flora of tropical east africa - woodsiaceae (2003) (flora of tropical east africa)

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LIST OF ABBREVIATIONS AVP=OHedberg Afroalpine Vascular Plants BJBB=Bulletin du Jardin Botanique de lrsquoEtat Bruxelles Bulletin du Jardin Botanique Nationale de Belgique BSBB=Bulletin de la Socieacuteteacute Royale de Botanique de Belgique CFA= Conspectus Florae Angolensis EJ=AEngler Botanische Jahrbuumlcher fuumlr Systematik Pflanzengeschichte und Pflanzengeographie EM=AEngler Monographieen Afrikanischer Pflanzen-Familien und Gattungen EP=AEngler Das Pflanzenreich EPA=GCufodontis Enumeratio Plantarum Aethiopiae Spermatophyta in BJBB 23 Suppl (1953) et seq E amp P Pf=AEngler amp KPrantl Die Natuumlrlichen Pflanzenfamilien FAC=Flore drsquoAfrique Centrale (formerly FCB) FCB=Flore du Congo Belge et du Ruanda-Urundi Flore du Congo du Rwanda et du Burundi FD-OA=APeter Flora von Deutsch-Ostafrika FFNR=FWhite Forest Flora of Northern Rhodesia FPNA=W Robyns Flores des Spermatophytes du Pare National Albert FPS=FWAndrews Flowering Plants of the Anglo-Egyptian Sudan or Flowering Plants of the Sudan FPU=ELind amp ATallantire Some Common Flowering Plants of Uganda FR=FFedde Repertorium Speciorum Novarum Regni Vegetabilis FSA=Flora of Southern Africa FTA =Flora of Tropical Africa FWTA=Flora of West Tropical Africa FZ=Flora Zambesiaca GFP=JHutchinson The Genera of Flowering Plants GP=GBentham amp JDHooker Genera Plantarum GT=DMNapper Grasses of Tanganyika IGU=KWHarker amp DMNapper An Illustrated Guide to the Grasses of Uganda ITU=WJEggeling Indigenous Trees of the Uganda Protectorate JB=Journal of Botany JLS=Journal of the Linnean Society of London Botany KB=Kew Bulletin or Bulletin of Miscellaneous Information Kew KTS=IDale amp PJGreenway Kenya Trees and Shrubs KTSL=HJBeentje Kenya Trees Shrubs and Lianas LTA=EGBaker Leguminosae of Tropical Africa NBGB=Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem POA=AEngler Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete RKG=AVBogdan A Revised List of Kenya Grasses TSK=EBattiscombe Trees and Shrubs of Kenya Colony TTCL=JPMBrenan Check-lists of the Forest Trees and Shrubs of the British Empire no 5 part II Tanganyika Territory UKWF=ADQAgnew (or for ed 2 ADQAgnew amp SAgnew) Upland Kenya Wild Flowers UOPZ=ROWilliams Useful and Ornamental Plants in Zanzibar and Pemba VE=AEngler amp O Drude Die Vegetation der Erde IX Pflanzenwelt Afrikas WFK=AJJex-Blake Some Wild Flowers of Kenya ZAE= Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907ndash1908 2 (Botanik)

FAMILIES OF VASCULAR PLANTS REPRESENTED IN THE FLORA OF TROPICAL EAST AFRICA

The family system used in the Flora has diverged in some respects from that now in use at Kew and the herbaria in East Africa The accepted family name of a synonym or alternative is indicated by the word ldquoseerdquo Included family names are referred to the one used in the Flora by ldquoinrdquo if in accordance with the current system and ldquoasrdquo if not Where two families are included in one fascicle the subsidiary family is referred to the main family by ldquowithrdquo Foreword and preface (pound300) Glossary (pound2350) Index of Collecting Localities (pound1850)

PTERIDOPHYTA Actiniopteridaceae (pound230) Grammitidaceae Polypodiaceae (pound1000)

Adiantaceae (pound1500) Hymenophyllaceae Psilotaceae (pound170)

Aspleniaceae Isoetaceae Pteridaceae (pound850)

Azollaceae (pound170) Lomariopsidaceae (pound1000) Salviniaceae (pound140)

Blechnaceae Lycopodiaceae Schizaeaceae (pound340)

Cyatheaceae Marattiaceae (pound170) Selaginellaceae

Davalliaceae (pound170) Marsileaceae (pound600) Thelypteridaceae

Dennstaedtiaceae (pound650) Oleandraceae (pound600) Vittariaceae (pound295)

Dryopteridaceae Ophioglossaceae (pound700) Woodsiaceae (pound800)

Equisetaceae (pound170) Osmundaceae (pound170)

Gleicheniaceae (pound230) Parkeriaceae (pound170)

GYMNOSPERMAE (pound300) Cupressaceae Cycadaceae Podocarpaceae

ANGIOSPERMAE Acanthaceae Aponogetonaceae (pound390)

Agavaceae Aquifoliaceae (pound150)

Aizoaceae (pound450) Araceae (pound1020)

Alangiaceae (pound150) Araliaceae (pound300)

Alismataceae (pound300) Arecaceaemdashsee Palmae

Alliaceae (pound400) Aristolochiaceae (pound300)

Aloaceae (pound1450) Asclepiadaceaemdashsee Apocynaceae

Amaranthaceae (pound2200) Asparagaceae

Amaryllidaceae (pound510) Asphodelaceae (pound600)

Anacardiaceae (pound850) Asteraceaemdashsee Compositae

Ancistrocladaceae (pound185) Avicenniaceaemdashas Verbenaceae

Anisophyllaceaemdashas Rhizophoraceae

Annonaceae (pound1050) Balanitaceae (pound600)

Anthericaceae (pound1150) Balanophoraceae (pound195)

Apiaceaemdashsee Umbelliferae Balsaminaceae (pound1180)

Apocynaceae Basellaceae (pound150)

Part 1 (pound2050) Begoniaceae

Part 2 Berberidaceae (pound150)

copy 2003 Royal Botanic Gardens Kew

FLORA OF TROPICAL EAST AFRICA WOODSIACEAE

BERNARD VERDCOURT Terrestrial with erect or less often creeping rhizome sometimes forming a short

caudex stipe not articulate (except in Woodsia) often densely scaly particularly near base with 2 vascular strands united and U-shaped above Frondsplusmnspaced or tufted lamina 1ndash3 pinnate the costal groove with raised often almost winged edges veins free or anastomosing Sori round elliptic to linear or J-shaped or U-shaped with inconspicuous to obvious entire or fimbriate indusium or indusium sometimes lacking Paraphyses present or absent Spores monolete with perispore

A cosmopolitan family (previously called Athyriaceae but Woodsiaceae has priority by 7 years) included within an extended Dryopteridaceae by Kramer in Kubitzki Fam Gen Vasc Plants As treated here it contains about 15 genera and 500 species with the Onocleaceae kept separate and Hypodematium Kunze treated under Dryopteridaceae 1 Soriplusmnround or ovate 2

Sori at least in part elliptic to elongate or J-shaped to U-shaped but some reduced sori towards apices of pinnule-lobes can be plusmnround

3

2 Fronds large 06ndash2 m long with pinnules of middle pinnae 45ndash7 cm long indusium rounded-reniform sometimes almost absent

1 Dryoathyrium

Fronds smaller up to 40 cm long with pinnules of middle pinnae 05ndash2 cm long indusium bladder-shaped

2 Cystopteris (p 4)

3 At least some sori J-or U-shaped and furthest end crossing the vein to which sorus is attached

3 Athyrium (p 8)

Sori all elliptic to linear never crossing the vein 4

4 Trichomes of margins of rhizome-scales distinctly bifid veins free or anastomosing

4 Callipteris (p 13)

Trichomes of margins of rhizomes-scales spiniform not distinctly bifid veins free

5 Diplazium (p 16)

1 DRYOATHYRIUM

Ching in Bull Fan Mem Inst Biol Bot 1179 (1941)

Rhizome erect forming a short caudex with non-peltate rhizome-scales Fronds tufted stipes not articulated Lamina large bipinnate with pinnules deeply pinnatifid into again pinnatifid lobes so as to appear tripinnate glabrous except for minute short hairs along the narrowly and evenly winged costules herbaceous veins free Sori superficial round with minute rounded-reniform indusia not apparent in mature sori sometimes almost absent

A tropical genus of about 10 species distributed from tropical Africa and Madagascar to China and Japan only one species occurs in continental Africa

Occasionally in Diplazium trichomes can appear minutely bifurcate at extreme apex due to a fine hair below their tips

D boryanum (Willd) Ching in Bull Fan Mem Inst Biol Bot 1181 (1941) Schelpe FZ Pterid 207 t 59 (1970) Faden in UKWF 57 fig (on 59) (1974) De Vol amp Kuo in Fl Taiwan ed1 1470 (1975) Schelpe amp Diniz Fl Moccedilamb 223 (1979) Jacobsen Ferns S Afr 411 t 308 (1983) Pic Serm in BJBB 55155 (1985) Schelpe amp Anthony FSA Pterid 229 fig 771 (1986) Type Reacuteunion Bory de St Vincent (B-W 19831 holo microfiche)

Rhizome erect ascending or horizontal with upturned apex forming a short thick fleshy caudex with pale brown concolorous subentire ovate to lanceolate rhizome-scales 5ndash12(ndash20) mm long 5ndash7 mm wide acuminate Fronds tufted arching herbaceous 06ndash2 m tall stipe matt greenish brown up to 1 m long glabrous except for pale brown scales about the base similar to those on the rhizome Lamina narrowly ovate in outline up to 1times064 m acute basal pinnae hardly reduced bipinnate with 15ndash22 pairs of pinnules so deeply pinnatifid as to appear tripinnate pinnae oblong-lanceolate-acuminate in outline 13ndash44 cm long 7ndash22 cm wide attenuate shortly petiolate developed equally acroscopically and basiscopically the basal pinnules somewhat reduced pinnules oblong-acuminate 3ndash125 cm long 08ndash3 cm wide base truncate deeply pinnatifid into narrowly oblong obtuse deeply crenate-serrate lobes 4ndash16 mm long 2ndash5 mm wide with an angular sinus between them glabrous on both surfaces except for minute short blunt brownish hairs along the costules and veins veins free rachis pale brown glabrous secondary rachises pale brown with scattered minute blunt hairs and with a narrow laminar green wing at least about the insertion of the pinnules Sori circular 4ndash13(ndash16) per pinnule-lobe plusmn06 mm in diameter indusium membranous minute or up to 08 mm wide subentire Fig 1 (page 3)

UGANDA Kigezi District Ishasha Gorge 6 km SW of Kirima 21 Sept 1969 Faden et al 691202 Toro District Ruwenzori Mts Kazinga to Bwamba Pass 23 Jan 1932 Hazel 124 Mbale District Mt Elgon 24 May 1923 Snowden 777 778

KENYA Naivasha District Kimakia Forest Station towards South Kinangop 13 July 1969 Faden 69908 Mt Kenya West Kenya Forest Station 5 Jan 1922 RE amp TCE Fries 742 Masai District Narok District about 48 km from Olokurto edge of Olenguerone Settlement 16 May 1961 Glover et al 1167

TANZANIA Arusha District 8 km N of Arusha S slope of Mt Meru above Eastern sawmills 29 Feb 1953 Drummond amp Hemsley 1298 Mpanda District Mahali Mts Sisaga 29 Aug 1958 Newbould amp Jefford 1915 amp 1916 Morogoro District Uluguru Mts Mwera Valley 26 Sep 1970 Faden et al 70586

DISTR U 2 3 K 3ndash6 T 2 4 6 7 Ivory Coast (fide Tardieu) S Nigeria Cameroon Bioko E Congo (Kinshasa) Rwanda Mozambique Zimbabwe South Africa Madagascar Reunion and Comoro Is also widespread in tropical and temperate Asia to NE Himalayas and Malesia (to Luzon)

HAB Evergreen forest often by streams also in bamboo zone and in mist forest 1350ndash2700 m

SYN Aspidium boryanum Willd Sp Pl ed 4 5285 (1810) Lastrea boryana (Willd) Moore Ind Fil 86 (1858) Nephrodium catopteron (Kunze) Hook var glabrum Hook Sp Fil 4137 (1862)

quoad syntype Bioko [Fernando Po] Mann sn (K syn) N boryanum (Willd) Bak Syn Fil 284 (1867) Aspidium glabratum Kuhn Fil Afr 133 (1868) Type Bioko [Fernando Po] Mann

sn (K holo) Nephrodium glabratum (Kuhn) Bak Syn Fil 300 (1874) Dryopteris glabrata (Kuhn) Kuntze Rev Gen Pl 2812 (1891) Aspidium kiboschense Hieron in POA C 85 (1895) Type Tanzania Kilimanjaro

Kibosho [Kiboscho] Volkens 1554 (B holo BM fragment) Dryopteris kiboschensis (Hieron) C Chr Ind Fil 273 (1906) D boryana (Willd) C Chr Ind Fil 255 (1906) Nephrodium catopteron sensu Peter FD-OA 57 (1929) non (Kunze) Hook

Tardieu-Blot states that the Mann type is at the BM Kuhn would not have seen the Kew Mann specimenmdashbut he cites Hooker

Flora of tropical East Africa 2

FIG 1 DRYOATHYRIUM BORYANUMmdash1 pinnatimes⅔ 2 fertile pinnule segmenttimesfrac12 Both from Schelpe 5663 From FZ Drawn by Monika Shaffer-Fehre

Dryoathyrium 3

Athyrium boryanum (Willd) Tagawa in Act Phytotax Geobot 4144 (1935) Shieh et al in Fl Taiwan ed 2 1418 (1994)

Ctenitis boryana (Willd) Copel Gen Fil 123 (1947) Thelypteris glabrata (Kuhn) Tardieu in Not Syst 14344 (1952) amp in Meacutem IFAN

28120 (1953) pro parte T glabrata (Kuhn) Tardieu var hirsuta Tardieu in Not Syst 14344 (1952) Type

Ivory Coast no specimens cited Athyrium glabratum (Kuhn) Alston in Bol Soc Brot seacuter 2A 3011 (1956) amp Ferns

WTA 64 (1959) Tardieu Fl Cameroun 3232 t 411ndash2 (1964) Cornopteris boryana (Willd) Tardieu in Meacutem Sc Madag 730 t 2 fig 1ndash4 (1956)

nom invalid amp in Amer Fern Journ 4832 (1958) amp in Fl Madag 5(1) 257 fig 331ndash5 (1958)

Parathyrium boryanum (Willd) Holttum in KB 13449 (1959) Lunathyrium boryanum (Willd) HOhba Sci Rep Yokosaka City Mus 1153 (1965) Deparia glabrata (Kuhn) Kato in Bot Mag Tokyo 9036 (1977) D boryana (Willd) Kato in J Fac Sci Univ Tokyo III 13386 (1984) NOTE There is considerable argument over the correct generic placing of this fern

Kramer in Kubitzki (Fam Gen Vasc Pl 136 (1990)) following Katorsquos revision includes Dryoathyrium in Deparia Hook f amp Grev but divides the latter into 4 quite distinct sections one of which is Dryoathyrium (Ching) Kato For the purpose of this flora there seems little point in changing a long used name


2 CYSTOPTERIS

Bernh in Schrad Neues J Bot 1(2) 5 26 (1805) (lsquo1806rsquo) Blasdell in Mem Torrey Bot Club 21(4) 1ndash102 (1963) Kato amp Kramer in Kubitzki Fam Gen Vasc Pl 1138 (1990) Fraser-Jenkins New species syndrome in Indian Pteridology amp the ferns of Nepal 95ndash100 (1997)

Small to medium-sized ferns terrestrial or often growing on rocks Rhizome short and suberect to long-creeping with entire membranous scales Fronds tufted or spaced stipe grooved Lamina rather thin to thinly coriaceous (membranous in one species) usually well-dissected 1ndash3-pinnate and pinnatifid rachis glabrous or scaly occasionally with bulbils (not in Africa) grooved veins free Sori round at first covered by ovate to oblong lanceolate or cup-shaped inflated bladder-like indusia attached below the sori spores echinate or rugose

About a dozen species C fragilis (L) Bernh in Schrad Neues J Bot 1(2) 27 (1805) Pirotta in Il

Ruwenzori Parte Scient 1428 (1909) (sensu lato) Sim Ferns S Afr ed 288 t 8 (1915) (sensu lato) AVP 27 (1957) (sensu lato) Alston FWTA Pterid 64 (1959) (sensu lato) Blasdell in Mem Torrey Bot Club 21(4) 38 (1963) (sensu stricto) WJacobsen Ferns S Afr 402 t 302 (1983) (sensu lato) Schelpe amp NC Anthony FSA Pterid 229 fig 762 (1986) (sensu lato) Derrick Crabbe amp Jermy in Sommerfeltia 662 (1987) (sensu stricto) JBurrows SAfr Ferns 279 t 465 fig 65284 284a (1990) (sensu lato) Crabbe amp Jermy in Fl Eur ed 2 124 (1993) (sensu stricto) Faden in UKWF ed 233 (1994) Type Plukenet Almagestum botanicum 150 t 180 fig 5 (1696) (lecto chosen by Weatherby in Rhodora 37376 (1935))

Schelpe amp NCAnthony (FSAPterid 229 (1986)) gives about 18 Schelpe gives the type as Europe Herb Sloane (HS 96 Fol 40) (BM holo) but Linnaeus would not have seen the specimen It could not be a holotype since Linnaeus cites several references There is also a specimen in the Linnean Herbarium 125151 but there appears to be doubt it was available to Linnaeus in 1753

Rhizome shortly creeping or plusmnerect with reddish brown glabrous lanceolate scales 3 mm long 06 mm wide sometimes with marginal glands Fronds plusmntufted or closely spaced stipe usually shorter than the lamina straw-coloured to dark brown 3ndash20 cm long with lanceolate scales at the base Lamina lanceolate to ovate-lanceolate 5ndash30 cm long 15ndash14 cm wide 1ndash3-pinnate and 3(ndash4)-pinnatifid rachis and axes with scattered multicellular glandular hairs pinnae in 4ndash20 pairs ovate to ovate-lanceolate 06ndash18 cm long 05ndash4 cm wide basal secondary pinnae shortly petiolate remainder adnate ovate to lanceolate obtuse to acute with apices of the lobes with rounded or acute teeth veins directed into teeth or sinuses Sori round discreet to overlapping at maturity indusia bladder-like ovate to oblong or cup-shaped dentate to lacerate

SYN Polypodium fragile L Sp Pl ed 11091 (1753) (as PF fragile) NOTE Blasdell records C fragilis var fragilis from eastern Africa (op cit p 39) but

cites nothing in his list of specimens Fronds 10ndash20 cm long simply pinnate with 4ndash8 pairs of pinnae veins mostly running into lobes or teeth but sometimes into sinuses spines of spores plusmnfurther apart than length of spines

subsp A

Fronds 75ndash37 cm long essentially 2-pinnate with 9ndash20 pairs of pinnae veins mostly running into the sinuses spines of spores densely packed

subsp B

subsp A Fronds 10ndash20 cm tall with stipe 3ndash10 cm long very slender Lamina lanceolate in

outline 5ndash95 cm long 15ndash3 cm wide simply pinnate with 4ndash8 pairs of pinnae and apical lobed segment pinnae and apical segment ovate or triangular in outline 06ndash15 cm long 04ndash11 cm wide shallowly to deeply lobed veins mostly running into lobes or teeth but sometimes into sinuses Spines on spores more sparse plusmnfurther apart than the length of the spines Fig 21 2 (page 6)

UGANDA Mbale District Mt Elgon in crater N slope of Jacksonrsquos Peak 17 May 1948 Hedberg 979 (photocopy)

TANZANIA Moshi District Kilimanjaro Kikafu R gorge 1 Mar 1997 Hemp 1623 amp stream near NW edge of Shira Plateau 8 Nov 1968 Bigger 2281 amp on same plateau Simba Camp area 5 Sep 1993 Grimshaw 93ndash616

DISTR U 3 T 2 HAB Crevices in rocks on rocks overhanging streams hanging from roofs of caves

and rock overhangs 3400ndash4750 m NOTE I at first thought this might be Cystopteris nivalis (Pirotta) Pic Serm judging

by some of Pichi Sermollirsquos comments (see note after subsp B) Discovery of topotypic material in BM (Osmaston 1740 Uganda Bujuku Valley Bigo July 1972) shows this is not so The morphology of the frond and densely echinate spores are the same as in subsp B

Although in East Africa this alpine form seems distinctive similar forms which appear almost identical can be found in S Africa and in the UK

Linnaeus actually gave the name as Polypodium F fragile in Sp Pl ed 11091 (1753) and in Syst Veg ed 101327 (1759) and several authors have suggested that the correct name for the species should be C filix-fragilis (L) Borbaacutes ABalaton Floacuteraacuteja 2 314 (1900) (without hyphen) Chiov in Ann Bot 210 (1903) Becherer in Ber Schweiz Bot Ges 43 (1) 39 (1934) This was discussed by Merrill in Am Fern Journ 25127 (1935) and by Weatherby in Am Fern Journ 2727 (1937) in which I think he conclusively shows the F was an error which Linnaeus corrected in manuscript in his own copy of Sp Pl ed 1 and in Fl Suec ed 2374 (1755) and Sp Pl ed 21553 (1763) This needs reiterating since Jones in Fl Austr 48419 (1998) has given the type of the genus as C filix-fragilis (L) Bernh Example 19 to Article 233 of the Code of Botanical Nomenclature states the name is to be treated as P fragile L My thanks are due to PJEdwards for preparing scanning electron micrographs from my stubs


FIG 2 CYSTOPTERIS FRAGILISmdashsubsp A 1 habittimes⅔ 2 fertile pinna enlarged subsp B 3 habittimes⅔ 4 fertile pinna enlarged 1ndash2 from Bigger 22811 3ndash4 from Battiscombe 978 Drawn by Pat Halliday

Cystopteris 7

subsp B Fronds 75ndash37 cm tall with stipe 35ndash20 cm long Lamina lanceolate or plusmntriangular in

outline 4ndash25 cm long 15ndash10 cm wide essentially 2-pinnate with 9ndash20 pairs of pinnae but pinnules narrowly decurrent deeply pinnatifid sometimes truly 2-pinnate veins mostly running into the sinuses Spores with densely packed spines Fig 23 4 (page 6)

UGANDA Toro District Ruwenzori Namwamba Valley 7 Jan 1935 G Taylor 2984 Kigezi District Virunga Mts Sabinio 24 Nov 1934 G Taylor 2036 Mbale District Mt Elgon Sasa trail Dirigana R 20 June 1997 Wesche 1462

KENYA Nakuru District Mau Forest near Elburgon HMGardner in FD 978 North Nyeri District West Mt Kenya Forest Station 5 Jan 1922 RE amp Th CE Fries 762 Kericho District SW Mau Forest along Kiptiget (Chepkoisi) R plusmn16 km SSE of Kericho Salt Lick Falls 12 June 1972 Faden et al 72331

TANZANIA Masai District Ngorogoro near Old Boma 2 Mar 1961 Newbould 5683 Arusha District Mt Meru crater 8 Mar 1971 Richards 26713 Moshi District Kilimanjaro Marangu R Himo Nov 1964 Beesley 58 Mbeya District Kikando 22 Sep 1956 Richards 6713

DISTR U 2 3 K 1 3ndash5 T 2 7 Cameroon Mt Congo (Kinshasa) (Kivu) Sudan Ethiopia Natal Mediterranean part of Europe Cape Verde Is Canary Is

HAB Mostly on damp rocks rock faces and cliffs often near rivers chiefly in evergreen forests eg Hagenia Podocarpus AlbiziandashMacarangandashEagarandashSyzygiumndashCroton or XymalosndashPrunusndashMaesa ndashAfrocrania also on moist loamy banks and in alpine zone 1400ndash3800 m

SYN Aspidium viridulum Desv Mag Nat Berl 5321 (1811) Type Tenerife collector (P)

Cystopteris viridula (Desv) Desv in Meacutem Soc Linn Paris 6264 (1827) Woodsia nivalis Pirotta in Ann Bot Roma 7173 (1908) amp in Il Ruwenzori Parte

Scient 1478 (1909) Type Uganda Toro District Ruwenzori Mobuku Valley Bujongolo 3800 m (sphalm 3000 m) Cavalli-Molinelli amp Roccati sn (TO holo)

[Cystopteris fragilis L subsp diaphana sensu Litard in Bull Geacuteogr Bot 21 (No 255) 20 (9 Jan 1911)) amp in Bull Soc Bot Deux-Sevres 88 (1911ndash12) Zeiller in Bull Soc Bot Fr 59800 (1912) (abstr) Fraser-Jenkins New species syndrome in Indian Pteridology amp the ferns of Nepal 97 (1997) non (Bory) Litard sensu stricto]

[C diaphana sensu Blasdell in Mem Torrey Bot Club 21 (4) 47 (1963) Derrick Crabbe amp Jermy in Sommerfeltia 662 (1987) Crabbe amp Jermy in Fl Europea ed 224 (1993) non (Bory) Blasdell sensu stricto]

C nivalis (Pirotta) Pic Serm in Webbia 23173 (1968) NOTE I at first thought this could be identified as subsp diaphana (Bory) Litard as

had been suggested by Blasdell and others the venation suggests this Jermy Flora Europaea ed 2 states that the spores are densely echinate their spines almost touching at the base exactly as in the above There is however a photograph of the type of Polypodium diaphanum Bory (Voy Iles Mers Afr 1328 (1804) Type Reunion Plaine des Chicots Bory St Vincent (P holo)) at the BM and attached to it a SEM photograph of spores showing spore echination which confuses the whole issue Desvauxrsquos name cited above probably provides the correct epithet for this subspecies Spores derived from specimens collected in Tenerife have dense echination


Crabbe amp Jermy (op cit) retain this as a species but admit that C fragilis is a widespread polymorphic polyploid complex and I agree with Fraser-Jenkins in regarding it as a subspecies Vida in his cytological paper (Act Bot Acad Sci Hungaricae 20 (1ndash2) 181ndash192 (1974)) also suggest this is the best solution Fraser-Jenkins states that the difference between the spores of subsp fragilis and subsp diaphanum [auctt] is a 100 reliable character However one specimen of subsp B Gardner in FD 978 cited above is somewhat intermediate in spore characters but not in frond shape

Pichi Sermolli (who examined the type) gives a number of characters by which he claims C nivalis differs from typical C fragilis including stouter rhizome slightly thicker rhizome scales mixed with hairs and all with red-castaneous pyriform glands similar hairs occur on the rachis costae and surface of the lamina the indusia are wide broadly cyathiform attached around one-half of the circumference of the receptacle coarsely and irregularly dentate-fimbriate dilacerate when sorus is mature not glandular spores with bacula and spinulae much more spaced and shorter (about one third) 17ndash25 micro long the bacula are more numerous than the spinulae and the latter are often hooked Some hairs can be found on all Cystopteris laminae but the rhizome scales of the Kilimanjaro material cited above do not end in glands Pichi Sermolli suggested that C nivalis would occur on other mountains Fraser-Jenkins (op cit) treats C nivalis as a synonym of C fragilis subsp fragilis

Cystopteris 9

3 ATHYRIUM

Roth Tent Fl Germ 331 58 (1799) Kato amp Kramer in Kubitzki Fam Gen Vasc Pl 1132 (1990)

Rhizome branched creeping or erect often plusmnfleshy with dense non-peltate scales Fronds tufted or closely spaced Stipe sulcate often pink Lamina 1ndash3-pinnate to 4-pinnatifid rarely simple (not in Africa) mostly glabrous veins free (save in one non-African species) Sori superficial mostly J-shaped or U-shaped (but round or oblong in many European species) and furthest end crossing the vein to which the sorus is attached indusium present and fimbriate or rudimentary or even lacking (not in Africa)

About 180 species cosmopolitan but mostly in the north temperate zone 1 Lamina essentially trifid the apical part pinnate with 12 or more pairs of deeply

pinnatifid pinnae 2

Lamina not as above regularly 2ndash3-pinnate 3

2 Lamina appearing plusmn5-partite since lowest basiscopic pinna of lateral parts very well developed apices of all parts distinctly narrowly attenuate

4 A lewalleanum

Lamina strictly tripinnate the lateral parts without well-developed basiscopic pinnae apices of all parts not narrowly attenuate

5 A rondoense

3 Fronds 16ndash20 cm tall with pinnae up to 25 cm long pinnules oblong-ovate plusmn7 mm long with broadly rounded lobes the stomata beneath appear as dense slightly paler minute spots (T7)

3 A sp

Fronds not as above usually much larger 4

4 Rhizome creeping with fronds closely spaced lamina 2-pinnate never truly 3-pinnate and even when pinnules very deeply pinnatifid the lobes are decurrent and axis winged lowest pair of pinnae often somewhat distant and much reduced (often not so noticeable in Southern African material)

1 A schimperi

Rhizome erect with fronds tufted lamina 2ndash3-pinnate lowest pair of pinnae only slightly reduced

2 A scandicinum

1 A schimperi Feacutee Meacutem Fougegraveres 5 Gen Fil 187 (1852) VE 223 (1908) Sim Ferns S Afr ed 2133 t 41 (1915) Tardieu Meacutem IFAN 28161 t 30 fig 3 4 (sphalm 5 6) (1953) Alston Ferns WTA 64 (1959) Tardieu Fl Cameroun 3229 t 35 fig 3 4 (1964) Schelpe FZPterid 202 t 57b (1970) amp CFA Pterid 162 (1977) Kornaś Distr Ecol Pterid Zambia 105 (1979) WJacobsen Ferns SAfr 404 fig 303 (1983) Pic Serm in BJBB 55155 (1983) Schelpe amp NCAnthony FSA Pterid 223 fig 75 (1986) JBurrows S Afr Ferns 273 t 461 fig 65278 278a (1970) Schippers in Fern Gaz 14 (6) 205 (1993) Faden in UKWF ed 233 (1994) Type Ethiopia Debra Eski ldquo3000 mrdquo Schimper 239 (B syn K photo US iso)

Rhizome creeping 6 mm in diameter with reddish brown lanceolate and acuminate scales 7 mm long 1 mm wide but often some much narrower Fronds closely spaced up

to plusmn 1 m tall stipe dark brown at base 14ndash38 cm long glabrous but with scales similar to rhizome scales at base Lamina oblong-lanceolate in outline 23ndash70 cm long (6ndash)10ndash28 cm wide acute to acuminate virtually to quite bipinnate the pinnules mostly narrowly decurrent pinnae in 15ndash20 pairs lanceolate in outline 4ndash18 cm long 15ndash5 cm wide attenuate the basal pair usually markedly smaller and

FIG 3 ATHYRIUM SCHIMPERImdash1 rhizome stipe and lower part of laminatimes⅔ 2 pinna times⅔ 3 lower surface of pinnule showing soritimes4 All from Chase 3774 From FZ Drawn by Lura Mason Ripley

Athyrium 11

distant rhachis straw-coloured pinnules in 7ndash18 pairs narrowly oblong 5ndash22 mm long 2ndash8 mm wide sharply serrate-dentate or deeply lobed and lobes serrate Sori oblong curved or J-shaped 1ndash15 mm long 1ndash7 per pinnule-lobe and 14ndash70 per pinnule indusia pale brown membranous erose Fig 3 (page 9)

UGANDA Karamoja District Mt Kadam Apr 1959 Wilson 788 KENYA Trans-Nzoia District Mt Elgon Suam Saw Mill track edge of Kiptogot

River Valley 12 June 1971 Faden et al 71452 amp Kiptogot waterfall Aug 1959 Tweedie 1882 amp Oct 1961 Tweedie 1882A

TANZANIA Ufipa District Nsanga Mt Malonje Plateau 13 Mar 1959 Richards 12119 Mbeya District Mbeya Range Nov 1957 Watermeyer 12 Rungwe District Kyimbila Mbaka 13 Apr 1912 Stolz 1220

DISTR U 1 K 3 T 1 (see note) 4 7 Ghana Nigeria Cameroon Congo (Kinshasa) Rwanda Burundi Sudan Ethiopia Angola Zambia Malawi Zimbabwe and eastern S Africa also India (Himalayas)

HAB Rocky forest margins with Olinia and Juniperus crevices of rock outcrops in shade 2100ndash2400 m

SYN Asplenium schimperi (Feacutee) ABraun in Schweinf Beitr Fl Aethiop 1224 (1867)

NOTE Many specimens lack rhizomes The record from U 1 needs confirming from material with a rhizome Gillman 363 from T 1 (Bukoba District Kaagya) also without a rhizome is probably this species

2 A scandicinum (Willd) Presl Tent Pterid 98 (1836) amp in Abh Koumlnigl Boumlhm Ges Wiss ser 4 598 (1837) Tardieu Fl Madag 5 (1) 267 fig 369ndash11 (1958) (as lsquoscandinicumrsquo) Type Reacuteunion Bory de St Vincent sn (B-W 19832 holo microfiche)

Rhizome erect plusmnfleshy 5ndash8 mm wide but the branches forming a caudex up to 25 cm wide rhizome scales brown lanceolate-attenuate 7ndash10 mm long 1ndash2 mm wide Fronds tufted 015ndash2 m tall stipe usually reddish or pink when living 75ndash55 cm tall glabrous save for scales at base but young stipes of undeveloped fronds often scaly throughout Lamina ovate-lanceolate in outline 20ndash50 cm long 10ndash34 cm wide acute to acuminate-attenuate at the apex 2ndash3-pinnate with plusmn18 pairs of pinnae the basal pair somewhat reduced pinnae lanceolate in outline acuminate to caudate at the apex 15ndash20 cm long 08ndash8 cm wide with 10ndash20 pairs of pinnules pinnules oblong-ovate to oblong-lanceolate 1ndash5 cm long 03ndash2 cm wide sometimes cut to the base into secondary pinnules the lamina then distinctly 3-pinnate but often only 2-pmnate-pinnatifid the pinnules crenate to divided into narrow lobes pinnule-lobes or secondary pinnules 3ndash10 mm long 1ndash7 mm wide soft spinules up to 1 mm long often present where wings of rhachis or rhachilla are interrupted by pinnule-bases and some short indumentum sometimes present Sori oblong curved or J-shaped up to 2 mm long 5ndash7 per pinnule-lobe or secondary pinnule indusium shortly to deeply fimbriate or subentire (fide Schelpe)

SYN Aspidium scandicinum Willd Sp Pl ed 4 5285 (1810) NOTE I agree with Pichi Sermolli that African material is certainly not identical with

typical material from Reunion which has much more delicately cut fronds with the pinnule-lobes much smaller I have therefore divided it into two subspecies admittedly not too well defined

subsp newtonii (Bak) Verdc comb nov Type Satildeo Tomeacute Newton 2 amp Quintas 9 (K syn COI isosyn)


It is clear from Bakerrsquos description that he saw both sheets although he merely cites lsquoIsland of St Thomas Newtonrsquo Exell (Cat Vasc Pl STomeacute 73 (1944)) gives Newton 2 and Newton 88 as being at Kew but there is no Newton 88 it is Quintas 9 collected on lsquo688rsquo (June 1888] I am certain Baker had also misread it as Newton and am treating the two sheets as syntypes Exell adds ldquoalso from Ruwenzorirdquo

Pinnule-lobes or secondary pinnules ovate or oblong typically shortly and obtusely crenate

UGANDA Toro District Ruwenzori Mijusi Valley 31 Mar 1948 Hedberg 615 amp Mihunga 14 Jan 1939 Loveridge 365 Ankole District Kalinzu Forest 4 km NW of Saw Mill W of Rubuzigye 19 Sep 1969 Faden et al 691140

KENYA Aberdare Mts Ramsden sn Fort Hall District Kyama R below Mbugiti School 13 July 1969 Faden amp Evans 69890 Kericho District 8 km NW of Kericho Kimugung R 10 June 1972 Faden et al 72292

TANZANIA Moshi District Kilimanjaro ravine E of Maundi Crater 12 Oct 1993 Grimshaw 93783 Lushoto District W Usambaras Mangula 18 Sep 1981 Mtui amp Sigara 64 Iringa District Kilombero Mwanihana Forest Reserve above Sanje village 10 Oct 1984 DW Thomas 3864

DISTR U 2 4 K 3ndash5 T 2ndash4 6 7 Satildeo Tomeacute Sudan Malawi Mozambique Zimbabwe Swaziland South Africa (the Madagascan material probably belongs here)

HAB Forest eg Parinari excelsa PolysdasndashMacarangandashFagarandashAlbiziandashNeoboutonia extending into ericaceous belt often by streams on marshy ground 1150ndash3500 m

SYN Athyrium newtonii Bak in Ann Bot 5307 (1891) Exell Cat Vasc Pl S Tomeacute 73 (1944) Schippers in Fern Gaz 14205 (1993)

A laxum Pappe amp Raws Syn Fil Afr Austr 16 (1859) non Schumach Type South Africa Natal Gueinzius sn (ubi)

[Athyrium scandicinum sensu Hieron in POA C 84 (1895) VE 23 (1908) Sims Ferns S Afr ed 2133 t 42 (1915) Schelpe FZPterid 204 (1970) Schelpe amp Diniz Fl Moccedilamb Pterid 220 (1979) WJacobsen Ferns S Afr 404 (1983) Pic Serm in BJBB 55154 (1985) Schelpe amp NCAnthony FSAPterid 223 (1986) Schippers in Fern Gaz 14205 (1993) Faden in UKWF ed 233 (1994) pro parte non (Willd) Presl sensu stricto]

Athyrium scandicinum (Willd) Presl var scandicinum sensu Schelpe in FZPterid 204 (1970) Schelpe amp Diniz Fl Moccedilamb 22 (1979) Jacobsen Ferns S Afr 405 fig 304a 304b (1983) Burrows S Afr Ferns 273 t 456 fig 65279 279a 279b (1990)

NOTE RB amp AJ Faden 74412 (Morogoro District Uluguru Mts Bondwa summit 2120 m in PodocarpusndashAllanblackiandashPolysdasndashBalthasariandashSyzygium forest) has erect rhizomes but the frond architecture is similar to that of A schimperi Attempts to divide the mainland subspecies further have been unsuccessful It was at first thought that the southern African population might form a taxon equivalent to A laxum but the variation overlaps too much with that of material further north to make it practical to do so Clair Thompson mentions (on a label) it can be epiphytic but there is no confirmation of this

Schelpe described A scandicinum var rhodesianum (Bol Soc Brot Seacuter 2 41211 (1967) FZPterid 204 t 57 fig a (1970) Type Zimbabwe Pungwe Gorge Schelpe 5722 (BOL holo BM iso)) but it seems doubtfully different and he makes no mention

Athyrium 13

of it in the FSAPterid account Burrows (op cit) and Jacobsen (op cit) keep it distinct

Without really adequate rhizome data A schimperi and A scandicinum can be difficult to distinguish particularly in southern Africa and some field notes mention creeping rhizomes yet say fronds are tufted

3 A sp Rhizome erect with spreading roots scales chestnut lanceolate 3 mm long 1 mm

wide attenuate Fronds tufted 16ndash20 cm tall stipe 5ndash6 cm long scaly at base Lamina oblong-lanceolate in outline 9ndash12 cm long 4ndash5 cm wide with 10ndash15 pairs of pinnae pinnae oblong in outline 1ndash25 cm long 03ndash13 cm wide subacute and not attenuate at apex with 4ndash5 pairs of pinnules and short apical lobed part pinnules oblong-ovate up to about 7 mm long 5 mm wide 3ndash5-lobed and plusmncrenate with sparse flattened trichomes beneath Sori one per pinnule about 1 mm long indusia deeply fimbriate

TANZANIA Iringa District Luhomero Massif South Msisimwana ridge 19 Aug 1985 Rodgers amp Hall 4558

DISTR T 7 HAB Rock outcrop 2300 m NOTE It was suggested that this was a young plant of A scandicinum but it is fertile

No other material has been seen which resembles it and until more is collected in the same area its status is uncertain

4 A lewalleanum Pic Serm in Webbia 27440 f 18 (1973) (lsquo1972rsquo) Type Burundi Bubana road towards Mabaye RNyamagana Lewalle 3433 (Herb Pic Ser 24934 holo Herb Lewalle iso)

Rhizome long-creeping slender scales pale brown or dark-marked triangular 02ndash12 mm long 03ndash04 mm wide acuminate cordate at the base mixed with multicellular hairs Fronds few paired or solitary 15ndash40 cm tall stipe 12ndash245 cm long minutely pubescent or plusmn glabrous or with few scales at base Lamina plusmn triangular-hastate in outline 115ndash22 cm long 12ndash26 cm wide with scattered flattened white multicellular hairs essentially trifid the apical part pinnate with 12 or more pairs of pinnae pinnae lanceolate up to 5 cm long 18 cm wide attenuate at the apex deeply pinnatifid with plusmn10 pairs of pinnatifid segments about 2ndash7 mm wide lateral parts similar to apical but with lowest basiscopic pinna well developed up to 10 cm long rachis shortly pubescent Sori round plusmn1 mm in diameter 1ndash10 per segment indusium sub-reniform fixed on one side only persistent or plusmn disintegrating and scarcely visible

TANZANIA Kigoma District Kasye Forest 18 Mar 1994 Bidgood et al 2788 amp same locality 19 Mar 1994 Bidgood et al 2806

DISTR T 4 Burundi Zambia (see note) HAB Evergreen forest along streams and rivers and on valley slopes Baikiaeandash

Julbernardiandash MonopetalanthusndashParkiandashTessmanniandashMaesopsisndashPycnanthus 900 m NOTE Kornaś (in KB 33101 (1978)) has described a very similar plant A annae

(Type Zambia Kalungwishi R gorge below Lumangwe Falls Kornaś amp MedweckandashKornaś 3746 (KRA holo K Herb Pic Serm iso) differing in a number of small points I suspect it is conspecific Frazer-Jenkins has annotated the Kew isotype suggesting the centro-basally attached indusium is wrong for Athyrium and that the species might be a Ctenitis or Hypodematium but it does not seem to fit well in either


5 A rondoense Verdc sp nov affinis A lewalleani Pic Serm lamina stricte tripartita pinnis basiscopicis partium lateralium haud anguste caudatis pinnis pinnulis lobisque pinnularum majoribus differt Typus Tanzania Rondo Plateau Bidgood et al 1550 (K holo BR C DSM EA MO NHT P WAG iso)

Rhizome creeping with blackish or black-veined clathrate scales plusmn15 mm long fronds single 15ndash40 cm tall stipe slender 10ndash20 cm long with few scales and dense very short hairs Lamina up to 20 cm long 15 cm wide tripartite the parts plusmn confocal each simply pinnate but pinnae very deeply divided so as to appear bipinnate main parts broadly lanceolate in outline up to 18 cm long 6 cm wide acuminate but not long-caudate at the apex each part with 6ndash9 pairs of pinnae up to plusmn5 cm long 2 cm wide the apical ones reduced toothed only at the apex running together and narrowly decurrent into each other resembling a series of fish-tails lower pinnae deeply divided into plusmn6 elliptic lobes the largest plusmn15 mm long 8 mm wide plusmnentire to bluntly or subacutely lobed or toothed main lobes of largest pinnae decurrent separated by narrowly winged midrib 1ndash2 mm wide Sori 1ndash4 per pinnalobe plusmn1 mm wide indusium fimbriate and hairy

TANZANIA Lindi District Rondo Plateau Rondo Forest Reserve 14 Feb 1991 Bidgood et al 1550

DISTR T 8 only known from the type HAB Evergreen forest of MiliciandashAlbiziandashDialium in small gully and amphitheatre

around well on escarpment 650 m NOTE This had been confused with Arachniodes foliosa (C Chr) Schelpe

(Dryopteridaceae) and is superficially similar but the pinnule-lobes are not aristate-dentate

Athyrium 15

4 CALLIPTERIS

Bory Voy Quatre princip Iles Afrique 1282 (1804) Pacheco amp Moran in Brittonia 51343ndash388 (1999)

Rhizome erect stout usually mucilaginous scaly scales brown eventually with dark brown to black edges with marginal trichomes bifid Fronds large Stipe often tuberculate to spinulose scaly at the base Lamina 1ndash2-pinnate (entire in one S American species) thin to fleshy the distal portion often only pinnatifid often proliferous lateral veins parallel with lateral veinlets anastomosing regularly with those of adjacent groups or free (see note below) Sori elongate along the veinlets numerous often in U-shaped pairs indusium elongate usually membranous

Genus often restricted to 3 species in Africa to Malesia Australia and Polynesia but as redefined by Pacheco amp Moran (op cit) with many species also in the Neotropics and others in the Old World which need transferring to the genus which may then contain over 30 species I am not completely convinced of this so far as the Old World species are concerned

Pacheco amp Moran (op cit) have revised some of the New World species They define Callipteris by its rhizome scales having dark chestnut to black margins and bifid marginal teeth not all species have anastomosing veins but the type does and the group of 15 species they have revised They further indicate that several old world species apart from the type should be transferred to Callipteris I have found difficulties with this however apart from the fact that most specimens including nearly all types do not have rhizomes preserved there is variation in some species where trichomes can be minutely bifid or uniformly undivided and scales are not black-margined They mention that Bory de Saint-Vincentrsquos original conception of the genus contained only species with anastomosing veins but this is not so Fronds simply pinnate veins anastomosing 1 C prolifera

Fronds bipinnate veins not anastomosing 2 C ulugurica

1 C prolifera (Lam) Bory Voy Quatre princip Iles Afrique 1283 (1804) DL Jones in Fl Australia 48422 figs 88 139dndashe Type La Reunion Commerson sn (PndashLAM syn microfiche 7423ndash4 BM photo Morton Neg 2765)

Rhizome ascending to erect fleshy scales pale brown lanceolate 7ndash10 mm long attenuate 1ndash2 mm wide at base pale to eventually black at margins and with stiff bifid trichomes Fronds tufted 15ndash21 m tall top of stipes and rachis sometimes with short stiff spinules or in some forms strongly muricate but usually plusmnsmooth in East Africa bulbils in upper 3 pinna axils and sometimes below in 7th producing young plants in situ stipe 15ndash90 cm tall Lamina oblong-lanceolate 15ndash120 cm long 26ndash44 cm wide with terminal segments 10ndash21 cm long 7ndash10 cm wide with teeth or lobes up to 25 cm long 15 cm wide pinnae alternate oblong 65ndash24 cm long 3ndash65 cm wide tapering at apex truncate emarginate or subcordate (in young plants) at the base the lowest the smallest

coarsely toothed or crenate the teeth or crenae 4times6 mm venation anastomosing Sori brown numerous narrow appearing like contiguous branched trees from each side of the midribs of the pinnae in each area bounded by a lobe with 7ndash8 pairs of sori per tree the whole gives a series of parallel zig-zag lines along the length of the pinna indusia linear Fig 4 (page 14)

UGANDA Bunyoro District Budongo Forest Reserve nature reserve close to Sonso R 31 Dec 1995 Poulsen et al 910 Toro District Bwamba Kabanga 21 Nov 1935 ASThomas 1499 amp Bwamba Forest Reserve 28 July 1960 Paulo 614

TANZANIA Lushoto District Amani July 1913 Grote in AH 5792 amp Kwamgumi Forest Reserve NW of Mhinduro peak 11 Nov 1986 Farkas amp Poacutecs 86606 amp Mwesini [Mwezi] to Wugu [Iwugu] near Bwiti June 1917 Peter 20487

DISTR U 2 4 T 3 Guinea to Nigeria Cameroon Satildeo Tomeacute Principe Bioko Gabon Congo (Kinshasa) Sudan Angola Mascarene Is Malesia Queensland Polynesia

HAB Riverine forest of Cynometra alexandri Khaya anthotheca etc swamps with Raphia sometimes in shallow water rain-forest rock-faces (in Usambaras) 400ndash1200 m

Callipteris 17

FIG 4 CALLIPTERIS PROLIFERAmdash1 rhizometimes⅔ 2 frond 3 part of frond with bulbils enlarged 4 plantlettimes⅔ 5 scaletimes8 6 hairs on scale margin enlarged 1 from Cheek 5864 2 5 6 from Poulsen 910 3 from Leeuwenberg 2336 4 from Paylo 614 Drawn by Pat Halliday


SYN Asplenium proliferum Lam Encycl Meacuteth 2807 (1786) Hieron in POA C 83 (1895)

A decussatum Sw in J Bot (Schrad) 1800 (2) 51 (1800) amp in Syn Fil 76 260 (1806) Willd Sp Pl ed 5 5310 (1810) Type Mauritius coll ignot ( S B-W 19877 iso microfiche)

Diplazium proliferum (Lam) Kaulf Enum Fil 182 (1824) Hieron in VE 224 fig 20dndashf (1908) Bonap Notes Pteacuterid 178 (1915) Alston J Bot 72 Suppl Pterid 5 (1934) amp Ferns WTA 65 (1959) Schelpe CFA Pterid 163 (1972) Johns in Kew Mag 8(3) 128 t 178 fig 5 (1991) Schippers in Fern Gaz 14205 (1993)

D incisum Schum Beskr Guin Pl 458 (1827) amp in KDanske Vid Selsk Nat Mat Afh 4232 (1829) Type Ghana Akwapim Isert sn Thonning sn (C syn)

D stratum Schum Beskr Guin Pl 459 (1827) amp in K Danske Vid Selsk Nat Mat Afh 4233 (1829) Type Ghana Akwapim Thonning sn (C holo)

Athyrium proliferum (Lam) Milde in Bot Zeit 1870353 (1870) Tardieu Meacutem IFAN 28163 t 31 figs 3ndash4 (1953)

Diplazium decussatum sensu Peter FD-OA 167 (1929) (Peter attributes this to (Sw) J Sm in Bot Mag 72 comp 28 (1846) but Smith does not mention Swartz but a Wallich specimen from Nepal)

NOTE The species has an extensive synonymy I have excluded some specimens from the description which appear to be different eg Ceylon material with pinnae 33ndash42times8ndash11 cm the lowest part with lobes cut almost to the midrib 45ndash7times15ndash25 cm

2 C ulugurica Verdc sp nov Diplazio arborescenti (Bory) Sw similis squamis rhizomatis nigromarginatis trichomatibus marginalibus nigris distincte apice bifurcatis a C prolifera (Lam) Bory frondis bipinnatis pinnulis pinnarum inferiorum usque 25 cm longis 22 cm latis venatione haud anastomosanti differt Typus Tanzania Morogoro District NE Uluguru Mts Kinole Faden et al 70681 (EA holo K EA iso)

Rhizome erect scales with black margins linear-lanceolate 16ndash24 cm long 08ndash2 mm wide at the base very attenuate dull the marginal trichomes black and nearly all bifid Fronds tufted 05ndash15 m tall somewhat fleshy stipe straw-coloured 40ndash45 cm long densely scaly near base or for some distance Lamina oblong-lanceolate mostly bipinnate 21ndash60 cm long 32(ndash60) cm wide the apical part 7ndash11 cm long 55ndash6 cm wide long-attenuate divided into oblong lobes 1ndash3 cm long 06ndash1 cm wide crenate upper pinnae in 6ndash7 pairs narrowly oblong-triangular 45ndash13 cm long 12ndash33 cm wide plusmntruncate at base narrowly attenuate and crenate at the apex the upper ones sessile shallowly lobed the lower stalked with up to 12 pairs of oblong lobes up to 2 cm long 09 cm wide lower pinnae in 4 pairs oblong-attenuate 13ndash28 cm long 6ndash11 cm wide with (3ndash)8ndash9(ndash11) pairs of pinnules up to 54ndash75 cm long 14ndash22 cm wide very shallowly lobed or crenate pinnae apices similar to frond apices venation not anastomosing Sori brown linear curved 4ndash7 mm long diverging from midrib appearing different according to the rank of the segment involved in the lobes of apices of fronds and pinnae and the ultimate pinnules there are 2ndash7 pairs of sori in the upper pinnae where lobes are not divided to the base the lobes have 1ndash3 pairs according to position the main sori are on veins ending in sinuses of the marginal divisions but occasionally there are much shorter sori on lateral veins capitate paraphysesplusmn numerous often with dark heads indusia very narrow unilaterally attached

Callipteris 19

TANZANIA Morogoro District NE Uluguru Mts Kinole 28 Sept 1970 Faden et al 70681 amp 30 Apr 1970 Poacutecs amp Harris 6165P amp E slope of Lupanga 11 Oct 1971 Poacutecs amp Mwanjabe 6470A Ulanga District SW of Mahenge Muhulu Mts 24 Feb 1932 Schlieben 1829

DISTR T 6 not known elsewhere HAB Intermediate rain-forest with Allanblackia Parinari Cylicomorpha

Myrianthus 900ndash1050 (ndash1500) m

Sometimes attributed to Thouars in Fl Trist drsquo Ac 35 (1804) but I can find no mention in that work The specimens are confusingmdashsee Hepper W Afr Herb Isert amp Thonning 160 (1976)

SYN Diplazium sp B Schippers in Fern Gaz 14205 (1993) NOTE Poacutecs amp Chambuko 6223X (N Uluguru Mts below Manga peak 16 Aug

1970) is the same but many of the trichomes are not bifid Poacutecs et al 87201N (Ulanga District Mahenge Plateau Mawenge Forest Reserve above Isongo village 10 Oct 1987 at 1150ndash1250 m) is probably also the same but I have not seen the rhizome and the lamina architecture differs The specimen seen is not adequate but a main pinna appears to be plusmn40 cm long with at least 10 pairs of pinnules about 10times25 cm divided for ⅔ndashfrac34 distance to costa into plusmn12 crenate lobes each with 8 sori in 4 angled pairs Black-headed paraphyses are numerous More material is needed


5 DIPLAZIUM

Sw in J Bot (Schrad) 1800 (2) 61 (1801) Kato amp Kramer in Kubitzki Fam Gen Vasc Pl 1133 (1990)

Mostly rather large ferns with creeping or erect rhizome sometimes forming a trunk-like caudex up to 1 m tall scales non-peltate dark entire or with spine-like or tooth-like emergences Lamina simple or 1ndash4-pinnate veins free Sori superficial elliptic to linear and elongate the indusium either attached along one side or all round or where a pair of sori are on either side of a vein the linear indusia are set back to back indusium rarely small and fugacious

Pantropical with about 370 species The species are difficult and this account is no more than a framework for further work It is very important to assess characters from the same part of the frond architecture which is sometimes difficult to ascertain in poor herbarium specimens when the apex of a frond can be mistaken for a pinna and undeveloped pinnae for true pinnules when the junction of the pinna with the main rhachis is preserved there is no difficulty The rhizome should always be collected or at least a portion showing scales 1 Fronds simply pinnate pinnae 4ndash16 cm long 08ndash18 cm wide crenate-

toothed to shallowly squarely pinnatifid (T 6 7) 1 D pseudoporrectum

Fronds bipinnate or almost tripinnate pinnae distinctly divided into pinnules

2

2 Fronds with gemmae at andor below apex 3

Fronds without gemmae 4

3 Pinnae 35ndash42times14ndash18 cm pinnules 4ndash10times12ndash5 cm with strongly pinnatifid lobes 08ndash25 cmtimes3ndash7 mm (U 2 T 4)

6 D humbertii

Pinnae up to 25times10 cm pinnules 15ndash5times07ndash17 cm with very shallowly crenate almost subentire lobes 5ndash8times5ndash6 mm (T 6)

5 D ulugurense

4 Indusium narrow attached to one side of sorus lsquoasplenioidrsquo sometimes difficult to see

5

Indusium wider attached all around the sorus lsquoallantodioidrsquo bursting irregularly across top or sometimes along one edge often leaving fragments all around the sorus but very obvious before dehiscence

6

Contiguous sori on either side of a usually basal vein have their indusia back to back and this can resemble a single sorus appearing to have a centrally split indusium) 5 Sori 3ndash4(ndash5) mm long usually 1ndash6 per pinnule-lobe (see note after species)

often only 1 (on lowest acroscopic vein of the venation in each pinnule lobe) pinnules shallowly to deeply cut but lamina not appearing tripinnate pinnule-lobes mostly crenulate only at the broadly rounded apex rhizome-scales 2 D nemorale

broader and shorter 9times2ndash5 mm paraphyses present

Sori 1ndash3(ndash4 on basal vein) mm long 7ndash(11ndash23) per pinnule-lobe (see note after generic description) pinnules very deeply cut so that lamina appears almost tripinnate pinnule-lobes crenate-lobed to serrate rhizome-scales long and narrow up to 32 cm long and 05ndash15(ndash2) mm wide paraphyses absent

3 D zanzibaricum

6 Lamina bipinnate with pinnules divided into 6ndash9 pairs of lobes for only frac12 to ⅔ of width not cut to near base and lobes plusmnentire all round or toothed or crenate at apex paraphyses short and few (K 5 Kakamega Forest) 8 D sp B

Lamina virtually tripinnate with pinnules divided into 10ndash15 pairs of lobes almost or quite cut to the base but still decurrent and not entirely free lobes mostly deeply crenate all round paraphyses mostly obvious and numerous (U 2 K 5 T 4 including Kakamega Forest) (forms of D humbertii without gemmae will key heremdashthey are even closer to being tripinnate with the lobes more separate as well as more deeply cut)

7 D velaminosum

1 D pseudoporrectum Hieron in EJ 28342 (1900) amp in VE 224 (1908) FD-OA 167 (1929) Types Tanzania Morogoro District SE Uluguru Mts Nghweme [Ngrsquoheme] Stuhlmann 8796 8810 (B syn)

Rhizome erect or ascending woody scales brown lanceolate 5ndash7 mm long 12ndash2 mm wide reticulate the cell walls darker and strongly raised margins with short blunt emergences Fronds tufted 50ndash90 cm tall simply pinnate stipe 22ndash40 cm long scaly at base Lamina oblong-lanceolate 33ndash68 cm long 105ndash29 cm wide with 14ndash31 pairs of pinnae and apical triangular segments 85ndash12 cm long 4 cm wide toothed at apex and pinnatifid beneath with plusmn5 lobes on each side pinnae stipitate plusmnfalcate narrowly oblong-lanceolate 4ndash16 cm long 08ndash18 cm wide forwardly directed crenate-toothed to shallowly squarely pinnatifid the lobes plusmn crenate up to 30 lobes on each side and pinnule apex narrowly attenuate crenulate stipes of pinnae 4ndash5 mm long Sori 1ndash3 on each side of pinna costa for each lobe diverging unequal the longer collateral pair 6ndash8 mm long the shorter 3 mm long indusium elongate Fig 5 (page 18)

TANZANIA Kilosa District Ukaguru Mts NNE slope of Mamiwa ridge below Mnyera peak 30 July 1972 Poacutecs amp Mabberley 6739A Morogoro District Uluguru Mts Morogoro to Lupanga Peak track 16 Aug 1951 Greenway amp Eggeling 8615 Iringa District Mwanihana Forest Reserve above Sanje village 10 Oct 1984 DWThomas 3857

See note after generic description A specimen from the Sese Is swamp forests will key here but has a more developed indusiummdashsee species 4 D sp A A fragment of type material is preserved at the BM (CChristensen Herb 4889) but it is not stated from which Stuhlmann sheet it came


FIG 5 DIPLAZIUM PSEUDOPORRECTUMmdash1 rhizometimes⅔ 2 frond showing both surfaces times⅓ 3 pinnatimes1 4 venation enlarged and diagrammatic 5 indusium enlarged and diagrammatic All from Faden 74386 Drawn by Pat Halliday

Diplazium 23

DISTR T 6 7 not known elsewhere (see note) HAB Montane forest including mist forest AllanblackiandashCussoniandashDasylepisndash

Ocotea CyatheandashOcoteandashPodocarpus and ParinarindashNewtoniandashOcoteandashMacaranga 1400ndash2100 m

SYN D pseudoporrectum Hieron var angustipinnatum Reimers in NBGB 11922 (1933) Schippers in Fern Gaz 14205 (1933) Type Tanzania Morogoro District NW Uluguru Mts Schlieben 3108 (B holo BM iso)

NOTE In Schlieben 3108 all the pinnae have retained the very shallowly toothed margins of the uppermost pinnae but new material shows variations in this respect Some sheets of the species had been determined as D silvaticum (Bory) Sw and some Burmese specimens of that are very similar It would be foolish to make suggestions without a revision of the Old World species

2 D nemorale (Bak) Schelpe in Bol Soc Brot seacuter 2 41212 (1967) amp in FZ Pterid 205 t 58a (1970) Schelpe amp Diniz Fl Moccedilamb Pterid 221 (1979) W Jacobsen Ferns S Afr 408 t 306 (1983) J Burrows S Afr Ferns 278 t 463 fig 66282 282a (1990) Faden in UKWF ed 233 (1994) Type Madagascar Antananarivo Pool sn (K holo)

Rhizome erect or ascending forming caudex up to 20(ndash30) cm long 7ndash20 cm in diameter scales shiny bronze-brown 9 mm long 2ndash5 mm wide the edges not or narrowly dark and with few undivided trichomes Fronds tufted 05ndash18 m tall stipe brown 18ndash90 cm long grooved above swollen and slightly scaly at the base Lamina plusmnovate 45ndash95 cm long 30ndash70 cm wide bipinnate-pinnatifid apical segment 7ndash12 cm long 35ndash5 cm wide attenuate and crenate at the tip lobed beneath pinnae in 10ndash15 pairs the plusmn 5 below the apical segment pinnatifid 9 cm long 2 cm wide the lobes crenate rest of pinnae oblong to oblong-ovate pinnate 20ndash50 cm long 35ndash22 cm wide with (7ndash)10ndash16 pairs of triangular-lanceolate pinnules 25ndash115 cm long 1ndash27 cm wide shallowly to very deeply pinnatifid according to position narrowly acuminate and crenulate at the apex pinna-stalks 0ndash15(ndash3) cm long venation not anastomosing Sori brown linear 3ndash4(ndash5) mm long usually one on the lowest acroscopic vein of the branching system in each pinnule-lobe but there are usually other shorter ones present 1ndash6(ndash9) in the basal lobes but usually only 1 in the upper lobes the appearance of the son varies according to which part of the frond is examined paraphyses present indusium very narrow unilaterally attached

KENYA Embu District SE Mt Kenya Irangi Forest Station 30 Apr 1972 Faden et al 72193

TANZANIA Lushoto District Shagayu Forest valley N of Shagein along tributary to Umba R 21 Oct 1986 Schippers 1589 Morogoro District Uluguru Mts Ululu Falls near Bunduki fishing camp 27 Nov 1974 Balslev 356 Iringa District Mwanihana Forest Reserve above Sanje Village 10 Oct 1984 DWThomas 3863

DISTR K 4 T 3 5ndash7 Satildeo Tomeacute Malawi Mozambique E Zimbabwe HAB Montane and lower montane forest eg OcoteandashEnsetendashDracaenandashCyatheandash

Parinarindash StrombosiandashSyzygiumndashMyrianthusndashFicalhoa in swampy places and on steep slopes (1100ndash)1350ndash1900m

SYN Asplenium nemorale Bak in JLS 15417 (1876) A hylophilum Hieron in POA C 84 (1895) Type Tanzania Lushoto District W

Usambaras Mbaramu Holst 2480 (B holo BM fragment amp photo K iso) (see note)


Diplazium hylophilum (Hieron) C Chr Index Fil 233 (1905) Faden in UKWF ed 157 (1974) Schippers in Fern Gaz 14205 (1993)

D stolzii Brause in EJ 53381 (1915) Type Tanzania Rungwe District Rungwe Stolz 1233 (B holo BM fragment amp photo K iso)

D arborescens (Bory) Sw var nemorale (Bak) C Chr in Perrier Cat Fl Madag Pteacuterid 36 (1932)

Schippers gives the type as Stuhlmann 8796 but this is not correct It is difficult to sort out the exact ecology of the collected material from field notes which give general ranges for a large area

NOTE I at first agreed with Christensen that D nemorale was not more than a local variety of D arborescens and that D hylophilum could be looked on as another variety but since the situation is not resolved I have not disturbed Schelpersquos treatment of the species

Two sheets of the eight making up the EA gathering of Faden et al 72193 are a young plant with the largest pinnules only 25times1 cm and not deeply pinnatifid The isotype of D hylophilum preserved at Kew consists of the top of a frond with similar small pinnules so I believe it has been interpreted correctly but mature specimens with rhizomes are needed from the Usambaras The type of D nemorale consists of a single pinna and it does not seem certain it is a pinna with pinnatifid pinnules rather than the apex of a frond with pinnatifid pinnae The somewhat cordate base of the pinnules is a minor variation not matched elsewhere D arborescens (Bory) Sw Syn Fil 92 (1806) Willd Sp Pl ed 5 5354 (1810) (Type Reunion by R St Denis Bory de St Vincent sn (P holo B-W 19950 iso (microfiche)) differs in having 6ndash15 sori in all the pinnules and they are joined basally in pairs at the mid vein Material from the Comoro Is is particularly characteristic There are however specimens from Zimbabwe which are deceptively similar eg BSFisher 1331 from Umtali Vumba 18 July 1947 Bory describes it as having a big trunk A further problem is the identity of material from West Africa cited by Alston (FWTA Pterid 65 (1959))

Schippers 1130 (Iringa District Mufindi slope towards Kigogo R 1630 m 9 Nov 1985) is probably only a form of D nemorale with rather different pinna architecture and more sori a lower pinnule is 40times19 cm with largest pinnule-lobes up to 20times9 mm with up to 14 sori More material with rhizome is needed to decide its status This is D sp C of Schippers in Fern Gaz 14205 (1993)

3 D zanzibaricum (Bak) C Chr Ind Fil 241 (1905) Schelpe in FZ Pterid 205 t 58b (1970) Schelpe amp Diniz Fl Moccedilamb Pterid 222 (1979) WJacobsen Ferns SAfr 410 t 307 (1983) Schelpe amp NCAnthony in FSA Pterid 227 (1986) JEBurrows SAfrFerns 276 t 4641 fig 66281 281a (1990) Schippers in Fern Gaz 14205 (1993) Faden in UKWF ed 233 (1994) Type Tanzania mainland W of Zanzibar probably Nguru Mts Last 261 (K holo)

Rhizome large erect scales pale chestnut brown linear to linear-lanceolate 15ndash32 cm long 05ndash15(ndash2) mm wide finely attenuate not black-margined or rarely partly so (see note) with numerous stiff spiniform trichomes very narrowly striate with raised walls of the elongate reticulation trunk 04ndash12 m long 15 cm wide Fronds tufted 1ndash25 m tall stipes up to 15 m tall with 2 C-shaped bundles very densely covered with scales similar to rhizome scales at base Lamina ovate-lanceolate 13ndash16 m long 09ndash11 m

Diplazium 25

wide bipinnate with about 15 pairs of pinnae pinnae (5ndash)42ndash68 cm long (1ndash)11ndash25 cm wide stalk up to 25 cm long pinnules in 12 to 27 pairs oblong-lanceolate to oblong-triangular (1ndash)7ndash13 cm long (03ndash)15ndash4 cm wide attenuated to a crenate apex deeply cut into 13ndash15 pairs of lobes so that lamina is almost tripinnate lobes narrowly oblong 05ndash2 cm long 2ndash8 mm wide crenate-lobed to serrate including the apex costae often with multicellular plusmnserrate hairs up to 15 mm but sometimes entirely glabrous venation free the veins often pale above towards margin Sori 1ndash3 mm (ndash4 mm on basal vein) long (7ndash)11ndash23 per pinnule-lobe indusium narrow on one side only or when two sori are paired and contiguous on basal vein the two indusia are persistent between them young sori not covered with wide indusium all round paraphyses absent

UGANDA Toro District E Ruwenzori Bwamba Pass July 1940 Eggeling 4007 (see note) amp Bwamba Pass 3 Feb 1934 Longfield 39 Kigezi District Kinaba Gap Dec 1938 Chandler amp Hancock 2540

KENYA Meru District NE Mt Kenya Ithanguni Forest road along base of volcanic cone Kirui and 15 km from Nkubu 22 June 1969 Faden et al 69764 amp Nyambeni Hills above Kiegoi 1 June 1969 Faden et al 69665

TANZANIA Pare District N Pare Mts Kindoroko Forest Reserve 24 Feb 1987 Schippers 1771 Morogoro District Uluguru Mts Bondwa 2 May 1970 Poacutecs 6183E Rungwe District Rungwe Forest Tukuyu June 1958 Watermeyer 39

DISTR U 2 K 4 T 2 3 6 7 W Congo (Kinshasa) Zambia Malawi Mozambique Zimbabwe amp South Africa also recorded from Madagascar and Comoro Is

HAB Evergreen forest eg PodocarpusndashLasianthusndashPsychotriandashPauridianthandashXymalosCyathea forest bamboo forest occasionally in swampy areas 1450ndash2550 m

SYN Asplenium zanzibaricum Bak in Ann Bot 5311 (1891) FD-OA 180 (1929) Aspidium sulcinervium Hieron in POA C 85 (1895) Type Tanzania Kilimanjaro

Marangu 2300 m Volkens 1492 (B holo BM iso (pro parte)) Diplazium sulcinervium (Hieron) C Chr in Perrier Cat Pl Madag Pterid 37 (1932) D sp D Schippers in Fern Gaz 14205 (1993) NOTE In Eggeling 4007 a few areas of the scale margin are black with black

trichomes and a very few have a minute bifurcation at the apex The indusium is almost invisible and flattened into the lobe surface More material is needed

4 D sp A Scales on stipe-base black-edged with trichomes showing a trace of bifurcation

Fronds up to 12 m tall stipe 56 cm long slender Lamina narrowly triangular 60 cm long 45 cm wide with plusmn13 pairs of pinnae up to 20 cm long 8 cm wide with long crenate apex pinnules up to 15 pairs lanceolate serrate at the tip 15ndash65 cm long 04ndash18 cm wide pinnatifid for about ⅓ndashfrac34 into up to 10 pairs of oblong lobes subentire or slightly crenate 3ndash8 mm long 2ndash5 mm wide Sori linear to oblong 1ndash3 mm long 1ndash7 per lobe those nearest the pinnule costule being the longest paraphyses lacking indusium wider than in D zanzibaricum but splitting and remaining on one side of sorus with dark reticulation

UGANDA Masaka District Sese Is Towa Forest 30 June 1935 ASThomas 1350 DISTR U 4 HAB Primary forest 1200 m NOTE This is close to D zanzibaricum in the lack of paraphyses but the indusium is

more developed and the pinnule-lobes less crenate than in typical material It is the only


specimen of Diplazium seen from the Sese swamp forest and may be distinct It resembles much W African material probably misnamed D hylophilum

5 D ulugurense Verdc sp nov affinis D humbertii (C Chr) Pic Serm pinnis pinnulis et lobis ultimis pinnularum minoribus lobis ipsis magis quadratis breviter crenato-dentatis vel subintegris haud profunde pinnatifidis differt Typus Tanzania Morogoro District Uluguru Mts E slope of Bondwa Faden et al 70635 (K holo iso EA iso)

Rhizome thick and fleshy erect to ascending scales pale chestnut narrowly lanceolate 8ndash12 mm long 12ndash2 mm wide attenuate not black-margined and with few simple trichomes prominently reticulately nerved Fronds tufted 12 m tall stipe 46 cm long scaly at base Lamina triangular-ovate plusmn70 cm long 48 cm wide bipinnate gemmiferous at apices of some pinnae and probably from near apex of frond also pinnae in plusmn13 pairs oblong-lanceolate up to 25 cm long 10 cm wide deeply pinnatifid at apex and with plusmn6ndash12 free pinnules beneath according to position pinnules oblong 15ndash5 cm long 07ndash17 cm wide shortly narrowed at the apex crenate to distinctly pinnatifid ultimate lobes of largest pinnae in plusmn7 pairs plusmn square up to 8 mm long 6 mm wide crenate-toothed Sori elliptic to linear 15ndash3 mm long one on acroscopic basal nerve of some ultimate lobes but only about 2ndash5 per pinnule the upper sori quite small indusium breaking irregularly spotted and streaked inside with brown paraphyses evident

A sheet labelled Zanguebar Sir John Kirk recd 10[18] 82rsquo is undoubtedly not from Zanzibar but from somewhere in T 6

TANZANIA Morogoro District Uluguru Mts E slope of Bondwa 26 Sept 1970 Faden et al 70635

DISTR T 6 HAB Montane evergreen rainforest 1650 m SYN D sp A Schippers in Fern Gaz 14205 (1993) NOTE Material has been annotated with a specific epithet based on lsquoUlugurursquo but this

has not been published (see Johns Pterid TEA 84 (1991) I have therefore taken up the name

6 D humbertii (C Chr) Pic Serm in Webbia 27444 (1973) (lsquo1972rsquo) Type Congo (Kinshasa) mountains W of Lake Kivu Humbert 7497 (BM lecto fragment and photo)

Rhizome ascending thick fleshy up to 7 cm in diameter elongate with scales dark chestnut linear 7ndash23 mm long 15ndash2 mm wide reticulate mostly black-edged entire not filiform at apex Fronds tufted 13ndash15 m tall sometimes gemmiferous in upper axils stipe greyish straw-coloured 30ndash112 cm long deeply trisulcate above with dense scales at base but plusmnglabrous Lamina triangular-ovate up to 1 m long 82 cm wide bipinnate-pinnatifid to virtually tripinnate with decurrent secondary pinnules rachis straw-coloured pinnae in plusmn 16 pairs oblong-lanceolate (8ndash)35ndash42 cm long (3ndash)14ndash185 cm wide with stalks 1ndash2 cm long and 10ndash16 pairs of pinnules pinnules oblong (1ndash)4ndash10 cm long (05ndash)12ndash5 cm wide with 9ndash12 pairs of narrowly oblong-lanceolate to oblong lobes 08ndash25 cm long 3ndash7 mm wide lobes themselves mostly deeply crenate-lobed (up to plusmn3 mm) but those on apical pinnules and towards apex of basal pinnules are plusmnentire Sori almost round to elliptic 1ndash15 mm long 1ndash18 per pinnule lobe (one at base of each

Diplazium 27

secondary lobe of the lobe) paraphyses dense indusium covering and attached all round the sorus bursting irregularly into lobes and leaving a cup and remnants or dorsally dehiscent

UGANDA Toro District Ruwenzori Nyamgasani Valley Jan 1935 Synge 1470 Ankole District Kalinzu Forest 4 km NW of saw mill W of Rubuzigye 19 Sep 1969 Faden et al 691135 amp 691136 amp 691135 amp Kasyoha-Kitomi Forest Reserve NE of Kyambura R 9 June 1994 Poulsen et al 560 (see note)

TANZANIA Mpanda District Mahali Mts Sisaga 29 Aug 1958 Newbould amp Jefford 1930

DISTR U 2 T 4 Congo (Kinshasa) (Kivu) HAB Lower montane forest Parinari excelsandashCarapandashPolyscias bamboo forest and

mist forest in swampy places and by rivers 1300ndash1450 m (Uganda) and 2250ndash2500 m (Tanzania)

SYN Athyrium humbertii C Chr in Dansk Bot Arkiv 9 (3) 54 t 6 7ndash9 (1937) NOTE Poulsen et al 560 cited above and Poulsen et al 607 from the same forest

appear not to be gemmiferous The Tanzanian specimen also has no gemmae but is closely similar to typical D humbertii Where some characters differ from the description above the extra information is available as followsmdashrhizome horizontal (fide collector) with scales up to 23 mm long attenuate fronds up to 2 m long with lamina 1 m long and 82 cm wide pinnae in plusmn 16 pairs 8ndash42 cm long 3ndash15 cm wide pinnules in 10ndash15 pairs sori 05ndash12 (-2) mm wide It is quite possible that this will prove only a form of D velaminosum Populations of both need examining for detailed frond architecture and presence of gemmae

7 D velaminosum (Diels) Pic Serm in Webbia 27443 (1973) (lsquo1972rsquo) Faden in UKWF 57 59 (1974) Type Cameroon Buea lsquoin Unter-Buea nahe dem Ost-Endersquo Preuss 910 (B syn BM isosyn)

The two sheets mentioned by Pichi Sermolli could not be found at the BM I do not know why Alston did not find the type material at Berlin only an empty covermdashit may have been on loan or misplaced in post-war chaosmdashbut even more inexplicable is the fact that a duplicate of Preuss 910 was in the BM and was overlooked

Rhizome ascending to erect stout to 40 cm long 10 mm diameter with mostly black-edged linear-lanceolate scales up to 3 cm long 1ndash2 mm wide scales with fine-tipped marginal trichomes Fronds tufted 12ndash18 m tall not proliferous stipe 64ndash100 cm long Lamina up to 90(ndash140) cm wide (not seen complete) bipinnate pinnae triangular-lanceolate in outline 14ndash44(ndash70) cm long 11ndash21 cm wide with attenuate crenate apex and 9ndash23 pairs of pinnules pinnules triangular-lanceolate 35ndash11 cm long and 1ndash3 cm wide with attenuate crenate apex and divided almost to the base (in lower pinnules) into 10ndash15 oblong lobes 05ndash25 cm long 3ndash8 mm wide which are subentire to shallowly pinnatifid venation often with scattered long multicellular hairs Sori elliptic 1ndash12 per lobe 05ndash3(ndash4) mm long the 2 lowest often contiguous lengthwise paraphyses mostly numerous and obvious but few and short in some specimens indusium wide and obvious before dehiscence attached all round the sorus and often bursting irregularly across the top or sometimes only along one edge or leaving fragments all around the sorus

UGANDA Toro District Kibale National Park near Kanyawara 13 July 1994 Poulsen et al 656 amp Kibale [Kebale] Forest 14 Dec 1957 BEGAllen 3711 Kigezi


District Bwindi National Park N Sector [Kayonza] Ishasha Gorge 28 Mar 1995 Poulsen et al 818

KENYA N Kavirondo District Kakamega Forest Kibiri Block S side of Yala R 20 Jun 1970 Faden et al 7010

DISTR U 2 3 K 5 Guinea Liberia Satildeo Tomeacute Cameroon HAB Forest Croton megalocarpusndashBequaertiodendronndashDracaena etc often in

swampy places 1200ndash1600 m SYN Hemitelia velaminosa Diels in E amp P Pf 1 (4) 132 (1899) Alston in

FWTA Pterid 29 (1959) Cyathea velaminosa (Diels) Domin Pterid 264 (1929) [Diplazium zanzibaricum sensu Alston FWTA Pterid 64 (1959) pro parte

Tardieu Fl Cam 3 Pterid 236 (1964) pro parte non (Bak) C Chr] 8 D sp B Rhizome erect fleshy scales lanceolate-attenuate 16 mm long 15ndash2 mm wide dark-

edged with few trichomes Fronds tufted plusmn1 m tall bipinnate not gemmiferous stipe green grooved black and swollen at base 25ndash35 cm tall pinnae in 11ndash15 pairs oblong-lanceolate up to 28 cm long 95 cm wide with pinnatifid apical segment and 11ndash15 pairs of pinnules pinnules oblong-lanceolate 25ndash7 cm long 08ndash28 cm wide with crenate apex and divided into 6ndash9 pairs of lobes for about frac12ndash⅔ of width free part of lobes oblong 3ndash8 mm long 3ndash6 mm wide often falcate near apex plusmnentire all round or toothed or crenate at apex Sori elliptic to oblong 15ndash3 mm long 1ndash5 per lobe paraphyses short and few indusium covering and fixed all round the sorus breaking irregularly but sometimes little or none left in mature sori

KENYA N Kavirondo district Kakamega Forest NE of Forest Station 24 Nov 1969 Faden et al 691976 amp along Yala R about 48 km SE of Forest Station 25 Nov 1969 Faden et al 692023

DISTR K 5 HAB Evergreen forest of CeltisndashCroton and Sapotaceae by stream 1650ndash1700 m SYN [D velaminosum sensu Faden in UKWF ed 157 (1974) pro parte] D sp sensu Faden in UKWF ed 233 (1994) NOTE Faden (1994) points out that this taxon has the broader less divided lobes of D

nemorale but small sori

UNIDENTIFIED WOODSIACEAE Rhizome not seen Fronds about 1 m tall stipe not seen Lamina plusmnovate up to 48 cm long 24 cm wide 2-pinnate with pinnules deeply pinnatifid so that frond appears 3-pinnate but costulae are very narrowly winged by the decurrent ultimate segment-bases pinnae in at least 15 pairs lanceolate up to 14 cm long 6 cm wide with 15ndash17 pairs of pinnules pinnules oblong-lanceolate up to 4 cm long 1ndash3 cm wide with 5ndash9 pairs of segments up to 7 mm long 4 mm wide segments with 1ndash2 bifid lateral lobes on each side and terminal 2ndash4-fid lobe the teeth subacute costulae with rather sparse multicellular hairs up to plusmn1mm long Sori not seen

TANZANIA Lushoto District W Usambaras Shume-Magamba Forest Reserve 2 May 1987 Kisena 577 pro parte

DISTR T 3 HAB Presumably forest 1680m

Diplazium 29

NOTE Sheets 3 4 amp 5 out of 5 are this species 1 amp 2 being a Dryopteris species Sheets 3ndash5 had been named Athyrium scandicinum but are not that species and show more resemblance to Dryoathyrium but are not a good match for the species treated in this part Possibly this also is a Dryopteris


INDEX TO WOODSIACEAE

Arachnoides foliosa (C Chr) Schelpe 12 Aspidium boryanum Willd 2 Aspidium glabratum Kuhn 2 Aspidium kiboschense Hieron 2 Aspidium scandicinum Willd 10 Aspidium sulcinervium Hieron 21 Aspidium viridulum Desv 7 Asplenium decussatum Sw 15 Asplenium hylophilum Hieron 19 Asplenium nemorale Bak 19 Asplenium proliferum Lam 15 Asplenium schimperi (Feacutee) ABraun 10 Asplenium zanzibaricum Bak 21 ATHYRIUM Roth 8 Athyrium annae Kornaś 12 Athyrium boryanum (Willd) Tagawa 4 Athyrium glabratum (Kuhn) Alston 4 Athyrium humbertii C Chr 22 Athyrium laxum Pappe amp Raws 11 Athyrium lewalleanum Pic Serm 12 Athyrium newtonii Bak 11 Athyrium proliferum (Lam) Milde 15 Athyrium rondoense Verdc 12 Athyrium scandicinum sensu auct 11 Athyrium scandicinum (Willd) Presl 10 11 12

subsp newtonii (Bak) Verdc 10 var rhodesianum Schelpe 11 var scandicum sensu Schelpe 11

Athyrium schimperi Feacutee 8 9 11 Athyrium sp 11

CALLIPTERIS Bory 13 Callipteris prolifera (Lam) Bory 13 14 15 Callipteris ulugurica Verdc 15 Cornopteris boryana (Willd) Tardieu 4 Ctenitis boryana (Willd) Copel 4 Cyathea velaminosa (Diels) Domin 23 CYSTOPTERIS Bernh 4 Cystopteris diaphana (Bory) Blasdell 7 Cystopteris filix-fragilis sensu auctt 5 Cystopteris fragilis (L) Bernh 4 6

subsp diaphana (Bory) Litard 7

subsp diaphana sensu auct 7 subsp A 5 subsp B 7 var fragilis 5

Cystopteris nivalis (Pirotta) Pic Serm 7 5 Cystopteris viridula (Desv) Desv 7

Deparia boryana (Willd) Kato 4 Deparia glabrata (Kuhn) Kato 4 DIPLAZIUM Sw 16 1 Diplazium arborescens (Bory) Sw 15 20

var nemorale (Bak) C Chr 19 Diplazium decussatum sensu auct 15 Diplazium humbertii (C Chr) Pic Serm 22 21 Diplazium hylophilum (Hieron) C Chr 19 20 21 Diplazium incisum Schum 15 Diplazium nemorale (Bak) Schelpe 19 20 23 Diplazium proliferum (Lam) Kaulf 15 Diplazium pseudoporrectum Hieron 17 18

var angustipinnatum Reimers 19 Diplazium serratum Schum 15 Diplazium silvaticum (Bory) Sw 19 Diplazium stolzii Brause 19 Diplazium sulcinervium (Hieron) C Chr 21 Diplazium sp 21 23 Diplazium sp A of Schippers 22 Diplazium sp B of Schippers 16 Diplazium sp C of Schippers 20 Diplazium sp D of Schippers 21 Diplazium sp sensu Faden in UKWF 23 Diplazium ulugurense Verdc 21 Diplazium velaminosum (Diels) Pic Serm 22 Diplazium velaminosum sensu Faden 23 Diplazium zanzibaricum (Bak) C Chr 20 21 23 Diplazium zanzibaricum sensu auct 23 DRYOATHYRIUM Ching 1 Dryoathyrium boryanum (Willd) Ching 2 3 Dryopteris boryana (Willd) C Chr 2 Dryopteris glabrata (Kuhn) Kuntze 2 Dryopteris kiboschensis (Hieron) C Chr 2

Hemitelia velaminosa Diels 23

Lastrea boryana (Willd) Moore 2 Lunathyrium boryanum (Willd) HOhba 4

Nephrodium boryanum (Willd) Bak 2 Nephrodium catopteron (Kunze) Hook 2

var glabrum Hook 2 Nephrodium catopteron sensu auct 2

Index to woodsiaceae 32

Nephrodium glabratum (Kuhn) Bak 2

Parathyrium boryanum (Willd) Holttum 4 Polypodium fragile L 5 Polypodium diaphanum Bory 7

Thelypteris glabrata (Kuhn) Tardieu 4

var hirsuta Tardieu 4

Woodsia nivalis Pirotta 7

New names validated in this part

Athyrium rondoense Verdc

Athyrium scandicinum (Willd) Presl subsp newtonii (Bak) Verdc

Callipteris ulugurica Verdc

Diplazium ulugurense Verdc


GEOGRAPHICAL DIVISIONS OF THE FLORA

Bignoniaceae Bischofiaceaemdashin Euphorbiaceae Bixaceae (pound150) Bombacaceae (pound390) Boraginaceae (pound1480) Brassicaceaemdashsee Cruciferae Brexiaceae (pound150) Buddlejaceaemdashas Loganiaceae Burmanniaceae (pound300) Burseraceae (pound1330) Butomaceae (pound150) Buxaceae (pound150) Cabombaceae (pound150) Cactaceae (pound150) Caesalpiniaceaemdashin Leguminosae Callitrichaceae (pound400) Campanulaceae (pound450) Canellaceae (pound150) Cannabaceae (pound150) Cannaceaemdashwith Musaceae Capparaceae (pound750) Caprifoliaceae (pound150) Caricaceae (pound150) Caryophyllaceae (pound300) Casuarinaceae (pound200) Cecropiaceaemdashwith Moraceae Celastraceae (pound1300) Ceratophyllaceae (pound150) Chenopodiaceae (pound300) Chrysobalanaceaemdashas Rosaceae Clusiaceaemdashsee Guttiferae Cochlospermaceae (pound150) Colchicaceae Combretaceae (pound890) Commelinaceae Compositae Part 1 (pound3200) Part 2 (pound3500) Part 3 Connaraceae (pound300) Convolvulaceae (pound1300) Cornaceae (pound150) Costaceaemdashas Zingiberaceae Crassulaceae (pound1100) Cruciferae (pound1100) Cucurbitaceae (pound1300)


Cyanastraceaemdashin Tecophilaeaceae Cyclocheilaceae (pound175) Cymodoceaceae (pound400) Cyperaceae Cyphiaceaemdashas Lobeliaceae Dichapetalaceae (pound370) Dilleniaceae (pound150) Dioscoreaceae (pound300) Dipsacaceae (pound300) Dipterocarpaceae (pound390) Dracaenaceae Droseraceae (pound150) Ebenaceae (pound1100) Elatinaceae (pound150) Ericaceae Eriocaulaceae (pound650) Eriospermaceae (pound200) Erythroxylaceae (pound300) Escalloniaceae (pound150) Euphorbiaceae Part 1 (pound3150) Part 2 (pound2200) Fabaceaemdashsee Leguminosae Flacourtiaceae (pound590) Flagellariaceae (pound150) Fumariaceae (pound150) Gentianaceae (pound1500) Geraniaceae (pound300) Gesneriaceae Gisekiaceaemdashas Aizoaceae Goodeniaceae (pound150) Gramineae (pound7400) Part 1 (pound1300) Part 2 (pound3800) Part 3 (pound5450) Gunneraceaemdashas Haloragaceae Guttiferae (pound450) Haloragaceae (pound300) Hamamelidaceae (pound150) Hernandiaceae (pound300) Hippocrateaceaemdashin Celastraceae Hugoniaceaemdashin Linaceae Hyacinthaceae (pound600) Hydnoraceae (pound400) Hydrocharitaceae (pound325) Hydrophyllaceae (pound185)


Hydrostachyaceae (pound300) Hymenocardiaceaemdashwith Euphorbiac Hypericaceae (pound300)mdashsee also Guttif Hypoxidaceae Icacinaceae (pound300) Illecebraceaemdashas Caryophyllaceae Iridaceae (pound1500) Irvingiaceaemdashas Ixonanthaceae Ixonanthaceae (pound300) Juncaceae (pound300) Juncaginaceae (pound150) Labiatae Lamiaceaemdashsee Labiatae Lauraceae (pound400) Lecythidaceae (pound150) Leeaceaemdashwith Vitaceae Leguminosae (pound7400) Part 1 Mimosoideae (pound1300) Part 2 Caesalpinioideae (pound1850) Part 3 Papilionoideae (pound5900) Part 4 Papilionoideae (pound5900) Lemnaceae (pound300) Lentibulariaceae (pound300) Limnocharitaceaemdashas Butomaceae Linaceae (pound300) Lobeliaceae (pound830) Loganiaceae (pound450) Loranthaceae (pound1275) Lythraceae (pound1120) Malpighiaceae (pound300) Malvaceae Marantaceae (pound300) Melastomataceae (pound900) Meliaceae (pound1100) Melianthaceae (pound150) Menispermaceae (pound300) Menyanthaceae (pound200) Mimosaceaemdashin Leguminosae Molluginaceaemdashas Aizoaceae Monimiaceae (pound150) Montiniaceae (pound150) Moraceae (pound1425) Moringaceae (pound275) Muntingiaceaemdashwith Tiliaceae Musaceae (pound390) Myricaceae (pound300)


Myristicaceae (pound270) Myrothamnaceae (pound180) Myrsinaceae (pound430) Myrtaceae (pound1700) Najadaceae (pound390) Nectaropetalaceaemdashin Erythroxylaceae Nesogenaceae (pound150) Nyctaginaceae (pound400) Nymphaeaceae (pound390) Ochnaceae Octoknemaceaemdashin Olacaceae Olacaceae (pound300) Oleaceae (pound300) Oliniaceae (pound150) Onagraceae (pound300) Opiliaceae (pound150) Orchidaceae Part 1 Orchideae (pound1850) Part 2 Neottieae Epidendreae (pound2200) Part 3 Epidendreae Vandeae (pound2400) Orobanchaceae (pound150) Oxalidaceae (pound300) Palmae (pound850) Pandaceaemdashwith Euphorbiaceae Pandanaceae (pound390) Papaveraceae (pound150) Papilionaceaemdashin Leguminosae Passifloraceae (pound600) Pedaliaceae (pound300) Periplocaceaemdashsee Apocynaceae Phytolaccaceae (pound150) Piperaceae (pound500) Pittosporaceae (pound300) Plantaginaceae (pound150) Plumbaginaceae (pound300) Poaceaemdashsee Gramineae Podostemaceae Polygalaceae Polygonaceae (pound450) Pontederiaceae (pound150) Portulacaceae (pound1000) Potamogetonaceae Primulaceae (pound300) Proteaceae (pound325) Ptaeroxylaceae (pound200) Rafflesiaceae (pound200)


Ranunculaceae (pound300) Resedaceae (pound150) Restionaceae Rhamnaceae (pound450) Rhizophoraceae (pound300) Rosaceae (pound600) Rubiaceae Part 1 (pound2200) Part 2 (pound3150) Part 3 (pound2370) Ruppiaceae (pound195) Rutaceae (pound775) Salicaceae (pound205) Salvadoraceae (pound300) Santalaceae Sapindaceae (pound1500) Sapotaceae (pound750) Scrophulariaceae Scytopetalaceae (pound150) Selaginaceaemdashin Scrophulariaceae Simaroubaceae (pound340) Smilacaceae (pound185) Solanaceae Sonneratiaceae (pound150) Sphenocleaceae (pound150) Sterculiaceae Strychnaceaemdashin Loganiaceae Surianaceae (pound200) Taccaceae (pound150) Tamaricaceae (pound150) Tecophilaeaceae (pound150) Ternstroemiaceaemdashin Theaceae Tetragoniaceaemdashin Aizoaceae Theaceae (pound150) Thismiaceaemdashin Burmanniaceae Thymelaeaceae (pound450) Tiliaceae (pound2050) Trapaceae (pound150) Tribulaceaemdashin Zygophyllaceae Triuridaceae (pound185) Turneraceae (pound300) Typhaceae (pound150) Uapacaceaemdashin Euphorbiaceae Ulmaceae (pound300) Umbelliferae (pound1480) Urticaceae (pound1100)


Vacciniaceaemdashin Ericaceae Vahliaceae (pound150) Valerianaceae (pound300) Velloziaceae (pound300) Verbenaceae (pound1780) Violaceae (pound630) Viscaceae (pound620) Vitaceae (pound1780) Xyridaceae (pound620) Zannichelliaceae (pound140) Zingiberaceae (pound575) Zosteraceae (pound170) Zygophyllaceae (pound315)

Adviser on Linnaean types CJarvis

Assistant editor Jonathan Stansbie

Typeset by Margaret Newman

Parts of this Flora are obtainable from AABalkema PO Box 825 2160 SZ Lisse The Netherlands httpbalkcmaimanl Royal Botanic Gardens Kew Richmond Surrey

TW9 3AB England (older parts only) wwwkeworg


BOOKCOVER

LIST OF ABBREVATIONS

TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM


LIST OF ABBREVIATIONS AVP=OHedberg Afroalpine Vascular Plants BJBB=Bulletin du Jardin Botanique de lrsquoEtat Bruxelles Bulletin du Jardin Botanique Nationale de Belgique BSBB=Bulletin de la Socieacuteteacute Royale de Botanique de Belgique CFA= Conspectus Florae Angolensis EJ=AEngler Botanische Jahrbuumlcher fuumlr Systematik Pflanzengeschichte und Pflanzengeographie EM=AEngler Monographieen Afrikanischer Pflanzen-Familien und Gattungen EP=AEngler Das Pflanzenreich EPA=GCufodontis Enumeratio Plantarum Aethiopiae Spermatophyta in BJBB 23 Suppl (1953) et seq E amp P Pf=AEngler amp KPrantl Die Natuumlrlichen Pflanzenfamilien FAC=Flore drsquoAfrique Centrale (formerly FCB) FCB=Flore du Congo Belge et du Ruanda-Urundi Flore du Congo du Rwanda et du Burundi FD-OA=APeter Flora von Deutsch-Ostafrika FFNR=FWhite Forest Flora of Northern Rhodesia FPNA=W Robyns Flores des Spermatophytes du Pare National Albert FPS=FWAndrews Flowering Plants of the Anglo-Egyptian Sudan or Flowering Plants of the Sudan FPU=ELind amp ATallantire Some Common Flowering Plants of Uganda FR=FFedde Repertorium Speciorum Novarum Regni Vegetabilis FSA=Flora of Southern Africa FTA =Flora of Tropical Africa FWTA=Flora of West Tropical Africa FZ=Flora Zambesiaca GFP=JHutchinson The Genera of Flowering Plants GP=GBentham amp JDHooker Genera Plantarum GT=DMNapper Grasses of Tanganyika IGU=KWHarker amp DMNapper An Illustrated Guide to the Grasses of Uganda ITU=WJEggeling Indigenous Trees of the Uganda Protectorate JB=Journal of Botany JLS=Journal of the Linnean Society of London Botany KB=Kew Bulletin or Bulletin of Miscellaneous Information Kew KTS=IDale amp PJGreenway Kenya Trees and Shrubs KTSL=HJBeentje Kenya Trees Shrubs and Lianas LTA=EGBaker Leguminosae of Tropical Africa NBGB=Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem POA=AEngler Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete RKG=AVBogdan A Revised List of Kenya Grasses TSK=EBattiscombe Trees and Shrubs of Kenya Colony TTCL=JPMBrenan Check-lists of the Forest Trees and Shrubs of the British Empire no 5 part II Tanganyika Territory UKWF=ADQAgnew (or for ed 2 ADQAgnew amp SAgnew) Upland Kenya Wild Flowers UOPZ=ROWilliams Useful and Ornamental Plants in Zanzibar and Pemba VE=AEngler amp O Drude Die Vegetation der Erde IX Pflanzenwelt Afrikas WFK=AJJex-Blake Some Wild Flowers of Kenya ZAE= Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907ndash1908 2 (Botanik)







































3


1 Dryoathyrium


2 Cystopteris (p 4)


3 Athyrium (p 8)





5 Diplazium (p 16)

1 DRYOATHYRIUM



















Dryoathyrium 3










2 CYSTOPTERIS









subsp A


subsp B












Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM








































3


1 Dryoathyrium


2 Cystopteris (p 4)


3 Athyrium (p 8)





5 Diplazium (p 16)

1 DRYOATHYRIUM



















Dryoathyrium 3










2 CYSTOPTERIS









subsp A


subsp B












Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM


1 DRYOATHYRIUM



















Dryoathyrium 3










2 CYSTOPTERIS









subsp A


subsp B












Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM













Dryoathyrium 3










2 CYSTOPTERIS









subsp A


subsp B












Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM











2 CYSTOPTERIS









subsp A


subsp B












Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM




subsp A


subsp B












Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM



Cystopteris 7


















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM



















Cystopteris 9

3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM


3 ATHYRIUM




pinnatifid pinnae 2



4 A lewalleanum


5 A rondoense


3 A sp



1 A schimperi


2 A scandicinum





Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM




Athyrium 11




























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM





























Athyrium 13





















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM






















Athyrium 15

4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM


4 CALLIPTERIS













Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM







Callipteris 17













Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM














Callipteris 19








5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM









5 DIPLAZIUM






2




6 D humbertii


5 D ulugurense


5


6





3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM




3 D zanzibaricum



7 D velaminosum







Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM



Diplazium 23























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM
























Diplazium 25
























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM

























Diplazium 27


























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM



























Diplazium 29
















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM

















































BOOKCOVER


TITLE

1 DRYOATHYRIUM

2 CYSTOPTERIS

3 ATHYRIUM

4 CALLIPTERIS

5 DIPLAZIUM


flora of tropical east africa - woodsiaceae (2003) (flora of tropical east africa)

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