiense is con- fined to the eastern parts of South Africa and northern Swaziland. This region receives its rain largely during the summer months (September-March). It is an exclusively forest-dwelling species, often growing in very wet conditions. In the southern limits of its distribution at Port St Johns, P. transkeiense grows in Typical Coast Belt Forest that occurs from near sea-level to approximately the 450 m contour. This region receives 900-1 500 mm of rain per annum. To the north and somewhat inland it occurs in forests of the Pondoland Coastal Plateau Sourveld where it occupies a plateau to 450 m above the sea. The forests are mainly found in protected places along the escarpment such as gorges and valleys below cliffs. Rainfall in this region is high; 1150-1300 mm of precipitation is measured per annum. In the KwaZulu-Natal midlands P. transkeiense occurs in forests of the 'Ngongoni Veld, extending between 450 and 900 m above the sea and receives on average 750-1300 mm of rainfall per annum. Nkandla, Qudeni and Weza are among the most notable forests occurring in this vegetation type. In northern KwaZulu-Natal Afromontane forests with slightly more tropical affinity occur on the inland mountains. The extensive Ngome Forest, where P. transkeiense is common, is an example of this forest type. Forests of this type occur northwards to the mountains south and west of Barberton. POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA Rainfall in this region is high, ranging between 900-1950 mm per annum (Acocks, 1988). Jacobsen ( 1 978) considered 'its tendency to grow isolated in deep shade and not in large clusters. . .' as a diagnostic feature of Polystichum transkeiense. My observations, however, do not con- form with this statement. Polystichum transkeiense does grow as isolated plants, but generally forms continuous, dominant stands, especially in the Weza, Karkloof, Nkandla and Ngoye forests. 12. Polystichum magnificum F. Ballard in Kew Bull. 12: 48, f. 1 (1957). Type: Uganda, Mount Elgon, in the crater (alpine region) in a small sheltered cleft on the ridge north of Maji ya moto, 3750 m, Hedberg 965 (K!-holotype; K!-isotype). Fig. 12. Plants terrestrial. Rhizome short-decumbent, branched, to 12 mm in diameter, set with roots, closely spaced persistent stipe bases, and paleae; paleae broadly attached, ferrugineous, chartaceous, linear, cordate, the margins subentire or with small, widely spaced, straight or curved outgrowths, the apex terminating in an acicular cell, to 30 x 2 mm long. Fronds 8-12 per plant, erect to suberect, to 1.13 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 470 mm long x 9 mm in diameter, densely paleated; paleae broadly attached, ferrugineous, chartaceous, those on the proximal part of stipe narrowly ovate, cordate, the margin with small widely spaced projections, the apex terminating in an acicular cell, to 40 x 2 mm, those on the distal part of stipe more variable in size; the larger broadly attached, narrowly ovate, ovate to narrowly oblong, cordate, with the margins variously set with short and/or long, simple or forked recurved outgrowths, the apex terminating in an acicular cell, to 28 x 8 mm; the smaller short-stalked, narrowly triangular, cordate, often somewhat auricled, the margins proxi- mally with short and/or long angular or curved outgrowths, distally entire or with small projections, the apex terminating in an acicular cell: lamina 2-pinnate to 2-pinnate-pinnatifid, coriaceous, adaxially dark green, abaxially slightly paler, narrowly ovate to narrowly oblong, to 660 mm long, reduced towards the base, often with a single proliferous bud near the apex, the bud paleae ferrugineous: rachis stramineous, adaxially sulcate, densely paleated; paleae stramineous to ferrugineous, the larger broadly attached, ovate to narrowly ovate, cordate, the margins proximally with long and/or short, straight and/or curved outgrowths, distally entire, the apex terminating in an acicular cell, the smaller short-stalked, narrowly triangular, the margins proximally with curved or angular outgrowths, distally entire, the apex terminating in an acicular cell: pinnae mostly somewhat overlapping, narrowly ovate to oblong, not sig- nificantly developed acroscopically, to 130 mm long: pinna-rachis stramineous, adaxially sulcate, densely set with paleae similar to but smaller than those on the rachis; pinnules proximally closely spaced, distally alternate, mostly somewhat imbricate, inaequilateral, ovate, lobate, crenate, the proximal pinnules acroscopically incised to or near to the adaxially sulcate costa, the segments unequally rhomboid to obovate, adaxially densely paleated; paleae short-stalked, ferrugi- neous, chartaceous, subulate, simple or proximally with short marginal outgrowths, often twisted, the apex always terminating in an acicular cell, to 5 mm long, abaxially densely paleated; paleae short-stalked, ferrugineous, subulate, straight or twisted, proximally with short marginal outgrowths, the apex terminating in an acicular cell, to 4 mm long. Venation raised. Sori circular, to 2.2 mm in diameter, uniseriate, discrete at maturity, terminal or near-terminal on abbreviated vein branches: sporangium with 1 1-( 13)-17 indurated annulus cells; stalk eglandular: indusium peltate, circular, erose, the maximum radius 0.63-(0.94)-1.14 mm, persistent, brown. Spores 63 dark brown, 64 per sporangium, the perispore relatively smooth, echinulate, closely perforated, the exospore 36-(47.78)-58 x 26- (33. 37)^0 urn. Chromosome number unknown. MATERIAL EXAMINED ETHIOPIA: Bale Region, Dello Awraja, in Harrena Forest c. 3.3 km N. of Rira, 3040 m, Mesfin 5077 (ETH); Bale Mountains, E. of Kara Deema, 4140 m, Miehe 1497 (ETH); Bale Mountains, E. of Kara Deema, 4200 m, Miehe 1541 (ETH); Bale Mountains, Mendoyn Anraja, in Harrena Forest, c. 1-2 km S. of Riva village, 2780-2850 m, Mesfin 5355 (ETH); Bale Region, Dello Awraja, c. 3.4 km N. of Rira village, 3120 m, Mesfin 5332 (ETH); Arussi, Juniper forest, 9000 ft, Thomerson 550 (ETH, K); Darra, bamboo forest, 9000 ft, Mulvany 48 (K); Bale Province, Rira, 20 miles SW of Goba, 10 800 ft, Mooney 7192 (K); Mount Tola, Gamu Highlands, 13 000 ft, Mulvany 1 (K); Bale Province, c. 30 miles S. of Goba, Saneti Plateau, 2720 m, Ash 3567 (BR). KENYA: Mount Elgon, 12 500 ft, Tweedie s.n. (K). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Most striking is the large stature of the plants occurring at these high elevations. Also diagnostic are the densely paleated fronds, rounded lobes, and the proliferous bud borne on the rachis near the frond apex. The affinity of Polystichum magnificum is uncertain. Super- ficially it appears to belong to section Lasiopolystichum but the palea structure and the short, branched decumbent rhizome does not support such an affinity. DISTRIBUTION AND ECOLOGY. Polystichum magnificum is only known from Mount Elgon, on the border between Uganda and Kenya, and the Bale, Arussi, and Gamu Gofa regions in southern Ethiopia. On Mount Elgon Polystichum magnificum occurs at elevations ranging from 3700 to 3800 m. In Ethiopia it occurs from 2700 to 4200 m and grows in a wide range of afromontane vegetation communities. These include afromontane forests of the Hagenia abyssinica (Bruce) J.F. Gmel. and Juniperus procera Hochst. ex Endl. types, afromontane bamboo zone, afromontane scrubland and afromontane grassland types as defined by White (1983). The species has been reported to form compact patches up to 1.8 m in diameter at 3000 m on Mount Tola and Mount Gughe in Ethiopia. At higher elevations Polystichum magnificum tends to be restricted to rock crevices where it is protected from wind and fire. 13. Polystichum zambesiacum Schelpe in Bol. Soc. Brot. ser. 2, 41: 215(1 967). Type: Rhodesia (Zimbabwe), Umtali District, Henkels Nek, Stapleford, Schelpe 5751 (BOL!-holotype; BOL!-isotype). Fig. 13. Plants terrestrial or epilithic. Rhizome short-decumbent to suberect, short-branched, to 25 mm in diameter, set with roots and closely spaced persistent stipe bases, the older parts nude, the apical part densely paleated; paleae broadly attached, rugose, ferrugineous, linear, truncate to cordate, the margins variously fimbriate, often also with a few long, straight, recurved, filiform outgrowths, to 30 x 2 mm. Fronds caespitose, 5-8 per plant, arching, to 1.8m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 840 mm long x 9 mm in diameter, proximally densely paleated; larger paleae more widely spaced and becoming smaller distally, membranous, ferrugineous throughout or with a narrow membra- nous margin and a dark, nitid centre, ovate, cordate, the margins fimbriate, the apex cuspidate or flagelliform, terminating in a thin- walled cell or a filiform cell, to 16 x 10 mm; smaller paleae short-stalked, narrowly ovate to narrowly triangular, cordate to cordate-imbricate, the margins proximally with irregular angular outgrowths, often also with long, twisted, filiform outgrowths often terminating in a small thin-walled cell, sparsely fimbriate distally, 64 J.P. ROUX 2 mm 5 mm Fig. 12 Polystichum magnificum. A, middle pinnae of lamina; B, adaxial surface of pinnule; C, abaxial surface of fertile pinnule; D, section of abaxial surface of rachis. All drawn from Hedberg 965 (K). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 65 Fig. 13 Polystichum zambesiacum. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A & C, drawn from Schelpe 5751 (BOL); B, drawn from Burrows 2935 (PRE). 66 J.P. ROUX the apex flagelliform, terminating in a small thin-walled cell or a filiform cell; lamina 2-pinnate to 3-pinnate, with up to 30 free pinna pairs, firmly herbaceous, narrowly ovate to ovate, up to 890 mm long, the proximal pinna pair often slightly reduced and often somewhat deflexed: rachis stramineous, adaxially sulcate, often somewhat flexuous towards the apex, sparsely paleated; paleae short-stalked, chartaceous, stramineous to ferrugineous, narrowly linear, narrowly ovate to narrowly triangular, cordate to cordate- imbricate, the margins proximally with straight or angular outgrowths or with long, twisted, filiform outgrowths, distally entire or sparsely fimbriate, the apex terminating in a small thin-walled cell or a filiform cell, to 1 .8 mm long: pinnae 1 -pinnate to 2-pinnate, with up to 27 free pinnule pairs, generally widely spaced but often imbricate, lanceolate, narrowly ovate to oblong-attenuate, the proximal pinnae to 268 mm long: pinna-rachis stramineous, adaxially sulcate, set with paleae similar to but smaller than those on the rachis: pinnules opposite to alternate, widely spaced or imbricate, asymmetric, ovate-oblong, narrowly ovate to narrowly triangular, acroscopically auricled, usually somewhat reclinate, lobate-serrate, obtuse, the proximal acroscopic pinnules on basal pinnae often reduced, be- coming larger towards middle of lamina, the proximal pinnules simple or incised to or near to costa, to 45 mm long; costa proximally adaxially sulcate, glabrous or with a few short-stalked, stramineous, twisted paleae; paleae filiform, simple or proximally with a few twisted outgrowths, the apex terminating in a small thin-walled cell or a filiform cell, to 1 .2 mm long, adaxially sparsely paleated; paleae short-stalked, narrowly triangular to narrowly ovate, cordate, the margins proximally with short or long angular, often filiform outgrowths, the apex entire or sparsely fimbriate, terminating in a small thin-walled cell or a filiform cell, to 0.8 mm long. Venation raised or obscure. Sort circular, terminal, medial or inframedial on unabbreviated or abbreviated vein branches, discrete at maturity, c. 1 mm in diameter: sporangium with 9-(14)-20 indurated annulus cells; stalk eglandular; indusium peltate, circular to reniform, repand to erose, the maximum radius 0.19-(0.33)-0.5 mm, persistent, brown. Spores 64 per sporangium, brown, the perispore folded to form inflated reticulate ridges and tubercles, the ridges and areas between variously perforated, echinate to echinulate, the exospore 30-(37.77)-54 x 22-(27.49)-36 urn. Chromosome number un- known. MATERIAL EXAMINED MALAWI: Nyika Plateau, 2250 m, Brass 17255 (K, PRE, SRGH); Mount Mulanje, Lichenya Plateau, Nessa Path Forest, 1800 m, Chapman 8262 (MAL, PRE); Zomba Plateau, W.-facing cliff edge below Malumbe Peak, 1900 m, Berrie 602 (MAL); Zomba Plateau, Malumbe Peak, Roux 2928 (NBG); Mulanje Mountains, Lichenya Plateau, Pawek 3815 (K); Mount Mulanje, Lichenya Plateau, 1 820 m, Brass 1 6566 (K, SRGH); Nyika Plateau, Kasaramba Peak, 8400 ft, Simon et al. 1730 (K, SRGH); Mount Mulanje, Lichenya Plateau, 1890m,fira 16820 (K, SRGH); Mount Mulanje, Lichenya Plateau, 1950 m, Richards 16556 (K); Mount Mulanje, Nayawani Forest, 6400 ft, Newman & Whitmore 547 (BR, SRGH); Mt. Mulanje, L. Ruo Plateau, 6400 ft, Newman & Whitmore 399 (BR). MOZAMBIQUE: Manica & Sofala, Penhalonga, 4500 ft, Schelpe 5325 (BOL); Manica & Sofala, Gorongosa Mountains, Gogogo Peak, 5000 ft, Schelpe 5518 (BOL); Manica & Sofala, Gorongosa, Serra da Gorongosa, vertente do monte Nhandare, 1750 m, Torre & Pereira 12515 (BOL, BR, SRGH); Namuli, Makua country, Last s.n. (K). TANZANIA: Uluguru Mountains, S. of Bunduki, NEedge of Lukwangule Plateau, on rocky outcrops of Muisile Hill, 2400-2450 m, Pocs et al. 86141/ A (WAG); W. Usambaras, towards Mount Kwashenhambu, 1 850 m, Schippers T 1578 A (WAG); W. Usambara Mountains, N. slopes of Mount Shegein, Shasayo Forest, 1850 m, Schippers T1604A (WAG); Morogoro, Ukaguru Mountains in Kilosa District, W. Mamiwa Ridge, 2100-2200 m, Pocs et al. 86100/B (WAG); Tanga Region, Lushoto District, W. Usambara Mountains, Shagayu Forest Reserve, NW slope of the summit 2.5 km ENE of Shagayu sawmill, 1850-1950 m, Borhidi et al. 84847 (ETH); Morogoro District, Uluguru Mountains, W. part of Lukwangule Plateau, 2400-2500 m, Harris et al. 3726 (K); Usagara: Itumba, Wood s.n. (K); Uluguru Mountains, above Morogoro, NE ridge of Bondwa between Morningside and Mwere Valley, 1050 m, P6cs 6537/C (PIC.SERM.); Uluguru Mountains, NW slope of Bondwa, along road to Morningside, Faden et al. 70/654 (BM, BOL); Uluguru Mountains, Morningside to Bondwa, Faden 70/316 (BOL, K); mainland W. of Zanzibar, Last s.n. (K); N. Uluguru Forest Reserve, Lupunga Peak, W. side, 2000 m, Hall s.n. (K); Morogoro: Uluguru gebirge, Lupanga, 2100 m, Schlieben 2977 (B, BR). ZIMBABWE: Inyanga, Pungwe Gorge, 6000 ft, Schelpe 5699 (BOL); Inyanga, circular drive on N. rim of Pungwe Gorge, 7000 ft, Mitchell 148 (BOL); Umtali, Stapleford Forest Reserve, W. of Rupere Peak, 5500 ft, Chase 7429 (BOL, K); Inyanga, Mitchell 1082 (BOL, K, NU, SRGH); Umtali, 'Cloudlands', Vumba, 5200 ft, Schelpe 5365 (BOL); Inyanga, ad dejectum fluminis Pungwe, c. 1700 m, Fries et al. 3795 (BOL, K); Umtali District, Vumba Mountains, 'Cloudlands', forest E. of Cripps Grid, Chase 8345 (BOL, K); Umtali District, on Lords Head property, Vumba Mountains, 5300 ft, Chase 8343, 8344 (BOL, K, SRGH); Melsetter, forest in gully on W. side of N. end of Bundi Valley, Mitchell 514 (BOL, K, SRGH); Umtali District, Stapleford Forest Reserve, 5600 ft, Chase 8373 (BOL, K, SRGH); Stapleford Forest Reserve, lower part of road to Henkels Nek, 5600 ft, Chase 8371 (BOL, SRGH); Umtali District, Vumba Mountains, Williams VUM18 (BOL); Vumba Mountains, Williams VUM1, VUM10, VUM11, VUM12, VUM13 (BOL); Chimanimani Mountains, Gwasha, Williams STP1, STP2 (BOL); Inyanga, 6500 ft, Chase 5100 (BOL); Inyanga, Pungwe Gorge, Inyangani Mountains, 6500 ft, Chase 5240 (BOL); Umtali, Vumba Moun- tains below Castle Beacon, 5600 ft, Chase 7489 (BOL); Inyanga, circular drive below Inyangani Mountain, 7500 ft, Mitchell 135 A (BOL); Umtali, Imbeza Forest Estate, Zuwanne indigenous forest, 5150 ft, Jacobsen 3838 (BOL, SRGH); Stapleford Forest Station, 6000 ft, Taylor 3234 (BOL, SRGH); Vumba Mountains, SE slope of Castle Beacon, 1675 m, Burrows 2944, 2945 (PRE); Inyanga, S. tip of Mount Inyangani, 2040 m, Burrows 2935 (PRE); Pungwe Gorge, in drier part of ravine, Schweikerdt 2412 (M, PRE); Umtali, Vumba, Castle Beacon, 6000 ft, Fisher 1638 (NU, PRE); Vumba Mountains, Chase 1 102 (PRE); W. slope of S. tip of Mount Inyangani, 2000 m, Burrows 2940 (PRE); Inyanga, new beacon on Mount Inyangani, 2540 m, Burrows 2828 (PRE); Inyanga, montane forest, 6000 ft, Chase 4024 (PRE); Vumba Mountains, 5400 ft, Chase 8347 (K); Vumba Mountains, Bunga Forest, Jacobsen 3037 (SRGH); Inyanga, lower E. slope of Inyangani, c. 1480 m,Miiller32\4 (SRGH); lower SE slope of Mt.Pene, \350m,Muller 2798 (SRGH); Inyanga, top of escarpment 2 km N. of Honde View, c. 1 840 m, Muller 3243 (SRGH); Melsetter, Chimanimani, Bundi River, 5500 ft, Whellan 2184 (SRGH); Vumba Mountains, forest on E. slope of Castle Mountain, Jacobsen 3033 (SRGH); Stapleford Forest Reserve, Ruperi Peak, 6100 ft, Chase 4656 (SRGH); Stapleford Estate, montane forest, Roux 2828 (NBG); SE edge of Vumba Hotel forest, Chase 4022 (SRGH); Stapleford Forest Reserve, Chase 8372 (SRGH); Melsetter District, Orange Grove, Chase 3088 (BR, NU, S, SRGH); Inyanga District, source of Inyahupina River, Romneydale, 6100 ft, Chase 2089 (NU, SRGH); 30 miles S. of Umtali, 6000 ft, Grout 33 (NU); Umtali, Pioneer farm, Fisher & Schweickerdt 306 (NU); Umtali District, Vumba Mountains, Natseland, Chase 3428 (NU); Umtali District, Nyagari farm, N. of Zwitembo, Chase 3156 (NU); Inyanga District, Chase 3206 (NU); Inyanga District, Pungwe Falls, 6000 ft, Chase 3197 (NU); Umtali District, Stapleford Forest Reserve, Chase 4520 (NU); Umtali District, Penhalonga, Chase 3132 (NU); Inyanga, Pungwe view point, Chase 3206 (NU); Umtali, Chase s.n. (NU); a 15 km a 1'E. d'Inyanga, sommet du mont Mimunzi, 1814'S, 3253'E, 1950 m, Bamps et al. 272 (BR); Vumba Mountains, Umtali District, Cloudlands, Jackson 8 (GRA). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum zambesiacum can be easily recognized among African members of the genus by its long and narrow rugose rhizome paleae, long- attenuate pinnae and (although variable) obtuse pinnule lobes, conspicuously small indusia, and the palea morphology. Polystichum zambesiacum is not considered to be related to any other taxon in the study area. POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 67 VARIATION. Schippers (1993) suggested that the Tanzanian plants may be a different but related species to Polystichum zambesiacum. He noted the Usambara Mountain plant to differ from that of the southern highlands in the more shiny fronds, the more deeply divided pinnae, and the segment margins having more clearly aristate teeth. Polystichum zambesiacum is clearly a very variable taxon with lamina division ranging between 2-pinnate and 3-pin- nate. Most apparent is the degree to which the size, shape and incision of the pinnules vary. This, however, cannot be linked to distribution as 3-pinnate plants have been recorded from the Pungwe Gorge in Zimbabwe [Chase 5240 (BOL)] and the Ukagura Moun- tains in Tanzania [Pocs et al. 86100/B (WAG)]. In the Uluguru Mountains, like in the rest of its distribution, the pinnules are mostly variously incised and lobate-dentate to ovate-serrate. One collection from this region [Faden et al. 70/316 (BOL, K)] shows hardly any incision of the pinnules which are poorly lobate. Lamina division can thus not be considered in subdividing the taxon. In some plants the proximal pinnule of the basal pinnae is reduced in size and thus not auriculate as the distal pinnae. The basiscopic pinnules of these pinnae are often more significantly reduced than the acroscopic pinnules. In others, however, the pinnules show no reduction of size on the lower pinnae. Stipe paleae also show variation. In some plants the larger paleae are variously impregnated with the central part of the paleae often dark brown to black and nitid with a narrow membranous margin. These paleae are generally persistent for a long time, resulting in the non-impregnated margin being worn away. In others the paleae are reddish brown and remain membranous. DISTRIBUTION AND ECOLOGY. Polystichum zambesiacum ranges from the eastern highlands of Zimbabwe and adjacent Mozambique to Mount Mulanje and the Nyika Plateau in Malawi and the Uluguru, Usambara and Pare Mountains in Tanzania. Polystichum zambesiacum is a terrestrial or epilithic species occurring in high-altitude, evergreen, montane mist forests, along forest margins and streambanks in forests. It has been reported on Mount Inyangani, Zimbabwe, from open montane grassland, and in the Uluguru Mountains from rock outcrops. Being a high altitude species, Polystichum zambesiacum is re- stricted to the often isolated high mountains of east Africa. In Zimbabwe and Mozambique it is confined to the Chimanimani and Nyanga Mountain ranges occurring between 1370 and 2540 m. In Malawi the fern is only known from Mount Mulanje in the south (1800-1950 m) and the Nyika Plateau in the north where it occurs between 1800 and 2550 m. In the Uluguru Mountains, Tanzania, it occurs at elevations ranging between 1050 and 2500 m. Further north, in the Usambara and Pare Mountains it occurs at 1 850 to 1950 m, but is reported to be rare (Schippers, 1993). H.Polystichum monticola N.C. Anthony & Schelpe in Bothalia 15: 554 (1985); Fl. Sthn. Afr., Pterid.: 257 (1986); Burrows, Sthn. Afr. ferns and fern allies: 3 14, f. 75, t. 320a-c (1990). Type: Cape Peninsula, Table Mountain, Dark Gorge, below saddle SE side, sheltered gully, dry in summer, on steep rocky slopes, Esterhuysen 26685 (BOL!-holotype; B, C, CHR, G, GH, K, M, MO, NBG!, NU, P, PR, PRE!-isotypes). Fig. 14. Plants terrestrial or epilithic. Rhizome short-decumbent, closely branched, to 14 mm in diameter, set with roots, crowded persistent stipe bases, and paleae; paleae broadly attached, ferrugineous, membranous, narrowly oblong to narrowly linear, cordate, the margins entire or with widely spaced, short, apically or basally directed outgrowths, the apex acicular or with a thin-walled apical cell, to 26 x 1 .8 mm. Fronds closely spaced, 5-8 per plant, suberect to arching, to 840 mm long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 385 mm long, to 5 mm in diameter, densely paleated; larger paleae broadly attached, concolorous or bicolorous, stramineous or with a castaneous central section, these occurring on the proximal third of the stipe, lanceolate to broadly ovate, cordate to cordate-imbricate, the margins with closely to widely spaced, short, straight or curved, often branched fimbriae, the apex acicular or with a thin-walled apical cell, to 20 x 7 mm; smaller paleae short-stalked, narrowly lanceolate to narrowly trian- gular, cordate to cordate-imbricate, the margins proximally with closely spaced, short and long, often branched fimbriae, the apex entire or with short, firm, widely spaced fimbriae terminating in an acicular cell or with a thin-walled cell: lamina 2-pinnate to 2- pinnate-pinnatifid, firmly herbaceous, dark green adaxially, paler abaxially, narrowly ovate to elliptic, to 510 mm long, proximally somewhat reduced, the proximal pinnae usually somewhat deflexed: rachis stramineous, adaxially sulcate, densely paleated; paleae short- stalked, ferrugineous, chartaceous, narrowly lanceolate, narrowly ovate or narrowly triangular, cordate to cordate-imbricate, the mar- gins proximally with long and short, straight or twisted marginal outgrowths becoming simple towards the apex, the apex usually acicular, rarely terminating in a thin-walled apical cell, to 1 1 .5 x 2.5 mm: pinnae often widely spaced proximally, somewhat overlapping distally, oblong-attenuate to narrowly oblong-attenuate, to 170 x 30 mm: pinna-rachis stramineous, adaxially sulcate, densely set with paleae similar to but smaller than those on the rachis, to 7 x 1 .5 mm: pinnules opposite to alternate, closely spaced, often imbricate, asymmetric, narrowly ovate to ovate, acroscopically auricled, ser- rate to lobate-serrate, sharp-tipped to aristate, the proximal acroscopic pinnule the largest, the proximal pinnules often acroscopically incised to or near to costa, to 22 mm long; adaxially with a few simple and filiform or proximally hastate paleae along the proximal part of the costa, the apical cell always acicular, to 4.5 mm long; abaxially sparsely set with ferrugineous, short-stalked, narrowly triangular to subulate paleae, proximally usually with a few short or long, straight or curved outgrowths, the apical cell always acicular, to 4 x 0.9 mm long. Venation immersed. Sori circular, c. 1 .5 mm in diameter, essentially uniseriate, terminal or near terminal on abbre- viated vein branches: sporangium with 7-( 1 2)-2 1 indurated annulus cells; stalk eglandular: indusium stramineous to castaneous, often black-centred, peltate, circular, entire to repand, the maximum radius 0.31-(0.61)-1.04 mm, persistent. Spores brown, 32 per spo- rangium, the perispore variable, folded to form a reticulum of inflated or compressed ridges and/or tubercles, the crests and areas between echinulate to spiculate, the areas between ridges perfo- rated, the exospore 38-(59.07)-74 x 22-(43.76)-60 Jim. Chromosome number 2n=246, apogamous. MATERIAL EXAMINED LESOTHO. 2828 (Bethlehem): Leribe (CC), Dieterlen 167, in part (B); Mafeteng, Mont Ka-majapela, Dieterlen s.n. (BR); Leribe, 5000-6000 ft, Dieterlen 695 (NBG, PRE, SAM). 2927 (Maseru): Mamathes District, 5850 ft (BB), Guillarmod 835 (PRE); Roma, 5550 ft (BC), Ruch 1909 B-only (PRE); Roma, W. ravine, Schmitz 399 (PRE); Roma, SE-facing slope, Schmitz 402 (PRE), Roma, Schmitz 6888 (PRE); Morija (DA), Dieterlen 1309 A-only (PRE). 2929 (Underberg): Sehlabathebe National Park, Mofoqoi, 2350 m, Hoener 1479 (BOL). SOUTH AFRICA. 2730 (Vryheid): Hlobane, Johnstone 295 (NH). 2828 (Bethlehem): Clarens, mountain N. of Leibrandt Kloof (CB), Roux 937 (NBG); Golden Gate National Park, NE of Glen Reenen Camp, 6200 ft (DA), Liebenberg 7498 (PRE); Golden Gate National Park, Wonderhoek, Gertenbach & Groenewald9\ 67 (PRE): Golden Gate National Park, Roberts 3235 (PRE); Golden Gate National Park, Rossouw 406 (BOL); N. of Mont- aux-Sources, Witsieshoek area, 1800 m (DB), Junod 14 (P); Royal Natal 68 J.P. ROUX 10 mm 10 mm Fig. 14 Polystichum monticola. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A, B, drawn from Roux 2581 (NBG); C, drawn from Van Jaarsveld 6494 (NBG). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 69 National Park, Gudu Forest, Edwards 325 (NU, PRE); Royal Natal National Park, Gudu Forest, Roux 25 1 3 (NBG); shady gully on the Lion facing the Tugelana Valley, c. 6300 ft, Esterhuysen 35645 (B, BOL, NBG, NU, PRE); Witsieshoek, 6000-7000 ft, Thode s.n. (NBG); Royal Natal National Park, Gudu Forest, 5000 ft, Schelpe 1455 (NU); Mont-aux-Sources, Bottomley s.n. (PRE); Mont-aux-Sources, Tugela River, near gorge, Hutchinson 41 (PRE); Mont-aux-Sources, Schweickerdt 760B (PRE); Royal Natal National Park, West 1 284 (PRE); Quaqua mountains, Lefika, Rustenberg s.n. (P); versant N. du Mont-aux-Sources, region de Witsies Hoek, c. 1 800 m, Junod 1 4 (P). 2829 (Harrismith): Swinburne, Boschhoek,E. slopeof Manyenyeza(AC),Jacofaz 4711 (PRE); Harrismith, Platberg, valley below Monkey Point, Roux 2520 (NBG); Platberg, 1600 m, NBG Exped. s.n. (BOL); Harrismith, Platberg, 1600 m, sine coll. 151191 (NBG); Harrismith, Platberg, dolerite cliffs left of summit of Donkey's Pass, Roux 2528 (NBG); Platberg, in shade of concrete furrow leading to Hawkins Dam, 2220 m, Roux 2527 (NBG); Harrismith, Platberg, Roux 2523 (NBG); Harrismith, Kerkenberg, Schelpe 7278 (BOL); Cathedral Peak area, nKdhlanhla Forest, Schelpe p.47 (NU); Cathedral Peak area, 5000 ft, Brass 76 (NU); Cathedral Peak area, Indumeni Forest, 5 1 00 ft, Schelpe 737 (NU); Van Reenen, 6800 ft (AD), Schlechter 6932 (B, NBG, PRE); Oliviershoek Pass, forests S. of Seheletwane (CA), Roux 25 1 8 (NBG); Oliviershoek, 6000-7000 ft, Thode s.n. (NBG); Cathedral Peak Hotel, along Cathedral Peak path, 1500 m (CC), Goetghebeur4563 (BR, PRE); Cathedral Peak area, Brass 67, 79, 82, 96, 98 (NU); Cathedral Peak Forest Station, 6050 ft, Killick 1 134 (NU); Cathedral Peak area, 2000-3000 ft, Wilker 69 (NU); Drakensberg, MnWeni Pass, c. 8500-9000 ft, Esterhuysen 27830 (PRE). 2831 (Nkanhla): Ngoye, 2-3000 ft (DC), Medley-Wood 10886 (PRE). 2926 (Bloemfontein): Thaba 'Nchu Mountain, 6500 ft (BB), Roberts 2998 (PRE). 2929 (Underberg): Giants Castle Nature Reserve, forests above Hillside campsite, c. 1600 m (AB), Roux 2499, 2501, 2506, 2507 (NBG); Injasuti area, 5000 ft, Esterhuysen 26041 (BOL); Kamberg area, Storms Heights, c. 7000 ft (BC), Milliard & Burn 11795 (NU, PRE); Mpendhle District, Mulangane Ridge, above Carter's Nek, 7000-7300 ft, Milliard & Burn 1 695 1 (BOL); 18406 (NBG, PRE); Kamberg, 'Game Pass', 6100 ft, Gordon-Gray 85 (NU); Drakensberg Garden State Forest Reserve, near Mlambonja River (CA), Van Jaarsveld 6492, 6494 (NBG); Drakensberg Gardens, 6000 ft, Bronhead 59 (NU); Drakensberg Gardens, Dyer 73 (NU); Drakensberg Garden area, 6000 ft, Schelpe p54 (NU); Drakensberg Garden Hotel, up to Rhino Peak, along Mlambonja River, 2000 m, Goetghebeur45 1 9 (BR, PRE); Gxalingenwa Valley between Sani Pass and Polela Valley, c. 6700 ft (CB), Milliard & Burn 17076 (NU, PRE); Cobham Forest Station, Ndlovini, Troutbeck, c. 6000 ft, Milliard & Burn 13311 (NU), 13329 (BOL, NU); Cobham Forest Reserve, Sipongweni, 6500 ft, Milliard & Burn 141 35 (BOL, NU, PRE); Cobham Forest Reserve, 'Lakes' Cave area, c. 7800 ft, Manning et al. 1 59 1 8 (BOL, NU); Garden Castle area, 9000 ft (CD), Crooked 62 (NU); Garden Castle area, Elliott 31 (NH), 37 (NU); Boston, Impendhle, 5000 ft (DB), Randies 1 85 (NU); Impendhle, 5 200 ft, Clarkson 1 33 (NU); Impendhle, c. 5000 ft, Huntley 167 (NU); Bulwer, Marwaga Mountain, farm Sunset, 1810 m (DC), Roux 2309 (NBG); Farm Sunset, 5800 ft, Rennie 1441 (NU, PRE); Himeville District, 5000 ft, Webb 101 (NU); Farm Sunset, gully above dams, 6000 ft, Rennie 1055 (NU); Polela District, near Bulwer (DD), Henkel s.n. (NU); Bulwer, 5100 ft, Clarkson 182 (NH, NU, PRE); Bulwer, Allsopp 843 (NU). 2930 (Pietermaritzburg): Zwaartkop (CB), Sim s.n. (NU, PRE). 3018 (Kamiesberg): Kamiesberg, Rooiberg, 5500 ft (AC), Rourke 1684 (BOL, NBG, PRE); Rooiberg, farm Damsland, Roux 2453 (NBG). 3027 (Lady Grey): Herschel District, Majuba Nek, Hepburn 262, B-only (GRA). 3028 (Matatiele): Ongeluks Nek, c. 4 km from Lesotho border post, 2250 m (AB), Matthews 916 (NBG). 3029 (Kokstad): Mt. Currie, farm Highland Home (AB), Roux 2488, 2490 (NBG); Mt. Currie, Edwards 214a (NU); Mt. Currie, 5200 ft, Edwards 44 (NU); Kokstad (CB), Mogg 1927 (PRE). 3030 (Port Shepstone): Ixopo (AA), Hancock s.n. (NU). 3124 (Hanover): Compassberg, near top, c. 2440 m. (DC), Trollip s.n. (PRE); top of Compassberg, Coetzee s.n. (PRE); Compassberg, farm Grootkop, 6300 ft, Acocks 23447 (PRE). 3126 (Queenstown): Broughton, near Molteno (AD), Flanagan s.n. (SAM); Broughton near Molteno, Flanagan 1681 (PRE); Broughton, Molteno, 6300 ft, Flanagan 527 (PRE); Queenstown, Hangklip Mountain, 6600 ft (DD), Roberts 20 1 2 (PRE). 3128 (Umtata): Maclear, farm Woodcliffs (AB), Roux 2481, 2484, 2485 (NBG). 3218 (Clanwilliam): Clanwilliam Division, between Tafelberg and Spout, 6000 ft (BB), Schelpe 1960 (BOL, K). 3219 (Wuppertal): Gideon's Kop, 1500 m (CB), Burrows 1235 (BOL). 3224 (Graaff-Reinet): Mount Oudeberg near Graaff-Reinet (DD), sine coll. 96897 (B); in fissuris rupium in monte Oudeberg prope Graaff-Reinet, 4800 ft, Bolus 1 736 (BOL, K). 3225 (Somerset East): Montis Boschberg (DA), MacOwen 870 (BR). 3226 (Fort Beaufort): Katberg, 3500-4000 ft (BC), Baur 865 (B, GRA); Katberg Pass summit, farm Pleasant View, Roux 2697 (NBG); Hogsback, Elandsberg summit (DB); Roux 2688 (NBG); Hogsback, Zincucha Forest, Roux 2683 (NBG). 3318 (Cape Town): in numerosis umbrosis montis Tafelberg, Paradys (CD), MacOwen s.n. (P); Table Mountain, Dark Gorge, below Saddle, SE side, Esterhuysen 26563 (B, BOL, PRE); Stellenbosch, Jonkershoek, Langrivierkloof (DD), Roux 2580, 2581, 2582, 2583 (NBG); Stellenbosch, Simonsberg, 3000 ft, Esterhuysen 25453 (BOL); Stellenbosch, Helderberg, Disa Gorge, Esterhuysen 28475 (BOL). 3319 (Worcester): Great Winterhoek Mountains, 4000-5000 ft (AA), Esterhuysen 26982 (B, BOL, NBG, PRE); Tulbagh, W. slopes of Swartgat Peak, Witzenberge, 4000 ft (AC), Esterhuysen 1 69 14 (BOL, NBG); Tulbagh, Great Winterhoek Mountains, 5500 ft, Esterhuysen 19787 (BOL, NBG); Ceres, Baviaansberg (BA), Stokoe s.n. (NBG, SAM); shale band below Milner Peak, Hex River Mountains, 5000 ft (AD), Esterhuysen 14264 (BOL), 14885 (BOL, NBG); Milner Ridge Peak and Buffels Dome, 5000 ft, Esterhuysen 28708B (BOL); Ceres, Slab Peak, 1310m, Winter 431 (NBG); Hex River Mountains, Moraine Kloof, c. 4000 ft, Ashton 352 (BOL); Goudini, Waaihoek Mountains, 4000 ft (CB), Barnard s.n. (SAM). 3320 (Montagu): Boesmansbos (DD), Adamson s.n. (BOL). 3321 (Ladismith): Swartberg, near Ladismith, Toverkop, 6500 ft (AC), Esterhuysen 26698 (B, BOL, NBG, NU, PRE); below Toverkop on S. slope of Swartberg, 5000- 6000 ft, Esterhuysen 14013 (BOL, PRE); cliffs at base of Toverkop, 6500 ft, Esterhuysen 18511 (BOL). 3322 (Oudtshoorn): Prince Albert Division, Swartberg Mountains (AC), Stokoe 9410 (NBG, SAM); Prince Albert, Popta s.n. (L); Blesberg, 6000 ft (BC), Esterhuysen 24920 (BOL); Montagu Pass (CD), Schweickerdt 4705 (PRE); Kammanassie Mountains, Mannetjiesberg, 4200 ft (DB), Matthews 1023 (NBG); S. slope of Mannetjiesberg, Kammanassie Mts, 5000 ft, Esterhuysen 1 8396 (BOL). 3323 (Willowmore): Hoopsberg, S. slope, 5000 ft (CB), Esterhuysen 6557 (BOL). 3324 (Steytlerville): S. slopes of Kouga Peak near Joubertina (CA), Esterhuysen 16280 (BOL, NBG); Uitenhage, Cockscomb, Great Winterhoek mountains, 4800 ft (DB), Esterhuysen 27090 (BOL, PRE). ZIMBABWE: Victoria Falls, Sim s.n. (PRE). WITHOUT EXACT LOCALITY: Orange Free State, Wittebergen, ad Caledonrivier, Rehmann 3938, 3978 (B); Natal, sine coll. s.n. (NBG); Blinkwater Bush, Graham 84 (NU); Transvaal, Repton 5B (PRE); in summo monte Koudveld, 6500 ft, Tyson 140 (PRE); in sylvis umbrosis faecium montium Hott. Holland., sine coll. 341 (S); Malappa's Place, Rustenberg s.n. (P); Natal, sine coll. s.n. (BR); Basotholand, Koopowitz s.n. (GRA); Bushmans Cave, Lubke s.n. (GRA). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum monticola is characterized by its largely montane habitat, the usually large stands it forms, the short decumbent rhizome clothed by long, narrowly linear to cuspidate paleae, the closely spaced, firmly herbaceous fronds, smallish pinnules, bicolorous larger paleae on the proximal part of the stipe, short-stalked paleae with proximal margins that bear stiff, often branched outgrowths, and with an apical cell that is not thin-walled, the relatively large repand to entire, persistent indusia, the 32 spores borne by each sporangium, and the somatic chromosome number of 2n=246. In palea morphology Polystichum monticola exhibits features of sections Lasiopolystichum and Metapolystichum. Both of these sections are characterized by short, unbranched, suberect to erect rhizomes resulting in the plants growing as individuals. In P. monticola, however, the rhizome is decumbent and mostly repeat- edly branched: as a result plants form large clonal stands. In spore morphology it is closer to section Metapolystichum than to section Lasiopolystichum . VARIATION. Depending on habitat, Polystichum monticola shows considerable variation in frond size, lamina texture, pinnae arrange- ment and pinnule size. Plants from xeric habitats have short fronds with closely spaced pinnae and small imbricate pinnules that are 70 J.P. ROUX firm-herbaceous in texture. Some collections proved to have unusu- ally large indusia, with a maximum radius up to 1.04 mm having been measured on a plant from Milner Peak in the Hex River Mountains [Esterhuysen 14885 (NBG, BOL)]. Indusia of the type collection are unusually small with the mean radius being merely 0.36 mm (n=6). Collections with these unusually large indusia occur at random throughout the distribution of the species and cannot be ascribed to any environmental condition. The same pattern applies to the often black-centred indusia. Although the number of indurated annulus cells per sporangium is relatively uniform in the species, unusually low and high numbers have been recorded. A plant from Leribe, Lesotho [Dieterlen 695 (SAM)] has sporangia with the number of indurated annulus cells varying between 7 and 13 (x=10.66; n=50), whereas a plant from Platberg, Harrismith [Roux 2523 (NBG)] has sporangia with the number of indurated annulus cells varying between 13 and 17 (x=14.7; n=50). Plants from forest habitats generally have larger fronds that are softer in texture to those occurring in more exposed habitats where the fronds are smaller and firm-herbaceous to subcoriaceous. DISTRIBUTION AND ECOLOGY. Polystichum monticola is confined to southern Africa where it has been recorded from the Northern Cape, Western Cape, Eastern Cape, KwaZulu-Natal and Free State, as well as in Lesotho, with one isolated record from the Victoria Falls in Zimbabwe that needs to be confirmed. The species is largely confined to the mountains ranging from the Cape Peninsula, along the southern Cape mountains to the Drakensberg Escarpment as far north as Platberg in the northeastern Free State. Plants have also been recorded from outlying locations such as the Kamiesberg in the Northern Cape Province and from Thaba 'Nchu Mountain in the eastern Free State. This apomictic taxon occurs at elevations ranging from 600 to 2740 m in often xeric environments. The habitat includes rock crevices in screes, cliff bases, streambanks, forest margins and forest floors. Often growing in exposed habitats where it forms large masses, the species is frequently exposed to veld fires, from which it soon recovers. 15. Polystichum dracomontanum Schelpe & N.C. Anthony in Contr. Bolus Herb. 10: 145 (1982). Type: Natal, Bergville Divi- sion, on banks above stream in side kloof west of main kloof, shortly above the Singati Cave (E. of Mont-aux-Sources), in unburnt sparse bush or small trees or in the open, c. 6000 ft, Esterhuysen 35646 (BOL!-holotype; B!, BOL!, C, GH, K, M, MO, NU!, P, PRE!, S-isotypes). Fig. 15. Plants terrestrial or epilithic. Rhizome decumbent, stoloniferous, to 10 mm in diameter, densely set with roots, persistent stipe bases, and paleae; paleae ferrugineous, broadly attached, chartaceous, linear- attenuate, entire, the apex terminating in a small thin-walled cell, to 28 x 2 mm. Fronds closely spaced, 5-7 per plant, erect or arching, to 1.15 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 540 mm long x 6 mm in diameter, initially densely paleated, becoming glabrous with age, proximally with paleae similar to those on the rhizome, the paleae distally of two types; larger paleae broadly attached, ferrugineous to castaneous throughout or stramineous to ferrugineous with a castaneous central region or apex, chartaceous, narrowly ovate to ovate, cordate, the margins with short, straight or curved projections, the apex often flagelliform, terminating in a thin-walled cell, to 25 x 6 mm; smaller paleae ferrugineous to stramineous, membranous, narrowly ovate to lanceolate, cordate to cordate-imbricate, the margins closely set with short and/or long, straight or curved, often branched outgrowths, the apex flagelliform, acicular or terminating in an oblong to clavate thin-walled cell, to 1 1 .5 x 1.8 mm: lamina 2-pinnate to 2-pinnate- pinnatifid, with up to 24 free pinna pairs, ovate, to 610 mm long: rachis stramineous to greenish, adaxialy sulcate, initially densely paleated; paleae short-stalked, membranous, narrowly lanceolate to narrowly ovate, cordate to cordate-imbricate, the margins proxi- mally with long and/or short, straight or curved, often branched outgrowths, the apex acicular or flagelliform and terminating in a thin-walled cell, to 7.3 x 1.6 mm: pinnae proximally short-stalked, 1 -pinnate to 1-pinnate-pinnatifid, with up to 18 free pinnule pairs, proximally widely spaced, often slightly reduced, distally often overlapping, ovate to narrowly ovate, to 105 mm long: pinna-rachis stramineous, adaxially sulcate, initially densely paleated; paleae stramineous, membranous, narrowly oblong to narrowly triangular, cordate to cordate-imbricate, the margins proximally with long and/ or short, curved, often branched outgrowths, the apex acicular, to 9.75 x 2 mm: pinnules opposite to alternate, firmly herbaceous to coriaceous, dark green adaxially, paler abaxially, proximally widely spaced to imbricate, asymmetric, ovate to ovate-rhomboid, often somewhat falcate, acroscopically auricled, serrate to doubly-serrate, sharp-tipped to strongly aristate, the margins somewhat revolute, the proximal acroscopic pinnule usually slightly longer than the next, to 22 mm long; adaxially glabrous or with a few stramineous, membranous, filiform, acicular paleae along proximal part of the costa, to 9.5 mm long, abaxially sparsely paleated; paleae short- stalked, stramineous, membranous, filiform, narrowly oblong to narrowly triangular, cordate, the margins proximally with short and/ or long, straight or curved outgrowths, the apex terminating in an acicular cell or with a thin-walled cell, to 8 mm long. Venation immersed or raised. Son circular, c. 2 mm in diameter, terminal or near-terminal on abbreviated vein branches, uniseriate, discrete to confluent at maturity: sporangium with 10-(13)-21 indurated annu- lus cells; stalk eglandular: indusium ferrugineous to stramineous, chartaceous, persistent, peltate, circular, with or without central processes, repand to weakly erose, the maximum radius 0.63- (0.81)-1.09 mm. Spores 64 per sporangium, brown, the perispore globose or folded to form inflated or narrow reticulate ridges, minutely and sparsely perforated, the ridges and areas between echinate to echinulate, the exospore 25-(54.24)-70 x 28-(41.04)- 58 (im. Chromosome number unknown. MATERIAL EXAMINED LESOTHO. 2927 (Maseru): pente de la montagne de Mathatha district de Mafeteng (CC), Dieterlen s.n. (P). SOUTH AFRICA. 2828 (Bethlehem): Tugela Gorge above chain ladder, 6200 ft (DB), Milliard & Burtt 15445 (BOL, NU); Royal Natal National Park, Tugela Gorge, scrub above chain ladder, Roux 27 1 5 (NBG); tributary flowing into the Singati, E. of Mont-aux-Sources (DD), Esterhuysen 35644 (B, BOL, NBG, PRE); Bergville, Mont-aux-Sources, Schweickerdt 760 (PRE); Mbunduni (MnWeni area), 9000 ft, Esterhuysen 27811 (BOL); MnWeni Pass, 8000 ft, Esterhuysen 27839 (BOL, PRE). 2829 (Harrismith): Farm Bosch Hoek (AD), Roux 896, 897, 898 (NBG); MnWeni area, foot of Rockies Pass, 5500 ft, (CB), Esterhuysen 21656 (BOL); Rockies Pass, 8000 ft, Edwards 2145 (NU); Cathedral Peak area, shady side of kloof (CC), Esterhuysen 15486 (BOL, NBG); Cathedral Peak, sheltered slopes below Cleft Peak, 8000 ft, Esterhuysen 101 99a (BOL, PRE); between Cathedral Peak and Royal Natal National Park, MnWeni Pass, 9000 ft, Edwards 85 1 (NU, PRE); SE slope of The Camel, 8700 ft, Everson 73, 74 (BOL); NE facing slope of The Camel, 8700 ft, Everson 75, 76 (BOL); Bergville, Orange Peel Gap, 7200 ft, Everson 71, 72 (BOL); NE slope of The Camel, 7000 ft, Schelpe 756 (NU); Orange Peel Gap, 2420 m, Crouch 647 (NU); Cathedral Peak, Nixon s.n. (NU); Cathedral Peak area, Umbonbonja River, 6000 ft, Schelpe 1096 (NU). 2929 (Underberg): Injasuti Valley, Solitude (AB), Malan 1 (BOL, NBG); Injasuti Nature Reserve, Leucosidea scrub at camp- site, Roux 2721 (NBG); Ndedema Valley, 6000 ft, Noel 1 28 1 (GRA); Injasuti POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 71 10 mm Fig. 15 Polystichum dracomontanum. A, proximal part of lamina; B, rhizome; C, fertile pinnule. All drawn from Esterhuysen 35646 (BOL). 72 J.P. ROUX area, 6500 ft, Esterhuysen 35219 (BOL); Injasuti area, 6500-8000 ft, Esterhuysen 26039 (BOL); Injasuti area, 7000 ft, Esterhuysen 26050 (K); Tabamhlope Mountain, 6000 ft (BA), West 184 (PRE); Mulangane Ridge, above Carter's Nek, 7000-7300 ft (BC), Milliard & Burtt 17032, 17529 (BOL, NU, PRE); 5-7 miles NNW of farm Castle View, headwaters of Mlahlangubo River, 6700 ft (CB), Milliard & Burtt 15188 (BOL, K, NU); Cathedral Peak, Ndumeni Valley, 1950 m (CC), Farrell2\ (NH). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum dracomontanum may be confused with P. monticola, which occurs in the same region and often in similar habitats. It can, however, be separated from it in the decumbent rhizome that produces slender stoloniferous branches, in the stipe that becomes near glabrous with age, and in the ovate lamina. Other diagnostic features are the entire rhizome paleae that evidently always terminate in a short thin- walled cell, in contrast to the often acicular apical cell in P. monticola. Paleae from the stipe, rachis, pinna-rachis and abaxial lamina sur- face appear to terminate more often in a thin- walled cell in contrast to those of P. monticola, where the apical cell appears to be largely acicular. The adaxial surface of the pinnules is largely glabrous but a few filiform paleae may occur proximally along the costa. These paleae always terminate in an acicular apical cell. Paleae from the abaxial surface of the pinnules largely terminate in an acicular apex, but paleae terminating in a short thin-walled cell are not unknown. The smaller paleae in P. dracomontanum are stramineous and membranous. The coriaceous lamina and somewhat revolute pinnule margins are also diagnostic. Micromorphological characters sepa- rating P. dracomontanum from other taxa are the small, almost square, adaxial epidermal cells with almost straight anticlinal walls and the almost circular stomata that are visible at a x 1 2 magnifica- tion. Considering the rhizome and palea morphology, P. dracomontanum is allied to the P. pungens group of species. VARIATION. Relatively little variation occurs within the species. Variation was observed in the colour of the larger stipe paleae. In most cases they are stramineous to ferrugineous throughout, but in a few collections they are centrally castaneous with a narrow stramineous margin. The paleae also show some variation in that some terminate in a long acicular cell whereas others may terminate in a short, thin-walled cell. Pinnules vary in size and in the margins that may be sharp-tipped to aristate. DISTRIBUTION AND ECOLOGY. Polystichum dracomontanum is largely confined to the northern Drakensberg Escarpment between Lesotho and KwaZulu-Natal where it occurs on both the lower Clarens Sandstone and the upper basalt formations. The species also occurs further northwards along the escarpment between the Free State and KwaZulu-Natal. Isolated populations have also been reported from the Mafeteng District in southeastern Lesotho. It occurs at elevations ranging from 1675 to 2745 m. Within this distribution the species is restricted to two vegetation types as defined by Acocks (1988). Along the high Drakensberg escarpment it occurs in Themeda-Festuca Alpine Veld that receives an annual precipitation of 600-1 900 mm. Polystichum dracomontanum occurs in grasslands and scrub forests associated with this vegetation type. To the north, along the escarpment between the Free State and KwaZulu-Natal, dominated by the Clarens Sandstone formation, the species occurs in sheltered forests of the Highland Sourveld type. Precipitation in this region measures between 750 and 1500 mm. The habitat includes streambanks, boulder bases, screes and scrub, and rarely also forests. Polystichum dracomontanum prefers moist cool slopes in shaded gullies and kloofs where it often forms large stands. At certain sites the species is subjected to regular veld fires, but this appears to have no adverse effect on plants. 1 6. Polystichum incongruum J.P. Roux in Bot. J. Linn. Soc. 125: 36 (1997). Type: South Africa, 3320 (Montagu): Swellendam, Marloth Nature Reserve, Koloniesbos, in scree on dry E.-facing slope (CD), Roux 2377 (NBG!-holotype). Fig. 16. Plants terrestrial. Rhizome decumbent, stout, to 16 mm in diameter, sparsely branched, set with roots, crowded stipe bases, and castaneous to ferrugineous, chartaceous paleae. Fronds crowded, to 8 per plant, suberect to arching, to 1.8 m long: stipe proximally castaneous, stramineous for most of its length, adaxially sulcate, to 930 mm long x 7 mm in diameter, moderately to densely paleated; paleae at stipe base of two types, the larger broadly attached, ferrugineous to stramineous, membranous, narrowly to broadly ovate-acuminate, rarely with unicellular clavate cells on the paleae surface, cordate to cordate-imbricate, the margins with short close-set outgrowths, the apex often flagelliform, terminating in an acicular cell or an oblong thin-walled cell, to 20 x 6 mm, the smaller sessile to short-stalked, stramineous, membranous, narrowly ovate-acuminate to narrowly triangular, truncate to cordate-imbricate, with short and/or long, straight or curved, somewhat lacerate outgrowths proximally, the apex flagelliform, entire and twisted, mostly terminating in an acicular cell but often also in a thin- walled cell: lamina 2-pinnate to 3-pinnate, with up to 27 free pinna pairs, ovate to broadly ovate, to 870 mm long, the pinnae more widely spaced proximally, the distal pinnae often imbricate, the proximal pinnae not to strongly reduced, not or slightly deflexed: rachis stramineous, adaxially sulcate, moderately to densely paleated; paleae sessile to short-stalked, ferrugineous to stramineous, membranous, narrowly ovate-acumi- nate to narrowly triangular-acuminate, truncate, cordate or cordate-imbricate, the margins proximally with short and/or long, curved, often branched outgrowths reduced in size and number towards the apex, the apex often flagelliform, twisted, terminating in an acicular cell or an oblong thin-walled cell, to 12x3 mm: pinnae 1 -pinnate to 2-pinnate, with up to 21 free pinnule pairs, narrowly oblong-attenuate to narrowly ovate-attenuate, themiddle pinnae to 265 mm long, to 95 mm wide: pinna-rachis stramineous, adaxially sulcate, moderately to densely paleated; paleae sessile to short- stalked, ferrugineous to stramineous, membranous, narrowly ovate-acuminate, narrowly triangular-acuminate to subulate, cor- date to cordate-imbricate, the proximal margins with short and/or long, often branched outgrowths reduced in size and number to- wards the apex, the apex often flagelliform, twisted, largely terminating in an acicular cell but often also in an oblong thin- walled cell, to 7 x 2 mm: pinnules opposite to alternate, closely to widely spaced, often slightly imbricate, firmly herbaceous to coriaceous, dark green adaxially, slightly paler abaxially, inaequilateral, ovate to narrowly trullate, often somewhat falcate, basiscopically cuneate, acroscopically cuneate to truncate and auriculate, the larger pinnules commonly deeply incised to form free or nearly free, narrowly ovate, narrowly elliptic to narrowly obovate ultimate segments, the margins serrate to doubly serrate, rarely aristate, the proximal acroscopic pinnule to 60 mm long, to 13 mm wide, often overlap- ping the pinna-rachis above; adaxially sparsely set with stramineous, membranous, twisted paleae chiefly along the costa, these filiform, narrowly linear to narrowly linear-hastate, simple or with few short marginal outgrowths proximally, the apex usually terminating in an acicular cell but often also in an oblong thin-walled cell, to 4.5 mm long, abaxially moderately paleated; paleae stramineous, membra- nous, subulate to narrowly triangular, twisted, short-stalked, truncate to cordate, proximally with short and/or long, often branched, somewhat lacerate marginal outgrowths, the apex usually termin- ating in an acicular cell, up to 3.5 mm long. Venation adaxially POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 73 10 mm Fig. 16 Polystichum incongruum. A, middle pinnae of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2377 (NBG). 74 obscure, raised abaxially. Sori circular, up to 1 .5 mm in diameter, terminal or near terminal on abbreviated vein branches, essentially uniseriate, discrete: sporangium with 1 1-(13)-21 indurated annulus cells; stalk glandular or eglandular: indusium stramineous or stramineous and black-centred, persistent, circular, simple or with central processes, the margins entire, minutely undulate to erose, often bearing clavate unicellular glands, the maximum radius 0.22- (0.46)-0.82 mm. Spores stramineous, the perispore folded to form narrow and broad reticulate ridges, the ridges crested, the crests and areas between minutely echinulate, porate, the exospore 30-(43.84)- 72 x 22-(33.22)-52 urn. Chromosome number 2n=164, n=82. MATERIAL EXAMINED SOUTH AFRICA. 3226 (Fort Beaufort): summit of Katberg Pass (BC), flow* 2426 (NBG); Katberg forests, 3000 ft, Adams 113, 118, 134, 147(NU); Hogsback, Zingcuka Forest (DB), Roux 2409, 2412 (NBG); Hogsback, Elandsberg, SE slope above scree, 4900 ft, Furness & Phillipson 49 (PRE); Hogsback, Auckland Forest Station, Roux 2418, 2419, 2420, 2421 (NBG); Hogsback Forest Reserve, 750 m, Dahlstrand 2926 (NBG); Hogsback Forest Reserve 'Fern Walk', 800 m, Dahlstrand 1550 (PRE); Auckland Forest, Hogsback, 750 m, Dahlstrand 28 1 9 (NBG); Hogsback, 3900 ft, Griffen 1 852 (PRE); Hogsback, Buckner s.n. (NU); Hogsback, Dahlstrand 1853 (NU); Hogsback, 4000 ft, Hoal 36 (NU); Middle Drift District, Hogsback, 3900 ft, Schelpe 6556 (B, PRE). 3227 (Stutterheim): Keiskamma Hoek, Gxulu Mountain, 5500 ft (BA), Story 3507 (PRE); Keiskammahoek, Gxulu Kop, 4500 ft (CA), Wells 3343 (PRE); Perie Forest (CB), Sim s.n. (PRE); Stutterheim, Evelyn Valley, Taylor 4203 (NBG); Mount Kemp, 5000 ft, Sim s.n. (PRE); Perie Forest, 4000 ft, Sim s.n. (PRE); Amabele (DA), Hardcastle 293 (NBG, PRE); 1 km past Amabele station on road to Stutterheim, Roux 2678 (NBG). 3320 (Montagu): Swellendam, Koloniesbos (CD), Roux 2373, 2377, 2589, 2590, 2591 (NBG); Swellendam, Duiwelsbos, along trail lead- ing to Die Plaat, Roux 2592, 2593, 2595 (NBG); Swellendam, Wamakersbos, Roux 2594 (NBG); Heidelberg, Grootvadersbos, Safraandraai (DD), Roux 2598 (NBG); Grootvadersbosch West, 1200 ft, Kruger 1322 (NBG, PRE); Grootvadersbos, Plantkunde Dept. Univ. van Stellenbosch s.n. (NBG); Swellendam, Strawberry Hill, Esterhuysen 10371 (BOL). 3322 (Oudtshoorn): Mossel Bay, Robinson Pass, Ruitersbos, Boesmansrivier, 1200 m (CC), Roux 2603, 2604, 2605, 2606 (NBG); Mossel Bay, Ruitersbos Forest Station, Roux 2381 (NBG); Groot Brakrivier, Jonkersberg Forest Station, Langbos, 380 m, Roux 2607, 2608, 2609, 2610, 2611, 2612 (NBG); Robinson Pass, 1700 ft, Schelpe 4989 (BOL); George, 1 km from turnoff to Witklippen Forest on old George-Knysna road (CD), Roux 2391, 2392 (NBG); George, lower circular drive, 275 m, Cameron 90 (PIC-SERM); George, Paterson 1239 (BOL); George, Schlechter 525 (PRE); George District, 200 m, Humbert 9834 (PRE); George, Montagu Pass, forest at summit of pass, Roux 2613 (NBG); Montagu Pass, Rehmann 118 (B); Woodville Forest, 440 m (DA), Roux 2622, 2623, 2624, 2625 (NBG); George, on road to Bergplaas Forest Station, 1 km past turnoff to Woodville hiking trail (DC), Roux 2395, 2396, 2397, 2398 (NBG); George, Saasveld Forest Station, Groenkop Research Area, Roux 2434 (NBG); George, Groenkop, Swartrivier, 300 m, Van Daalen 167 (BOL); old road between George and Knysna, above Touws River, Knysna side, 260 m, Roux 2620, 2621 (NBG); George, Groenkop Forest, Geldenhuys 394 (BOL); Wilderness, Levyns s.n. (BOL); George, Wilderness, Mogg 11656 (PRE); George, Schlechter s.n. TM525 (PRE); in silvis pr. George, 300 m, Schlechter 2441 (B); George, Wilderness, Jacobsen 2292 (PRE); George, Saasveld, forest above the reservoir, Roux 2384, 2385 (NBG); George, 1 .2 km from Saasveld turnoff on road to George, Roux 2437, 2438, 2439, 2440, 2441, 2614, 2615, 2616, 2617, 2618 (NBG); old road between George and Knysna, above Touwsriver, George side, 210 m, Roux 2619 (NBG); Goudveld Forest Reserve, Jubilee Creek forest walk, 340 m (DD), Roux 2627, 2629 (NBG); Farleigh Forest Station, forest above Platbos hut, Roux 2401 (NBG). 3323 (Willowmore): Concordia Forest Station, near Witklipdraai (CC), Roux 2637 (NBG); Knysna, Kom-se-Pad, Gouna Forest, Grootdraai picnic site, Roux 2638 (NBG); Knysna, Kom-se-Pad, 2.2 km from T-junction to Diepwalle, Roux 2405 (NBG); Knysna, Kleinbos, Buffelsnek, Schelpe 4312 (BOL); Knysna, Paardekop, Steyn 720 (NBG); Knysna, Deepwalls, Schonau 318 (BOL); Knysna, Gouna, Schelpe s.n. (BOL); Keurbooms River Forest J.P. ROUX Reserve, 200 m (CD), Dahlstrand 1355 (NBG); Bloukrans Pass, Varkrivier (DC), Roux 2649, 2650, 2651 (NBG); Bloukrans Forest Station, Platbos, along hiking trail, 260 m, Roux 2645 (NBG); Bloukrans Pass, 300 ft, Schelpe 4342 (BOL); Bloukrans Pass, Acocks 21298 (PRE); Tsitsikama Forest Re- serve (DD), Roux 2647 (NBG); Storms River Forest Reserve, 100 m, Dahlstrand 1693 (PRE); Stormsriver, 250 ft, Schlechter 5963 (PRE); Stormsriver Gorge, 400 ft, Jacobsen 2331 (PRE); in umbr. pr. Storms River, 80 m, Schlechter 5963 (B); Tsitsikama Forest Reserve, 260 m, Roux 2648 (NBG). 3423 (Knysna): Knysna, Kaffirkop Forest, 420 m (AA), Roux 2640, 264 1 , 2642 (NBG), near Knynsna, Mitchell s.n. (M, PRE); Knysna, Kaffirkop Forest, Roux 1994, 1995, 1996 (NBG); Knysna, Marloth 5710, 5711 (PRE); Knysna, Rex s.n. (PRE); Knysna, Mitchell s.n. (BOL); Tzitzikamma forest, 1 mile E. of Storms River village, 850 ft (BB), Schelpe 4372 (BOL); Storms River mouth, Maguire 507 (NBG). 3424 (Humansdorp): Hofman's Bosch (BB), Britten s.n. (PRE). WITHOUT PRECISE LOCALITY. Puspasvlei, Voormansbosch, Duivelsbosch and Keurboom, 1000-4000 m, Zeyher 4610 (PRE); George to Wilderness, Moss 6280 (PRE); Zuurberg, Rogers s.n. (PRE); loco incerto, Dahlstrand 1303 (NU); loco incerto, Zeyher s.n. B-97089 (B); loco incerto, Burchell 5200 (B); Kaffrarian forests, sine coll. 96893 (B); loco incerto, sine coll. B-96855 (B); Pr. b. sp, Zeyher 4610 (B); Cap. b. Sp., Krebs 360 (B); Prom. bon. spei, During s.n. (B); Cap. b. sp., Drege s.n. (B); Pr. b. sp., Ecklon & Zeyher 63 (B); Cap Colonie, Breutel s.n. (B); loco incerto, Herb. Lipzig, A & B only B-97050 (B); loco incerto, Herb. Lipzig 97051 (B); loco incerto, Braun s.n. (B); loco incerto, Gueinzius s.n. B-97069 (B); between Plettenberg Bay and Humansdorp, Rodin 1191 (BOL, PRE, S); loco incerto, d'Urban s.n. (B); loco incerto, Hort. bot. Berol. B-97049 (B); loco incerto, Hort. bot. Berol. B-97047 (B); Pr. b. spei, sine coll. B-97048 (B). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Differentiation in the field between this species and Polystichum pungens is not always easy because of their sympatric distribution, the variation within and among populations, and the absence of stable macromorphological characters. However, P. incongruum can be separated from P. pungens by its thicker, more stout rhizome. The pinnules in the former species are narrower, more slender and acuminate (often also slightly falcate), compared to the somewhat ovate to ovate-oblong, almost obtuse pinnules of P. pungens. A further character separating P. incongruum from P. pungens is the frequent occurrence of clavate unicellular glands along the sporangium stalk in the former species. Clavate unicellular glands occurring along the indusium margin have also been observed in some populations in the southern Cape, a feature never occurring in P. pungens. The former species is furthermore a sexual tetraploid (2n=164), whereas P. pungens is a sexual octoploid (2n=328). VARIATION. Variation in lamina morphology is perhaps the most apparent, hence the specific epithet (incongruens = inconsistent). Lamina division may vary between 2-pinnate to 3-pinnate, the pinnae being narrowly oblong-attenuate and widely spaced to nar- rowly ovate-attenuate and imbricate, with extreme variations often occurring within populations. Proximal pinnae may be reduced or not with the length ratio between these and the middle pinnae ranging between 1 : 1 and 1 :0.42. The most proximal pinna pair is often deflexed. Pinnules also show a large degree of variation in size and incision as illustrated by Roux (19976). Proximal acroscopic pinnules also show a large degree of variation in length. In some plants these pinnules are only slightly longer than the next pinnule, but in others the pinnules may extend beyond the pinna-rachis of the pinna above. These variations do not appear to be environmentally induced, but the variation in frond size and lamina texture is clearly influenced by the environment. Plants occurring in exposed habitats in the Amatola Mountains have short erect fronds, a coriaceous lamina, and sprout from a stout rhizome, whereas plants growing in shaded habitats have long and slender arching fronds with a herba- ceous lamina, and the rhizome is slender and branches freely. Palea POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA morphology shows minor variations between plants from the south- ern and eastern part of the distribution. In the southern part of the distribution the apices of the generally long marginal outgrowths of the paleae tend to split leaving a somewhat lacerated appearance, whereas in the eastern part of the distribution the outgrowths are short and tend not to split at the apices. Indusia vary in size, shape, the absence or presence of central processes, and in the occurrence of clavate unicellular glands along the margin. When glandular cells are present along the indusium margin of a specific collection not all indusia will bear them. Glandular cells along the sporangium stalk may be present or absent. Since these variations occur randomly they are not considered to be environmentally induced. DISTRIBUTION AND ECOLOGY. Polystichum incongruum is confined to the Western and Eastern Cape Provinces of South Africa. The species has a somewhat disjunct distribution, with a southern centre ranging from Swellendam to Hofmans Bosch and an eastern region centred in the Amatola Mountains. In the Swellendam region the species occurs in forests of the 'Ngongoni veld type (Acocks, 1988). These forests are small, isolated, and confined to deep sheltered ravines and steep slopes below the south-facing cliffs. In the south- ern Cape the species is confined to the Knysna forest type, which is more extensive. This region receives a well-distributed rainfall that ranges between 460-1250 mm per annum. Soil in this region is sandy and is largely derived from weathered Table Mountain Sand- stone. In the eastern part of its distribution the species occurs at elevations ranging between 600 and 1350 m where forests of the Dohne Sourveld type are predominant. In this region, however, the species is not confined to forested habitats but also occurs above the tree-line. The region receives an annual rainfall of 600-1000 mm with regular snowfalls during winter and the soil is predominantly of doleritic derivation. In the southern part of its distribution the species is confined to forests where it forms small or large clonal stands and is especially common in light shade along streambanks, road cuttings and forest clearings. It may occur in dryish or very wet conditions. In the eastern part of its distribution the species occurs in more varied habitats ranging from natural forests to pine plantations but also occurs above the tree line forming large stands along streams, on screes and at boulder and cliff bases. Plants in the latter habitats are exposed and generally stunted with short erect fronds. In this region the plants are frequently subjected to veld fires but this appears not to adversely affect them. IT.Polystichum pungens (Kaulf.) C. Presl, Tent, pterid.: 83 (1836); Schelpe & Anthony, Fl. Sthn. Afr, Pterid. : 254 ( 1 986), pro parte; Burrows, Sthn. Afr. ferns and fern allies: 312 (1990), pro parte. Type as for Aspidium pungens Kaulf. Fig. 17. Aspidium pungens Kaulf, Enum.fil: 242 (1824). Type: Habitat in Promentorio b. spei, Chamisso s.n. (LE-holotype; BOL!-photo- graph). Dryopteris pungens (Kaulf.) Kuntze, Rev. gen. pi. 2: 813 (1891). Plants terrestrial or epilithic. Rhizome decumbent, sparsely branched, to 370 mm long, to 20 mm in diameter, set with roots and closely to widely spaced persistent stipe bases, the older parts nude, the apical part densely paleated; paleae broadly attached, castaneous to ferru- gineous, chartaceous, narrowly lanceolate to narrowly ovate, truncate to cordate, the margins subentire to erose, the apex flagelliform, generally terminating in a small thin- walled cell, to 17 x 3 mm. Fronds 5-6 per plant, suberect to arching, to 1.4 m long: stipe proximally castaneous, stramineous distally, adaxially sulcate, to 75 685 mm long x 7 mm in diameter, sparsely to densely set with conspicuous larger and smaller paleae; larger paleae more frequent proximally, widely spaced and smaller distally, proximally castaneous, broadly attached, distally ferrugineous, chartaceous, narrowly to broadly ovate, often oblique, cordate to cordate-imbri- cate, the margins minutely erose to short-fimbriate, the apex flagelliform, terminating in a long acicular cell or a small oblong thin- walled cell, to 2 1 x 6 mm; smaller paleae short-stalked, ferrugi- neous to stramineous, chartaceous to membranous, narrowly triangular, narrowly lanceolate to narrowly ovate, cordate to cor- date-imbricate, the margins proximally erose or with short and/or long, straight or curved outgrowths, the apex entire, flagelliform, terminating in a long acicular cell or a small oblong thin-walled cell: lamina 2-pinnate to 2-pinnate-pinnatifid, with up to 21 free pinna pairs, firmly herbaceous, adaxially dark green, abaxially slightly paler, ovate to broadly ovate, to 704 mm long, the pinnae often slightly imbricate distally, more widely spaced proximally, the most proximal pinna pair slightly reduced, often somewhat deflexed: rachis stramineous to greenish, adaxially sulcate, sparsely to densely paleated; paleae short-stalked, ferrugineous to stramineous, mem- branous, ovate, narrowly ovate to narrowly triangular, sessile or short-stalked, cordate to cordate-imbricate, the proximal margins erose to sparsely fimbriate or with short and/or long, curved or angular, often branched outgrowths that reduce in size and number towards the apex, the apex flagelliform, terminating in a long acicular cell or a small thin-walled cell: pinnae 1 -pinnate to 1- pinnate-pinnatifid, with up to 24 free pinnule pairs, narrowly ovate-attenuate to narrowly oblong-attenuate, to 272 x 48 mm; pinna-rachis stramineous, adaxially sulcate, moderately to densely paleate; paleae ferrugineous to stramineous, membranous, narrowly triangular, short-stalked, cordate-imbricate, the proximal margins with short and/or long, often branched outgrowths reduced in size and number towards the apex, the apex flagelliform, twisted, termin- ating in an acicular cell or a small thin-walled cell, to 4.5 mm long, each pinna often subtended by one or more large, often bullate, broadly ovate, cordate, minutely erose to fimbriate paleae: pinnules opposite to alternate, widely spaced to slightly imbricate, the proxi- mal acroscopic pinnule the largest, often significantly longer than the next, especially towards the middle of the lamina, each sub- tended by one or more large, often bullate, broadly ovate paleae, similar to but smaller than those on the rachis, inaequilateral, ovate, ovate-oblong to ovate-rhomboid or trullate, often somewhat falcate, basiscopically cuneate, acroscopically cuneate to truncate and au- riculate, the larger pinnules commonly deeply incised near to the costa forming a nearly free auricle acroscopically, the margins serrate to lobate-serrate, sharp-tipped, rarely aristate, the costa proximally adaxially sulcate, most proximal acroscopic pinnule to 50 mm long, to 19 mm wide, often reaching beyond pinna-rachis above; adaxially sparsely set with paleae chiefly along costa, fili- form to taeniform, the margins entire or proximally with a few short curved or long angular outgrowths, the apex terminating in a long acicular cell or a small thin-walled cell, to 6 mm long; abaxially sparsely to moderately paleate, the paleae stramineous, membra- nous, short-stalked, subulate, narrowly triangular to narrowly ovate, cordate to cordate-imbricate, the margins proximally with short and/ or long, angular outgrowths, the apex entire, filiform, terminating in a long acicular cell or a small thin-walled cell, to 3.7 mm long. Venation adaxially obscure, raised abaxially. Sori circular, c. 1 mm in diameter, terminal or near terminal on abbreviated vein branches, essentially uniseriate, discrete at maturity: sporangium with 10- ( 1 2.8)- 1 9 indurated annulus cells; stalk eglandular: indusium peltate, stramineous, castaneous or black, nitid, amorphous to circular, entire to repand, the maximum radius 0.26-(0.5)-0.8 mm in diameter, 76 J.P. ROUX 10 mm Fig. 17 Polystichum pungens. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2367 (NBG). POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 77 persistent, brown. Spores pale brown, 64 per sporangium, the perispore folded to form inflated reticulate ridges, the ridges crested, the ridges and areas between sparsely to densely echinulate, vari- ously porate, the exospore 30-(49.31)-62 x 28-(38.08)-56 prn. Chromosome number 2n=328. MATERIAL EXAMINED SOUTH AFRICA. 2330 (Tzaneen): Woodbush (CC), Wager s.n. CH7461 (PRE); De Hoek, Woodbush, Schweickerdt s.n. TM1852C (PRE). 2430 (Pilgrim's Rest): Pilgrim's Rest, Mount Sheba Nature Reserve, Waterfall trail (DC), Roux 2554 (NBG). 2730 (Vryheid): Hlobane, Mto'.a forest (DB), Johnstone 295 (NU). 2828 (Bethlehem): Royal Natal National Park, Gudu forest, Gudu waterfall (DB), Roux 2512 (NBG); Royal Natal National Park, Gudu forest, Schelpe 1454 (NU). 2829 (Harrismith): Farm Whitestones (CA), Roux 1901 (NBG, PRE); Bezuidenhouts Pass, farm Whitestones, Roux 1684 (NBG, PRE); Oliviershoek Pass, forests S. of Seheletwane, Roux25\9 (NBG); Cathedral Peak area, Indumeni forest, 5100 ft (CC), Schelpe 781 (NU). 2831 (Nkandla): Eshowe (CD), Laura s.n. CH6421 (PRE). 2929 (Underberg): Giants Castle Nature Reserve, forest above Hillside camping site (AB), Roux 2500, 2502, 2505 (NBG); Lions River District, Lions Bush, 5000 ft (BD), Moll 832 (PRE). 2930 (Pietermaritzburg): Buccleuch (AD), Sim s.n. CH3641 (PRE); Hilton Road (CB), Wager s.n. B-only (NU); Hilton District, Cedara Dam, 3200 ft, Churcher s.n. (NU); Zwaartkop, Sim s.n. PRE- 11026 (PRE); Sim s.n. CH387 (PRE); Sim s.n. TM1239C (PRE). 2931 (Stanger): prope Mapumulo (AA), Abraham s.n. (B). 3029 (Kokstad): Insizwa forest (CC), Strey 10749 (PRE). 3030 (Port Shepstone): Paddock, Burntwood (CC), Strey 5994 (PRE). 3128 (Umtata): Tsolo District, Ngadu, 3200 ft (BC), Von Breitenbach 1330 (PRE). 3129 (Port St Johns): Port St Johns (DA), Wager s.n. CH2996 (PRE). 3225 (Somerset East): Boschberg (DA), MacOwen s.n. (P); forest kloofs of the Boschberg mountain, sine coll. s.n. (P); in sylvis ad ped. Mont. Boschberg, 3000 ft (DA), MacOwen s.n. (B). 3226 (Fort Beaufort): Katberg forests (BC), Adams 168, 175 (NU); Hogsback, Auckland forest (DB), Roux 510 (NBG); Hogsback forest, 4250 ft, Schirach 280 (NBG); Hogsback, Stirton 6267 (PRE). 3227 (Stutterheim): Fort Cunninghame, 3300 ft (AD), Galpin 2446 (PRE); Isidenge forest (CA), Roux 1981 (NBG); Keiskamma Hoek, 2000 ft, Ely s.n. (PRE); Cathcart, Fort Cunningham forest Reserve (CB), Roux 2428, 2429, 2432 (NBG); Frankfort, Sim s.n. (PRE); Pirie, Sim s.n. (PRE); Dohne Hill, Sim s.n. (PRE); Komgha (DB), Flanagan s.n. (PRE). 3318 (Cape Town): Nursery Gorge (CD), Schelpe s.n. BOL-35933 (BOL); head of Nursery Gorge, Esterhuysen 2585 1 (BOL); top of Nursery Gorge, Esterhuysen 15355 (BOL); Devils Peak, Dark Gorge, Esterhuysen 26564 (BOL, NBG); Table Mountain, Hiddingh Ravine, Esterhuysen 25862 (BOL); Kirstenbosch, sine coll. BOL-55808 (BOL); Devils Peak, waterfall, Wolley-Dod 915 (BOL); Table Mountain, Skeleton Gorge, Schelpe s.n. (BOL); SE slopes of Devils Peak, Pillans 2694 (BOL); Skeleton Gorge, Esterhuysen 26674 (BOL); Skeleton Gorge, Roux91 (BOL); mountain woods at back of Newlands, sine coll. s.n. (P); Newlands Ravine above contour path, Roux 2370, 2371 (NBG); Lubberts Gift, Roux 2372 (NBG); Kirstenbosch, contour path, Compton 14629 (NBG); Window Gorge, Roux 36 (NBG); Dark Gorge, Roux 2367, 2368a, 2369 (NBG); Window Gorge, Wasserfall 84, 156 (NBG); Newlands, Paradise, Rawson s.n. (SAM). 3320 (Montagu): Heidelberg, Grootvadersbosch (DD), Roux 2596 (NBG); Grootvadersbosch, Safraandraai, Roux 2597, 2599 (NBG); Grootvadersbosch, Stinkhout hiking trail, Roux 2378, 2379, 2380 (NBG); Grootvadersbosch, Taylor 1228 (BOL); Grootvadersbosch, near end of road running past red- woods, Roux 2600 (NBG). 3322 (Oudtshoorn): George, Wildernes (DC), Compton 14305 (NBG); Goudveld Forest Station, Krisjan se Nek picnic site (DD), Roux 2626 (NBG). 3325 (Port Elizabeth): Enon (BC), sine coll. B- 97063 (B). 3326 (Grahamstown): in sylvis prope Grahamstown (BC), MacOwan s.n. (P); kloofs near Grahamstown, Holland s.n. (NBG). 3418 (Simonstown): Diepgat, kloof below SW Triplets (BB), Esterhuysen 27060 (BOL); ravines of the Helderberg, Parker 4311 (BOL). 3419 (Caledon): Riviersonderend, Oubos (BB), Roux 2586 (NBG). 3423 (Knysna): Knysna (AA),Marloth 1901 (L). SWAZILAND. 2631 (Mbabane): Millers Falls, 4500 ft (AC), Compton 25967 (NBG); 5 km NW of Mbabane, 1200 m, Kemp 896 (PRE). WITHOUT EXACT LOCALITY: loco incerto, sine coll. BOL-55877 (BOL); Albany District, Cooper 1415 (P); Africa austral, Drege s.n. (P); Cap. b. spei, Bojer s.n. (P); Cap. b. spei, Herb Musei Palat. Vindob. 126 (P); in umbrosis montium Hottentots Hollandiae, Zeyher s.n. (SAM); Drakensberg, Bottomley s.n. CH5018 (PRE); Fort Beaufort District, Myburg s.n. (NBG); Bedford District, Van Rensburg s.n. (NBG); Katberg, Young s.n. (PRE); Natal, Pondoland and Zululand midlands, Watt & Brandwyk 336 (PRE); district of Albany, Cooper 1415 (PRE); loco incerto, Flanagan s.n. (PRE); Kaffirland, St. Augustine, Baur 215, B-only (B); Afr. austr., sine coll. s.n. (B); Prom. b. spei, Krebs 360 (B), Natalia, Buchanan 85 (B); Natalia, in sylva ad fr. Tugela, Gueinzius s.n. (B). Controversy as to the correct name for this taxon has existed for a long time. Sim (1892) initially labelled this species as Aspidium aculeatum var. pungens, but by 1915 he realized that two entities could be recognized, a forest dwelling species that he referred to Polystichum aculeatum and a montane form that he referred to P. pungens. Becherer (1937), however, proposed the name P. lucidum (Burm.f.) Becherer (= Asplenium lucidum Burm.f.) for the forest growing species, a name that became well entrenched (Schelpe, 1969; Roux, 1979; Jacobsen, 1983). Following a reinterpretation of the types, Anthony & Schelpe (1985) concluded that Asplenium lucidum Burm.f. is synonymous with Asplenium adiantum-nigrum L. This largely follows the view of C.V. Morton who distributed photographs of what he believed to be the type of the species. A review of these anomalous typifications has been provided by Roux (1994). Since it was concluded that A. lucidum is synonymous with A. adiantum-nigrum, a new name was required for the forest species. The next available name for the species is P. pungens (Kaulf.) C. Presl (= Aspidium pungens Kaulf.). DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum pungens forms part of a species group characterized by decumbent rhizomes and paleae that usually terminate in a long flagelliform apex. It can, however, be separated from other taxa in the group by not having glandular cells on the sporangium stalk and by the longer, more slender rhizome. Furthermore, P. pungens has a somatic chromo- some number of 2n=328, versus 2n=164 in P. incongruum with which it may be confused and to which it evidently is related. VARIATION. Variation in pinnule size and shape may be influenced by numerous environmental factors. Smaller pinnules may be ovate in outline and shallowly lobate-dentate. As the pinnules increase in size, they become more deeply lobate in the proximal part of the pinnule, often extending to the costa, resulting in the proximal acroscopic segment being short-stalked. The proximal acroscopic pinnule is generally longer than the following pinnule. The length ratio of the proximal and the following pinnule taken from the central part of the lamina ranges between 1:0.91 [Compton 14629 (NBG)] and 1:0.56 [Roux236Sa (NBG)]. Paleae vary mostly in shape and in the degree to which the margin is sculptured. Although most paleae terminate in a long filiform apical cell, some do terminate in a short, thin-walled cell. In one collection [Esterhuysen 26564 (BOL, NBG)] unicellular glandular cells also occur along the palea margin as well as on the surface of the larger rhizome paleae. Indusium size and outline vary considerably within the species. Although indusia are generally peltate, often some are reniform or have the flange not fully 360 developed. Both conditions are frequent within a single plant. Although the general outline of the indusium may be considered circular, it is often irregular with the margins varying from subentire to repand. Indusia appear to increase in size from the western part of the distributional range to the east. The plant with the smallest mean indusium radius was recorded from Table Mountain [Compton 14629 (NBG), x=0.3 mm; n=6] and the plant with the largest mean radial length is from the George region [Roux 2626 (NBG), x=0.7 1 mm; n=6]. The maximum radius of the indusia varies between 0.26 and 0.78 mm. Plants as far east as 78 J.P.ROUX Port Elizabeth usually have stramineous indusia, whereas plants ranging from the Boschberg farther north have dark, almost black indusia, and are uniform in outline. DISTRIBUTION AND ECOLOGY. Polystichum pungens is restricted to South Africa and Swaziland. In this region the species occurs from Table Mountain on the Cape Peninsula to the Hottentots Holland Mountains, along the Riviersonderend, Langeberg, Outeniqua and Great Winterhoek Mountains to Port Elizabeth and Grahamstown. Inland it occurs from the Boschberg at Somerset East to the Amatola Mountains and along the Drakensberg foothills to the Wolkberg in the Northern Province. In the eastern part of its distribution P. pungens is restricted to isolated climax forest patches largely restricted to the southern mountain aspects and sheltered ravines. This region, and the more extensive Knysna forest complex, is subject to a high, well-distrib- uted rainfall and acidic sandy soils. At Grahamstown and on the Boschberg the species occurs in temperate scrub forest subject to more seasonal precipitation. In the Amatolas the species occurs in forests of the Dohne Sourveld type that lie between 600-1350 m above sea level. From here the distribution extends to the eastern slopes and foothills of the Drakensberg. Forests in this region are of the Highland Sourveld type, which is largely confined to the deep gorges and protected mountain slopes occurring at elevations rang- ing from 1350-2150 m. To the north the species occurs in forests of the Northeastern Mountain Sourveld. In the Eastern Cape and southern KwaZulu-Natal it has been reported from forests of the Pondoland Coastal Plateau Sourveld that are found at an elevation of 300-450 m. These forests are largely confined to the escarpment, gorges and valleys below krantzes. In the KwaZulu-Natal midlands it is confined to forests of the Mist Belt 'Ngongoni Veld, whereas in northern KwaZulu-Natal it occurs in 'Ngongoni Veld. Polystichum pungens is a terrestrial or epilithic species occurring as isolated individuals or often also as large clones on dryish or moist slopes in partially to deeply shaded conditions. 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Sledge, W.A. 1973. The dryopteroid ferns of Ceylon. Bulletin of the British Museum (Natural History), Botany 5: 1-43. Tagawa, M. 1940. Polystichum of Japan, Korea, and Formosa I. Acta Phytotaxonomica Geobotanica 9: 119-138. Tardieu-Blot, M.L. 1964. Pteridophytes. In A. Aubrdville (Ed.), Flore du Cameroun, 3: Paris. Tryon, R.M. 1985. Fern speciation and biogeography. Proceedings of the Royal Society of Edinburgh, 86B: 353-360. & Tryon, A.F. 1982. Ferns and allied plants with special reference to tropical America. New York. Verwoerd, W.J. 1 97 1 . Geology. In E.M. van Zinderen Bakker, Sr., J.M. Winterbottom & R.A. Dyer (Eds), Marion and Prince Edward Islands: 40-62. Cape Town. Vida, G. & Reichstein, T. 1975. Taxonomic problems in the fern genus Polystichum caused by hybridization. In S.M. Walters (Ed.), European floristic and taxonomic studies: 126-135. England. Wagner, D.H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1: 1-64. White, F. 1978. The Afromontane region. In M.J.A. Werger (Ed.), Biogeography and ecology of southern Africa: 463-513. The Hague 1983. The vegetation of Africa. Paris. Yatskievych, G. 1996. A revision of the fern genus Phanerophlebia (Dryopteridaceae). Annals of the Missouri Botanical Garden 83: 168-199. Zhang, L. & Kung, H. 1995. A taxonomic study on the fem genus Polystichum Roth sect. Metapolystichum Tagawa from China (I). Acta Phytotaxonomica Sinica 33: 469^75. 1996o. A taxonomic study on the fern genus Polystichum Roth sect. Metapolystichum Tagawa from China (II). Acta Phytotaxonomica Sinica 34: 68- 76. 19966. A taxonomic study on the fern genus Polystichum Roth sect. Metapolystichum Tagawa from China (III). Acta Phytotaxonomica Sinica 34: 194- 203. SYSTEMATIC INDEX Accepted names are in roman and synonyms in italics. Acropelta Nakai 35 Acropelta omeiensis (C. Chr.) Nakai 35 Aetopteron House 35 Aspidium aculeatum (L.) Sw. 45 Aspidium aculeatum subsp. angulare (Kit. ex Willd.) Asch. 47 Aspidium angulare Kit. ex Willd. 47 Aspidium hastulatum Ten. 47 Aspidium lobatum (Huds.) Sw. 45 Aspidium luctuosum Kunze 39 Aspidium macleae Baker 36 Aspidium pungens Kaulf. 75 Aspidium tsus-simense Hook. 39 Aspidium volkensii Hieron. 41 Cheilanthes speciosissima Kunze 35 Dryopteris aculeata (L.) Kuntze 45 Dryopteris aculeata subsp. angularis (Kit. ex Willd.) Schinz & Thell. 47 Dryopteris pungens (Kaulf.) Kuntze 75 Dryopteris setifera (Forssk.) Woyn. ex Schinz & Thell. 47 Dryopteris setifera subsp. angularis (Kit. ex Willd.) Maire 47 Dryopteris setifera subsp. lobata (Huds.) Maire 45 Hemesteum H. Lev. 35 Hypopeltis Michx. 35 Hypopeltis lobulata Bory 35 Papuapteris C. Chr. 35 Papuapteris linearis C. Chr. 35 Plecosorus Fee 35 Plecosorus mexicanus Fee 35 Polypodium aculeatum L. 45 Polypodium lobatum Huds. 45 Polypodium lonchitis L. 35 Polypodium setiferum Forssk. 47 Polystichum Roth 35 Polystichum aculeatum (L.) Roth 35, 36, 45, 46 Polystichum aculeatum subsp. angulare (Kit. ex Willd.) Vollm. 47 Polystichum aculeatum van mildbraedii Brause 53 Polystichum aculeatum var. rubescens Bonap. 53 Polystichum aculeatum var. stenophyllon Bonap. 53 Polystichum alticola Schelpe & N.C. Anthony 53 Polystichum angulare (Kit. ex Willd.) C. Presl 47 Polystichum barbatum C. Chr. 41 Polystichum x bicknellii (H. Christ) Hahne 47 Polystichum coursii Tardieu 45 Polystichum cystostegia (Hook.) J.B. Armstr. 60 Polystichum discretum (D. Don) J. Sm. 56 Polystichum dracomontanum Schelpe & N.C. Anthony 36,70, 71 Polystichum elegans J. Remy 60 Polystichum falcinellum (Sw.) C. Presl 37 Polystichum fuscopaleaceum Alston 53 Polystichum fuscopaleaceum var. ruwensoriense (Pirotta) Pic.Serm. 53 Polystichum glaciale H. Christ 35 Polystichum incongruum J.P. Roux 36, 72, 73 Polystichum kalambatitrense Tardieu 37 Polystichum kilimanjaricum Pic.Serm. 36, 42, 44 Polystichum lemmonii Underw. 60 Polystichum lineare (C. Chr.) Copel. 35 Polystichum lobatum (Huds.) Bastard 45 Polystichum lobatum var. luctuosum (Kunze) H. Christ 39 Polystichum lobatum var. ruwensoriense Pirotta 53 Polystichum lonchitis (L.) Roth 35 Polystichum luctuosum (Kunze) T. Moore 36, 37, 40, 42 Polystichum macleae (Baker) Diels 36, 38 Polystichum magnificum F. Ballard 36, 63, 64 Polystichum marionense Alston & Schelpe 36, 58, 59 Polystichum mohrioides (Bory) C. Presl 60 Polystichum monticola N.C. Anthony & Schelpe 36, 67,68 Polystichum munitum (Kaulf.) C. Presl 37 Polystichum nigropaleaceum (H. Christ) Diels 56 Polystichum omeiense C. Chr. 35 Polystichum pauciaculeatum Bonap. 45 Polystichum plicatum (Poepp.) Hicken 60 Polystichum pungens (Kaulf.) C. Presl 36, 75, 76 Polystichum x saltum J.P. Roux 36, 56, 57 Polystichum scopulinum (R.J. Eaton) Maxon 60 Polystichum setiferum (Forssk.) T. Moore ex Woyn. 36,47,48 Polystichum setiferum var. fuscopaleaceum (Alston) Schelpe 53 Polystichum setiferum var. nigropaleaceum (H. Christ) Sledge 56 Polystichum speciosissimum (Kunze) R.M. Tryon & A.F. Tryon 35 Polystichum transkeiense W. Jacobsen 36, 60, 61 Polystichum transvaalense N.C. Anthony 36, 49, 50 Polystichum tsaratananense Tardieu 45 Polystichum tsus-simense (Hook.) J. Sm. 39 Polystichum volkensii (Hieron.) C. Chr. 36, 41, 43 Polystichum wilsonii H. Christ 36, 53, 54 Polystichum zambesiacum Schelpe 36, 63, 65 section Lasiopolystichum Daigobo 45, 52, 56, 63, 69 section Metapolystichum Daigobo 47, 69 section Xiphopolystichum Daigobo 41 Sorolepidium H. Christ 35 Sorolepidium glaciale (H. Christ) H. Christ 35 Bull. nat. Hist. Mus. Lond. (Bot.) 30(2): 81-99 Issued 30 November 2000 Recent records of pteridophytes for Belize, Central America xy O s DAVID A. SUTTON Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD ANGELA HUGHES International Centre for Underutilized Crops, Institute of Irrigation and Development Studies, University of Southampton, Southampton SO 17 1BJ BARBARA BULMER-THOMAS 55 Maze Hill, Greenwich, London, SEW 8XQ CONTENTS Introduction 82 Listoftaxa 83 Aspleniaceae 83 Blechnaceae 84 Cyatheaceae 84 Davalliaceae 84 Dennstaedtiaceae 85 Dicksoniaceae 85 Dryopteridaceae 85 Gleicheniaceae 86 Grammitidaceae 86 Hymenophyllaceae 86 Isoetaceae 87 Lomariopsidaceae 87 Lophosoriaceae 88 Lycopodiaceae 88 Marattiaceae 88 Metaxyaceae 89 Ophioglossaceae 89 Polypodiaceae 89 Psilotaceae 90 Pteridaceae 90 Salviniaceae 91 Schizaeaceae 91 Selaginellaceae 92 Tectariaceae 92 Thelypteridaceae 93 Vittariaceae 95 Woodsiaceae 95 Discussion 96 References ... 98 SYNOPSIS. Belize has one of the richest pteridophyte floras in the world, expressed on an area basis. Analysis of recent collections and the widely scattered literature for the country reveals that the number of species has risen by approximately 20% since the publication of the major regional flora, Flora Mesoamericana, in 1995. Investigation of the dates of discovery for each taxon produces a graph that indicates many more taxa are yet to be discovered in the country. A preliminary assessment is made of areas in Belize likely to reveal new records. The Natural History Museum, 2000 82 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS INTRODUCTION The vascular plants of Belize have never been treated adequately in a national flora. Early botanical investigation during the nineteenth century was focussed very much on economically important species, especially timber, latex and fruit, though the list produced for the now defunct Botanic Station by Campbell ( 1 899) included a number of native ferns and orchids. Other early accounts of the flora (Blake, 1917; Record, 1925), based on exploration and forestry, excluded pteridophytes. Standley & Record (1936) provided a good floristic grounding for the country with the Forests and flora of British Honduras, though this lacks some of the elements of a regular flora and the content is biased strongly towards trees both in species coverage and detail. Herbaceous angiosperms and pteridophytes are merely listed, except where they are believed to be new species. However, this single publication provided the earliest published record for almost 25% of species in the entire flora, far exceeding the significance of any other publication for Belize in this respect. A number of checklists have appeared subsequently for the country. Those by Dwyer and others (Spellman et al., 1975; Dwyer & Spellman, 1981) excluded pteridophytes, but some records appear in the catalogue by Schipp (1934). The catalogue of the Belize National Herbarium (BRH) by Vargas & Shawe (1997) includes a number of plant records for the country not published elsewhere. Fig. 1 Districts of Belize. Shaded areas represent 300 m and 600 m contours. RECENT RECORDS OF PTERIDOPHYTES 83 Recent expedition reports and surveys (for example Parker et al., 1993) include many of the new records of pteridophytes for the country. The regional Flora of Guatemala expressly included Belize (as British Honduras) in its remit (for example, Standley & Steyermark 1958) though for the majority of pteridophyte taxa listed (Mickel, 1981; Smith, 19816; Stolze, 1976, 1981, 1983; Stolze & Hickey, 1983; 011gaard, 1983) there is no explicit mention of the presence in or absence from Belize. For most vascular plants, this publication still provides the most complete coverage of the flora for Belize to date, though for pteridophyes it has been superseded by the regional Flora Mesoamericana (Davidse et al., 1994, 1995). This latter project will provide a comprehensive coverage of the species of Belize on its eventual completion and it has a unique strength in the development of a live link to the TROPICOS (Solomon et al., 2000) database in 1998, so that the Internet version of the flora is updated on a continual basis. Since the publication of the volume of Flora Mesoamericana, there has been considerable interest in the pteridophytes of Belize arising out of a number of projects (for example, Hughes, 1998) and recent fieldwork. A list of current names used for Belize, with an indication of their status is given below. The report includes an analysis of over 2500 pteridophyte specimen records from The Natural History Museum in London (BM) and other herbaria at AAU, B, BR, BRH, CR, DS, F, G, GH, K, LAGU, MEXU, MICH, MO, NY, S, SEL, TEX, UAMIZ, UC, UCWI, US and Z. Full use has been made of information sources on the Internet, and the authors particularly express their gratitude to Missouri Botanical Garden for making TROPICOS available on the World Wide Web. LIST OF TAXA The following list is arranged alphabetically by family following the family delimitation used in Flora Mesoamericana (Davidse et al., 1 995). Taxa previously recorded in Flora Mesoamericana are shown in italic and the superscript reference indicates the account and page number. Representative material is shown with one voucher for each district (Fig. 1) where the species has been recorded, except where the taxon has particular significance as a new record or endemic. Preference in citation is given to specimens cited in a monograph, revision or a recent regional flora. A superscript reference number or numbers following a specimen is to the source included in the bibliography where that specimen is listed. For new records not included in Flora Mesoamericana, the name is shown in bold and additional specimens are listed where known. An indication is given of where the taxon was first listed for Belize. ASPLENIACEAE Asplenium abscissum Willd., Sp. pi. 5: 321 (1810) 2:295 Toledo, G. Davidse & D.L. Holland 36836 (MO 115 ). Asplenium auritum Sw. in J. Bot. (Schrader) 1800(2): 52 (1801) 2:2 - Cayo, A. Hughes 64 (BM-000557717! 22 ); Toledo, T.B. Croat 24260 (MO 2 ). Asplenium cirrhatum Rich, ex Willd., Sp. pi. 5: 321 (1810) 2:2< Toledo, B.H. Allen 15443 (MO 2 ' 3 ' 115 ). Asplenium cristatum Lam., Encycl 2(1): 310 (1786) 2:3( Cayo, A. Hughes 98 (BM-000557710! 22 ); Stann Creek, W.A. Schipp 50 (BM 2 , BRH 125 , NY); Toledo, B.K. Hoist 4397 (MO 91 ). Asplenium delitescens (Maxon) L.D. Gomez in Brenesia 8: 52 (1976^301 Cayo, A. Hughes 12 (BM-000531878!, BM-000557708! 22 , BRH!); Stann Creek, P.H. Gentle 2714 (F, GH, K 2 , MICH, NY); Toledo, P.H. Gentle 6261 (F, G, NY, S, UC). Asplenium dentatum L., Sp. pi. 2: 1080-1081 (1753) Stann Creek, P.H. Gentle 2712 (NY); Toledo, T.E. Hawkins 1631 (MO 115 ). A new name for Asplenium trichomanes-dentatum L., listed for Belize in Flora Mesoamericana (Adams, 19956: 322) based on extralimital distribution in Proctor (1985: 375) but without refer- ence to a specimen. The name is automatically corrected according to the International Code of Botanical Nomenclature (Tokyo Code 23.8, ex. 14, Greuter et al., 1998). Vargas & Shawe (1997: 22) listed a specimen (W.A. Schipp 1066, BRH) under the name A. dentatum L., but this material has also been identified as A. macilentum by Adams (19956: 309) based on a duplicate at K. Asplenium formosum Willd., Sp. pi. 5: 329 (1810) 2:304 Cayo, G.R. Proctor 29902 (BM 2 ); Stann Creek, R. Rivera et al. 2576 (BRH 125 ); Toledo, G. Davidse 35782 (MO 115 ). Asplenium heterochroum Kunze in Linnaea 9: 67 ( 1 834) 2: 306 Cayo, A. Hughes 148 (BM-000557731! 22 ); Stann Creek, W.A. Schipp 211 (BM 2 , BRH 125 ); Toledo, G. Davidse & A.E. Brant 32131 (MO 115 ). Asplenium juglandifolium Lam., Encycl. 2(1): 307 (1786) 2 3( Toledo, B.K. Hoist 4027 (MO 91 ' 115 ). Asplenium laetum Sw., Syn.fil: 79, 271 (1806) 2:308 Stann Creek, WA. Schipp S-277 (B 115 ); Toledo, P.H. Gentle 7378 (BM 2 ' 115 , F 115 , G 115 , MICH 115 , NY 115 , S" 5 , UC 115 ). Asplenium macilentum Kunze ex Klotzsch in Linnaea 20: 351 (1 8 47)2: 309 Toledo, W.A. Schipp 1066 (BRH 125 , K 2 , NY). Adams (1995: 309-310) considered that material cited as A. monodon Liebm. was more closely related to A. macilentum than A. auritum, although it was included as a synonym in the latter taxon by Stolze ( 198 1 : 60). It is not clear if A. monodon occurs in Central America and the name may have been misapplied. Asplenium palmeri Maxon in Contr. U.S. Natl. Herb. 13: 39 (1909) 2 313 First recorded for Belize in extralimital distribution for the Pteridophyte flora of Oaxaca, Mexico (Mickel & Beitel, 1988: 65) but without explicit reference to specimens. Vargas & Shawe (1997: 22) list a specimen collected in 1929 from the Stann Creek District ( W.A. Schipp 211, BRH) under this name, but a duplicate at BM has been identified as A. heterochroum by Adams (19956: 306). Material needs to be located to support the record for Belize. Asplenium pteropus Kaulf., Enum. filic. : 1 70 ( 1 824) 2: 3I4 Stann Creek, WA. Schipp 365 (MO 2 , NY); Toledo, B.K. Hoist et al. 5 195 (MO 115 ). Asplenium pumilum Sw., Prodr.: 129 (1788) 2;315 Cayo, A. Hughes 136 (BM!); Toledo, T. Arnason & J. Lambert 17182 (MO 2 ), G. Davidse & B.K. Hoist 36181 (MO 115 ). Asplenium riparium Liebm. in Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 5, 1: 244 (1849) Toledo, G. Davidse & D.L Holland 36837 (MO 115 ), B.K. Hoist 5332 (MO 115 ). Not previously recorded from Belize in the Central American literature, but material collected from Toledo District in 1 996 and 1997 is listed in TROPICOS and identified as this taxon by A.R. Smith in 1997. Asplenium salicifolium L., Sp. pi- 2: 1080 (1753) 2:318 Asplenium salicifolium var. salicifolium 1 3 Cayo, T.B. Croat 23773 (MO 2 ). Asplenium serra Langsd. & Fisch., PL Voy. Russes monde 1: 16, 1. 19 (1810) 2:319 84 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS Cayo, B.H. Allen 15177 (BM-000543291!, MO 2 - 3 ' 115 ). Asplenium serratum L., Sp. pi. 2: 1079 (1753) 2:319 Cayo, C.L. Lundell 6235 (MICH 28 , NY); Stann Creek, R. Rivera et al. 2602 (BRH 125 ); Toledo, C. Whitefoord 1625 (BM- 000543328 ! 2 ). Asplenium uniseriale Raddi in Opusc. Sci. 3: 291 (1819) Toledo, B.K. Hoist 5321 (MO 115 , UC" 5 ). Not listed for Belize in the published version of Flora Mesoamericana (Adams, 19956: 323), but included in the Internet version based on a specimen collected in 1996 from Toledo District. BLECHNACEAE HI cell n 11 in x ant Ulan u in Proctor in Brit. Fern Gaz. 9: 214 (1965) Cayo, G.R. Proctor 29904 (BRH 125 ). Interpreted in Flora Mesoamericana (Moran, I995ae: 329) as a hybrid between B. meridense and B. glandulosum and expected from Mesoamerica. Vargas & Shawe (1997: 31) listed an unpub- lished herbarium name with the epithet 'antillense' attributed to G.R. Proctor and based on a specimen (cited above, collected in 1968 from Rio Frio Caves in Cayo District) which apparently equates to Proctor's Blechnum x antillanum. However, of the putative parent species, only B. meridense has been recorded from Belize and the status of this specimen is uncertain. Blechnum x caudatum Cav., Descr. pi: 262 (1802) 67:326 Cayo, A. Hughes 34 (BM-000531849!). Blechnum ensiforme (Liebm.) C. Chr., Index filic.: 153 (1905) Toledo, G. Davidse & D.L. Holland 36781 (MO 115 , UC" 5 ). Not recorded from Belize in the published version of Flora Mesoamericana (Moran, \995ae: 327), but included in the Internet version based on a specimen (G. Davidse & D.L. Hol- land 36781, UC) collected from Toledo District in 1997 and identified by A.R. Smith in the same year. Blechnum fragile (Liebm.) C.V. Morton & Lellinger in Amer. Fern J. 57:68(1967) 67:328 Cayo, B.H. Allen 15204 (MO 3 - 67 - " 5 ); Toledo, B.K. Hoist et al. 5303 (MO 115 ). Blechnum gracile Kaulf., Enum. filic.: 158 (1824) 67:328 Toledo, G. Davidse & A.E. Brant 31915 (MO 67 115 ). Blechnum meridense Klotzsch in Linnaea 20: 349 (1847) Toledo, G. Davidse 36979 (MO 115 , UC), D.L. Holland65 (MO 115 ), D.L. Holland & B. Kid 100 (MO 115 ). Not recorded from Belize in the published version of Flora Mesoamericana (Moran, \995ae: 329), but included in the Internet version based on a specimen (G. Davidse 36979, UC) collected from Toledo District in 1997 and identified by A.R. Smith in the same year. Blechnum occidentale L., Sp. pi. 2: 1077 (1753) 67:329 Cayo, A. Hughes93 (BM-000531830!, BM-000531837!); Stann Creek, W.A. Schipp 162 (BRH 125 , NY); Toledo, G. Davidse & A.E. Brant 32304 (MO 67 - 115 ). Blechnum polypodioides Raddi in Opusc. Sci. 3: 294 (1819) 67:329 Cayo, H.H. Bartlett 11643 (US). Moran (\995af: 329) did not base his record for this taxon on a specimen, but on extralimital distribution in Flora of Chiapas (Smith, 198 la: 60). The Bartlett specimen cited above is at US, and a duplicate is to be expected at MICH. Blechnum serrulatum Rich, in Actes Soc. Hist. Nat. Paris 1: 114 (1 792)67: 330 Belize, G. Davidse & A.E. Brant 32999 (MO 67 - " 5 ); Orange Walk, G.R. Proctor 35794 (BRH 125 ); Stann Creek, P.H. Gentle 8227 (NY); Toledo, G. Davidse & A.E. Brant 32496 (MO 115 ). Salpichlaena volubilis (Kaulf.) J. Sm. in Hook. & Bauer, Gen.fil.: t. 93(1841) 68:332 Toledo, B.K. Hoist 4287 (MO 6 "-"- 115 ). CYATHEACEAE Alsophila firma (Baker) D.S. Conant in J. Arnold Arbor. 64: 372 (1983) IO.:89 Cayo, B.H. Allen 15221 (MO 3 - l01 - 115 ); Stann Creek, A.M. Gentry 7958 (MO 115 ); Toledo, B.K. Hoist et al. 5301 (MO 115 ). Alsophila salvinii Hook, in Hook. & Baker, Syn.fil.: 36 (1866) Toledo, B.K. Hoist 5854 (MO 115 ). This Central American endemic was not recorded for Belize in Flora Mesoamericana (Riba, 1995c: 90), though it has been recorded from all surrounding countries and is included here based on a specimen collected in 1997 from Toledo District. Further material is to be expected from the wetter parts of the Maya Mountain divide and the Liquidambar forests. Cyathea costaricensis (Mett. ex Kuhn) Domin in Acta Bot. Bohem. 9: 107(1930) 40:96 Stann Creek, J.D. Dwyer et al. 583 (MO 40 - 115 ). Cyathea divergens Kunze in Linnaea 9: 100 (1834) 40197 Cyathea divergens var. tuerckheimii (Maxon) R.M. Tryon in Contr. Gray Herb. 206: 56 (1976) 40:97 Cayo, B.H. Allen 15224 (MO 3 - 40 - 115 ); Toledo, B.K. Hoist et al. 5245 (MO 115 ). Cyathea microdonta (Desv.) Domin in Pteridophyta: 263 (1929) 40:98 Stann Creek, J.D. Dwyer et al. 549 (MO 40 - " 5 ); Toledo, P.H. Gentle 1933 (NY). Cyathea multiflora Sm. in Mem. Acad. Roy. Sci. (Turin) 5(1790- 1791): 416 (1793) 40:99 Cayo, J. Meave 1037 (MO 40 - 115 ); Toledo, M.J. Balick et al. 2543 (BRH 125 , MO 115 , NY). Cyathea myosuroides (Liebm.) Domin in Pteridophyta: 263 (1929) 40199 Cayo,A.//wg/u>sl32a(BM-000531752!,BM-000531908!,BM- 000531909!); Stann Creek, W.A. Schipp 191 (BRH 125 , NY); Toledo, B.K. Hoist 4285 (MO 91 - 115 ). Cyathea schiedeana (C. Presl) Domin in Pteridophyta: 263 (1929) 401 101 Cayo, J.D. Dwyer 11431 (MO 40 - 115 ); Stann Creek, P.H. Gentle 8257 (NY); Toledo, B.H. Allen 15430 (MO 3 - 115 ). Cyathea ursina (Maxon) Lellinger in Amer. FernJ. 77: 101 (1987) 40: 102 Stann Creek, P.H. Gentle 3 197 (LL 40 , MICH 23 , MO-photograph 40 - " 5 , US-holotype 23 - 30 - 120 - 124 ). Reported only from the type collected from Antelope Ridge in the Stann Creek Valley of Belize, this Central American endemic occurs also in Guatemala and from Nicaragua to Panama. Sphaeropteris horrida (Liebm.) R.M. Tryon in Contr. Gray Herb. 200:20(1970) 89:104 Toledo, B.K. Hoist 4213 (MO 89 - 91 - 115 ). DAVALLIACEAE Nephrolepis biserrata (Sw.) Schott, Gen.fil.: t. 3 (1834) 80:286 Cayo, A. Hughes 130 (BM-000531755! 22 , BM-000531910!); Stann Creek, WA. Schipp 394 (BRH 125 , NY 80 ); Toledo, G. Davidse & A.E. Brant 32193 (MO 115 ). Nephrolepis cordifolia (L.) C. Presl, Tent, pterid.: 79 (1836) 80:287 Cayo, G.R. Praetor 29881 (BRH 125 ); Toledo, D.E. Breedlove & D.C. McClintock 23670 (DS 80 ). Nephrolepis multiflora (Roxb.) P.M. Jarrett ex C.V. Morton in Contr. U.S. Natl. Herb. 38: 309 (1974) 80:287 Cayo, A. Hughes 67 (BM-000531799!, BM-000557718! 22 ); Stann RECENT RECORDS OF PTERIDOPHYTES Creek, R. Rivera et al. 2536 (BRH 125 ); Toledo, B.K. Hoist 4507 (MO 80 ' 91 -" 5 ). Nephrolepis pendula (Raddi) J. Sm. in J. Bot. (Hooker) 4: 197 (1841) 80:288 Cayo, C.L. Lundell 6305 (MICH 28 , NY 80 ); Toledo, M.E. Peck 538 (NY). Nephrolepis rivularis (Vahl) Mett. ex Krug in Bot. Jahrb. Syst. 24: 122(1897) Toledo, G. Davidse 36876 (MO 115 , UC), T.E. Hawkins 1425 (MO 115 ). Not recorded for Belize in the published version of Flora Mesoamericana (Nauman, 1995: 288), although included in the Internet version on the basis of material collected in 1997 from Toledo District and identified by A.R Smith in 1997 and 1998. Nephrolepis undulata (Afzel. ex Sw.) J. Sm. in Bot. Mag. 72(Com- panion): 35 bis (1846) Cayo, A. Hughes 104 (BM-000532008!). Not recorded for Belize in Flora Mesoamericana (Nauman, 1995: 288), although recorded from all surrounding countries and to be expected in Belize. The material listed here from Cayo District is not fully fertile and confirmation of the identification is required from further collections. Oleandra articulata (Sw.) C. Presl, Tent, pterid.: 78, t. 2, f. 12 (1836) 90:289 Toledo, T.E. Hawkins 1542 (MO 115 ). DENNSTAEDTIACEAE Dennstaedtia bipinnata (Cav.) Maxon in Proc. Biol. Soc. Wash. 51: 39(1938) 47:152 Toledo, B.K. Hoist 4474 (MO 47 ' 91 ' 115 ). Dennstaedtia cicutaria (Sw.) T. Moore, Index fil: xcvii (1857) 47: 152 Toledo, W.A. Schipp S-802 (GH 47 - 115 ). Dennstaedtia dissecta (Sw.) T. Moore, Index fil.: 305 (1861) 47: 152 Stann Creek, R. Rivero et al. 2587A (BRH 125 ); Toledo, W.A. Schipp S-921 (GH 47 ' 115 ). Hypolepis repens (L.) C. Presl, Tent, pterid.: 162 (1836) 48: 156 Stann Creek, R. Rivero et al. 2532 (BRH 125 ); Toledo, W.A. Schipp 258 (GH 48 ' 115 , NY). Lindsaea klotzschiana Moritz in Ettingsh., Farnkr. Jetztw.: 212, t. 145, f. 1-2(1865) 49:158 Toledo, B.K. Hoist 4265 (MO 49 ' 91 ' 115 ). Lindsaea lancea (L.) Bedd., Suppl. ferns S. Ind.: 6 (1876) 49: 158 Lindsaea lancea var. lancea 49 159 Stann Creek, W.A. Schipp 100 (BRH 125 , NY); Toledo, C. Whitefoord 1310 (BM, MO 49 - 115 ). Lindsaea quadrangularis Raddi in Opusc. Sci. 3: 294 (1819) 49: 159 Lindsaea quadrangularis subsp. subalata K.U. Kramer in Acta Bot. Neerl.6: 190 (1957) 49: 159 Stann Creek, W.A. Schipp 200 (BRH 125 , F 49 - " 5 , US); Toledo, G. Davidse 36247 (MO 115 ). Lindsaea stricta (Sw.) Dryand. in Trans. Linn. Soc. London 3: 42 (1 797)49: 159 Lindsaea stricta var. stricta 49 159 Belize, P.H. Gentle 1536 (K, NY); Cayo, G. Davidse & A.E. Brant 33024 (MO 49 ' 115 ); Toledo, G. Davidse & A.E. Brant 31958 (MO 115 ). Lonchitis hirsuta L., Sp. pi. 2: 1078 (1753) Toledo, G. Davidse & D.L. Holland 36478 (MO 115 ), G. Davidse 36888 (MO 115 ), G. Davidse 36983 (MO 115 , UC), B.K. Hoist et al. 55 16 (MO 115 ). Not listed for Belize in the published version of Flora Mesoamericana Moran ( 1 995n: 1 60), but included in the Internet 85 version on the basis of a collection from Toledo District (G. Davidse 36983, UC) in 1997 and identified by A.R. Smith in December 1997. This is the first record of the genus for Belize. Odontosoria schlechtendalii (C. Presl) C. Chr., Index filic. : 209 (1905) 51:161 Toledo, G. Davidse & A.E. Brant 31986 (MO 51 - " 5 , US). Pteridium caudatum (L.) Maxon, in Proc. U.S. Natl. Mus. 23( 1226): 631 (1901) 52:162 Belize, C. Whitefoord 2452 (BM); Cayo, A. Hughes 65 (BM- 000531794!, BM-000557716!); Stann Creek, W.A. Schipp 190 (BRH 125 , NY); Toledo, C. Whitefoord 2204 (MO 52 115 ). Saccoloma elegans Kaulf. in Berlin. Jahrb. Pharm. Verbundenen Wiss. 21:51 (1827) 53:163 Saccoloma elegans subsp. chartaceum G.B. Nair ex Cremers & K.U. Kramer, Bot. Helvet. 99: 47 (1989) 53: 163 Cayo, T.B. Croat 24577 (MO 53 - 115 ); Stann Creek, W.A. Schipp 90 (BRH 125 , NY); Toledo, P.H. Gentle 9315 (NY). Saccoloma inaequale (Kunze) Mett. in Ann. Sci. Nat., Bot. ser. 4, 15: 80(1861) 53:163 Toledo, W.A. Schipp 315 (MO 53 " 5 , US). DICKSONIACEAE Cibotium regale Verschaff. & Lem. in ///. Hort. 15: t. 548 (1868) Cayo, A.K. Monro & S. Cafferty 2639 (BM-000543340!, BM- 000543341!, BM-000543342!, BM-000543343!, BRH, LAGU, MEXU, MO). Known formerly from Chiapas (including the type), Guatemala and Honduras to El Salvador (Perez-Garcia, I995b: 86-87), this Central American endemic is restricted to mixed woodland of Pinus, Quercus and Liquidambar. The vegetation class occurs in Belize in the southern part of the Chiquibul Forest and probably will be found in other little-explored parts of the Maya Mountain Divide. Though the presence of Liquidambar styraciflua L. in Belize has been reported since the first half of the twentieth century (Stevenson, 1928) and Standley & Record (1936: 147) considered that it occurred 'in some abundance in the higher parts of the Cockscomb Mountains', material was not collected until very recently. A specimen can be found in the Belize National Herbarium (Belize: Cayo; J.C. Meerman s.n. (BRH), cf. Vargas & Shawe, 1997: 67). General collections from this class of vegetation were not made until 1998 by Monro and Cafferty, who provide the first record of Cibotium regale. DRYOPTERIDACEAE Arachniodes denticulata (Sw.) Ching in Acta Bot. Sin. 10: 260 (1962) 59:2M Toledo, B.H. Allen 15391 (AAU 59 - 115 , MO 3 ' 115 ). Didymochlaena truncatula (Sw.) J. Sm. in J. Bot. (Hooker) 4: 196 (1841) 60:212 Stann Creek, W.A. Schipp 406 (BRH 125 , MO 60 115 ); Toledo, B.K. Hoist et al. 5160 (BRH 115 , MO 115 , SEL). Olfersia cervina (L.) Kunze in Flora 7: 312 (1824) 61:214 Stann Creek, W.A. Schipp 402 (BRH 125 , F 36 - 61 - 115 , GH 36 , MICH 36 , NY 36 , UC 36 , US 36 , Z 36 ); Toledo, G. Davidse & D.L. Holland 36737 (MO 115 ). Polybotrya caudata Kunze in Linnaea 9: 23 (1834) 62 217 Cayo, C.L. Lundell 6416 (MICH 28 ); Stann Creek, W.A. Schipp 273 (BRH 125 , F 62 - 115 ). Polybotrya osmundacea Humb. & Bonpl. ex Willd., Sp. pi 5: 99 (1810) Toledo, B.K. Hoist 4048 (MO 91 - 115 ), B.K. Hoist 4049 (MO 91 - " 5 ). Not recorded for Belize in Flora Mesoamericana (Moran, 1 995z: 86 218), although the species was recorded for Toledo District by Parker et al. (1993) and material bearing this identification is at MO and listed in TROPICOS. Polybotrya polybotryoides (Baker) H. Christ in Bull. Herb. Boiss. ser. 2, 1:70(1901) 62:218 Cayo, T.B. Croat 24566 (MO 62 ' ll5 ); Toledo, B.K. Hoist 4047 (MO 91 '" 5 ). Stigmatopteris sordida (Maxon) C. Chr., Index filic., Suppl. 3: 175 (1934) Toledo, B.K. Hoist 5897 (MO 115 , UC). Not recorded for Belize in the published version of Flora Meso- americana (Moran, \995aa: 226), but included in the Internet version based on a specimen from Toldeo District (B.K. Hoist 5897, UC) collected in 1997 and identified by A.R. Smith in December of the same year. This is also the first record of the genus from Belize. GLEICHENIACEAE Dicranopteris flexuosa (Schrad.) Underw. in Bull. Torrey Bot. Club 34:254(1907) 38:58 Belize, P.H. Gentle 9558 (MO 38 115 , US). Dicranopteris pectinata (Willd.) Underw. in Bull. Torrey Bot. Club 34:260(1907) 38:58 Cayo, C.L. Lundell 6603 (MICH 28 , NY, US); Stann Creek, W.A. Schipp 350 (BRH 125 , K!, NY, US); Toledo, D.L. Spellman &W.W. Newey 1648 (MO 38 ' 115 ). Sticherus palmatus (J.H. Schaffn. ex Underw.) Copel., Gen. fil. (Ann. Cryptog. Phytopathol. 5): 28 (1947) 39:61 Toledo, B.K. Hoist 4270 (MO 39 ' 91 - 115 ). GRAMMITIDACEAE Cochlidium linearifolium (Desv.) Maxon ex C. Chr. in Dansk Botanisk Arkiv 6(3): 23 (1929) 6:371 Cayo, G. Davidse & A.E. Brant 33087 (MO 115 ); Toledo, B.H. Allen 1 5304 (MO 3 ' 6 - 115 ). Cochlidium serrulatum (Sw.) L.E. Bishop in Amer. Fern J. 68: 80 Cayo, G. Davidse & A.E. Brant 33088 (MO 115 ); Toledo, W.A. Schipp 213 (N\, UC 6 ). Enterosora ecostata (Sodiro) L.E. Bishop in Syst. Bot. 17(3): 348, f. 1A-C(1992) 1I2:373 Toledo, W.A. Schipp S-801[a] (F 117 , GH 129 ). Lellingeria mitchellae (Baker ex Hemsl.) A.R. Sm. & R.C. Moran in Amer. Fern J. 81: 85 (1991) 77:378 Cayo, A. Hughes 48 (BM-000531771 !); Orange Walk, Mitchell s.n. (K-holotype 10 - 19 ' 77 ' 106 ); Toledo, G. Davidse 36126 (MO 115 ). Based on a Belizean type, this Central American endemic occurs from Chiapas to Panama and is unique in having setulose sporangia capsules. Micropolypodium taenifolium (Jenman) A.R. Sm. in Novon 2: 423 (1992) Toledo, G. Davidse & D.L. Holland 36701 (MO 115 ), 36782 (MO 115 ), T.E. Hawkins 1526 (MO 115 ), B.K. Hoist et al. 5248 (MO 115 ). Not included in Flora Mesoamericana (Smith, 1995ft: 384) but included here on the basis of material collected from Toledo District in 1996-1997 and identified by A.R. Smith in 1997- 1998. This is the first record of the genus from Belize. Terpsichore asplenifolia (L.) A.R. Sm. in Novon 3: 485 (1993)" 4: 387 Cayo, B.H. Allen 15174 (MO 3 ' " 4 - l15 ); Toledo, B.K. Hoist et al. 5 189 (MO 115 ). D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS Terpsichore lehmanniana (Hieron.) A.R. Sm. in Novon 3: 487 (1 993).. 4: 389 Cayo, B.H. Allen 15215 (MO 3 ' 114 ' 115 ); Toledo, G. Davidse & D.L. Holland 36738 (MO 115 ). Terpsichore mollissima (Fee) A.R. Sm. in Novon 3: 487 (1993) 114:39 Toledo, B.K. Hoist 4040 (MO 91 ' 115 ). HYMENOPHYLLACEAE Hymenophyllum abruptum Hook., Sp.fil. 1: 88, t. 3 IB (1844) 84:64 Cayo, H.H. Bartlett 11751 (F 84 ' 115 ). Hymenophyllum fucoides (Sw.) Sw. in J. Bot. (Schroder) 1800(2): 99(1801) 84:66 Toledo, B.H. Allen 15305 (MO 3 ' 84 - 115 ). Hymenophyllum hirsutum (L.) Sw. in J. Bot. (Schrader) 1800(2): 99 (1801 )84:66 Cayo, B.H. Allen 15203A (MO 3 ' 84 - 115 ); Toledo, B.K. Hoist et al. 5304 (MO 115 ). Hymenophyllum poly anihos (Sw.) Sw. in/. Bot. (Schrader) 1800(2): 102(1801) 84:68 Cayo, C. Whitefoord 9\4l (BM); Stann Creek, W.A. Schipp 114 (F 84 - 115 ); Toledo, G. Davidse 36124 (MO 115 ). Hymenophyllum pulchellum Schltdl. & Cham, in Linnaea 5: 618 (1830) 84:68 Toledo, B.K. Hoist 4041 (MO 84 ' 91 ' 115 ). Hymenophyllum sieberi (C. Presl) Boschin in Ned. Kruidk. Arch. 4: 414(1858). Toledo, B.K. Hoist el al. 5296 (BRH 115 , MO 115 , UC 115 ), B.K. Hoist 5974 (MO 115 ). Not listed for Belize in the published version of Flora Mesoamericana (Pacheco, 1995a: 69), H. sieberi is included in the Internet version on the basis of a collection from Toledo District (B.K. Hoist et al. 5296, UC) made in 1996 and identified by A.R. Smith in May 1997. Trichomanes ankersii C. Parker ex Hook. & Grev., Icon, filic. 2(11): t. 201(1831) No material has been collected from this far north in Central America according to Flora Mesoamericana (Pacheco, 1995ft: 73), though it is recorded from Nicaragua to Panama. Standley & Record (1936: 60) listed the taxon for Belize but without explicit reference to a specimen. There may be material at F, though this needs further investigation to verify the identification. Trichomanes capillaceum L., Sp. pi. 2: 1099 (1753) 85:73 Trichomanes capillaceum var. capillaceum* 5 74 Toledo, B.H. Allen 15411 (MO 3 - 85 - 115 ). Trichomanes collariatum Bosch in Ned. Kruidk. Arch. 4: 368 (1858) 85:74 Stann Creek, W.A. Schipp 364 (BM-000543308!, K!, NY); Toledo, M.E. Peck 601 (F 85 - " 5 , K!). Trichomanes crispum L., Sp. pi. 2: 1097 (1753) 85:74 Cayo, B.H. Allen 15172 (MO 3 ' l15 ); Toledo, B.K. Hoist 4023 (MO 85 - 91 ' 115 ). Trichomanes curtii Rosenst. in Repert. Spec. Nov. Regni Veg. 22(606-608): 5 (1925) 85:75 No specimen was listed for Belize in Flora Mesoamericana (Pacheco, 1995ft: 75), but reference is made to Flora of Guate- mala (Stolze, 1976: 78). The latter author also did not cite material, but made explicit reference to 'British Honduras'. There may be supporting material at F. Trichomanes curtii is also recorded from Chiapas and Guatemala, extending southeast from Nicaragua to Colombia. Trichomanes diaphanum Kunth in Humb., Bonpl. & Kunth, Nov. Gen.Sp. 1:25(1816) 85;75 Toledo, B.K. Hoist et al. 5214 (MO 115 ). RECENT RECORDS OF PTERIDOPHYTES 87 Trichomanes diversifrons (Bory) Mett. ex Sadeb. in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 108 (1899) 85:75 Belize, C. Whitefoord 1210 (BM-000543306!); Stann Creek, W.A. Schipp 237 (K!, NY); Toledo, T.B. Croat 24514 (P 5 - " 5 ). Trichomanes ekmanii Wess. Boer in Acta Bot. Neerl. 11: 319, f. 33 (1 962 )85: 75 Stann Creek, W.A. Schipp 324 (BM-000543311!, BRH 125 , K, MO 85 - 115 , NY). Trichomanes elegans Rich. in Actes Soc. Hist. Nat. Paris 1: 114 (1792) Toledo, E.J.F. Campbell s.n. (K!), Mitchell s.n. (K!). First collected from Belize in 1875 (Mitchell s.n., K!) but not recorded in any subsequent publication for the country either as an accepted name, a synonym or as a misapplied name. Trichomanes elegans has previously been recorded in Central America from Honduras to Panama (Pacheco, 1995ft: 75), from South America and the Caribbean region, and is to be expected from Belize. Trichomanes galeottii E. Fourn. in Bull. Soc. Bot. France 15: 147- 148(1868) 85:76 Cayo, H.H. Bartlett 11640 (MO 85 ' " 5 ); Stann Creek, W.A. Schipp 482 (BM-000543305!, BRH 125 , NY); Toledo, G. Davidse&A.E. Brant 32238 (MO 115 ). Trichomanes godmanii Hook, in Baker in J. Linn. Soc., Bot. 9: 337, t. 8A(1866) 85:76 Cayo, C.L. Lundell 6187 (MICH 28 , NY); Toledo, F. Boutin & Sc/z/oer5023(MO 85 - 115 ). Trichomanes hymenoides Hedw., Fil. gen. sp.: t. 3, f. 3 (1799) 85:77 No specimen was listed for Belize in Flora Mesoamericana (Pacheco, 1995ft: 77), but reference is made to Stolze (1976: 81- 82) who explicitly cited 'British Honduras' in the distribution for Flora of Guatemala, but did not mention any material. There may be a specimen at F to verify the occurrence in Belize. Trichomanes krausii Hook. & Grev., Icon.filic. 2: t. 149 (1830) Cayo, T.E. Hawkins 1179 (MO 115 ); Orange Walk, C. Whitefoord 8095 (BM-000543310!); Toledo, C. Whitefoord 1984 (BM- 000543309!). Recorded for Belize since Forests and flora of British Honduras (Standley & Record, 1936: 61). Stolze (1976: 83) gave the distribution as 'Mexico to Panama', which may be taken as an implicit reference to Belize though no material was listed for 'British Honduras'. Pacheco (1995ft: 78) did not list the species for Belize in the published version of Flora Mesoamericana, though several specimens have been collected recently and appear on TROPICOS. Trichomanes membranaceum L., Sp. pi. 2: 1097 (1753) 85: Stann Creek, WA. Schipp S-108 (F 85 - 115 ). Trichomanes ovale (E. Fourn.) Wess. Boer in Acta Bot. Neerl. 11: 296(1962) Toledo, C. Whitefoord 1244 (BM, MO 115 ). Not recorded for Belize in the published version of Flora Mesoamericana (Pacheco, 1995ft: 79) but material collected in 1976 (C. Whitefoord 1244, MO) is added to the Internet version as an extended range record. Trichomanes pinnatum Hedw., Fil. gen. sp.: t. 4, f. 1 (1799) 85:8 Cayo, H.H. Bartlett 11726 (NY); Stann Creek, W.A. Schipp 359 (K!, MO 85 ' " 5 , NY); Toledo, E.J.F. Campbell 86 (K!). Trichomanes polypodioides L., Sp. pi 2: 1098 (1753) 85:8 Stann Creek, WA. Schipp S-83 (F 5 - 115 , K!); Toledo, B.K. Hoist et al. 5 193 (MO 115 ). Trichomanes punctatum Poir. in Lam., Encycl. 8: 64 (1808) 85:8 Trichomanes punctatum subsp. sphenoides (Kunze) Wess. Boer in Acta Bot. Neerl. 11: 301 (1962) 85:81 Cayo, A. Hughes 129 (BM-000557729!); Toledo, B.H. Allen 15442 (MO 3 - 85 -" 5 ). Trichomanes pyxidiferum L., Sp. pi. 2: 1098 (1753) 85:81 Cayo, A. Hughes 126 (BM-000557730!); Toledo, B.K. Hoist 43 10 (MO 85 - 91 -" 5 ). Trichomanes radicans Sw. in J. Bot. (Schroder) 1800(2): 97 (1801) Toledo, B.K. Hoist 5815 (MO 115 , UC), B.K. Hoist 5816 (MO 115 ). Not recorded for Belize in the published version of Flora Mesoamericana (Pacheco, 1 995ft: 8 1 ) but included in the Internet version. The first record for the country is material collected from Toledo District on vertical rocks in waterfall spray (B.K. Hoist 5815, UC) in 1997, and identified by A.R. Smith in December of the same year. Trichomanes rigidum Sw., Prodr.: 137 (1788) 85:82 Cayo, B.H. Allen 15186 (MO 3 - 115 ); Toledo, B.K. Hoist ela\. 5197 (MO 115 ). Trichomanes tuerckheimii H. Christ in Hedwigia 44: 361 (1905) 85:! Belize, C. Whitefoord 1298 (BM-000543313!, MO 115 ); Stann Creek, W.A. Schipp S-21 (F 5 - " 5 , US); Toledo, G. Davidse & M. Meadows 35841 (MO 115 ). ISOETACEAE Isoetes cubana Engelm. ex Baker in J. Bot. 18: 110 (1880) 21:42 Toledo, M.E. Peck 420 (GH 21 , NY). LOMARIOPSIDACEAE Bolbitis bernoullii (Kuhn ex H. Christ) Ching in C. Chr., Index filic., Suppl. 3:47(1934) 20:248 Toledo, WA. Schipp S-776 (GH 20 - 115 ). Bolbitis hastata (E. Fourn.) Hennipman in Amer. Fern J. 65: 1975 (1975) Toledo, B.K. Hoist 5757 (MO 115 , UC). Not recorded for Belize in the published version of Flora Mesoamericana (Hennipman & Moran, 1995: 248), but included in the Internet version based on a specimen (B.K. Hoist 5757, UC) collected from Toledo District in 1997 and identified by A.R. Smith later that year. Bolbitis hemiotis (Maxon) Ching in C. Chr., Index filic., Suppl. 3: 48 (1 934)20: 248 Toledo, B.H. Allen 15441 (MO 3 - 20 - 115 ). Bolbitis pergamentacea (Maxon) Ching in C. Chr., Index filic., Suppl. 3:49(1934) 20:249 Toledo, WA. Schipp S-764 (GH 20 - 115 ). Bolbitis portoricensis (Spreng.) Hennipman in Amer. Fern J. 65: 30 (1 975 )20: 249 Stann Creek, W.A. Schipp 526 (BRH 125 , NY); Toledo, G. Davidse & A.E. Brant 32381 (MO 20 - 115 ). Elaphoglossum christianeae Mickel in Novon 2: 371 (1992) Toledo, G. Davidse & H.B. Buchanan 36941 (MO 115 ). Not recorded for Belize in Flora Mesoamericana (Mickel, 1995a: 263), this Central American endemic has been recorded formerly from Costa Rica and Panama but to that distribution can be added a specimen collected in Toledo District of Belize in 1997 and identified by A.F. Rojas-Alvarado in 1998. Elaphoglossum decursivum Mickel in Brittonia 32: 334 (1980) Toledo, G. Davidse & D.L. Holland 36729 (MO 115 ), D.L Hol- land & B. Kid 90 (MO 115 ), B.K. Hoist 5924 (MO 115 ). This taxon is not recorded for Central America in the published version of Flora Mesoamericana (Mickel, 1995a), but appended to the Internet version on the basis of recent identifications by A.R. Smith of material from Belize, Honduras and Costa Rica. 88 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS All of the Belize material listed above was collected in 1997 from Toledo District. Elaphoglossum erinaceum (Fee) T. Moore, Index fil: 9 (1857) Toledo, G. Davidse & D.L. Holland 36764 (MO 115 ). Not included in the published version of Flora Mesoamericana (Mickel, 1995a: 265) but listed for Belize in the Internet version based on the collection listed above from Toledo District made in 1997 and identified by A.R. Smith in 1998. Elaphoglossum eximiiforme Mickel in Novon 2: 374 (1992) Toledo, B.K. Hoist 5672 (MO 115 ). Not recorded for Belize in Flora Mesoamericana (Mickel, 1 995a: 265), this Central American endemic was formerly known only from Costa Rica and Panama. The material included here was originally identified as E. latifolium (Sw.) Sm. by A.R. Smith but re-determined as E. eximiiforme by A.F. Rojas-Alvarado in October 1998. Elaphoglossum glaucum T. Moore, Index fii: 10 (1857) Toledo, B.K. Hoist 3869 (MO 91 115 ). Not recorded for Belize in Flora Mesoamericana (Mickel, 1 995a: 267), though recorded from the neighbouring countries of Mexico, Guatemala and Honduras. The record here is tentative as it is based on material from Toledo District identifed by R. Moran in 1992 and cited by Parker et al. (1993: 40), yet excluded from the flora. Elaphoglossum guatemalense (Klotzsch) T. Moore in Parker's Cat. (1858) 32:268 Cayo, B.H. Allen 15175 (MO 3 ' 115 ); Toledo, G. Davidse & A.E. Brant 32239 (MO 3 - 115 ). Elaphoglossum herminieri (Bory ex Fee) T. Moore, Index fil.: xvi (1 8 57)32:268 Toledo, F. Boutin & Schlosser 5088 (NY 32 ). Elaphoglossum herrerae A. Rojas in Brenesia 45-46: 13, f. 5 (1996) Toledo, B.K. Hoist 5670 (MO 115 ). A new species provisionally included in the Internet version of Flora Mesoamericana based on a specimen from Toledo District listed above and identified by A.F. Rojas-Alvarado in October 1998. This taxon was originally considered to be endemic to the Cordillera de Guanacaste in Costa Rica. Elaphoglossum latifolium (Sw.) J. Sm. in London J. Bot. 1: 197 Toledo, P.H. Gentle 3756 (NY 32 ). Elaphoglossum latum (Mickel) Atehortiia ex Mickel in Fieldiana, Bot.n.s.,27: 123 (1991) 32:271 Toledo, T.B. Croat 24313 (MO 32 - 115 ). Lomariopsis japurensis (Mart.) Sm., Hist, fil.: 140 (1875) 6 * 284 Toledo, T.B. Croat 24393 (MO 66 - 115 ). Lomariopsis recurvata Fee, Mem. Foug. 2: 68 (1845) 66:284 Cayo, T.B. Croat 23800 (MO 66 ); Stann Creek, R. Rivero et al. 2524 (BRH 125 ); Toledo, B.K. Hoist 4417 (MO 91 ' 115 ). Lomariopsis vestita E. Fourn. in Bull. Soc. Bot. France 19: 250 (1 872)66: 284 Toledo, F. Boutin & Schlosser 5093 (MO 66 - 115 ). Peltapterispeltata (Sw.)C.V. Morton in Amer. FernJ. 45: 13 (1955) Cayo, B.H. Allen 15275 (BM-000543333, MO 3 - 115 ); Toledo, C. Whitefoord 1727 (BM-000543338!). Curiously omitted for Belize in Flora Mesoamericana (Mickel, 1 995b: 285) although it was cited under the name Elaphoglossum peltatum (Sw.) Urb. by Parker et al. (1993: 40) and there are many specimens listed under the same synonym in TROPICOS. The earliest collection from Belize seen to date (C. W. Whitefoord 1727, BM) was collected in May 1979. Mickel listed four separate forms of this species, but the northern material from Mexico, Guatemala and Honduras all falls within typical forma peltata. LOPHOSORIACEAE Lophosoria quadripinnata (J.F. Gmel.) C. Chr., Nat. Hist. Juan Fernandez 2: 16(1920) Lophosoria quadripinnata var. quadripinnata Toledo, T.E. Hawkins 1540 (MO 115 , UC). Not listed for Belize in the published version of Flora Mesoamericana (Riba, 1995a: 85), but included in the Internet version on the basis of a collection from Toledo District (T.E. Hawkins 1540, UC) made in 1996 and identified by A.R. Smith in January 1998. This is the first record of both the genus and the family for Belize. LYCOPODIACEAE Huperzia dichaeoides (Maxon) Holub in Folia Geobot. Phytotax. 20:72(1985) 82:11 Toledo, B.H. Allen 15384 (BM-000543283!, MO 3 - 82 - 115 ). Huperzia dichotoma (Jacq.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 (1874) 82:11 Toledo, P.H. Gentle 3034 (GH 82 - 115 ). Huperzia linifolia (L.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 (1 874)82: 13 Huperzia linifolia var. linifolia* 2 13 Cayo, B.H. Allen 15280 (MO 3 - 115 ); Toledo, W.A. Schipp 811 (US 82 - 115 ). Huperzia pithyoides (Schltdl. & Cham.) Holub in Folia Geobot. Phytotax. 20:76(1985) Toledo, B.K. Hoist 4381 (MO 91 ). Not listed for Belize in Flora Mesoamericana (011gaard, 1995a: 15) though it was recorded from all surrounding countries and to be expected from Belize. Huperzia pithyoides was first cited for the country by Parker et al. (1993: 40) based on the specimen listed above collected from the Columbia River Forest Reserve in Toledo District. Huperzia reflexa (Lam.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 (! 874)82: 16 Huperzia reflexa var. reflexa* 2 - 16 Toledo, M.C. Carlson 2617 (F 82 - 115 ). Huperzia taxifolia (Sw.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 (1874) 82:17 Cayo, C.L. Lundell 6258 (MICH 28 , NY 82 - 115 ); Toledo, G. Davidse 36384 (MO 115 ). Lycopodiella caroliniana (L.) Pic. Serm., Webbia 23: 165 (1968) 83: 19 Lycopodiella caroliniana var. meridionalis (Underw. & F.E. Lloyd) B. 011g. & P.O. Windisch in Bradea 5: 27 (1987) 83: 19 Belize, C. Whitefoord 2405 (BM-000543290!); Cayo, G. Davidse &A.E. Brant 33079 (BM-000543288!, BM-000543289!, MO 115 ); Stann Creek, WA. Schipp51S (BM-000543286!, MO 115 , NY, S 83 - 115 ). Lycopodiella cernua (L.) Pic. Serm. in Webbia 23: 166 (1968) 83: 19 Belize, C. Mute/o0frf2570(BM-000543294!); Cayo, J.N. Hedger 127 (BM-000543297!); Stann Creek, W.A. Schipp 234 (BM- 000543243!, NY); Toledo, P.H. Gentle 6781 (BM-000543244!, NY). MARATTIACEAE Danaea elliptica Sm. in Rees, Cycl. 11: Danaea no. 2 (1808) 9:49 Cayo, C. Whitefoord 1215 (BM-000543246!); Stann Creek, T.B. Croat 24525 (MO 9 - 115 ); Toledo, B.K. Hoist 4305 (MO 91 - " 5 ). RECENT RECORDS OF PTERIDOPHYTES 89 Danaea nodosa (L.) Sm. in Mem. Acad. Roy. Sci. (Turin) 5(1790- 1791): 420, t. 9, f. 11 (1793) 9:49 Cayo, B.H. Allen 15 185 (MO 3 ' " 5 ); Stann Creek, W.A. Schipp 422 (F- " 5 , US); Toledo, B.K. Hoist et al. 5503 (MO 115 ). Marattia excavata Underw. in Britton, N. Amer.fl. 16(1): 22 (1909) Toledo, B.K. Hoist et al. 5300 (BRH 115 , MO 115 , UC" 5 ). Not listed for Belize in the published version of Flora Mesoamericana, (Perez-Garcia, 1995a: 50) but included in the Internet version on the basis of a collection from Toledo District (B.K. Hoist et al. 5300, UC) made in 1996 and identified by A.R. Smith in April 1997. This is the first record of the genus in Belize. The specimen is cited as 'B (Hoist et al. 5300, US)' in the Internet description but it is not clear from the TROPICOS account if there is a duplicate at US or if this is an error for UC. METAXYACEAE Metaxya rostrata (Kunth) C. Presl, Tent, pterid.: 60, t. 1, f. 5 (1836) 100:86 Cayo, D. Burch 5881 (MO 115 , NY); Stann Creek, W.A. Schipp 89 (BRH 125 , NY); Toledo, P.H. Gentle 2631 (GH 100 ). OPHIOGLOSSACEAE Cheiroglossa palmata (L.) C. Presl, Suppl. tent, pterid.: 57 (1845) 126:46 Cayo, B.H. Allen 15170 (AAU 126 , MO 3 - 115 ); Toledo, B.K. Hoist 5935 (MO 115 ). Ophioglossum nudicaule L.f., Suppl. pi: 433 (1782) 126:47 Belize, C. Whitefoord 2605 (BM-000543245!); Stann Creek, P.H. Gentle 2997 (MO 115 - 126 ). POLYPODIACEAE Campyloneurum angustifolium (Sw.) Fee, Mem. Foug. 5: 257 (1852) 25:335 Cayo, D.R. Hunt 7054 (K!); Toledo, T.E. Hawkins 1348 (MO 115 ). Campyloneurum aphanophlebium (Kunze) T. Moore, Index fil. : 223 (1861) 25:335 Toledo, P.H. Gentle 7327 (US 25 ). Campyloneurum brevifolium (Lodd. ex Link) Link, Fil. spec.: 124 (1841) 25:335 Cayo, A. Hughes 96 (BM-000531759!, BM-000531835!); Stann Creek, W.A. Schipp 527 (BRH 125 , K); Toledo, P.H. Gentle 7835 (F 5 , US). Campyloneurum costatum (Kunze) C. Presl, Tent, pterid.: 190 Toledo, P.H. Gentle 6792 (US 25 ). Campyloneurum fasciale (Humb. & Bonpl. ex Willd.) C. Presl, Tent. pterid.: 190 (1836) 25:336 Cayo, C. Whitefoord 2044 (BM, MO 25 ); Toledo, G. Davidse 36003 (MO 115 ). Campyloneurum phyllitidis (L.) C. Presl, Tent, pterid.: 190 (1836) 25:33 ~ Belize, C.L. Lundell s.n. (K!); Cayo, A. Hughes 25b (BM- 000531767!); Toledo, P.H. Gentle 1108 (F 25 , K). Campyloneurum repens (Aubl.) C. Presl, Tent, pterid.: 190 (1836) 25:331 Cayo, C.L. Lundell 6262 (MICH 28 , US); Toledo, T.B. Croat 24462 (MO 25 ). Campyloneurum xalapense Fee, Mem. Foug. 5: 258 (1852) 25: Toledo, P.H. Gentle 6515 (F 5 , US). Microgramma lycopodioides (L.) Copel., Gen. fil. (Ann. Cryptog. Phytopathol. 5): 185 (1947) 69:339 Cayo, C.L Lundell 6287 (MICH 28 , NY); Stann Creek, R. Rivero et al. 2555 (BRH 125 ); Toledo, C. Whitefoord 1863 (BM, CR 69 ). Microgramma nitida (J. Sm.) A.R. Sm. in Proc. Calif. Acad. Sci. ser. 4, 40(8): 230 (1975) 69:339 Orange Walk, G. Davidse & A.E. Brant 32765 (MO 115 ); Toledo, D.L. Spellman & W.W. Newey 2105 (MO 69 ). Microgramma percussa (Cav.) de la Sola in Physis (A, B & C) 44(106, Secc. C): 28 (1986) 69:339 Cayo, A. Hughes 77 (BM-000531808!, BM-000557711! 22 ); To- ledo, G. Davidse & A.E. Brant 32027 (MO 69 - " 5 ). Microgramma reptans (Cav.) A.R. Sm. in Proc. Calif. Acad. Sci. ser. 4,40(8):230(1975) 69:34 Stann Creek, W.A. Schipp 210 (BRH 125 , NY); Toledo, C. Whitefoord 1590 (BM, CR 69 ). Neurodium lanceolatum (L.) Fee, Mem. Foug. 3: 28 (1852) 87:341 Cayo, P.H. Gentle 2518 (MEXU 87 " 5 , NY); Toledo, G. Davidse & A.E. Brant 32346 (MO 115 ). Niphidium crassifolium (L.) Lellinger in Amer. Fern J. 62: 106 (1 972)70: 341 Cayo,A//wg/^21(BM-000531766! 22 ,BM-000531797!);Stann Creek, W.A. Schipp 88 (BRH 125 ); Toledo, P.H. Gentle 4964 (MO 70 - 115 ). Niphidium oblanceolatum A. Rojas in Brenesia 45-46: 28, f. 1 (1996) The protologue of this recently described species includes Belize in the distribution, and the taxon is provisionally accepted for the Internet version of Flora Mesoamericana. Pecluma atra (A.M. Evans) M.G. Price in Amer. Fern J. 73: 113 (1 983)7 1:342 Cayo, C.L. Lundell 6639 (GH, MICH 28 , US-1638286 15 - 71 ); To- ledo, M.E. Peck 820 (NY). Pecluma dispersa (A.M. Evans) M.G. Price in Amer. Fern J. 73: 114 (1983) 71:343 Cayo, A. Hughes 38 (BM-000531768! 22 , BM-000531855!); Toledo, T.B. Croat 24176 (MO 71 - 115 ). Pecluma divaricata (E. Fourn.) Mickel & Beitel in Mem. New York Bot. Card. 46: 269 (1988) 71:343 Toledo, B.K. Hoist 4460 (MO 71 - 91 - 115 ). Pecluma pectinata (L.) M.G. Price in Amer. Fern J. 73: 115 (1983) Cayo, D.R. Hunt 605 (BM), G.R. Proctor 29841 (BRH 125 ). Not recorded for Belize in Flora Mesoamericana (Moran, 1 995ai: 344) though specimens have been collected under the synonym Polypodiumpectinatum L. This record requires further investigation. Pecluma plumula (Humb. & Bonpl. ex Willd.) M.G. Price in Amer. Fern J. 73: 115 (1983) 71:345 Belize, C. Whitefoord 1242 (BM); Cayo, A. Hughes 120 (BM- 000557724! 22 ); Corozal, G. Davidse & A.E. Brant 32534 (MO 71 - I15 ); Orange Walk, C.L. Lundell 537 (NY); Toledo, T.E. Hawkins 1717 (MO 115 ). Phlebodium decumanum (Willd.) J. Sm. in J. Bot. (Hooker) 4: 59 (1 84 1)72: 345 Toledo, A.H. Gentry 88 (F 2 ). Pleopeltis astrolepis (Liebm.) E. Fourn., Mexic. pi. 1: 87 (1872) 26:347 Cayo, C.L. Lundell 6450 (MICH 28 , NY 26 ); Toledo, W.A. Schipp 923 (BRH 125 , MO 115 ). Pleopeltis crassinervata (Fee) T. Moore, Index fil.: 345 (1862) Not recorded from Belize in Flora Mesoamericana (Lorea Hernan- dez, 1995: 348). This taxon was originally described from Mexico and listed by Weatherby (1922) from Veracruz, Chiapas and Guatemala. Lorea Hernandez expanded this distribution to include Honduras, Nicaragua and Costa Rica. Seymour (1975: 167) had listed Weatherby's combination for the taxon in his notes on the genus Polypodium in Nicaragua, but with the incorrect epithet 'crassinervum'. It is clear from the exact reference to Weatherby that this taxon is intended, and Seymour listed Central American material including a specimen from 'British Honduras' at GH. This material needs locating and the identification verifying. 90 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS x Pleuroderris michleriana (D.C. Eaton) Maxon in J. Wash. Acad. Sci. 24: 551, f. 1-2 (1934), pro sp. Stann Creek, R. Rivero et al. 2571 (BRH 125 ), 2608 (BRH 125 ). Hybridization between Tectaria incisa andDictyoxiphiumpanam- ense produces the very variable intrageneric hybrid x Pleuroderris michleriana (Wagner et al., 1978). It was not recorded for Belize by Moran (1995f: 201, 1995v: 207) though both parental species are present in the area. Polypodium dissimile L., Syst. nat. 10th ed., 2: 1325 (1759) 73;355 Cayo, B.H. Allen 15173 (MO 3 - 115 ); Toledo, B.K. Hoist 4034 (MO 73 ' 91 ' 115 ). Polypodium dulce Poir. in Lam., Encycl. 5: 523 (1804) 73:352 Toledo, F. Boutin & Schlosser 5022 (MO 73 ' 115 ). Polypodium fallax Schltdl. & Cham, in Linnaea 5: 609 (1830) 73:361 Cayo, C.L. Lundell 6216 (MICH 28 , NY); Toledo, C. Whitefoord 1745 (BM, NY). Polypodiumfraternum Schltdl. & Cham, in Linnaea 5: 608 (1830) 73:358 Cayo, A. Hughes 121 (BM-000532009!). Polypodium fraxinifolium Jacq., Collectanea 3: 187 (1791) Toledo, G. Davidse & D.L. Holland 36729A (MO 115 ), T.E. Hawkins 1427 (MO 115 ), 1429 (MO 115 ), B.K. Hoist et al. 5257 (MO 115 ). Not listed for Belize in the published version of Flora Mesoamericana (Moran, \995ak: 355), but included in the Internet version on the basis of a collection from Toledo District (Davidse & Holland 36729 A, MO) made in 1997 and identified by A.R. Smith in January 1998. Polypodium hispidulum Bartlett in Proc. Amer. Acad. Arts 43: 48 (1907) Cayo, T.E. Hawkins 1078 (MO 115 , UC); Toledo, T.E. Hawkins 1330 (MO 115 ). Not listed for Belize in the published version of Flora Mesoamericana (Moran, \995ak: 358), but included in the Internet version on the basis of a collection from Cayo District (T.E. Hawkins 1078, UC) made in 1996 and identified by A.R. Smith in December 1997. Polypodium lindenianum Kunze, Farrnkrduter 2: 83 (1849) 73:362 Cayo, A. Hughes 131 (BM-000557728! 22 ); Toledo, G. Davidse et al. 35704 (MO 115 ). Polypodium polypodioides (L.) Watt in Canad. Naturalist & Quart. J. Sci. ser. 2, 3: 158 (1867) 73:363 Polypodium polypodioides var. aciculare Weath. in Contr. Gray Herb. 124: 33 (1939) Not recorded for Belize in Flora Mesoamericana, though present in Chiapas, Guatemala and Honduras and to be expected from Belize. Seymour (1975: 160) cited material of P. polypodioides var. burchellii (Baker) Weath. from countries of Central America in his treatment of the genus Polypodium for Nicaragua, and included reference to 'British Honduras' based on material at GH. Moran (\995ak: 364) considered that var. burchellii did not occur in Central America, but there is a note describing the confusion between this South American variety and var. aciculare. It is evident that Seymour's concept of var. burchellii falls under var. aciculare in Flora Mesoamericana, though the Belizean material at GH should be located to verify the identity. Polypodium polypodioides var. polypodioides 1 ^ 364 Belize, C.L. Lundell 3884 (K!, NY); Cayo, J.D. Dwyer & R.L. Liesner 12092 (MO 73 ); Stann Creek, W.A. Schipp 349 (BRH 125 , K!, NY); Toledo, P.H. Gentle 906 (K!, NY). Polypodium triseriale Sw. in/ Bot. (Schroder) 1800(2): 26 (1801) 73:357 Cayo, A. Hughes 117 (BM-000531901!, BM-000557725! 22 ); Toledo, P.H. Gentle 5014 (MO 73 - l15 ). Pseudocolysis bradeorum (Rosenst.) L.D. Gomez in Brenesia 10- 11: 116(1977) 74:365 RecordedforBelizeinF/oraMesoam^ncana (Moran, 1995a/: 365) on the basis of the entry in Flora of Guatemala (Stolze, 1981:381) where the distribution explicitly included 'British Honduras'. Stolze did not cite any material but made reference to Evans & Mickel (1969). Material was cited at GH from 'British Honduras' by Seymour ( 1 975 : 1 67), but without detail. This or other material should be sought to verify the distributional record for Belize. PSILOTACEAE Psilotum nudum (L.) P. Beauv. , Prodr. Aetheogam. : 1 06, 1 1 2 ( 1 805) 88; 3 Belize, G. Davidse & D.L. Holland 37046 (MO 115 ); Corozal, G. Davidse & A.E. Brant 32533 (MO 115 ); Stann Creek, R. Rivero et al. 2613 (BRH 125 ); Toledo, W.A. Schipp S-261 (MO 88 - 115 ). PTERIDACEAE Acrostichum aureum L., Sp. pi: 1069 (1753) 41; 105 Belize, M.-H. Sachet & D.R. Stoddart 1631 (BRH 125 ); Stann Creek, F.R. Fosberg & M.-H. Sachet 53849 (BRH 125 , MO 41 - " 5 ); Toledo, D.L Spellman & D.R. Stoddart 2299 (BRH 125 ). Acrostichum danaeifolium Langsd. & Fisch., PI. Voy. Russes monde 1:5, t. 1 (1810) 41:105 Belize, J.D. Dwyer 11449 (MO 115 ); Toledo, D.L. Spellman & D.R. Stoddart 2477 (MO 41 - 115 ). Adiantopsis radiata (L.) Fee, Mem. Foug. 5: 145 (1852) 86: 106 Cayo, T.B. Croat 23775 (MO 86 - 115 ); Toledo, P.H. Gentle 6086 (MO 115 ). Adiantum capillus-veneris L., Sp. pi.: 1096 (1753) 78: 108 Toledo, B.K. Hoist 4014 (MO 78 91 - 115 ). Adiantum concinnum Humb. & Bonpl. ex Willd., Sp. pi. 5: 451 (1810 )78:109 Cayo, A. Hughes 146 (BM-000528104! 22 , BM-000531933!). Adiantum decoratum Maxon & Weath. in Amer. J. Bot. 19: 165 (1 932)78: 113 Toledo, T.B. Croat 24179 (MO 78 - 115 ). Adiantum fructuo sum Poepp. ex Spreng., Syst. veg. 16th ed., 4(1): 113(1827) 78;114 Toledo, T.B. Croat 24373 (BRH 125 , MO 78 - 115 ). Adiantum humile Kunze in Linnaea 9: 80 (1834) 78: " 4 Toledo, G. Davidse 36082 (MO 115 ). Adiantum latifolium Lam., Encycl. 1(1): 43 (1783) 78: 114 Cayo, C. Whitefoord 2096 (BM-000543184! 78 ' 115 ); Orange Walk, T. Arnason & J. Lambert 17176 (MO 115 ); Stann Creek, W.A. Schipp 77 (BM-000543 186!, BRH 125 , NY); Toledo, C. Whitefoord 1812 (BM-000543 188!, MEXU). Adiantum macrophyllum Sw., Prodr.: 135 (1788) 78: 112 Cayo, A. Hughes 14 (BM-000528105! 22 ); Stann Creek, W.A. Schipp 340 (BM-000543254!, BRH 125 , NY); Toledo, C. Whitefoord 1848 (BM-000543 190!, MO 78 - 115 ). Adiantum obliquum Willd., Sp. pi. 5: 429 (1810) 78: 112 Belize, C. Whitefoord 2366 (BM); Cayo, A. Hughes 70 (BM- 000528 106! 22 , BM-000531796!); Stann Creek, W.A. Schipp 275 (BM, BRH 125 , MO 115 , NY); Toledo, C. Whitefoord 1521 (BM- 000543253 ! 115 ). Adiantum petiolatum Desv. in Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 326-327 (1811) 78: 112 Belize, C. Whitefoord 2569 (BM); Cayo, A. Hughes 62 (BM- 000528 100! 22 , BM-000531788!); Stann Creek, R. Rivero et al. 2595 (BRH 125 ); Toledo, G.R. Proctor 36007 (BM-000543 194!). Adiantum princeps T. Moore in Card. Chron. n.s., 4: 197, f. 43^4 (1875) Cayo, M.J. Balick et al. 3140 (MO 115 , NY). RECENT RECORDS OF PTERIDOPHYTES 91 Not recorded for Belize by Moran in Flora Mesoamericana (Moran, Zimmer & Jermy 1995: 109), though the specimen listed here was collected in 1991 and identifed by J.T. Mickel in 1993. Adiantum pulverulentum L., Sp. pi. 2: 1096 (1753) 78: " 5 Cayo,A//Hg/2^56(BM-000528101! 22 ,BM-000531791!);Stann Creek, W.A. Schipp 241 (BM-000543 199!, BRH 125 , NY); Toledo, C. Whitefoord 1524 (NY, BM-000543196!). Adiantum tenerum Sw., Prodr.: 135 (1788) 78: " Cayo, DA. Sutton et al. 3 (BM-000543 1 80!); Orange Walk, M.J. Balick 3219 (BRH 125 , NY); Stann Creek, M.J. Balick 3091 (BRH 125 , NY); Toledo, G. Davidse&A.E. Brant 32302 (MO 78 ' 115 ). Adiantum terminatum Kunze ex Miq. in Verslagen Meded. Vier Kl. Kon. Inst. Wetensch. Letterk. Schoone Kunsten 1842: 187 (1843) 78: " 6 Toledo, G.R. Proctor 35922 (BRH 125 , F 8 - 115 ). Adiantum tetraphyllum Humb. & Bonpl. ex Willd., Sp. pi 5: 441 (1810) 78:116 Orange Walk, C.L. Lundell 404 (NY); Stann Creek, R. Rivero et al. 2523 (BRH 125 ); Toledo, B.K. Hoist 4360 (MO 91 - 115 ). Adiantum trapeziforme L., Sp. pi. 2: 1097 (1753) 78: " 6 Cayo, P.H. Gentle 2342 (F 8 - " 5 , MO 115 , NY); Toledo, G. Davidse 36198 (MO 115 ). Adiantum trichochlaenum Mickel & Beitel in Mem. New York Bot. Card. 46:29, f. 41L (1988) Toledo, B.K. Hoist 5873 (MO 115 ). Jermy did not record this taxon from Belize in the published version of Flora Mesoamericana (Moran et al., 1995: 116), but it was included in the Internet version based on a collection made in 1997 from Toledo District and identified by A.R. Smith in the same year. Adiantum tricholepis Fee, Mem. Foug. 8: 72 (1857) 78: uo Cayo, M. Brunt 2219 (BM-000543178!); Orange Walk, T. Arnason & J. Lambert 17177 (MO 78 - " 5 , NY); Toledo, T.E. Hawkins 1676 (MO 115 ). Adiantum villosum L., Syst. nat. 10th ed., 2: 1328 (1759) 78: 117 Cayo, A. Hughes 135 (BM-000528108! 22 , BM-000531916!); Orange Walk, M.J. Balick 3220 (BRH 125 , NY); Toledo, P.H. Genr/e2409(F 8 - 115 , NY). Adiantum wilesianum Hook., Sp.fil. 2: 50 (1851) 78: 117 Belize, P.H. Gentle 1569 (MO 115 , US); Cayo, A. Hughes 144b (BM-000528109!, BM-000531930!); Stann Creek, W.A. Schipp 48 (BM-000543267!, BRH 125 , US); Toledo, G. Davidse et al. 36450 (BM-B000531990, BRH, MO 115 ). Adiantum wilsonii Hook., Sp.fil. 2: 6 (1851) 78: 113 Toledo, C. Whitefoord 1855 (BM-000543 176! 78 -" 5 , NY). Cheilanthes microphylla (Sw.) Sw., Syn.fil.: 111 (1806) 131: 127 Cheilanthes microphylla var. microphylla 131 Cayo, C.L. Lundell 6542 (MICH 28 - 121 , US). Cheilanthes notholaenoides (Desv.) Maxon ex Weath. in Contr. Gray Herb. 114: 34 (1936) 131: 128 Cayo, A. Hughes 92 (BM-000557720! 22 , BRH, RNG). Hemionitis palmata L., Sp. pi.: 1077 (1753) 97: 132 Stann Creek, W.A. Schipp 250 (BM-000543336!, BRH 125 , MICH 97 , MO 115 ); Toledo, C. Whitefoord 1904 (BM- 000543266!). Pityrogramma calomelanos (L.) Link, Handbuch 3: 20 (1833) 42: Pityrogramma calomelanos var. calomelanos 42 Belize, P.H. Gentle 1548 (K, NY); Cayo, T.B. Croat 24862 (MO 42 - 115 ); Stann Creek, P.H. Gentle 8372 (BM-000543264!, NY); Toledo, B.K. Hoist et al. 35596 (MO 115 ). Pteris altissima Poir. in Lam., Encycl. 5: 722 (1804) 43: 141 Cayo, A. Hughes 72 (BM-000531762! 22 , BM-000531800!, BM- 000531801!); Stann Creek, /. Robertson23\ (BM-000543 191!); Toledo, T.B. Croat 23808 (MO 43 - 115 ). Pteris biaurita L., Sp. pi: 1076 (1753) 43: 141 Cayo, A. Hughes 114 (BM-000531903!, BM-000557721! 22 ); Toledo, P.H. Gentle 2376 (US). Pteris grandifolia L., Sp. pi. 2: 1073 (1753) 43 l41 Cayo, A. Hughes 37 (BM-000531758! 22 , BM-000531856!, BM- 000531857!); Orange Walk, C. Whitefoord 8169 (BM-000543166!); Toledo, P.H. Gentle 1523 (K). Pteris longifolia L., Sp. pi.: 1074 (1753) 43: 142 Cayo, A. Hughes 81 (BM-000557714! 22 ); Orange Walk, C. Whitefoord 8168 (BM-000543 171!); Stann Creek, W.A. Schipp 428 (BM-000543 173!, BRH 125 ); Toledo, J.D. Dwyer et al. 261 (MO 43 - 115 ). Pteris propinqua J. Agardh, Recens. spec. Pter.: 65 (1839) Toledo, D.L. Holland 14A (MO 115 , UC). Not included for Belize in the published version of Flora Mesoamericana (Moran, 1995g: 143) but included in the Internet version based on a collection from Toledo District (D.L. Holland 14A, UC) made in 1997 and determined by A.R. Smith. Pteris pungens Willd., Sp. pi. 5: 387 (1810) 43: 144 Cayo, D.R. Hunt 607 (BM-000543 170!); Stann Creek, W.A. Schipp 289 (BM-000543 167!, BRH 125 ); Toledo, C. Whitefoord 1849(BM-000543169!). Pteris quadriaurita Retz., Observ. bot. 6: 38 (1791) 43: 144 Toledo, B.K. Hoist 3900 (MO 43 - 91 - 115 ). SALVINIACEAE Salvinia auriculata Aubl., Hist. pi. Guiane 2: 969 (1775) 75: 3% Orange Walk, G. Davidse & A.E. Brant 32892 (MO 75 - 115 ). Salvinia minima Baker in J. Bot. 24: 98 (1886) 75:396 Belize, G. Davidse & A.E. Brant 33130 (MO 115 ); Toledo, C. Whitefoord 2145 (BM, MO 75 - 115 ). SCHIZAEACEAE Actinostachys germanii Fee, Mem. Foug. 11: 123 (1866) I02; Toledo, M.E. Peck 936 (GH 120 ). Anemia adiantifolia (L.) Sw., Syn.fil.: 157 (1806) 76:53 Belize, P.H. Gentle 1316 (K!, MO 76 - 115 ); Cayo, A. Hughes 83 (BM-000557706! 22 ); Toledo, G. Davidse & A.E. Brant 32322 (MO 115 ). Anemia bartlettii Mickel in Iowa State J. Sci. 36: 420 (1962) 76:54 Cayo, H.H. Bartlett 1 1898 (MICH-holotype 76 - " 5 , MICH-isotype, UC, US), D.R. Hunt 428 (BM, BRH, UCWI, US 120 ), J.R. Wiley 402 (MO 76 - 115 ). One of very few Belizean endemic taxa, this locally distributed fern from the Mountain Pine Ridge area in Cayo District was first collected in 1931. It was identified as A. flexuousa by Maxon (1944a: 17) and distributed under that name, but not recognized as a new species until 1962. The specimen cited in the distribu- tional statement in Flora Mesoamericana (Moran & Mickel, 1995: 54) as 'B (Wiley 4402, MO)' is incorrect as the collections made by J.R. Wiley in August 1970 were less than 500 numbers and another record on TROPICOS correctly lists this as 402. Anemia hirta (L.) Sw., Syn.fil.: 155 (1806) 76;54 Cayo, D.A. Sutton et al. 4 (BM-000543239!). Anemia mexicana Klotzsch in Linnaea 18: 526 (1844) 76:55 Anemia mexicana var. makrinii (Maxon) Mickel in Brittonia 33: 42 1 (1981) 76:55 Cayo, A. Hughes 99 (BM-000531834!, BM-000557705! 22 ); Toledo, G. Davidse & A.E. Brant 32127 (MO 76 - 115 ). Described originally from Mexico, this taxon has only been recorded from Belize for the Flora Mesoamericana area (Moran & Mickel, 1995: 55). 92 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS Anemia oblongifolia (Cav.) Sw., Syn.fil.: 156 (1806) 76:55 Cayo, J.N. Hedger 319 (BM-000543241!); Toledo, J.R. Wiley 351 (MO 76 - 115 ). Anemia pastinacaria Moritz ex Prantl, Unters. Morph. Gefasskrypt. 2: 110(1881) 76:55 Cayo, T.E. Hawkins 1031 (MO 115 ); Stann Creek, W.A. Schipp 366 (K!, MO 76 ' 115 , NY). Anemia speciosa C. Presl, Suppl. tent, pterid.: 89 (1846) 76:56 Cayo, C.L. Lundell 6906 (MICH 28 , MO 76 " 5 , NY); Toledo, B.K. Hoist Qtal 35855 (MO 115 ). Lygodium heterodoxum Kunze, Farrnkrduter 2: 32 (1849) 37:56 Cayo, A. Hughes 55 (BM-000531763! 22 ); Toledo, G. Davidse 36891 (BM-000531988!, MO 115 ). Lygodium venustum Sw. in/. Bot. 1801(2): 303 (1803) 37;57 Belize, P.P. Barlee s.n. (K!); Cayo, A. Hughes 16 (BM- 000531764! 22 , BM-000531880!); Corozal, M.J. Balick et al. 2190 (MO 115 ); Orange Walk, C. Whitefoord 8054 (BM- 000543272!, F); Stann Creek, G.R. Proctor 36566 (MO 115 ); Toledo, W.A. Schipp 328 (MO 76 ' 115 ). Lygodium volubile Sw. in /. Bot. 1801(2): 304 (1803) 37:57 Cayo, C.L. Lundell 6613 (MICH 28 , NY); Stann Creek, D.R. Hunt 355 (BM-000543271 !); Toledo, B.K. Hoist et al. 35597 (MO 115 ). Schizaea elegans (Vahl) Sm. in Mem. Acad. Roy. Sci. (Turin) 5(1790-1791): 419 (1793) 103:57 Stann Creek, PH. Gentle 3487 (MEXU 103 - 115 , NY, US); Toledo, E.J.F. Campbell 10 (K!). Schizaea poeppigiana J.W. Sturm in Mart., Fl. bras. 1(2): 181 (1859) 103:57 Toledo, G. Davidse & A.E. Brant 32268 (MO 103 ' 115 ). SELAGINELLACEAE Selaginella apoda (L.) Spring in Mart., Fl. bras. 1(2): 119(1 840), as 'Selaginella apus' Toledo, C. Whitefoord 3300 (BM-000543202!). Not recorded for Belize by Fraile in Flora Mesoamericana (Fraile et al., 1995: 29) but present in Chiapas and Guatemala. The specimen listed here was identified by Fraile in 1987 yet not included in the flora; further investigation is required. Selaginella cladorrhizans A. Braun in Ann. Sci. Nat., Bot. ser. 5, 3: 282(1865) 17:3 Cayo, H.H. Bartlett 11457 (BM 17 ' 115 ); Toledo, C. Whitefoord 8359(BM-000543203!). Selaginella diffusa (C. Presl) Spring in Bull. Acad. Roy. Sci. Bruxelles 10: 143(1843) 17:26 Cayo, B.H. Allen 15208 (BM-000543206!, MO 3 ' 17 ' 115 ). Selaginella eurynota A. Braun in Ann. Sci. Nat., Bot. ser. 5, 3: 293 (1865) Toledo, G. Davidse 35970 (MO 115 ), G.R. Proctor 36009 (BM- 000543200!). This Central American endemic was not recorded by Fraile for Belize in Flora Mesoamericana (Fraile et al., 1995: 26) but material was collected under this name in 1976 (G.R. Proctor 36009, BM!) and a subsequent collection in 1996 (Davidse 35970, MO) was determined as this taxon by A.R. Smith in 1997. Selaginella flagellata Spring in Bull. Acad. Roy. Sci. Bruxelles 10: 228 (1843) Toledo, B.K. Hoist et al. 5522 (MO 115 ). Fraile (Fraile et al., 1995: 32) did not record this taxon for Belize in Flora Mesoamericana, though it was listed from Chiapas and Guatemala. It is included here on the basis of a specimen col- lected from Toledo District in 1997 and determined by A.R. Smith in the same year. Selaginella guatemalensis Baker in J. Bot. 21: 243 (1883) I7:3 Toledo, C. Whitefoord 1664 (BM-000543249! 17 ' 115 , NY). Selaginella harrisii Underw. & Hieron. in Urb., Symb. antill. 7: 162 (1912) 17:33 Fraile (Fraile et al., 1995: 33) did not cite a specimen from Belize but referred to extralimital distribution in the Pteridophyte flora ofOaxaca (Mickel & Beitel, 1988: 341). Specimens need to be located to confirm the record. Selaginella hoffmannii Hieron. in Hedwigia 41: 41 (1902) 17:33 Cayo, C. Whitefoord 2059 (BM-000543204!). Selaginella huehuetenangensis Hieron. in Hedwigia 43: 32 (1904) 17;34 Belize, C. Whitefoord 2492 (BM-000543152!); Cayo, H.H. Bartlett 13032 (BM-000543154!); Toledo, M.E. Peck 634 (BM- 000543155!). Selaginella idiospora Alston in Bull. Brit. Mus. (Nat. Hist.), Bot. 1: 246, t. 6(1955) 17:34 Cayo, C. Whitefoord 1253 (BM-000543207! 17 - 1!5 ); Toledo, G. Davidse 36100 (MO 115 ). Selaginella microdendron Baker in /. Bot. 23: 116 (1885) 17:35 Stann Creek, W.A. Schipp 99 (BM-000543205! 17 ' 115 ); Toledo, J.R. Wiley 356 (MO 115 ). Selaginella mollis A. Braun in Ann. Sci. Nat., Bot. ser. 5, 3: 276 (1865) I7:36 Belize, C. Whitefoord 2765 (BM-000543201!); Toledo, W.A. Schipp 925 (NY). Selaginella ovifolia Baker in J. Bot. 22: 90 (1884) 17:37 Toledo, W.A. Schipp 924[a] (BM 17 - I15 ). Selaginella pallescens (C. Presl) Spring in Mart., Fl. bras. 1(2): 132 (1840) 17:37 Selaginella pallescens var. acutifolia Stolze in Amer. Fern 7. 71: 51 (1981) 17:37 Cayo, C. Whitefoord 1255 (BM-000543211! 17 ' 115 ). Selaginella pallescens var. pallescens 11 37 Belize, T.B. Croat 23835 (NY); Cayo, C. Whitefoord 1941 (BM- 000543212! 17 - 115 ); Toledo, B.K. Hoist 3881 (MO 91 - 115 ). Selaginella sertata Spring in Mem. Acad. Roy. Soc. Belgique 24: 104 (1 849) 17: 28 Belize, P.H. Gentle 1396 (BM 17 , NY); Cayo, C. Whitefoord 1091 (BM-000543209!); Orange Walk, C.L. Lundell 394 (NY); To- ledo, B.K. Hoist 4341 (MO 91 - 115 ). Selaginella silvestris Aspl in Ark. Bot. 20A(7): 30, f. 3-5 (1926) Cayo, G.R. Proctor 30105 (BM-000543226!); Toledo, C. Whitefoord 1983 (BM-000543213!, NY). Not recorded for Belize by Somers and Moran in the published version of Flora Mesoamericana (Fraile et al., 1995: 28), though many specimens have been collected recently from Toledo Dis- trict and material from as early as 1969 (G.R. Proctor 30105, BM!) is known from the country. Selaginella stellata Spring in Flora 21: 194 (1838) 17:28 Stann Creek, WA. Schipp 52 (BM-000543224!, MO 17 , NY); Toledo, C. Whitefoord 1776 (BM-000543223!). Selaginella umbrosa Lem. ex Hieron. in Engler & Prantl, Nat. Pflanzenfam. 1(4): 683, f. 404 (1901) 17:41 Belize, P.H. Gentle 1552 (K, NY); Cayo, D.J. Lewis 43 (BM- 00054322 1 !); Orange Walk, C.L. Lundell 395 (NY); Stann Creek, W.A. Schipp 51 (BM-000543216!, NY); Toledo, C. Whitefoord 1851 (BM, MEXU 17 , NY). TECTARIACEAE Ctenitis equestris (Kunze) Ching in Sunyatsenia 5(4): 250 (1940) 54: 197 Ctenitis equestris var. equestris 54 197 RECENT RECORDS OF PTERIDOPHYTES 93 Stann Creek, W.A. Schipp 276 (BM-000543331 !, BM- 000543332!, BRH 125 , MO 54 ' 115 ). Ctenitis excelsa (Desv.) Proctor in Rhodora 63: 34 (1961) 54: 198 Cayo, J.D. Dwyer 11200 (MO 54 ' 115 ); Toledo, G. Davidse 36202 (MO 115 ). Ctenitis interjecta (C. Chr.) Ching in Sunyatsenia 5(4): 250 (1940) 54: 198 Toledo, T.B. Croat 24223 (MO 54 ' 115 ). Ctenitis melanosticta (Kunze) Copel., Gen. fil. (Ann. Cryptog. Phytopathol. 5): 124 (1947) 54: '" Cayo, T.B. Croat 23320 (MO 54 - l15 ); Toledo, B.K. Hoist et al. 35883 (MO 115 ). Ctenitis nigrovenia (H. Christ) Copel., Gen. fil. (Ann. Cryptog. Phytopathol. 5): 124 (1947) 54:199 Cayo, D.A. Sutton et al. 7 (BM-000543335!, MO 54 - l15 ). Ctenitis salvinii (Baker) Stolze inAmer. Fern J. 67: 43 (1977) 54: '" Toledo, B.K. Hoist 3890 (MO 91 - 115 ), WA. Schipp S-773 (GH 54 115 ). Cyclopeltis semicordata (Sw.) Sm. in Bot. Mag. 72(Companion): 36 (1846) 55:20 Orange Walk, M.J. Balick 3222 (BRH 125 , NY); Toledo, J.D. Dwyer 9921 (MO 55 - 115 ). Dictyoxiphium panamense Hook., Gen. fil: 62 (1840) 56:201 Stann Creek, W.A. Schipp 228 (BRH 125 , NY); Toledo, G. Davidse &A.E. Brant 32211 (MO 56 - 115 ). Lastreopsis effusa (Sw.) Tindale in Victoria Naturalist 73: 184 (1 957)57:201 Lastreopsis effusa subsp. divergens (Willd. ex Schkuhr) Tindale in Contr. New South Wales Natl. Herb. 3: 299, t. 21 (1965) 57: 201 Cayo, A. Hughes 133 (BM-000557727! 22 ); Toledo, T.B. Croat 24463 (MO 57 ' 115 ). Lastreopsis exculta (Mett.) Tindale in Victoria Naturalist 73: 185 (1957) Lastreopsis exculta subsp. exculta Toledo, G. Davidse 35648 (MO 115 ), G. Davidse 35799 (MO 115 ), G. Davidse 36208 (MO 115 ). Recorded from Chiapas, Honduras and Guatemala in Flora Mesoamericana (Moran, 1995w: 202), but not from Belize. Included here on the basis of material collected from Toledo District and identified as L. exculta (Mett.) Tindale by A.R. Smith in 1997. Further material from Belize was identified as L. exculta subsp. guatemalensis (Baker) Tindale, though this name is included as a synonym of subsp. exculta by Moran. Megalastrum lunense (H. Christ) A.R. Sm. & R.C. Moran in Amer. Fern J. 77: 128(1987) Toledo, B.K. Hoist 5820 (MO 115 , UC). This Central American endemic was not recorded for Belize in the published version of Flora Mesoamericana (Smith & Moran, 19950: 203), but included in the Internet version on the basis of a collection from Toledo District (B.K. Hoist 5820, UC) made in February 1997 and identified by A.R. Smith later the same year. This is the first record of the genus from Belize. Tectaria heracleifolia (Willd.) Underw. in Bull. Torrey Bot. Club 33: 200(1906) 58:207 Tectaria heracleifolia var. heracleifolia 20 Belize, C. Whitefoord 1163 (BM-000543323!); Cayo, C. Whitefoord 2034 (BM, MO 58 - " 5 ); Orange Walk, C.L. Lundell 338 (NY); Toledo, G. Davidse & D.L. Holland 36495 (BM- 000531986!, MO 115 ). Tectaria incisa Cav., Descr. pi: 249 (1802) 58:207 Cayo, D.L. Spellman & W.W. Newey 1849 (MO 58 - l15 ); Stann Creek, W.A. Schipp 272 (BM-0005433 15!, BRH 125 , NY); Toledo, C. Whitefoord 1583 (BM-0005433 16!). Tectaria mexicana (Fee) C.V. Morton in Amer. Fern J. 56: 133 (1 966) 58:207 Belize, P.H. Gentle 1551 (K, NY); Cayo, A. Hughes 51 (BM- 000531 775 ! 22 , BM-000531780!); Toledo, G. Davidse & A.E. Brant 32371 (MO 58 ' 115 ). Tectaria nicotianifolia (Baker) C. Chr., Index filic., Suppl. 3: 182 (1934) Toledo, G. Davidse & D.L Holland 36603 (UC 115 ). Not recorded for Belize in Flora Mesoamericana (Moran, 1 995 v: 208), but present in Guatemala, Honduras and from Nicaragua to Panama. Included in the Internet version based on the specimen from Toledo District listed above collected in 1997 and deter- mined by A.R. Smith. Tectaria pilosa (Fee) R.C. Moran in Novon 2: 138 (1992) Toledo, G.R. Proctor 36155 (BM-0005433 14!). Only recorded for Costa Rica and Panama in Flora Mesoamericana (Moran, 1995v: 208), T. pilosa is listed here based on a collection from high forest in the Columbia Forest Reserve of Toledo, originally identified as T. incisa var. pilosa by G.R. Proctor. It differs from T. heracleifolia in the oblique or decurrent base to the terminal segment of the lamina and rather sparsely pilose upper and lower faces. Further investigation of this material is required. Tectaria plantaginea (Jacq.) Maxon in Contr. U.S. Natl. Herb. 10: 494(1908) 58:208 Stann Creek, W.A. Schipp 465 (BRH 125 , MO 58 - " 5 , NY). Tectaria rivalis (Mett. ex Kuhn) C. Chr., Index filic., Suppl. 3: 184 (1 934)58: 208 Cayo, T.E. Hawkins 1273 (MO 115 ); Stann Creek, W.A. Schipp S- 66 (BM-000543327!, F 58 - 115 , NY). Tectaria vivipara Jermy & T.G. Walker in Bull. Brit. Mus. (Nat. Hist.), Bot. 13: 274, f. 15 (1985) 58:209 Belize, P.H. Gentle 1542 (K, MO 58 " 5 , NY); Toledo, C. Whitefoord 1583 (BM, NY). THELYPTERIDACEAE Macrothelypteris torresiana (Gaud.) Ching in Acta Phytotax. Sin. 8:310(1963) Cayo, A. Hughes 143 (BM-000531924!, BM-000557726! 22 ); Toledo, G. Davidse 35918 (MO 115 ). Not recorded for Belize in Flora Mesoamericana (Smith, 1995a: 164). The species and genus were first listed for Belize by Hughes (1998) for the Chiquibul Forest Reserve and additional material is cited in TROPICOS based on identifications by A.R. Smith in 1997 and 1999. Thelypteris balbisii (Spreng.) Ching in Bull. Fan. Mem. Inst. Biol, Bot. 10:250(1941) 110:17 Stann Creek, P.H. Gentle 2720 (NY); Toledo, A.H. Gentry 7900 (UC 110 ). Thelypteris biolleyi (H. Christ) Proctor in Bull. Inst. Jamaica, Sci. Ser. 5: 58 (1953) 110 183 Toledo, G. Davidse & A.E. Brant 32208 (MO 115 , UC 11 115 ). Thelypteris blanda (Fee) C.F. Reed in Phytologia 17: 264 (1968) 110: 183 Cayo, A. Hughes 113 (BM-000531751! 22 ); Toledo, M.E. Peck s.n. (US 110 - 115 ). Thelypteris decussata (L.) Proctor in Bull. Inst. Jamaica, Sci. Ser. 5: 59(1953) Thelypteris decussata var. costaricensis A.R. Sm.in Univ. Calif. Publ. Bot. 76: 16(1980) Toledo, B.K. Hoist 5919 (MO 115 ). This Central American endemic was not recorded for Belize in 94 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS the published version of Flora Mesoamericana (Smith, 1995ft: 194), but was recorded from Honduras, Costa Rica and Panama. Belize was included in the Internet version based on a specimen collected from Toledo District listed here and identified by A.R. Smith in 1997. A second specimen (Hoist 5819, MO), identified by A.R. Smith as T. decussata (L.) Proctor, may also be referable to var. costaricensis. Thelypteris dentata (Forssk.) E.P. St. John in Amer. Fern J. 26: 44 (1936)110:181 Toledo, G. Davidse 36886 (MO 115 ), R. Rivero 2539 (UC). Not recorded for Belize in the published version of Flora Mesoamericana (Smith, 1995ft: 181) but included in the Internet version based on a collection (Rivero et al. 2539, UC) identified by A.R. Smith in March 1996. Thelypteris falcata (Liebm.) R.M. Tryon in Rhodora 69: 6 (1967)" 192 Stann Creek, W.A. Schipp 97 (BRH 125 , NY); Toledo, P.H. Gentle 8693 (NY, UC"). Thelypteris ghiesbreghtii (Hook.) C.V. Morton in Contr. U.S. Natl. //- cumulative % taxa recorded by each decade trendline (polynomial: y = 2.6442x' 5763 ) 80 70 60 50 40 30 20 10 97 1800- 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 Checklists 1990 Internet decade World Wide Web databases Spellmanetal. (1975) Dwyer & Spellman (1981) Vargas & Shawe ( 1997) Regional floristics Botany of the Maya area (1940-1961) Flora of Guatemala (1946-1985) Flora Neotropica (1968-) Flora Mesoamericana (1994-) National floristics Standley & Record (1936) Preliminary checklists Campbell (1899) Fig. 2 Growth of knowledge of the pteridophyte flora of Belize. 98 D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS documented. The project currently seeks to collate basic informa- tion for vascular plant taxa from Belize, supported by relevant specimens and literature, and some 8000 names, 7000 references and 32 000 specimens are in the database at the time of submission of the paper. It is estimated that minimal information on at least 45 000 specimens from Belize is comparatively readily available in the public domain, and that 75-100 000 specimens have been collected from the country. This material forms the basis for the cumulative knowledge on the flora of Belize, and provides a basis for further research. REFERENCES 1. Adams, C.D. 1995a. Diplazium Sw. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 228-246. 2. 19956. Asplenium L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 290-324. 3. Allen, B.H. 1995. Vegetation. In S. Matola (Ed.), Expedition to Doyle's Delight, Southern Maya Mountains, Belize. Forest Planning Management Project Occa- sional Series 5: 10-30. 4. Alston, A.H.G. 1955. The heterophyllous Selaginellae of continental North America, Bulletin of the British Museum (Natural History), Botany 1: 221-274, t. 5-6. 5. Arvigo, R. & Balick, MJ. 1993. Rainforest remedies. One hundred healing herbs of Belize. Twin Lakes. 6. Bishop, L.E. 1995. Cochlidium Kaulf. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 371-372. 7 . Blake, J. 1 9 1 7 . Descriptions of new Spermatophy tes, chiefly from the collections of Prof. M.E. Peck in British Honduras. Contributions from the Gray Herbarium of Harvard University 52: 59-106. 8. Campbell, E.J.F. 1899. List of plants under cultivation at the Belize Botanic Station, 1899: 1-8. [Unpublished list] 9. Camus, J.M. & Perez-Garcia, B. 1995. Danaea Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 48-50. 10. Copeland, E.B. 1952. American species of Xiphopteris. American Fern Journal 42:93-110. 11. Crane, E.H. 1997. A revised circumscription of the genera of the fern family Vittariaceae. Systematic Botany 22: 509-517. 12. Davidse, G., Sousa S., M. & Chater, A.O. 1994. Flora Mesoamericana 6. Mexico City. 13. & Knapp, S. 1995. Flora Mesoamericana. 1. Mexico City. 14. Dwyer, J.D. & Spellman, D.L. 1981. A list of the Dicotyledoneae of Belize. Rhodora 83: 161-236. 15. Evans, A.M. 1968. 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Moran, R.C. 1986. The neotropical fern genus Olfersia. American Fern Journal 76: 161-178. 37. 1995a. Lygodium Sw. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 56-57. 38. 19956. Dicranopteris Bernh. In G. Davidse etal. (Eds), Flora Mesoamericana 1:58. 39. 1995c. Sticherus C. Presl. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 59-62. 40. 1 995d. Cyathea Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 93- 103. 41. 1995e. Acrostichum L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 105-106. 42. 1 995/ Pityrogramma Link. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 137-140. 43. 1995g. Pteris L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 140- 144. 44. 1995/1. Anetium Splitg. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 145-146. 45. 1995/. Antrophyum Kaulf. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 146-147. 46. 1995/. Vittaria Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 148- 150. 47. 1 995k. Dennstaedtia Bernh. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 151-153. 48. 1995/. Hypolepis Bernh. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 153-157. 49. 1995m. Lindsaea Dryand. ex Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 157-160. 50. 1995n. Lonchitis L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 160. 51. 1995o. Odontosoria Fee. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 160-161. 52. 1995/7. Pteridium Gled. ex Scop. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 161-162. 53. 1995