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Full text of "Bulletin of the Natural History Museum"

ISSN 0968-044 



Bulletin of 

The Natural History 

Museum 



THE NATURAL 
HISTORY 



22 KOV 2000 

Q6NEKAI LIBRARY 




THE 

NATURAL 
HISTORY 
MUSEUM 



VOLUME 30 NUMBER 2 30 NOVEMBER 2000 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
(Natural History) ), instituted in 1949, is issued in four scientific series, Botany, 
Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Botany Series is edited in the Museum's Department of Botany 
Keeper of Botany: Dr R. Bateman 

Editor of Bulletin: Ms M.J. Short 



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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) 
The Natural History Museum, 2000 Botany Series 



ISSN 0968-0446 Vol. 30, No. 2, pp. 33-130 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 30 November 2000 

Typeset by Ann Buchan (Typesetters), Middlesex 

Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



Bull. nat. Hist. Mus. Land. (Bot.) 30(2): 33-79 



Issued 30 November 2000 



The genus Polystichum (Dryopteridaceae) in 
Africa . ./ " , \") 




JACOBUS P. ROUX 

National Botanical Institute, Compton Herbarium, Private Bag X7, Claremont 7735, South Africa 



22 NOV 



CONTENTS 



igfgKAL LJ6RARV 



Introduction 33 

Materials and methods 34 

Terminology 34 

Taxonomic treatment 35 

Key to the African species of Polystichum (including Marion and Prince Edward Islands) 36 

References 78 

Systematic index 79 



SYNOPSIS. Polystichum Roth is a fern genus of 1 60 to 200 species occurring throughout the temperate parts of the world and the 
montane tropics, but absent from the arid regions as well as the lowland tropics. Although floristic accounts of Polystichum exist 
for many parts of the world, the genus remains poorly understood and a taxonomic account for the group as a whole has never 
been undertaken. Polystichum is poorly represented in Africa (including the Marion Island group) with merely 16 species and one 
known hybrid having been recorded from the region. Most of the species are confined to sub-Saharan Africa, occurring mainly 
along the eastern mountain ranges and Mt. Cameroon in the west. In this review of the African Polystichum species detailed 
observations are presented especially on the paleae, as these structures were found to provide the best characters on which a 
subgeneric classification can be based. Diagnostic features and relationships, variation, and the distribution and ecology of each 
species are included. 



INTRODUCTION 



The genus Polystichum Roth consists of between 160 (Tryon & 
Tryon, 1982) and 200 (Daigobo, 1972) species. It occurs throughout 
the temperate parts of the world as well as the montane tropics, but 
is mostly absent from the lowland tropics. Within the range two 
distinct centres of diversity can be identified, namely a larger Asiatic 
centre with approximately 70 species and a tropical American centre 
with approximately 55 species. Both these regions fall within the 
mountainous tropics characterized by mild, moist climates that are 
hardly seasonal. They also correspond with the areas of highest 
species diversity for homosporous ferns (Tryon, 1985). 

Within the Dryopterideae Polystichum is most closely related to 
Arachniodes Blume, Cyrtomium C. Presl, Dryopteris Adans. and 
Phanerophlebia C. Presl. Cyrtomium and Phanerophlebia have 
often been included in Polystichum (Kramer, 1990), but it has 
subsequently been shown that both these genera have a closer 
affinity to Polystichum than to each other (Yatskievych, 1996). 

Polystichum in a strict sense is a natural and relatively homogene- 
ous group of plants characterized by 1 -pinnate to 3-pinnate laminae 
with acroscopically developed ultimate segments, anadromous free 
venation, uni- or biseriate circular sori positioned medially, termi- 
nally or near terminally on abbreviated or unabbreviated vein 
branches, and peltate indusia (a number of species are exindusiate). 
The receptacle appears to be nude in most species. 

Although Polystichum as a genus is easily recognized, many 
species are superficially very similar. The delimitation of species is 
further hampered by the fact that many, mostly common species are 



allopolyploids (Vida & Reichstein, 1975; Wagner, 1979). Also the 
frequent occurrence of Fl -hybrids in some groups obscures species 
limits. 

A formal subgeneric classification for the genus has thus far only 
been provided for the east Asian species by Tagawa (1940) and 
Daigobo (1972). Some of these sections have since been subdivided 
further by Zhang & Kung (1995, 1996a, b) and Kung & Zhang 
(1998) to make provision for some of the Chinese species. Since 
most species remain poorly known a phylogeny for the genus cannot 
be proposed. 

Although floristic accounts of the genus have been published for 
many parts of the world, some being very old and outdated, no single 
monographic treatment exists and most species remain poorly known. 
Some of the modern-day regional treatments provide no detailed 
observations that may suggest affinities. Within the study area 
floristic accounts for Polystichum are available for North Africa 
(Maire, 1952), West tropical Africa (Alston, 1959), Cameroon 
(Tardieu-Blot, 1964), Mozambique, Malawi, Zambia and Zimba- 
bwe (Schelpe, 1970), Rwanda, Burundi and Kivu (Democratic 
Republic of Congo) (Pichi Sermolli, 1985), southern Africa (Schelpe 
& Anthony, 1986) and Bioko (Benl, 1991). 

It is a well-known fact that Africa, when compared with other 
tropical parts of the world, supports a floristically impoverished 
vascular flora. This phenomenon is also reflected in the pteridophyte 
flora of the continent. The cause of this floristic poverty is ascribed 
to the isolation of Africa from the other continents since the mid- 
Cretaceous and the subsequent significant changes in the climate as 
a result of uplift, continental drift and aridification caused by extra- 
tropical glaciation. All these changes may well have resulted in a 



The Natural History Museum, 2000 



34 



J.P. ROUX 



progressive elimination of the once rich tropical and subtropical 
forests that existed towards the late Jurassic and the establishment of 
extensive deserts and semi-deserts by the early Pliocene (Coetzee, 
1993). 

Within the study area two regions can be identified: an African 
and a sub-Antarctic region. The origin, composition and floristic 
affinities of these regions differ markedly. The African region is the 
largest and today there are three fundamentally different floras or 
biogeographical subregions which can be identified: a southern 
African flora, a tropical African flora and a North African flora. 

The flora of the southern African subregion is believed to have 
evolved gradually since the mid-Tertiary, derived partly from an 
ancient southern African temperate flora and partly from a tropical 
African forest flora (Goldblatt, 1978). Elements of the southern 
African flora currently extend into tropical Africa along the eastern 
escarpment. The southern Cape forests are believed to be impover- 
ished remnants of the tropical African forest flora (Coetzee & 
Muller, 1984). Also the tropical African flora is believed to be an 
impoverished remnant of a once much richer tropical rainforest flora 
that extended over a far greater area than it currently occupies. 

North Africa has also experienced significant changes in its 
climate and vegetation. During the Palaeocene the present Sahara 
desert was clothed by a rich tropical lowland rainforest that also 
covered part of Europe (Greenway, 1973; Raven & Axelrod, 1974), 
but by the Oligo-Miocene it was replaced by a subtropical woodland 
savanna (Axelrod & Raven, 1978). From the Pliocene a desert 
climate established itself in the major part of the Sahara (Quezel, 
1978), serving as an effective barrier to migration from the south. 
The formation of glaciers on the high mountains during the 
Pleistocene permitted the establishment of circumboreal elements. 
Many of these elements are present in the North African flora since 
it is composed of relict elements of African origin as well as 
elements from Eurasia, not frequent in the present sub-Saharan 
flora. The mediterranean influence on the flora of North Africa 
justifies it being considered as a biogeographical subregion of its 
own. 

The Marion and Prince Edward Island group forms part of the 
sub- Antarctic region, a phytogeographical area completely different 
from the foregoing. This island group is of volcanic origin and is 
estimated to be 0.5 million years old (Verwoerd, 1971). Situated in 
the Southern Ocean some 1 800 km from Africa, its biota consists of 
taxa capable of long-distance dispersal and the ability to establish 
themselves in habitats not always favourable for plant growth. 

About two-thirds of the African pteridophy tes are limited in their 
occurrence to the continent (Kornas, 1993). The majority of these, 
however, are closely related to taxa in either tropical America and/ 
or southeast Asia. Pteridophytes of the sub-Sahara biogeographical 
region exhibit three discontinuous distribution patterns: an Ameri- 
can-African disjunction, an African-Madagascan disjunction and an 
African-Asian disjunction. 

Polystichum in Africa is largely confined to the Afromontane 
Phytochorion. White (1978) divided this montane archipelago into 
seven regional mountain systems. The North African Atlas moun- 
tain ranges are here added as an eighth. Although the sub-Saharan 
mountain ranges are sufficiently distinct, the systems are connected 
by a complex series of intermediate floras (Fig. 1 ). The Drakensberg 
system, with six Polystichum endemics, is the richest. This is also 
true for the angiosperms (White, 1978). The only other mountain 
systems with true endemics are the Imatongs-Usambara system with 
two endemics (P. kilimanjaricum & P. volkensii) and the Ethiopian 
system with one endemic (P. magnificum). Other African Polystichum 
species have wider distributions. Polystichum zambesiacum occurs 
in the Chimanimani, Uluguru-Mulanje, and Imatongs-Usambara 



mountain systems, whilst P. transvaalense and P. wilsonii are dis- 
tributed throughout seven mountain systems. The distribution of P. 
wilsonii, however, also extends along the Himalaya mountains to 
Bhutan, Japan and Taiwan. Polystichum luctuosum has an almost 
similar eastern distribution but is confined to the Drakensberg and 
Chimanimani mountain systems in Africa. Polystichum luctuosum 
and P. wilsonii also show a disjunct distribution with the Madagascan 
region. 

Based on observations taken from the systematic treatment, and 
judging from wide-ranging species, southern and tropical African 
Polystichum has a closer affinity with taxa from Asia than with those 
from the Americas. Polystichum in the North African phyto- 
geographical subregion shows a closer affinity with Polystichum 
from Europe than from Africa as P. aculeatum and P. setiferum are 
widespread in that region. Only P. marionense, endemic to Marion, 
Prince Edward and Crozet Islands, occurs in the sub-Antarctic 
phytogeographical region. 



MATERIALS AND METHODS 

This review is based on observations made during extensive field- 
work in southern Africa and on cultivated plants collected during 
these travels. The collections of several herbaria were also studied. 
These include: B, BM, BOL, BR, ETH, GRA, K, L, M, MAL, NBG, 
NH, NU, P, PRE, RAB, SAM, SRGH, WAG (abbreviations follow 
Holmgren et al., 1990) and the private herbarium of Prof. R.E.G. 
Pichi Sermolli (PIC.SERM.). 

Palea and indusium observations were made by removing a small 
number of these structures from selected specimens. These were 
cleaned and cleared in diluted household bleach, after which they 
were semi-permanently mounted in glycerine and the cover slips 
sealed with Entellan. Observations were made with an Olympus 
CH-2 light-microscope fitted with a drawing tube. 

The collections studied are all listed under 'Material examined'. 
These are arranged alphabetically according to country of origin. 
South African (including Lesotho and Swaziland) collections are 
further arranged according to the quarter-degree square-grid system 
(Edwards & Leistner, 1971). In this system each one-degree square 
is known by a standardized name, derived from a town or other 
feature of importance in the square. Each one-degree square is 
divided into four half-degree squares (30' x 30'), numbered A, B, C 
and D from left to right and top to bottom. Each half-degree square 
is again subdivided into quarter-degree squares (15' x 15'), again 
numbered A, B, C and D. By using these co-ordinates a geographical 
area can immediately be identified. 

Unless cited otherwise, the chromosome numbers provided here 
are based on the author's own observations and will be published 
elsewhere. 

Terminology 

This study is principally based on a detailed comparative morpho- 
logical analysis of the sporophyte, where palea structure proved to 
be most informative in suggesting species groups. The terms used to 
describe the apex or the apical cell of the paleae are defined as: 

- apex flagelliform: the apex of the palea terminates in a uniseriate 
series of slender cells. 

- subulate cell: the apical cell is less than 0.4 mm long and the apex 
is usually blunt. 

- acicular cell: the apical cell is slender, straight, more than 0.4 mm 
long, and the apex is usually sharp. 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 

1 



35 



r V 

VIII. NORTH AFRICAN 




U-y IV. IMATONGS-USAMBARA 
\ P. kilimanjaricum 



VII. ORAKENSBERG 

'. drakensbergense 
. incongruum 

. macleae 
. monticola 
. pungens 
. transkeiense 



Fig. 1 Distribution of Polystichum in Africa. 



filiform cell: the apical cell is slender, twisted, and more than 0.6 
mm long. 

thin-walled cell: these cells can vary considerably in size and the 
cell wall is conspicuously thinner than that of the surrounding 
cells, and is therefore often lost. In dry material the cellular 
contents are usually yellowish in colour and appear crystalline. 



TAXONOMIC TREATMENT 

Polystichum Roth, Tent.fl. Germ. 3: 31, 69 (1799). Type species: 
Polystichum lonchitis (L.) Roth (= Polypodium lonchitis L.). 

Hypopeltis Michx., Fl. bor.-amer. 2: 266 (1803). Type species: 
Hypopeltis lobulata Bory (= Polystichum aculeatum (L.) Roth). 

Plecosorus Fee, Mem.foug. 5: 1 50 ( 1 852). Type species: Plecosorus 
mexicanus Fee, nom. superfl. for Cheilanthes speciosissima Kunze 
(= Polystichum speciosissimum (Kunze) R.M. Tryon & A.F. 
Tryon). 

Sorolepidium H. Christ in Bot. Gaz. 51: 350 (1911). Type species: 
Sorolepidium glaciale (H. Christ) H. Christ (= Polystichum 
glaciate H. Christ). 



Hemesteum H. Lev., Fl. Kouy-Tcheou: 450, 496 (1915), non Newm. 

(1851). Type species: several Polystichum species are listed. 
Aetopteron House inAmer. Fern J. 10: 88 (1920), nom. nud. 
Papuapteris C. Chr. in Brittonia 2: 300 (1937). Type species: 

Papuapteris linearis C. Chr. (= Polystichum lineare (C. Chr.) 

Copel.). 
Acropelta Nakai in Bull. Natl. Sci. Mus. Tokyo 33: 5 (1953). Type 

species: Acropelta omeiensis (C. Chr.) Nakai (= Polystichum 

omeiense C. Chr.). 

Plants terrestrial, or epilithic, rarely low-level epiphytes. Rhizome 
erect to suberect and mostly unbranched, or creeping, or decumbent 
and branched; rarely stoloniferous; dictyostelic; set with roots, 
closely to widely spaced persistent stipe bases, and paleae. Fronds 
monomorphic, caespitose or closely to widely spaced, to 1 .8 m long: 
stipe proximally convex adaxially, becoming slightly to deeply 
sulcate distally; with two larger near-circular vascular bundles 
dorso-laterally, ventrally with three to five smaller circular vascular 
bundles; initially moderately to densely paleated, becoming near 
glabrous later, the paleae often appearing heteromorphous, variable: 
lamina 1 -pinnate to 3-pinnate, anadromous, sometimes bearing 1 to 
several paleated proliferous buds adaxially along the rachis near the 



36 



J.P. ROUX 



lamina apex: rachis adaxially shallowly to deeply sulcate, the sulcus Distal pinnae never folded ventrally along the rachis; spores aborted 

proximally not open to sulci of lower order axes, moderately to 9. P. x saltum 

densely paleated; paleae variable: pinnae short-stalked, opposite to 10 Rhizome to 10 mm in diameter; sori exindusiate 1 1 

alternate, closely to widely spaced, often imbricate, simple to 2- 

pinnate, acroscopically auricled: pinna-rachis adaxially sulcate, Rhizome more than 10 mm in diameter; sori indusiate 12 

open to sulci of costae, sparsely to densely paleated; paleae variable: u Rhizome to 5 mm in diameter; stipe and rachis paleae with long 

pinnules proximally mostly short-stalked, opposite to alternate, flagelliform outgrowths along the margin 10. P. marionense 

closely to widely spaced, often imbricate, the proximal acroscopic _ Rhizome to , Q mm - n diameter; stjpe ^ fachis pa , eae ^ Qr without 

pinnule mostly longer than the next in 2-pmnate or more dissected thin-walled cells along the margin 1 1 . P. transkeiense 

species, herbaceous to firmly coriaceous, inaequilateral, ovate to 

ovate-rhomboid or trullate, often somewhat falcate, mostly 12 Lamina with a proliferous bud along the rachis near the apex 

acroscopically auricled in 2-pinnate or more dissected species, 12. P. magm icum 

lobate, dentate or serrate, sharp-tipped or aristate; variously paleated. Lamina without proliferous buds along the rachis 13 

Venation free, pinnately branched, anadromous, terminating near or ._..,. 

, . . 13 Rhizome paleae conspicuously rugose, often with a few long filiform 

at the margin when sterile, immersed or raised. Son circular, essen- outgrowths along the margin; smaller stipe, rachis and pinna-rachis 

tially unisenate, borne medially on unabbreviated vein branches, or paleae basally with short and/or , ong flliform outgrowths along the 

near or at a vein ending of mostly anadromous vein branches: margin 13. P. zambesiacum 
sporangium with 8-(13)-30 indurated annulus cells; stalk with 

glandular cells or eglandular, 3-seriate below capsule: indusium Rhizome paleae not conspicuously rugose, mostly with short straight or 

curved marginal outgrowths; smaller stipe, rachis and pinna-rachis 

absent or present, peltate, mostly persistent, the margin variously ... f, , ., 

J h J paleae basally without short and/or long unisenate outgrowths along the 

sculptured, with or without gland-like cells. Spores monolete, the margin 14 

laesura 2 / 3 to 3 / 4 of the spore length, the perispore irregularly folded, 

mostly somewhat spinulose, often perforate. Chromosome number 14 Proximal acroscopic pinnule to 22 mm long; larger stipe base paleae 

n=41, 82, 164; 2n=82, 164, 328; apogamous 123, 246. often bicolorous r 

Proximal acroscopic pinnule usually more than 22 mm long; larger stipe 
Key tO the African Species Of PolystictlUm (includ- base paleae never bicolorous 16 

ing Marion and Prince Edward Islands) 

1 5 Rhizome short-decumbent with crowded stipe bases, closely branched; 

apogamous (32 spores per sporangium) .... 14. P. monticola 

1 Lamina 1 -pinnate (rarely 1 -pinnate-pmnatifid) l.P. macleae 

- , - Rhizome decumbent, stoloniferous; sexual (64 spores per sporangium) 

- Lamina 2-pinnate to 3-pmnate 2 

15. P. dracomontanum 

2 Rhizome short, erect to suberect, mostly unbranched 3 

16 Pinnules maequilaterally ovate to narrowly trullate, to 60 x 13 mm; 

Rhizome short-decumbent to widely creeping, mostly branched .... 10 sporangium stalk glandular or eglandular; indusium with or without 

unicellular thin-walled cells along the margin 16. P. incongruum 

3 Larger rhizome and stipe base paleae with long unisenate hairs along 

the margin and superficially 2. P. luctuosum Pinnules inaequilaterally ovate, ovate-oblong, ovate-rhomboid or 

trullate, to 50 x 1 9 mm; sporangium stalk and indusium always eglandular 
Larger rhizome and stipe base paleae without long unisenate hairs along ,7 p Dun o ens 

the margin and superficially 4 

4 Lamina with 1-3 paleated proliferous buds along the rachis near the 1. Polystichum macleae (Baker) Diels in Engl. & Prantl, Nat. 
lamina apex 5 Pflanzenfam. 1(4): 190(1902), as macleanii. Type: South Africa, 

- Lamina without proliferous buds along the rachis .... ... 6 in convallibus humidis - Drakensbergen prope 'Pilgrim's Rest 

Gold Fields', McLea 34 sub Bolus 3030 (Kl-lectotype, desig- 

5 Pinnule margins obtusely serrate to crenate, never aristate natedby Schelpe & Anthony (1986); BOL!, SAM!-isolectotypes). 

3. P. volkensii p- ^ 

Pinnule auricle and apex aristate 4. P. kilimanjaricum ... . \ti i/i /ioo/:\ 

Aspidium macleae Baker in Hook.f., Icon, pi.: t. 1654 (1886), as 

6 Apices of paleae terminating in a short subulate cell or a small thin- macleaii. 

walled cell, the margins set with short straight and/or angular outgrowths 

7 Plants terrestrial, epilithic, or rarely epiphytic. Rhizome decumbent, 

to 200 mm long x 20 mm in diameter, densely set with roots, 

Apices of paleae always terminating in an acicular cell, the margins set per sistent stipe bases, and paleae; paleae ferrugineous, membranous 
with long straight and/or long twisted emarginate to forked outgrowths tQ chartaceouSi narrowly ovate or lanceolate , to 7 x 2 mm. Fronds 

caespitose, to 7 per plant, arcuate, to 1.47 m long: stipe proximally 

7 Stipe and rachis moderately paleated; paleae mostly flat or irregularly castaneous, stramineous distally, adaxially sulcate, to 670 mm long 

folded 5. P. aculeatum x g mm j n diameter, proximally densely paleated; larger paleae 

Stipe and rachis densely paleated; paleae mostly helically twisted broadly attached, concolorous or bicolorous, the concolorous paleae 

6. P. setiferum ferrugineous, the bicolorous paleae centrally dark brown or black, 

8 Conspicuously larger paleae mostly confined to the stipe, rugose ... ovate ' narrowl y ovate ' or narrowl y oblon g' cordate ' entire or with 

7 P. transvaalense short and/or long marginal outgrowths proximally, the apex 

flagelliform, terminating in a long filiform cell or an oblong thin- 

:onspicuously larger stipe paleae extending to the rachis, never rugose walled cell to 37 x 8 mm; smaller paleae concolorous, ferrugineous, 

membranous, narrowly ovate, lanceolate, narrowly triangular, nar- 

9 Distal pinnae folded ventrally along the rachis (conduplicate); spores rowly oblong to acicular, cordate, cordate-imbricate, or short-stalked, 
not aborted 8. P. wilsonii proximally erose and/or with long twisted, filiform outgrowths, the 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



37 



apex flagelliform, terminating in a filiform cell or an oblong thin- 
walled cell: lamina 1 -pinnate (rarely 1-pinnate-pinnatifid), oblong 
to narrowly elliptic, with up to 37 free pinna pairs, to 840 mm long, 
the proximal pinnae slightly reduced, usually deflexed: rachis 
stramineous, adaxially sulcate, moderately paleated; paleae short- 
stalked, ferrugineous, membranous to chartaceous, narrowly ovate, 
narrowly lanceolate, or oblong to subulate, cordate, cordate-imbri- 
cate, proximally usually erose and/or with a few short or long 
twisted marginal outgrowths, the apex long-attenuate to flagelliform, 
terminating in an acicular cell or an oblong thin-walled cell, to 3 mm 
long: pinnae firmly herbaceous, olive-green adaxially, paler 
abaxially, generally not overlapping, short-stalked, narrowly ob- 
long-attenuate, straight, auriculate acroscopically, the base unequally 
broad-cuneate to truncate, doubly serrate, to 168 mm long x 16 mm 
wide, the acroscopic auricle on proximal pinnae often free, ovate to 
trullate, to 24 x 22 mm; costa adaxially sulcate, sparsely paleated, 
the paleae taeniform, sessile or short-stalked, entire, the apex ter- 
minating in an acicular cell or an oblong thin-walled cell, to 3 mm 
long, abaxially moderately to densely paleated, the paleae ferrugin- 
eous, membranous, narrowly lanceolate to narrowly trullate, often 
bullate, cordate to cordate-imbricate, the margin proximally with 
short and/or long irregular outgrowths, entire distally, the apex 
terminating in an acicular cell or an oblong thin-walled cell, to 2.4 
mm long. Venation raised. Sori circular, to 1.5 mm in diameter, 
variable in size, those closest to the costa largest, discrete at matu- 
rity, medial to inframedial on unabbreviated vein branches: 
sporangium with 12-(16)-28 indurated annulus cells; stalk 
eglandular: indusium brown, persistent, peltate, circular to irregular, 
repand to erose, often with flabellate central processes, the maxi- 
mum radius 0.29-(0.49)-0.7 mm. Spores 64 per sporangium, brown, 
the perispore folded to form a sparse reticulum of low compressed 
ridges, variously granulate, verruculate to echinulate, closely perfo- 
rated, the exospore 40-(51.31)-66 x 28-(37.89)-48 urn. 
Chromosome number 2n=164. 



MATERIAL EXAMINED 

SOUTH AFRICA. 2330 (Tzaneen): Tzaneen, Woodbush Forest Reserve 
(CC), Balsinhas 2166 (PRE); Wolkberg, Agatha Forest Reserve, 1500 m, 
Mutter 264 (PRE); Woodbush, Van Jaarsveld6\ 10 (BOL). 2430 (Pilgrim's 
Rest): Haffenden Heights, Zoutpansberg (AA), Junod 4069 (P, PRE); 
Mariepskop (DB), Van der Schijf 4305, (B, NU, PRE), 5597 (PRE); 
Mariepskop summit, 1800m, Van der Schijf '4861 (PRE); Mariepskop, below 
radar station, Krynauw 786 (PRE); Graskop, Erasmus Kop, Hardcastle 59 
(PRE); Mariepskop, Schweickerdt 4305 (BOL); Ohrigstad Nature Reserve, 
6000 ft (DC), Jacobsen 1556 (PRE); Pilgrim's Rest, Mount Sheba Nature 
Reserve, Roux 2555 (NBG); Mount Sheba Nature Reserve, Jacobsen 4436 
(PRE); Mount Sheba Nature Reserve, Crouch 633 (NU); Graskop, Cigar 
Rock (DD), Rauh & Schlieben 9744 (PRE); Graskop, Kowyn's Pass, Rauh & 
Schlieben 9725 (PRE); Graskop, Driekop Gorge, Wager 173 (PRE); Pil- 
grim's Rest, Rogers 14925, 14927 (PRE); Blyde Bosboustasie, Bredenkamp 
s.n. (PRE); Pilgrim's Rest, MacLea 170 (PRE); Kowyn's Pass, Schelpe 1641 
(BOL, NH, NU), 6092 (BOL); Graskop, Fairyland, Roux 2548, 2549 (NBG); 
Graskop, The Pinnacle, 4500 ft, Braithwaite 207 (BOL). 2530 (Lydenburg): 
Lydenburg, Hartbeesvlakte (BA), Kluge 2039, 2333 (PRE); Lydenburg, 
Hartbeesvlakte, 1 960 m, Mohle 288 (PRE); Pilgrim's Rest, Mount Anderson, 
Smuts 38 (PRE); Sabie, forest at Tweefontein (BB), Wager 53 (PRE); Sabie/ 
Lydenburg road, Roux 2242, 2561 (NBG); Witklip Staatsbos (BD), Kluge 
806 (PRE); Belfast (CA), Wager s.n. (PRE); Kaapse Hoop (DB), Van 
Jaarsveld2Q8%a (NBG, PRE), 3376 (BOL); Kaapse Hoop, Wager 73 (BOL), 
1496c (PRE). 2531 (Komatipoort): Barberton, Tiger Creek, 4500 ft (CC), 
Thorncroft96 (BR, P, Herb. PIC.SERM., PRE); Barberton, Maid of the Mist, 
Thorncroft 50 (P, PRE), 68 (NBG, P, PRE); 17 miles SE of Barberton towards 
Havelock, 5000 ft, Schelpe 41 15 (BOL, PRE); W. of Havelock, Songimvelo 
Game Reserve, on farm Josefsdal, 1640 m, Kunitz & Otto 15 (J, PRE). 
SWAZILAND. 2531 (Komatipoort): New Havelock, 12 miles from 



Havelock (CC), Schutte4 (BOL); Havelock Mine, Dyer51 (NU). 2632 (Bela 
Vista): Mbabane, Ngwenya Mountain (AA), Compton 31405 (NBG) 

WITHOUT EXACT LOCALITY: loco incerto, Bolus s.n. (PRE); South 
Africa, Wood s.n. (NU). 

The change of the specific epithet macleaii to macleae is in concord- 
ance with Article 60.11 (Recommendation 60C.l.a) of the 
International Code of Botanical Nomenclature (Greuter et al., 1994). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum macleae 
is the only 1 -pinnate (rarely 1-pinnate-pinnatifid) species in Africa 
and is quite similar to P. kalambatitrense Tardieu from Madagascar. 
Baker (1886) considered P. macleae to be related to P. munitum 
(Kaulf.) C. Presl from North America and P. falcinellum (Sw.) C. 
Presl from Madeira. This assumption was probably based on the 1- 
pinnate lamina morphology in these taxa. Both these species, 
however, have ciliated indusia. Also, the palea morphology of both 
these species differs from that in P. macleae. In P. macleae the 
marginal outgrowths of the lamina paleae are pluricellular whilst 
those of P. munitum and P. falcinellum are unicellular. 

VARIATION. Polystichum macleae shows considerable variation in 
the number of indurated annulus cells per sporangium and in indu- 
sium and pinna morphology. The number of indurated annulus cells 
per sporangium ranges from 1 2 to 28. The mean number of indurated 
annulus cells per sporangium is 16.67 (n=650, SD=0. 14) taken from 
13 populations throughout the species' distribution. Some popul- 
ations have a larger number of indurated annulus cells than others. 
Although no definite correlation could be made between habitat and 
the number of indurated annulus cells, plants collected from an 
exposed streambank on the Hartbeesvlakte near Lydenburg [Kluge 
2039 (PRE)] have a number of indurated annulus cells that ranges 
from 19 to 28 (x=23.86, SD=1.91, n=50), which is significantly 
higher than for a plant growing in a forest habitat [Thorncroft 68 
(NBG)], where the number of cells ranges between 12 and 16 
(x=13.28, SD=0.75, n=50). Intermediates between these extremes 
do occur. Indusia are mostly simple, but on some plants they may 
bear one or more small wings, whilst on others they may bear 
numerous flabellate central processes. The margins vary from repand 
to erose. 

A 1-pinnate-pinnatifid form of Polystichum macleae has been 
recorded from Mpumalanga with the central pinnae bearing up to 1 1 
nearly free pinnule pairs. Pinnules are inaequilaterally narrowly 
trullate to oblong-attenuate in outline with the margins obtusely 
serrate. In the distal pinnae the margins are merely lobed midway to 
the costa. Sori are uniseriate on either side of the costa and are borne 
inframedially. 

The size of the acroscopic auricle varies considerably, and on the 
proximal pinnae it is often detached from the rest of the pinna. The 
auricle sometimes overlaps with the pinna directly above. Pinna 
margins are generally obtusely serrated or doubly serrated but rarely 
the margins are also deeply lobed and serrated. 

DISTRIBUTION AND ECOLOGY. Polystichum macleae is confined to 
the Drakensberg Escarpment and Wolkberg in the Mpumalanga 
province of South Africa and the northern parts of Swaziland, 
occurring at elevations ranging between 1350 and 1960 m. The 
species is largely confined to forests where it grows on banks above 
streams, in forest margins, among rocks and often as a low-level 
epiphyte. Plants often form large stands in deep shade, but rarely 
also occur in exposed habitats. 

2. Polystichum luctuosum (Kunze) T. Moore, Ind.fil.: 95 (1858). 

Type as for Aspidium luctuosum Kunze. 
Fig. 3. 



38 



J.P. ROUX 




10 mm 



Fig. 2 Polystichum macleae. A, proximal part of lamina; B, rhizome; C, section of abaxial surface of fertile pinna. A & B, drawn from Roux 2548 (NBG); 
C, drawn from Roux 2242 (NBG). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



39 



Aspidium luctuosum Kunze in Linnaea 10: 548 (1836). Type: In 
monte Katriviersberg in sylvis, Ecklon s.n. (LZf-syntype); ad 
fontes fl. Katrivier prope Philipstown, in sylvis montium, Ecklon 
s.n. (LZt-syntype). 

Aspidium tsus-simense Hook., Sp. fil. 4: 16, t. 220 (1862). Type: 
Island of Tsus Sima, in the Straits of Korea, Wilford s.n. (K- 
holotype, 2 sheets; NBG! -photograph). 

Polystichum tsus-simense (Hook.) J. Sm., Hist, fil.: 219 (1875). 

Polystichum lobatum var. luctuosum (Kunze) H. Christ in Ber. 
Schweiz. Bot. Ges. 3: 34 (1893). 

Plants terrestrial, epilithic, or rarely low-level epiphytes. Rhizome 
short, erect to suberect, to 10 mm in diameter, densely set with roots, 
persistent stipe bases, and paleae; larger paleae broadly attached, 
castaneous, chartaceous, ovate, narrowly ovate, or lanceolate, cor- 
date, with long twisted uniseriate, gland-tipped hairs on the apical 
margin and surface, the apex flagelliform, terminating in an oblong 
thin-walled cell, to 10.5 x 3.3 mm; smaller paleae short-stalked, 
narrowly triangular to subulate, cordate, the margins proximally 
with numerous long and twisted uniseriate hairs, distally with 
widely spaced apically and basally directed marginal outgrowths 
that become smaller apically, the apex flagelliform, terminating in a 
small thin-walled cell. Fronds crowded, caespitose, 7-16 per plant, 
suberect to arching, to 0.93 m long: stipe proximally castaneous, 
stramineous distally, adaxially sulcate, to 450 mm long x 5 mm in 
diameter, densely paleated; proximal paleae broadly attached, 
castaneous, chartaceous, ovate, cordate, proximally entire or with a 
few short and/or long uniseriate hairs, distally with numerous 
multicellular hairs as for rhizome paleae; distal paleae short-stalked, 
narrowly oblong, narrowly triangular or subulate, cordate to hastate, 
the margins bearing a few long and/or short multicellular hairs 
proximally, distally with widely and irregularly spaced outgrowths 
reduced in size and number towards apex, the apex flagelliform, 
terminating in a small thin-walled cell, to 15 x 1.5 mm: lamina 2- 
pinnate to 2-pinnate-pinnatifid, with up to 25 free pinna pairs, to 480 
mm long, firmly herbaceous to coriaceous, olive-green adaxially, 
paler abaxially, narrowly ovate to ovate, the proximal pinnae slightly 
reduced, often somewhat deflexed: rachis stramineous, adaxially 
sulcate, densely paleated; paleae short-stalked, dark brown to black, 
glossy, chartaceous to crustaceous, narrowly triangular to subulate, 
cordate to hastate, the auricles usually bearing long and twisted 
multicellular and uniseriate hairs some of which terminate in a thin- 
walled cell, the margins either distally with short, widely and 
irregularly spaced outgrowths that reduce in size and number to- 
wards the apex, or more or less entire in smaller paleae, to 7 mm 
long: pinnae short-stalked, 1 -pinnate to 1-pinnate-pinnatifid, with 
up to 12 free pinnule pairs, narrowly lanceolate, proximally widely 
spaced, distally often somewhat overlapping, to 173 mm long: 
pinna-rachis stramineous, adaxially sulcate, densely set with paleae 
similar to but less complex than those on the rachis: pinnules widely 
spaced to overlapping, the proximal acroscopic pinnule the largest, 
often significantly longer than the next, up to 40 mm long and 1 2 mm 
wide, inaequilateral, narrowly trullate to rhomboid, basiscopically 
cuneate, acroscopically truncate and auricled, often somewhat falcate, 
lobate-serrate, aristate; proximal pinnules short-stalked, often 
acroscopically incised to or nearly to the costa; costa adaxially 
sulcate, glabrous, abaxially sparsely paleated, the paleae castaneous, 
chartaceous, narrowly triangular-hastate to subulate-hastate, cor- 
date to cordate-imbricate, proximally with long and/or short filiform 
outgrowths often terminating in a thin-walled cell, the apex always 
terminating in a small thin-walled cell, to 0.3 mm long. Venation 
immersed. Sori circular, c. 1.2 mm in diameter, terminal or nearly 
terminal on abbreviated vein branches, essentially uniseriate: spor- 



angium with 10-(13)-19 indurated annulus cells; stalk eglandular: 
indusium peltate, circular, entire, repand or crenulate, persistent, 
brown, pale brown and often dark centred before drying, cupulate 
when dry, the maximum radius 0.5-(0.73)-0.95 mm. Spores 32 per 
sporangium, brown, the perispore unevenly folded to form narrow 
and broad reticulate ridges, the ridges and areas between ridges 
echinulate, spiculate or verruculate, the exospore 30-(38.84)-50 x 
22-(28.2)-36 urn. Chromosome number 2n=123, apogamous. 

MATERIAL EXAMINED 

LESOTHO. 2828 (Bethlehem): Leribe (CC), Phillips s.n. (SAM). 2927 
(Maseru): Roma Valley (BC), Schmitz 6963 (PRE); Roma, Ruch 1909 A- 
only (PRE). 

SOUTH AFRICA. 2430 (Pilgrim's Rest): The Downs (AA), Junod s.n. 
& 4044 (BR, P, PRE); Mt Sheba (DC), Kluge 2320 (NBG, PRE); Blyde 
Forest Reserve (DD), Jacobsen 4365, 4376 (PRE); Driekop Gorge, Graskop, 
Wager 178 (PRE); Pilgrim's Rest, Ponies Krantz, Braithwaite 229 (BOL); 
Sabie, just outside Ceylon Forest Reserve, Braithwaite 135 (BOL). 2530 
(Lydenburg): Lydenburg District, Spitzkop, Wilms 1781 (B, BM); Sabie 
Gorge (BB), Wager s.n. (PRE); Lone Creek Falls, Sabie, Burrows 1342 
(BOL); Sabie, Rogers 20379 (PRE); Sudwala Caves, 1500 m (BC), Kluge 
2463 (NBG, PRE); Lydenburg, Buffelskloof Nature Reserve, Burrows 3860 
(GRA); between Machadodorp and Badplaas (CD), Steel 242 (PRE). 2531 
(Komatipoort): Rimers Creek, Barberton (CC), Thorncroft 35 (P); Baberton, 
Pott-Leendertz 5574 (PRE); Lomati Falls behind Barberton, Wager 154 
(PRE); Barberton, Thorncroft 36, 104c (PRE); creeks near Barberton, 
Thorncroft 2475 (L). 2729 (Volksrust): Newcastle, Nkandu Reserve, 4900 ft 
(DD), Smith 64 (NU). 2730 (Vryburg): road to Luneburg (AD), Roux 2268 
(NBG); Pongola Bush Nature Reserve, 1500 m (BC), Glen 2390 (PRE); 
Utrecht, Donkerhoek, 5500 ft, Devenish 1 144 (PRE); Hlobane, Mtola Forest, 
Johnstone 296 (NU). 2828 (Bethlehem): Farm Boschkloof (DB), Roux 1228 
(NBG); Witsieshoek, Junod s.n. (P); Royal Natal National Park, Okell 60 
(NU). 2829 (Harrismith): Van Reenen, 5000 ft (AD), Schlechter 6718 (B, 
BM, GRA, PRE, SAM); Van Reenens Pass, Rehmann 7204 (P); Van Reenen, 
5000 ft, Lidey 42 (NU); Robinson's Bush, Oliviershoek Pass (CA), Schelpe 
7967 (BOL); near Cathedral Peak, Box 3371 (BM). 2929 (Underberg): 
Giants Castle Nature Reserve (AB), Roux 2503 (NBG); Injasuti Nature 
Reserve, below Cataract Valley, Roux 2718 (NBG); Champagne Castle, 
Bayer 1444 (NU); Cathedral Peak, bank of Kweliquala River, 4700 ft, 
Schelpe P4 (NU); Cathedral Peak, 5700 ft, Killick 1155 (NU, PRE); Cathe- 
dral Peak, Rainbow Gorge, 5500 ft, Cowan 96 (NU); Cathedral Peak, 1550 
m, Goetghebeur4552 (BR, PRE); Cathedral Peak, c. 5000 ft, MacGregor43 
(NU); Cathkin Park, Howlett 53 (NH); Estcourt, Nolema Forest, 4200 ft, 
Edwards 2685 (NU, PRE); Cathkin Peak, Ndema Forest, 4400 ft, Hillary 106 
& 107 (NU); Cathkin Park, Howlett 53 (NH); Injasuti area, 5000 ft, 
Esterhuysen 26034 (BOL, NBG, PRE); above Dalton Bridge, above Bushmans 
River, c. 4500 ft, Wright et al. 27 (NH, PRE); Mooi River, The Hoek, 4700 ft 
(BC), Bourquin 320 (NU); Polela District, Ndumduma, Glengariff (CB), 
Rennie 913, 940 (NU); Cobham Forest Station, Whale Rock, Hill 48 (GRA); 
Bulwer Mountain (DB), Van Jaarsveld 6468 (NBG, PRE); Bulwer (DD), 
Clarkson 111 (NH, NU); Bulwer, Sunset, 5200 ft, Rennie 546 (NU); Bulwer, 
Allsopp 839 (NU); near Bulwer, Schelpe P52 (NU). 2930 (Pietermaritzburg): 
Balgowan, farm Boschfontein, 4000 ft (AC), Fisher 638 (NH, NU), 642 (NH, 
NU); Lions River District, Dargle, Smook 624 (NU); Balgowan, Thomas 71 
(NU); Balgowan, Devlin 62 (NU); Balgowan, 4000 ft, Lindahl 107 (NU); 
Balgowan District, Thienel 109 (NU); Balgowan, 3500 ft, Bernele 1 13 (NU); 
Balgowan, 3500 ft, Crookes 105 (NU); Nottingham Road, McClean 899 
(NH, PRE); Nottingham Road, sine coll. NH-26790 (NH); Dargle, Griffin's 
Farm, 1500 m, Jones 20 (NH); Lions River, Dargle, Esterhuysen 26202 
(BOL); Balgowan, Bosch Hoek, 1400 m, Moll 905 (BOL, NU, PRE); Lions 
River, Lions Bush Forest, Moll 829 (BOL, NU); Pietermaritzburg (CB), 
Tyson s.n. (PRE); Pietermaritzburg, c. 2200 ft, Carnegie 692 (NU); 
Pietermaritzburg, Blackridge, F.G.C. 692 (NU); Inanda, Wood s.n. (B). 3029 
(Kokstad): Langewacht Forest Reserve near Kokstad, c. 1200 m (CB), De 
Joncheere s.n. (PRE); Mt Currie, Kokstad, Stephany 505 (BOL); Glen Hope, 
Jaconet & Jaconet 539 (BM). 3126 (Queenstown): Woodvale Forest, 
Gwatyn, 4200 ft (AA), Galpin 8203 (PRE). 3127 (Lady Frere): Engcobo 
(DB); McLoughlin 1022 (PRE); Engcobo, Flanagan 2781 (PRE). 3128 
(Umtata): Maclear, farm Woodcliffs (AB), Roux 2479 (NBG); Engcobo, 



40 



J.P.ROUX 




10 mm 



Fig. 3 Polystichum luctuosum. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2433 (NBG). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 

Ku-Hlophekazi Forest (AC), Cawe 111 (BOL); Tsolo, Gxalibomvu Forest 
(AD), Cawe 660 (BOL); Tsolo, Bele Forest (BC), Cawe 731 (BOL); Nqadu 
Forest, Hatchings 39 (BOL); Mount Baziya, Baur644 (B). 3225 (Somerset 
East): in sylvis ad pedem montis Boschberg (DA), Barkley s.n. (GRA, P, 
SAM); in sylvis ad pedem montis Boschberg, MacOwan 1884 (B, BOL); 
Boschberg, Bolus 95 (BOL). 3226 (Fort Beaufort): Katberg Forest (BC), 
Holland s.n. (NBG); Katberg, Hutton s.n. (B, L); Katberg Forests, c. 3000 ft, 
Adams 142 (NU); Hogsback, Madonna and Child Falls (DB); Greatheads.n. 
(SRGH); Hogsback Forest, Auckland Kloof, Griffen x46 (PRE); Brambledene, 
Menziesberg, Acocks 11112 (PRE); Hogsback, Gibbs-Russel 3832 (PRE). 
3227 (Stutterheim): Isidinge Forest (CA), Roux 1986 (NBG); Keiskamma 
Hoek, Gxulu Mountain, 5500 ft, Story 3509 (PRE); Kalogha Forest Station 
(CB), Roux 2433 (NBG); Pine Forest, along Amatola trail, Roux 2709 
(NBG); Pirie Forest, Flanagan 1758 (PRE); Fort Cunningham, Roux 2427 
(NBG); Pirie (CC), sine coll. s.n. (GRA); Amabele (DA), sine coll. s.n. 
(PRE); Pirie, Sim s.n. 505, 1727c (GRA, PRE); Komgha (DB), Flanagan s.n. 
(SAM). 

SWAZILAND. 2631 (Mbabane): Gobolo, c. 3500 ft (AC), Dlamini s.n. 
(NBG, NH, PRE); Stroma, c. 4000 ft, Campion 25822 (NBG, PRE). 

ZIMBABWE: Inyanga, Nyangani, 6000 ft, Chase 3813 (NU, SRGH); 
Gweni, Mt. Cashel, Chase 1083 (SRGH). 

WITHOUT EXACT LOCALITY: Gold Fields, Ayres s.n. (NH); Cap de 
Bonne Esperance, Drege s.n. (P); Johannesburg, Westeman s.n. (P); Cap b. 
Spei, Ecklon s.n. (P); Natal, sine coll. s.n. (P); Mor Bridge, Hill 692 (PRE); 
Drakensberg, Bottomley s.n. (PRE); Natal, Buchanan s.n. (BOL, M); O.F.S. 
TM1761c (PRE); Natal, Wood 504 (PRE); in vollbus montium seciis Katrivier, 
prope Philipstown, 3000-4000 ft, Ecklon & Zeyhers.n. (P); Natal, Buchanan 
21 A-only (M); Kaffrarian forests, Sim s.n. (B); Natalia, Buchanan 74, 84 
(B); CapTPr. b. sp., Ecklon & Zeyher 38.6 (B); ceded territory, Quellen des 
Katrivier, 3000-4000 ft, Drege s.n. (B); Prom. b. Spei, Drege s.n. B & C only 
(B); Natal, Wood s.n. (NU); Xumeri Forest, Rycroft 518 (NU); loco incerto, 
sine coll. s.n. NH-9785 (NH); below Mwndali, 5000 ft, Anderson s.n. (BM); 
Himalaya, Ravi Valey, Chanjii, 7000 ft, McDonnell 34 (BM); South Africa, 
Barckley 95 (GRA); sine coll. s.n. (L); loco incerto, sine coll. s.n. NH-26468 
(NH). 

The African Polystichum luctuosum (Kunze) T. Moore and the 
Asian P. tsus-simense (Hook.) J. Sm. have been considered either as 
distinct taxa(Mitui, 1965, 1968;Hirabayashi, 1969;Daigobo, 1973; 
Nakaike, 1975;Gibby, 1985;Punethaetal., 1988) or as synonymous 
(Hope, 1902; Hooker in Hooker & Baker, 1868). Plants occurring in 
the western Indo-Himalayan mountains have been ascribed to either 
P. tsus-simense (Dixit, 1983) or to P. luctuosum (Khullar, 1987; 
Punetha et al., 1988). Fraser- Jenkins (in Gibby, 1985) considers the 
two taxa as vicariants. I have studied material throughout the 
distribution range of these taxa and find them to be conspecific. 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Diagnostic of 
Polystichum luctuosum is the olive-green colour of the adaxial 
surface of the lamina and the darker veins seen in living plants. It is 
also separated from other taxa in the region by the usually very dark 
and narrow paleae occurring along the stipe and rachis. The larger 
rhizome and stipe base paleae bear long filiform outgrowths along 
the margin and palea surface. Indusia are large, persistent and entire, 
and take on a cupulate form when mature. Polystichum luctuosum is 
furthermore a triploid apomict with 32 spores per sporangium and 
has a somatic chromosome number of 2n=123. 

Within the study area Polystichum luctuosum is the only member 
belonging to section Xiphopolystichum Daigobo. 

VARIATION. Polystichum luctuosum shows little variation in stipe, 
lamina and basal pinna length within the study area. A comparison 
of these parts with Asian material shows that African (including 
Madagascar) plants are slightly larger than the plants from Asia. 
Guard-cell length in African material shows little variation, but in 
Asian plants the variation is pronounced. Asian plants also have 
larger guard cells than African plants. Sori may be uniseriate or 



41 

biseriate, variation that appears to be environmentally induced. 
Indusia show a large degree of variation in both African and Asian 
plants, with African plants having larger indusia than Asian plants. 
Also the number of indurated annulus cells per sporangium shows 
some variation. In African plants the number ranges from 10 to 19, 
whereas in Asian plants the number ranges between 10 and 21. 
Spores too show variation, with Asian plants having larger spores 
than African plants (Table 1). Although variations in palea colour 
occur, their morphology remains fairly stable throughout the distri- 
bution of the species. 

DISTRIBUTION AND ECOLOGY. In South Africa Polystichum 
luctuosum occurs from the Eastern Cape through KwaZulu-Natal to 
the northeastern parts of the Free State, Mpumalanga, and the 
Northern Province. It also extends to the lower elevations in the 
western parts of Lesotho, the higher-lying part of Swaziland, and 
with isolated populations occurring along the eastern escarpment in 
Zimbabwe. Outside of Africa the species occurs on Reunion and the 
central parts of Madagascar, extending to the Indian subcontinent, 
Pakistan, China, Vietnam, South Korea and Japan (Honsyu, Sikoku 
and Kyusyu). 

Polystichum luctuosum occurs in the eastern parts of the summer 
rainfall regions of southern Africa where it appears to be restricted 
to the drier forest types such as Dohne Sourveld in the Eastern Cape, 
Highland Sourveld along the Drakensberg foothills, 'Ngongoni 
Veld in the KwaZulu-Natal midlands and Northeastern Mountain 
Sourveld in Swaziland, and along the lowveld escarpment and 
Soutpansberg. In South Africa P. luctuosum occurs at elevations 
ranging from 670 m to 1740 m, whereas in Zimbabwe on Mount 
Nyangani it occurs at elevations as high as 1825 m. 

Polystichum luctuosum mostly grows on rocks along streams, but 
often also as a low-level epiphyte in moist forests. Plants often also 
grow on rocks away from water and in fairly dry conditions. 

3. Polystichum volkensii (Hieron.) C. Chr., Index filic.: 97 (1905). 

Type as for Aspidium volkensii Hieron. 
Fig. 4. 

Aspidium volkensii Hieron. in H.G. A. Engler, Pflanzenw. Ost-Afrikas: 
86 ( 1 895). Type: Tanzania, an der oberen Grenze des Waldes iiber 
Kiboscho, 3000 m, Volkens 1520 (B!-holotype). 

Polystichum barbatum C. Chr. in Notizbl. Bot. Gart. Berlin-Dahlem 
9: 178 (1924). Type: Kenya, Mt. Aberdare, pr. Kinangop, regio 
Hagenia abyssinica, c. 3300 m, Rob. E. & Th. C.E. Fries 2735 
(K!-holotype; B!-isotype). 

Plants terrestrial. Rhizome short, erect, to 10 mm in diameter, 
densely set with roots, persistent stipe bases, and paleae; paleae 
sessile or short- stalked, ferrugineous, chartaceous, narrowly lanceo- 
late, cordate, entire, the apex terminating in an acicular cell, to 15 
mm long. Fronds caespitose, to 14 per plant, erect, to 1.2 m long: 
stipe proximally castaneous, stramineous distally, adaxially sulcate, 
to 520 mm long x 10 mm in diameter, densely paleated; larger paleae 
short-stalked, ferrugineous, membranous, translucent, shrivelled, 
elliptic to ovate, cordate to cordate-imbricate, minutely fimbriate, 
the apex long, shrivelled, filiform, entire, terminating in an acicular 
cell, to 34 x 10 mm; smaller paleae short-stalked, convolute, ovate 
to narrowly ovate, cordate to cordate-imbricate, proximally with 
short straight or angular marginal outgrowths, becoming entire 
towards the apex, the apex subulate, terminating in an acicular cell: 
lamina 3-pinnate, herbaceous, narrowly elliptic, to 925 mm long, 
olive-green adaxially, slightly paler abaxially, with a single paleated 
proliferous bud on the rachis near the lamina apex, the proximal 
pinnae decrescent, often somewhat deflexed: rachis stramineous, 
adaxially sulcate, densely paleated; paleae short-stalked, convolute, 



42 



J.P. ROUX 



Table 1 Variation in metric characters for African and Asian Polystichum luctuosum (Kunze) T. Moore. 



Character 


X 


African 
Range 


n 


X 


Asian 
Range 


n 


Stipe length 


197mm 


90-450 mm 


35 


193mm 


98^50 mm 


21 


Lamina length 


271 mm 


158^48 mm 


46 


235 mm 


162-465 mm 


29 


Basal pinna length 


66.8 mm 


31-178 mm 


46 


51.8mm 


30-98 mm 


21 


Guard cell length 


41.16 urn 


30-52 urn 


580 


46.9 (am 


36-60 urn 


760 


Indusium size 


0.73 mm 


0.56-0.95 mm 


90 


0.58 mm 


0.31-0.80 mm 


115 


Number of indurated annulus cells 


13.52 


10-19 


600 


14.44 


10-21 


537 


Spore length 


38.84 (am 


30-50 urn 


275 


40.75 Mm 


30-52 Mm 


405 


Spore width 


28.2 urn 


22-36 urn 


275 


28.83 Mm 


22-40 Mm 


680 



stramineous to ferrugineous, narrowly ovate, narrowly lanceolate, 
or transversely elliptic, cordate to cordate-imbricate, proximally 
erose or with short straight or angular outgrowths, becoming entire 
towards the apex, the apex subulate, terminating in an acicular cell, 
the smaller paleae to 18x6 mm: pinnae generally not overlapping 
at the lamina base, overlapping towards middle of the lamina, 
oblong-attenuate, somewhat falcate, basal pinnae to 54 mm long, the 
middle pinnae to 190 x 40 mm, proximal acroscopic pinnule slightly 
enlarged; pinna-rachis stramineous, adaxially sulcate, densely 
paleated; paleae similar to but smaller than those on the rachis: 
pinnules opposite to alternate, asymmetric, acroscopically auricu- 
late, ovate, to 23 x 11 mm, deeply lobed, the acroscopic auricle 
ovate, cuneate, lobes oblong to narrowly oblong, serrate to crenate, 
adaxially moderately paleated; paleae castaneous to ferrugineous, 
chartaceous, convolute, filiform, to 15 mm long, abaxially moder- 
ately to densely paleated; paleae short-stalked, castaneous to 
ferrugineous, chartaceous, convolute, filiform, narrowly linear or 
subulate, cordate to cordate-imbricate, proximally with short angu- 
lar outgrowths, entire towards apex, the apex terminating in an 
acicular cell, to 16.5 mm long. Venation immersed. Sori circular, <1 
mm in diameter, essentially uniseriate, discrete at maturity, terminal 
or near-terminal on abbreviated vein branches, or dorsally on 
unabbreviated vein branches: sporangium with 1 2-( 1 4)- 1 9 indurated 
annulus cells; stalk eglandular: indusium ferrugineous to castaneous, 
peltate, circular, elliptic or irregular, coarsely erose, the maximum 
radius 0.48-(0.66)-0.92 mm. Spores 64 per sporangium, brown, the 
perispore folded to form a close reticulum of compressed ridges, the 
ridges and areas between granulate, verruculate or echinulate, vari- 
ously perforated, the exospore 34-(42.64)-52 x 24-(30.32)-38 (im. 
Chromosome number unknown. 

MATERIAL EXAMINED. 

KENYA: Mt. Nyandarua, forest belt, 10800-1 1000 ft, Rabb & Nightingale 1 
(K). 

TANZANIA: Kilimanjaro, highest forest above Kibosho, Uhlig 186 (B), 
Uhlig 242 (B, K); Kilimanjaro, cave above Moschi, Uhlig 76 (B, K); 
Kilimanjaro, forested area just below 1 st hut and also above Machame route, 
1820 m, Schippers T1452 (WAG); Kilimanjaro, Machame route, 3450 m, 
Pocs s.n. (WAG); Kilimanjaro, B-only, Brenner s.n. (P); Kilimanjaro, S. 
slope along the Mweka route, near Mweka base hut, 2850 m, Pocs 6718/A 
(K). 

WITHOUT EXACT LOCALITY: loco incerto, sine coll. BOL-5726 
(BOL). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum volkensii 
is unique among African Polystichum species in the narrowly 
elliptic lamina outline and the long-decrescent, deflexed or arcuate 
pinnae. The densely paleated stipe and rachis, and the single 
proliferous bud borne near the frond apex are also characteristic. 
The finely divided pinnules and the morphology of especially the 
smaller paleae ensure that it cannot be mistaken for any other species 
in the region. 



The affinity of Polystichum volkensii is yet to be determined. 

VARIATION. Polystichum volkensii shows little infraspecific mor- 
phological variation. Variation is largely restricted to pinnule size 
and pinna orientation and this may be ascribed to environmental 
influences. 

DISTRIBUTION AND ECOLOGY. Polystichum volkensii appears to be 
confined to Mount Kilimanjaro in Tanzania and the Aberdare Moun- 
tain Range in Kenya. At lower elevations (1820 m) on Mount 
Kilimanjaro it occurs in Undifferentiated Afromontane forests but 
higher up, at 3450 m, it occurs in the Ericaceous belt with Erica 
arborea and Podocarpus milanjianus. On the Aberdare Mountain 
Range the species occurs in Undifferentiated Afromontane forests 
but also in Single-dominant Afromontane forests such as Hagenia 
abyssinica-forests at elevations ranging between 3300 and 3610 m 
(White, 1983). 

4. Polystichum kilimanjaricum Pic.Serm. in Webbia 27: 445 
(1972). Type: Tanzania, Kilimanjaro, presso la Bismarck's Hut, 
terrestre, nel sottobosco rado nella parte piu alta della foresta 
umida montana a Podocarpus milanjianus, Hagenia abyssinica 
ed Ilex mitis, c. 2850 m, 8 July 1 956, Pichi Sermolli 5171 (Herb. 
PIC.SERM. 20640-holotype; Herb. PIC.SERM. 25150!, K!- 
isotypes). 

Fig. 5. 

Plants terrestrial. Rhizome erect to suberect, to 1 80 mm long, closely 
set with roots, persistent stipe bases, and paleae. Fronds 8-12 per 
plant, caespitose, suberect to arching, to 1.05 m long: stipe proxi- 
mally castaneous, stramineous distally, adaxially sulcate, to 430 mm 
long x 8 mm in diameter, proximally densely paleated; proximal 
paleae broadly attached, ferrugineous, crustaceous, narrowly trian- 
gular, truncate to cuneate, the margins irregularly set with large 
recurved outgrowths, the apex terminating in an acicular cell, to 9 x 
0.8 mm; distal paleae of two types, the larger broadly attached, 
bicolorous, with a central ebeneous to castaneous, glossy, crusta- 
ceous band, and a dull brown, chartaceous margin, narrowly ovate to 
broadly ovate, truncate to cuneate, the margins minutely fimbriate, 
the fimbriae straight or twisted, simple or apically forked, the apex 
terminating in an acicular cell, to 18x7 mm, the smaller short- 
stalked, concolorous, ferrugineous to stramineous, chartaceous, 
narrowly ovate to subulate, the margins minutely fimbriate, the 
subulate paleae always with long and/or short, simple or branched, 
often apically forked fimbriae at the base and widely spaced, 
recurved or apically directed outgrowths distally, the apex termin- 
ating in an acicular cell: lamina 2-pinnate, with up to 35 free pinna 
pairs, triangular to ovate, to 685 mm long, with 1-3 often widely 
spaced proliferous buds in pinna axils near the apex: rachis 
stramineous, adaxially sulcate, moderately to densely set with paleae 
similar to but smaller and paler than those on the stipe: pinnae 1- 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



43 




10 mm 



5 mm 



Fig. 4 Polystichum volkensii. A, middle pinnae of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A, drawn from Volkens 1520 (B); B, drawn 
from Radd & Nightingale 1 (K); C, drawn from Pocs s.n. (WAG). 



44 



J.P. ROUX 




Fig. 5 Polystichum kilimanjaricum. A, proximal pinnae of lamina; B, lamina apex showing proliferous bud; C, abaxial surface of fertile pinnule. All drawn 
from Pichi Sermolli5\l\ (Herb. PIC.SERM.). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



45 



pinnate, with up to 14 free pinnule pairs, not overlapping proxi- 
mally, the basal pinnae not or slightly reduced in size, often somewhat 
deflexed, ovate, narrowly ovate or oblong-attenuate, to 140 x 35 
mm: pinna-rachis stramineous, adaxially sulcate, set with acicular 
paleae with numerous long and often twisted outgrowths at the base, 
the apex terminating in an acicular cell, to 7 mm long: pinnules 
slightly imbricate, short-stalked proximally, firm-herbaceous to 
subcoriaceous, adaxially olive-green, somewhat paler abaxially, 
inaequilateral, ovate-rectangular to ovate-rhomboid, basiscopically 
cuneate, acroscopically truncate and weakly auriculate, shallowly 
undulate or serrate, the teeth and aristae bent inwards, the auricle and 
apex aristate, the proximal acroscopic and basiscopic pinnules on 
basal pinnae often slightly reduced in size, the proximal acroscopic 
and basiscopic pinnules on upper half of lamina slightly larger than 
the next, to 20 x 10 mm; adaxially sparsely set with short-stalked, 
acicular, somewhat twisted paleae, often with a few long straight or 
twisted, often branched outgrowths at the base, the apex terminating 
in an acicular cell, to 2.5 mm; abaxial surface with similar paleae but 
more densely set. Venation immersed or raised. Sort circular, c. 1 .4 
mm in diameter, terminal on abbreviated vein branches, uniseriate 
or biseriate on acroscopic auricle, discrete: sporangium with 8- 
(13)-20 indurated annulus cells; stalk eglandular: indusium peltate, 
subcircular to irregular, the maximum radius 0.73-(0.87)-1 .02 mm, 
persistent, brown. Spores brown, the perispore folded to form 
inflated or compressed tubercles, echinulate, verruculate to echinu- 
late, sparsely to closely perforated, the exospore 34-(43.44)-56 x 
26-(32.45)^M \im. Chromosome number unknown. 

MATERIAL EXAMINED 

TANZANIA : Kilimanjaro, below 1 st hut, Machame Route, 2950 m, Schippers 
T1465 (WAG); Kilimanjaro, above Mandate Hut, 1830 m, Schippers7l234A 
(WAG). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum 
kilimanjaricum is characterized by the proliferous buds on the 
lamina and the palea morphology. Paleae on the proximal part of the 
stipe are narrowly triangular and are either castaneous throughout, 
or may often have a very narrow paler brown margin. Larger paleae 
higher up the stipe have an ebeneous to castaneous centre and a 
broader ferrugineous to stramineous margin. These paleae are mostly 
oblique in outline. Larger paleae on the upper two-thirds of the stipe 
are ovate to broadly ovate in outline with only the central part of the 
apices being ferrugineous to castaneous in colour. 

Pichi Sermolli (1972) considered this species to closely resemble 
P. pauciaculeatum Bonap. (as P. coursii Tardieu) and P. 
tsaratananense Tardieu from Madagascar and went on to describe 
how P. kilimanjaricum can be distinguished from them. Morpho- 
logically P. kilimanjaricum is more similar to P. tsaratananense than 
it is to P. pauciaculeatum. Polystichum kilimanjaricum belongs to 
section Lasiopolystichum Daigobo. 

DISTRIBUTION AND ECOLOGY. Polystichum kilimanjaricum appears 
to be endemic to Mount Kilimanjaro in Tanzania, occurring at 
elevations ranging between 1830 and 2950 m. The species is evi- 
dently confined to the Hagenia abyssinica montane forests and 
thickets associated with the Ericaceous belt where it mostly occurs 
on the forest floor and on rocky streambanks. 

5. Polystichum aculeatum (L.) Roth, Tent. fl. Germ. 3(1): 79 

(1799). Type as for Polypodium aculeatum L. 
Fig. 6. 

Polypodium aculeatum L., Sp. pi.: 1090 (1753). Type: Habitat in 
Europa. H.L.B. 908,311.72 (L-lectotype, designated by Alston 
(1940)). 



Polypodium lobatum Huds., Fl. angl: 390 (1762). Type: Habitat in 

umbrosis et ad sepes. Haller, Hist, stirp. Helv. : 1 7 1 2 ( 1 768); Pluk., 

Phytographia: 180, f. 1 (1691); Ray, Syn. meth. stirp. brit.: 121 

(1690)-syntypes. 
Aspidium aculeatum (L.) Sw. in Jl. Bot. (Schroder) 1800(2): 37 

(1801). 
Aspidium lobatum (Huds.) Sw. in Jl. Bot. (Schroder) 1800(2): 37 

(1801). 
Polystichum lobatum (Huds.) Bastard, Essaifl. Maine et Loire: 367 

(1809). Chevall., Fl. Belg., Pterid.: 107 (1950). 
Dryopteris aculeata (L.) Kuntze, Revis. gen. pi. 2: 812 (1891). 
Dryopteris setifera subsp. lobata (Huds.) Maire in E. Jahandiez & 

R.C.J.E. Maire, Cat. pi. Maroc 1: 3 (1931). 

Plants terrestrial or epilithic. Rhizome short, erect to suberect, to 1 20 
mm long, to 15 mm in diameter, set with roots, closely spaced 
persistent stipe bases, and paleae. Fronds crowded, caespitose, 8-11 
per plant, erect to arching, to 935 mm long: stipe proximally 
castaneous, stramineous distally, adaxially sulcate, to 170 mm long 
x 6 mm in diameter, proximally densely set with conspicuously 
larger and smaller paleae, moderately paleated distally; larger paleae 
sessile, castaneous, chartaceous, broadly ovate, cordate, the margins 
proximally closely set with short curved outgrowths, the apex 
usually short-flagelliform, terminating in a small thin-walled cell, to 
15 x 9 mm; smaller paleae sessile, castaneous to stramineous, 
chartaceous, lanceolate or narrowly to broadly ovate, cordate to 
cordate-imbricate, the margins proximally with short curved 
outgrowths, the apex terminating in a subulate cell or a small thin- 
walled cell: lamina 2-pinnate, with up to 41 free pinna pairs, 
coriaceous, adaxially dark green, slightly paler abaxially, narrowly 
elliptic, to 770 mm long, closely spaced and often imbricate distally, 
proximally more widely spaced, the proximal pinnae reduced, often 
slightly deflexed: rachis stramineous, adaxially sulcate, moderately 
paleated; paleae sessile, ferrugineous, chartaceous, broadly ovate, 
ovate, narrowly elliptic or hastate, cordate to cordate-imbricate, the 
margins with short, somewhat curved outgrowths extending nearly 
to the apex, the apex terminating in a subulate cell, a long acicular 
cell, or a small thin-walled cell, to 6 x 3 mm: pinnae short-stalked, 
pinnatifid to 1 -pinnate, with up to 16 free pinnule pairs, narrowly 
oblong-attenuate, the middle pinnae to 110 mm long, the proximal 
pinnae to 88 mm long: pinna-rachis stramineous, adaxially sulcate, 
moderately paleated; paleae sessile or short-stalked, ferrugineous, 
chartaceous, ovate, narrowly lanceolate to hastate, cordate to cor- 
date-imbricate, the margins proximally with short or long, usually 
curved outgrowths, the apex terminating in a subulate cell or a small 
thin-walled cell, to 2 x 0.8 mm: pinnules opposite to alternate, 
somewhat imbricate, the proximal acroscopic pinnule usually slightly 
longer than the next, asymmetric, trullate to narrowly trullate, 
basiscopically cuneate, acroscopically cuneate to truncate and au- 
riculate, serrate to long-aristate, to 15 mm long; adaxially with a few 
membranous, filiform paleae terminating in a subulate or thin- 
walled cell confined to proximal part of pinnule, to 1 .75 mm long; 
abaxially moderately set with membranous, narrowly trullate or 
narrowly lanceolate paleae with a few short and straight marginal 
outgrowths, or the paleae filiform, short-stalked, with the apex 
terminating in a subulate cell or a small thin-walled cell, to 2.5 mm 
long. Venation immersed. Sori circular, to 1.5 mm in diameter, 
terminal or near terminal on abbreviated vein branches, essentially 
uniseriate, discrete to confluent at maturity: sporangium with 12- 
(13)-17 indurated annulus cells; stalk eglandular: indusium 
chartaceous, peltate, circular, entire to repand, the maximum radius 
0.63-(0.92)-1 .26 mm, persistent, brown. Spores brown, the perispore 
folded to form short echinate ridges or crests, the areas between 



46 



J.P. ROUX 




10 mm 



Fig. 6 Polystichum aculeatum. A, proximal part of lamina; B, fertile pinnae; C, rhizome. A & B drawn from Lindberg 2793 (B); C, drawn from Cosson s.n. 
(S). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



47 



fenestrate with pores of variable sizes, the exospore 32-(37.46) 48 
x 24-(27.93)-34 jam. Chromosome number 2n= 1 64 (Manton, 1 950). 

MATERIAL EXAMINED 

ALGERIA: Montagnes du Djurdjura, cercle de Dra el Mizan, Cosson s.n. 
(B); La Gourraya de la Bougie, Corzeillea Tere s.n. (S). 

MOROCCO: Haul Atlas, Ourika, 1400 m, Litardiere s.n. (M, P); Great 
Atlas Mountains, Si Chamharouch, 2280 m, Polunin 2 1 84 (BM); N. face G- 
bou Orionl, 2950 m, Newbould 109, 1 10 (BM); Arromiel, Balls 2972 (B, BM, 
S); Taddert, Marrakesh-Quarzazat road, High Atlas, 1600 m, Chatworth- 
Musters 362 (BM); Atlas Magnum, Amismiz, 1400 m, Lindberg 2793 (B, 
S); Meknes, Aguelmane Azigza, 1600 m, Casas et al. s.n. (B); Grand Atlas, 
Ourika, 1 300-1400 m, Maire s.n. (RAB); Haul- Atlas, Ourika, 2600 m, sine 
coll. s.n. 18633 (RAB); env. de la maison forestier de Khanolak-Anasar, 
Jovet-Astetal 13313 (RAB). 

Variation in Polystichum aculeatum and P. setiferum and the occur- 
rence of intermediate forms and hybrids (P. x bicknellii (H. Christ) 
Hahne) between these species have resulted in diverse interpreta- 
tions as to their delimitation. The result has been some nomenclatural 
confusion (Newman, 1844; Alston, 1940; Elliot, 1950; Meyer, 
1960). 

Hudson ( 1 762), unaware of the existence of the name Polypodium 
setiferum Forssk., recognized two forms in European P. aculeatum 
and divided plants into two species. The rigid and less divided form 
he named Polypodium lobatum, and the lax and more divided form 
he retained in P. aculeatum. Hudson's interpretation off. aculeatum 
is therefore synonymous with P. setiferum (Forssk.) T. Moore ex 
Woyn. and P. lobatum with P. aculeatum as now interpreted. Al- 
though the name P. lobatum never became well-established, 
arguments in favour of its retention were made by Meyer (1960). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum 
aculeatum is a fertile sexual species intermediate in morphology 
between P. lonchitis (L.) Roth and P. setiferum (Forssk.) T. Moore ex 
Woyn., although it is closer in appearance to the latter. Manton 
(1950) demonstrated P. aculeatum to be a tetraploid of hybrid origin 
between the putative parents P. lonchitis and P. setiferum. Daigobo 
(1972) placed this species and P. setiferum in different sections, but 
their palea morphology suggests them to be related. Both belong to 
section Metapolystichum Daigobo. 

Polystichum aculeatum is characterized by having a coriaceous 
lamina and smooth, shiny, dark green pinnules. The longest pinnae 
occur at or near the middle of the lamina with the most proximal 
pinna pair distinctly shorter than the middle pinnae. The stipe/ 
lamina ratio in P. aculeatum is 1:3.27 (n=12). Palea density and 
morphology are also diagnostic and differ from that of P. setiferum 
with which it may be confused. In P. aculeatum the stipe, rachis and 
pinna-rachis are moderately paleated, whereas in P. setiferum they 
are usually densely set with twisted paleae. Marginal outgrowths in 
larger paleae are short and curved, and gradually phase out towards 
the apex. Smaller paleae are mostly broad-based and have short 
curved marginal outgrowths, but distally they terminate abruptly in 
an almost simple subulate apex. Paleae occurring abaxially on the 
pinnules are short and proximally bear a few small, straight or 
curved, marginal outgrowths, with the apex terminating in a subulate 
cell or a small thin-walled cell. Polystichum aculeatum also differs 
from P. setiferum in a number of micromorphological characters, 
with the mean adaxial epidermal cell length, guard cell length, 
maximum radial length of the indusium, and the spores being larger 
than those in P. setiferum. 

VARIATION. Polystichum aculeatum varies in the degree to which 
the pinnae reduce in size towards the base of the lamina and in the 
length of the stipe in relation to the length of the lamina. European 
plants appear to have shorter stipes than plants from Africa. Pinnules 



of plants from the study area are remarkably stable and show little 
variation. 

DISTRIBUTION AND ECOLOGY. Polystichum aculeatum is widespread 
in Europe but in North Africa its distribution is more restricted. 
Hansen & Sunding (1993) and Derrick et al. (1987) considered the 
species to also occur on Madeira and the Canary Islands, but no 
material originating from Madeira could be traced by Manton et al. 
(1986), Gibby & Paul (1994), or myself. I have also not seen any 
material of this species from the Canary Islands. 

In Algeria and Morocco Polystichum aculeatum is restricted to 
the High Atlas Mountains. The lithology of the region consists 
largely of basement rock and unconsolidated clay marls (White, 
1983). The rainfall is low and seasonal at lower elevations but at 
higher elevations precipitation may occur throughout the year. The 
species occurs at elevations ranging between 1400 m and 2950 m, 
where it is confined to moist shaded rock crevices along streams and 
at waterfalls. 

6. Polystichum setiferum (Forssk.) T. Moore ex Woyn. in Mitt. 

Naturwiss. Vereines Steiermark 49: 181 (1913). Type as for 

Polypodium setiferum Forssk. 
Fig. 7. 

Polypodium setiferum Forssk., Fl. aegypt.-arab.: 185 (1775). Type: 

Turkey, Dardanelles ('Ad Dardanelles'), Forsskal 814 (C!- 

lectotype, designated by Hepper & Friis (1994)). 
Aspidium angulare Kit. ex Willd., Sp. pi 4, 5(1): 257 (1810). Type: 

Habitat in Hungaria, sine coll. s.n. (B-Willd.-holotype, NBG!- 

photograph). 
Aspidium hastulatum Ten., Semina 1830: 15 (1830). Type: In nostri 

regni nemoribus, et abunde in vallibus circa Neapolim, 5. Rocco, 

Ponti Rossi & Camaldoli s.n. (not located). 
Polystichum angulare (Kit. ex Willd.) C. Presl, Tent, pterid.: 83 

(1836). 
Aspidium aculeatum subsp. angulare (Kit. ex Willd.) Asch. in P.F.A. 

Ascherson & K.O.R.P.P. Graebner, Syn. mitteleur. Fl. 1: 39 

(1896). 
Polystichum aculeatum subsp. angulare (Kit. ex Willd.) Vollm., Fl. 

Bayern: 9 (1914). 
Dryopteris aculeata subsp. angularis (Kit. ex Willd.) Schinz & 

Thell. in H. Schinz & R. Keller, Fl. Schweiz 3rd ed., 2: 3 (1914). 
Dryopteris setifera (Forssk.) Woyn. ex Schinz & Thell., 

Vierteljahrsschr. Naturf. Ges. Zurich 60: 340 (1915). 
Dryopteris setifera subsp. angularis (Kit. ex Willd.) Maire in E. 

Jahandiez & R.C.J.E. Maire, Cat. pi. Maroc 1: 3 (1931). 

Plants terrestrial or epilithic. Rhizome erect to suberect, short, to 18 
mm in diameter, set with roots, closely spaced stipe bases, and 
paleae; paleae broadly attached, stramineous to ferrugineous, 
chartaceous, ovate to broadly ovate, often somewhat bullate, cor- 
date, the margins minutely fimbriate to erose, the apex generally 
entire, terminating in a subulate cell or a small thin-walled cell. 
Fronds 8-22 per plant, suberect to arching, to 1 .2 m long: stipe 
proximally castaneous, stramineous distally, adaxially sulcate, to 
520 mm long x 6 mm in diameter, densely paleated; paleae ferrugi- 
neous, chartaceous, the larger paleae sessile, narrowly ovate, ovate, 
or broadly ovate, often somewhat bullate, cordate, the margins 
finely fimbriate to erose, the apex entire, terminating in a subulate 
cell or a thin-walled cell, to 20 x 1 1 mm, the smaller paleae narrowly 
oblong, narrowly ovate, or subulate, mostly helically twisted, short- 
stalked, cordate-imbricate, the margins proximally with short and/or 
long outgrowths, the outgrowths straight, narrowly triangular, or 
angular, reduced in size towards the apex, the apex terminating in a 



48 



J.P. ROUX 




Fig. 7 Polystichum setiferum. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinna. A & C drawn from Mandon 291 (S); B, drawn 
from Tullgren 21 (S). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 

subulate cell or a small thin-walled cell: lamina 2-pinnate to 2- 
pinnate-pinnatifid, with up to 45 free pinna pairs, firmly herbaceous 
to coriaceous, dark green adaxially, somewhat paler abaxially, ovate 
to elliptic, to 830 mm long, the proximal pinnae reduced, often 
somewhat deflexed: rachis stramineous, adaxially sulcate, densely 
paleated; paleae narrowly oblong, narrowly ovate, narrowly lanceo- 
late, or subulate-hastate, helically twisted, short-stalked, cordate to 
cordate-imbricate, the margins proximally with short and/or long 
straight, curved or angular outgrowths that reduce in size towards a 
usually entire apex, the apex terminating in a subulate cell or a small 
thin-walled cell, to 6 x 2 mm: pinnae with up to 26 free pinnule pairs, 
closely to widely spaced, often overlapping towards the apex, 
narrowly oblong-attenuate, the middle pinnae to 162 mm long, the 
proximal pinnae to 1 40 mm long: pinna-rachis stramineous, adaxially 
sulcate, moderately to densely set with paleae similar to but smaller 
than those on the rachis: pinnules opposite to alternate, closely 
spaced, the proximal acroscopic pinnule not or slightly enlarged, 
inaequilateral, acroscopically auriculate, trullate or oblong-acumi- 
nate, lobate-dentate, aristate, to 13x5 mm; adaxially with a few 
twisted filiform paleae confined to the costa on the proximal part of 
the pinnule, these terminating in a subulate cell or a small thin- 
walled cell, to 4.5 mm long; abaxially moderately set with narrowly 
triangular, subulate-hastate, or filiform paleae, the larger paleae 
proximally usually with a few long marginal outgrowths, the apex 
entire, terminating in a subulate cell, to 2.9 mm long. Venation 
immersed. Sori circular, c. 1 mm in diameter, terminal or near 
terminal on abbreviated vein branches, essentially uniseriate, dis- 
crete to confluent at maturity: sporangium with 1 1 -( 14)-20 indurated 
annulus cells; stalk eglandular: indusium pale brown, persistent, 
peltate, circular, repand, the maximum radius 0.48-(0.85)-1.21 
mm. Spores 64 per sporangium, brown, the perispore folded to form 
inflated tubercles and ridges, echinulate to verruculate, sparsely 
perforated, the exospore 26-(34.52)-44 x 1 8-(25.92)-36 Jim. Chro- 
mosome number 2n=82 (Manton et al., 1986). 

MATERIAL EXAMINED 

ALGERIA: Mont Magnis, l500m,Reverchon371 (BM, P); Djebel Edough, 
Cosson s.n. (P); gorge de la Chiffa, Cosson s.n. (P); Djebel Marouf, petite 
Kabylie, Prov. de Constantine, Cosson s.n. (P); dans la fout du Dirah aux 
environs d' Aumale, Chaoy 834 (P); montagnes du Djurdjura, cerde de Dra el 
Mizan, Prov. d'Alger, Cosson s.n. (P); 3 miles W. of the Hotel Lambert, 
Adekar, c. 900 m, Alston & Simpson 37578 (BM); Djebel, Stephenson s.n. 
(BM); Remain, Nud el Kebin, Alwah, sine coll. s.n. (BM); Algeria, Eichard 
s.n. (WAG). 

MOROCCO: entre les rochers humides et umbrage du mont Amareza, 
Atlas, Bove s.n. (P); Al Hoceima. cerca de Ketama, subiendo al monte 
Koudiet Imougras, 1 880 m, Casas 7237 (B); Hafa-es-Sabbaba (Ben-Hosmar), 
ad 500 m, Quer s.n. (B, S). 

TUNISIA: NV d'ain-Draham, Cosson s.n. (P); Massif d' El-Fedja, Cosson 
& Duval s.n. (P); Ain Draham, open cork-oak forest, c. 900 m, Simpson 
38370 (BM); foret du Feidja, 20 km W. of Ghardimaou, 800 m, Jansen 462 
(WAG). 

WITHOUT EXACT LOCALITY: Herb. Luerssen 5242, sine loc. (P); 
loco incerto, sine coll. B-96812 (B); loco incerto, De Buck s.n. (B); Kaap de 
Goede Hoop [error, not a native of the Cape], sine coll. 9 (L). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Problems are fre- 
quently experienced in separating Polystichum setiferum from P. 
aculeatum. The former, however, has larger fronds that are softer in 
texture. Also the proximal pinnae are not usually markedly reduced 
in P. setiferum as they are in P. aculeatum. In P. setiferum the paleae 
are characteristically helically twisted and the apices more fre- 
quently terminate in a small thin-walled cell than those of P. 
aculeatum. Micro morphological characters separating the two taxa 
are reported under P. aculeatum. 



49 

VARIATION. Considering the wide geographical distribution of the 
species, it shows remarkably little variation. Dyce (1963) reported 
that a wide range of minor variations in shape and habit can be 
expected in any colony of this species. I found variations in the size 
and shape of the fronds to be most pronounced, but since no obvious 
geographic pattern was detected, it is here considered to be environ- 
mentally induced. Plants from drier areas, in particular the North 
African region, have fronds that are often merely 1 -pinnate or 1- 
pinnate-pinnatifid. In large specimens from moist areas, however, 
the lamina may be 2-pinnate-pinnatifid with the proximal acroscopic 
pinnule being 1 -pinnate and often twice as long as the next pinnule. 
Irrespective of habitat and environmental conditions, the palea 
structure shows little variation. 

DISTRIBUTION AND ECOLOGY. Polystichum setiferum is widespread 
in Britain, Europe south of 53 N latitude, the Crimea, Macaronesia 
(Azores, Canary Islands and Madeira), and Africa north of the 
Sahara. 

In North Africa P. setiferum occurs at elevations ranging between 
500 and 1 880 m in the Saharan Atlas-, High Atlas- and Anti-Atlas 
Mountain ranges in Tunisia, Algeria and Morocco. In this region of 
low rainfall plants are restricted to well-protected rock crevices and 
moist banks. In Tunisia, however, the species also occurs in open 
cork-oak (Quercus suber L.) forests. 

7. Polystichum transvaalense N.C. Anthony in Contr. Bolus Herb. 
10: 146 (1982). Type: South Africa, Transvaal (Northern Prov- 
ince), Pietersburg District, Woodbush Forest Reserve, 
Bredenkamp & Van Vuuren 450 (BOL!-holotype; PRE!-isotype). 

Fig. 8. 

Plants terrestrial or epilithic. Rhizome short, erect, to 8 mm in 
diameter, densely set with roots, persistent stipe bases, and paleae; 
paleae broadly attached, castaneous, chartaceous, narrowly linear to 
narrowly lanceolate, the margins proximally entire, distally with 
numerous short, apically or basally directed outgrowths, the apex 
terminating in an acicular cell, to 14 x 1 mm. Fronds caespitose, 5- 
17 per plant, suberect to arching, to 1.045 m long: stipe proximally 
castaneous, stramineous distally, adaxially sulcate, to 535 mm long 
x 5 mm in diameter, densely paleated, the paleae twisted; larger 
paleae mostly confined to the stipe, concolorous or bicolorous, 
castaneous to ferrugineous or with the central part castaneous to 
black, rugose, narrowly ovate-acuminate to ovate-acuminate, short- 
stalked, the margins irregularly lacerate-fimbriate, the apex 
terminating in an acicular cell, to 20 x 6 mm; smaller paleae short- 
stalked, narrowly ovate to narrowly lanceolate, the margins 
proximally lacerate, distally irregularly lacerate-fimbriate, the apex 
terminating in an acicular cell, to 6.5 x 1 .4 mm: lamina 2-pinnate, 
with up to 26 free pinna pairs, herbaceous, ovate to narrowly ovate, 
to 670 mm long, pale green adaxially, paler abaxially, the proximal 
pinnae often slightly reduced, often deflexed: rachis stramineous, 
adaxially sulcate, often flexuous distally, densely paleated; paleae 
short-stalked, twisted, castaneous to ferrugineous, narrowly ovate, 
narrowly triangular, or linear, the margins proximally lacerate, 
distally irregularly and widely fimbriate, the apex terminating in an 
acicular cell, to 4.5 mm long: pinnae 1 -pinnate, with up to 20 free 
pinnule pairs, proximally widely spaced, distally closely spaced and 
somewhat overlapping, oblong-attenuate, the basal pinnae to 140 
mm long x 28 mm wide, proximally often slightly reduced, the 
basalmost acroscopic pinnules longer towards the middle of the 
lamina: pinna-rachis stramineous, adaxially sulcate, densely 
paleated; paleae similar to but smaller than those on the rachis: 
pinnules opposite to alternate, inaequilateral, acroscopically auricu- 
late, ovate to obliquely transversely rhomboid, to 15 mm long. 



50 



J.P.ROUX 




Fig. 8 Polystichum transvaalense . A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2414 (NBG). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 

serrate to lobate-serrate, often short-aristate, the proximal pinnules 
often pinnatifid; adaxially subglabrous or with a few twisted, fili- 
form paleae proximally on costa, the apex terminating in an acicular 
cell, to 2 mm long; abaxially moderately paleated; paleae short- 
stalked, twisted, narrowly linear to narrowly triangular, the margins 
proximally long-lacerate or fimbriate, the apex terminating in an 
acicular cell, to 2.5 mm long. Venation immersed. Sori circular, c. 1 
mm in diameter, terminal or near terminal on abbreviated vein 
branches, discrete at maturity: sporangium with 10-(14)-22 
indurated annulus cells; stalk eglandular: indusium stramineous, 
peltate, circular, simple or often with a few long central processes, 
repand to erose, persistent, the maximum radius 0.48-(0.76)-0.97 
mm. Spores 64 per sporangium, brown, the perispore folded to form 
tubercles and inflated or compressed reticulate ridges, echinulate, 
closely perforated, the pores to 1 .5 urn in diameter, the exospore 30- 
(38.54)-48 x 22-(28.86)-38 urn. Chromosome number 2n= 1 64. 

MATERIAL EXAMINED 

BIOKO: Bioko (Fernando Po), 9000 ft, Mann 340 (K); carratera del pico 
Basile, km 18-19, nacimento del rio Cope, 32NMJ7597, 2470 m, Carvalho 
3682 (B, BR). 

CAMEROON: Pisted'Acha-Abaw au lac Oku, 40 km NE Bamenda, 
Letouzey 1 3439 (P); montane forest between hut 1 and hut 2, 1 950 m, Breteler 
et al. 266 (K, P, WAG); Buea, Preuss 719 (B); Mannsquell, Luckhardt 636 
(B); Mount Cameroon, Kalbreyer 133 (B); Uauenzuba, 1900 m, Schaeter90 
(B). 

DEMOCRATIC REPUBLIC OF CONGO: Mare de Kikeri, Volcan 
Hekeno, Jean-Louis 5 1 9 1 (BR, P, PRE); entre le Hikeno et le Heisfumangabo, 
Jean-Louis 5002 (BR, P, PRE); Kivu Province, Goma, petite mare de Kikeri, 
au pied du Mikeno Pare National Albert, 2200 m, Lebrun 7225, 7228 (BR, K, 
P); entre Kibumba et le Ngamuragira, Lebrun 7087 (BR, P); Kivu, Pare des 
Virunga, Karisimbi, vers le SW Zaire, 3465 m, Van der Veken PV 9130 (B); 
Kivu, Buhavu-Goma, Gupffert 169 (BR); 748'S, 2944'E, Marungu route 
Kieluzi-Mwela km 45 Ravin Lukole, 1 890 m, Bodenghiem & Malaisse 1459 
(BR); Pare National Albert, versant S. du Mikens, 2400-2600 m, Lebrun 
7307 (BR); Omigi, Mickule, Bequaert 6294 (BR); route Goma-Rwindi 30 
km, Breyne 1785 (BR); Volcan Niamlagyra, c. 2000 m, Germain 1381 (BR); 
Kivu, Rumangabo, 1525 m, Germain 3025 (BR); Numbi territory, Kalehe, 
2300 m, Leonard 4554 (BR); Mt. Kiniki territory, 1960 m, Gutzwiller 1236 
(BR); Mt. Kiniki, Wambalyro, 1960 m, Gutzwiller 1 106 (BR); Pare National 
Albert, Kalonge, Butahu, vallee du la Nyamwumba, 2010 m, Demaret5\92 
(BR). 

ERITREA: Eritrea- Assaorta: bosco del Caribozza, c. 2700 m, Pappi 
2812 (BOL, BR). 

ETHIOPIA: c. 5 km NW of Addis Ababa, c. 2500 m, De Wilde 598 1 (BR, 
ETH, WAG); Kaffa Province, 35 km W. of Bonga along the road to Shewa 
Ghimmira,c. 1 950 m, Fr/w et al. 2171 (ETH, K); near Wush-Wush,c. 20km 
NW of Bonga, c. 1800 m, De Wilde 7756 (BR, ETH, WAG); Gara Ades, 
Burger 2580 (K); Mount Wachacha, near Addis Ababa, 2400 m, Mooney 
7895 (K); Wofasha Forest, Shoa, Mooney 7003 (K); Bellete State Forest, 40 
km SW of Jimma, c. 2000 m, De Wilde 6999 (BR, WAG); Kaffa Province, 
village c. 2 hours walk NW of Maji, 2200 m, De Wilde 6194 (BR, WAG); W. 
slope of Mount Uociacia, c. 15 km W. of Addis Ababa, 2700 m, De Wilde 
9580 (WAG); Mount Uociacia, c. 15 km W. of Addis Ababa, c. 2600 m, De 
Wilde 8532 (WAG); Kaffa Province, Limmu, Monti Botor, c. 2250 m, Pichi 
Sermolli 7069 (SRGH); 717'N, 365'E, Kaffa Province, 35 km W. of Bonga 
along the road to Shewa Ghimmira, 1950 m, Friis 211 \ (BR). 

KENYA: Nyambeni Hills, base of Kirima, 6400 ft, Polhill & Verdcourt 
295 (BR, K, PRE); Aberdares, Cave Waterfall, Coe 794 (PRE); Molo, Mail 
Escarpment, 2440 m, Alluaud 55 (BR, P, PRE); Kiambu District, Katamayu 
River Forest, 2200-2250 m, Faden & Evans 69/236 (BOL); Meru District, 
Ngambeni Hills, above Kiegoi, 2250 m, Faden et al. 69/678 (K); Aberdares, 
S. Kinangop, 8600 ft, Molesworth-Allen 3637 (K); Kericho District, crossing 
at the Kitinges River, c. 8 km ENE of Kericho, 2060 m, Faden 72/302 (K); 
Kericho District. W. Mau Forest, SW of Mt. Blacket, Faden et al. 72/356 (K); 
Samburu District, Nyiro Mountain, 2400 m, Bono 23 (K); Taita Hills, Vuria 
Hill, 1920-2200 m, Faden 72/255 (BOL, K): Kinangop, Brown Trout Inn, 
9000 ft, Verdcourt 880 (K); S. Kinangop, near Brown Trout Inn, Molesworth- 



51 

Allen 3620 (K); Kinangop, above Isanga farm, 8500 ft, Andrews 4461 (K); 
Chyulu Hills, 6800 ft, Van Someren 7572 (K); Chyulu Hills, 2250 m, Bally 
1 163 (K); Mount Meru, 5000-6000 ft, Leighton s.n. (K); Aberdare Range, 
base of Mount Kenya, Dawson 96a (K); Taita Hills, Vuria Forest, c. 7000 ft, 
Schippers K27 1 (WAG); Thompson Falls, c. 7600 ft, Schippers K 1 7 (WAG); 
Kinangop, Brown Trout Inn, Verdcourt & Moggi 2486 (B, SRGH); Elgon 
Forest, Webster 9055 (K); prope 'West Kenia Forest Station', 2300 m, Friis 
594 (B, BR, S); Mount Elgon, 4300 ft, Barrele 92 (NU); Aberdare Range, 
near W. part of the Nyeri track, 3 1 00 m, Hedberg 1 533 (S); Samburu District, 
Mt. Nyiru, 8000 ft, Cameron 147 (BR). 

LESOTHO. 2927 (Maseru): gorge dans la montagne Ma-Khrarane, au 
dessus de la station missionaire de Morija (DA), Dieterlen 1309 (P, PRE). 

MALAWI: Nyika, Zovochipolo, 2225 m, La Croix 4634 (PRE); Nyika 
Plateau, Zovochipolo forest patches, 2200 m, Dowsett-Lemaire 297 (MAL); 
Kirk Range, Dzonze Forest, 1750-1800 m, Dowsett-Lemaire 1079 (K); 
Mwanembu Mountain, McClounie 6 (K); S. region, Malosa Mountains (N. of 
Zomba), 1900 m, Dowsett-Lemaire 973 (K); Mount Mulanje, Tuchila Pla- 
teau, 6000 ft, Newman & Whitemore 214 (SRGH). 

MOZAMBIQUE: Penhalonga Waterfall, Chase 3247 (NU, SRGH); 
Penhalonga Forest, Chase 3219 (SRGH). 

RWANDA: Plantation Gasiza au N. de Ruhengeri au pied Ngahinga et du 
Muhavura, 2350 m, Van der Veken PV 10265 (B, BR); Kirunga Vulcan, 2500 
m, Poetsen 8 1 (B); Dalinghi, Zappelli 262 (BR); Kissenyi, Sake, Zappelli 1 77 
(BR); Chaine des Birunga, pied SE du Gahinga, 2500 m, Lambinon 74/1 534 
(BR). 

SOUTH AFRICA. 2329 (Pietersburg): Louis Trichardt, Hanglip Forest 
Station (BB), Roux 2572 (NBG); Louis Trichardt, Zoutpansberg Suds, farm 
Rustfontein, c. 1400 m, Schlieben 7342 (BR); Tzaneen, Dap Naude Dam, 
Woodbush (DD), Burrows 3269 (BOL, PRE). 2330 (Tzaneen): Duiwelskloof, 
Westfalia Estate (CA), Scheepers 419 (PRE); De Hoek Forest Station (CC), 
Roux 2563 (NBG); Woodbush Forest Reserve, Grootbos, Roux 2564, 2570 
(NBG); Magoebaskloof near De Hoek Forest Station, Van Jaarsveld 6093 A 
(BOL, NBG); Woodbush, Jenkins s.n. TM 919c (PRE); Woodbush, Wager 
s.n. CH7464 (PRE); Woodbush, Reynolds s.n. CHI 0246 (PRE); De Hoek, 
Schweickerdt s.n. (NBG, PRE); Pietersburg, Woodbush, Schelpe 6050 (BOL). 
2430 (Pilgrim's Rest): Mariepskop, base of Klaserie Waterfall (DB); Bur- 
rows 3113 (BOL, PRE); Mount Sheba Nature Reserve (DC), Roux 2556 
(NBG); Mount Sheba, Kluge 2320 (NBG); Mount Sheba Nature Reserve, 
Jacobsen 4420, 4428 (PRE); Ohrigstad Nature Reserve, Jacobsen 1413 
(PRE); Pilgrims Rest (DD), Collins s.n. TM895c (PRE); Graskop, Blyde 
River Forest Reserve, Jacobsen 4363 (PRE). 2530 (Lydenburg): Lydenburg, 
Coromandel farm (AD), Roux & Burrows 13 (BOL); Coromandel farm, 
Burrows 1309 (BOL); Sabie, Tweefontein (BB), Wager 48 (PRE); Sabie 
Gorge, Wager 25 (PRE); Sudwala Caves, forest 2 km N. of caves, 1500 m 
(BC), Kluge 2465 (PRE); Sudwala Caves, Burrows 3193 (BOL); Nelspruit, 
Witklip Staatsbos (BD), Kluge 853 (PRE); Kaapsehoop (DB), Wager s.n. 
TM149c (PRE); Lydenburg, Clivia Pass (DD), Edwards 1149 (NU). 2531 
(Komatipoort): Lomati falls and kloof behind Barberton (CC), Wager 151 
(PRE); Barberton, Williams 104 (P); Baberton, Thorncroft 104 (GRA). 2630 
(Carolina): Marieriestad (CA), Pott-Leendertz 4848 (BOL, PRE). 2730 
(Vryheid): Piet Relief- Wakkerstroom road, 6 km from turnoff to Liineburg 
(AD), Roux 2269 (NBG); Wakkerstroom, Oshoek, Devenish 2 (PRE). 2828 
(Bethlehem): Royal Natal National Park, Goodoo Forest (DB), Doidge s.n. 
(PRE); Tugela Valley, Hafstrom & Acocks 1970 (PRE); Royal Natal National 
Park, Devils Hoek, 5000 ft, Schelpe 7973 (BOL). 2829 (Harrismith): Van 
Reenens Pass (AD), Rehmann 7205 (B, P); Oliviershoek Pass, Begonia Falls 
(CA), Roux 2514 (NBG); Qualeni Valley, 800 ft (CC), Schelpe 7270 (NU, 
PRE). 2929 (Underberg): Cathedral Peak Forest Research Station, 6050 ft 
(AB), Killick 1 134 (PRE); Lions River, Lions Bush (BD), Moll 829 (PRE). 
2930 (Pietermaritzburg): Lidgetton (AC), Mogg CHI 764 (PRE); Lions 
River District, Karkloof, 'Braco', 4300 ft, Schelpe 51 19 (BOL); Lidgetton, 
Roberts 87 1 (PRE); Zwaartkop (CB), Sim s.n. PRE-9045 (PRE); Zwaartkop, 
4500 ft, Sim s.n. (NU); Pietermaritzburg, Ferncliff Nature Reserve, Crouch 
593 (NU). 3029 (Kokstad): Kokstad (CB), McLoughlin 753 (BOL). 3127 
(Lady Frere): Cala (DA), Young 5 1 1 (PRE); Engcobo (DB), McLoughlin s.n. 
CH7677 (PRE). 3128 (Umtata): Maclean farm Woodcliffs (AB), Roux 2482 
(NBG). 3129 (Port St Johns): Port St Johns (DA), Wager s.n. CH2905 
(PRE). 3226 (Fort Beaufort): Katberg Forest Reserve (BC), Roux 2700 
(NBG): Hogsback Forest Reserve, Fern Walk, 800 m (DB), Dahltrand 1853 



J.P. ROUX 



(PRE); Hogsback, Zingcuka Forest, Roux 2414 (NBG). 3227 (Stutterheim): 
Stutterheim, Isidinge Forest (CA), Roux 1982 (NBG); Keiskamma Hoek, Ely 
526 (PRE); Cathcart, Fort Cunnyngham Forest Reserve (CB), Roux 243 1 
(NBG); Pirie, Sim s.n. TM5 14 (PRE); Kingwilliamstown, Pirie Forest along 
Amatola trail, Roux 2708 (NBG). 3325 (Port Elizabeth): Johana Kloof 
(BC), Breutel s.n. (L). 3419 (Caledon): Riviersonderend, farm 'Oubos' 
(BB), Roux 2585 (NBG). 

SUDAN: Gilo, Imatong Mountains, Ngairigi River, 5000 ft, McLeay 455 
(K). 

TANZANIA: Station Kyimbila, Fundort Rungwe, 1300 m, Stolz 889 (B, 
L, P, S, WAG); Kilimanjaro, environs de Kibosho, 2500 m, Daubenberger 
s.n. PRE-6788 (PRE); Kilimanjaro, Kibosho, 2000-4000 m, Daubenberger 
35 (B, BR, P, PRE, S); Mt. Meru, NE end of the caldeira wall, c. 8500 ft, 
Greenway & Fitzgerald 13613 (K, PRE); Kilimanjaro-Slid, c. 1900 m, 
Schlieben 4596 (BOL, BR, PRE, SRGH); region de Kilimanjaro, environs de 
Kibosho, Kilema-Machame, Daubenberger s.n. (BR, P); Kilimanjaro, 2800 
m, Alluaud 310 (P); British East- Africa, forets de Lamoru, Le Petit s.n. (P); 
forets de plateau Kikuyu, 2000 m, Le Petit s.n. (P); Usambara Mountains, 
Mahali Mountains, 6000 ft, Newbould & Jefford 1731 (K); Morogoro, 
Glover 268 (K); Marangu, SE Kilimanjaro, 4600 ft, Beesley 14 (K); Mbeya 
District, Kikondo camp, Poroto Mountains, 1950m, Richards 13972(B,BR, 
K); Moshi District, Kilimanjaro, c. 1900 m, Schlieben 4596 (K); Arumeru 
District, banks of Engare Olmotonyi River, c. 4 km N. of Olmotonyi Forestry 
Institute, Mtui 143 (K); Mount Meru, Engarenanyuki, 7600 ft, Vesey- 
FitzGerald 3031 (K); Mbeya District, Mount Kikondo, 6500 ft, M.R. 13972 
(K); Mount Meru, end of Olmotonyi, Schippers T778 (WAG); Mount Meru, 
2090 m, Schippers T729 (WAG); South Pare Mountains, Mugambo Forest 
Reserve, 1480 m, Schippers T95 1 (WAG); W. Usambara Mountains, on hill 
above Shume Forest Meteorological Station, 2050 m, Schippers T1506 
(WAG); Kilimanjaro, oberhalb Marangu, Volkens 1266 (B); Kilimanjaro, 
1900 m, Schlieben 4596 (B); Usambara, Lutindi, Liebush s.n. (B); Kondoa- 
Frangi, Ndiomeberg, 1800 m, Ledemann s.n. (S); Kilimanjaro, above 
Marungu, 2000 m, Pedersen 527 (BR); Morogoro Mountains, 2300 m, 
Chisongela 9 (BR). 

UGANDA: near Luhiza-Kigezi, 7000 ft, Rose 10311 & 10312 (K); forest 
near Mt. Debasien, 6000 ft, Eggeling 2683 (K); Ruwenzori Mountains, 7000 
ft, Hazel 1 14 (K); Luhiza-Kigezi, 7000 ft, Rose 10299B (K). 

ZAMBIA: Nyika Plateau, Chowe Forest, 2100 m, Dowsett-Lemaire 220 
(K). 

ZIMBABWE: near Umtali, Holland s.n. (NBG); Vumba Mountains, near 
Umtali, 6000 ft, Obermeyer 2099 (K, PRE); Melsetter, Bridal Veil Falls, 
Jacobsen 3087 (PRE); Melsetter, in gulley border of 'Skyline' & 'Thornton' 
areas, Chase 7482 (BOL, K); Umtali, Banti south, 5800 ft, Jacobsen 3864, 
3879 (SRGH); Inyanga, above Pungwe rest hut 2, 5300 ft, Chase 5655 (BOL, 
PRE, SRGH); Melsetter, Musapa mountain, Grosvenor 264 (BOL, SRGH); 
Melsetter, Gwendingwe, Muller 2880 (SRGH); Umtali District, Cashel, 
Black Mountain Inn, Chase 4021 (NU); Melsetter District, Bridal Veil Falls, 
Chase 4020 (NU); Inyanga, Patterson 24 (GRA); Inyanga, Pungwe Rest 
Huts, 5300 ft, Schelpe 5679 (BOL). 

WITHOUT EXACT LOCALITY: Natal, Tyson s.n. CH2168 (PRE); 
Natal, Gerrard 1931 (P); Zululand, Gerrard & McKen s.n. (P); Zimbabwe, 
Wild 1470 (K); loco incerto, Mann 2067 - pro parte (K); near Bamenda, 7500 
ft, Migeod3&3 (K); Natal, sine coll. s.n. (NBG); Natal, Buchanan 27 (B-only) 
(M); Natal, Plant 328 (B); Natal, Buchanan 75 (B); loco incerto, Bergius s.n. 
(B); Rebfall bei Gaffat, Haidner s.n. (B, S); Kissenye, Ninagongo, 2500- 
2900 m, Mildbraed 1341 (B); loco incerto, Hoist 3837 (B); loco incerto, Sim 
s.n. CH4171 (PRE); Gold Fields, Ayres s.n. (NH); Burungo, De Witte 1472 
(BR); Natal, Holub s.n. (BR); Kikuku, 1750 m, Ban 367 (BR); Natal, 
Buchanan s.n. (BOL). 

Polystichum transvaalense and P. wilsonii are often confused. Pichi 
Sermolli (1977, 1985) ascribed material of P. transvaalense to P. 
fuscopaleaceum Alston var.fuscopaleaceum, while Schelpe (1967, 
1970, in part) and Jacobsen (1978) ascribed material of this species 
to P. setiferum var. fuscopaleaceum (Alston) Schelpe. Jacobsen & 
Jacobsen (1989), however, considered P. fuscopaleaceum and P. 
transvaalense to be conspecific. 

Aware that two forms exist, Schelpe (1967) concluded that no 
clear differentiation at the specific level was possible and considered 



plants with dark stipe base paleae as P. setiferum var.fuscopaleaceum. 
This classification was largely followed by Jacobsen (1978), al- 
though he considered the high elevation collections a Drakensberg 
form. Although he ascribed several collections to this form he 
refrained from giving it any formal taxonomic status. Pichi Sermolli 
(1977), however, considered P. fuscopaleaceum distinct from P. 
setiferum. He also recognized two 'altitudinal vicariants' with P. 
fuscopaleaceum var.fuscopaleaceum occurring at lower elevations 
than P. fuscopaleaceum var. ruwensoriense, a subdivision he re- 
tained in 1985. I consider the two groups sufficiently distinct to 
warrant specific status, a conclusion supported by the discovery of 
a sterile hybrid between these putative parents. 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Diagnostic of 
Polystichum transvaalense is its confinement to moist forests, the 
presence of up to 17 caespitosely arranged, suberect to arching 
fronds that may reach a length of up to 1 .045 m on a short erect to 
suberect rhizome, the stipe, rachis and pinna-rachises bearing mostly 
ferrugineous, twisted and somewhat shrivelled, proximally lacerate 
paleae terminating in an acicular cell, and the mostly fimbriated but 
often erose and rarely repand indusium. The perispore is highly 
porate. 

Polystichum transvaalense appears to be most similar morpho- 
logically to P. wilsonii H. Christ; the two belong to section 
Lasiopolystichum. An analysis of the differences between these 
species is provided under P. wilsonii. 

VARIATION. Polystichum transvaalense shows considerable varia- 
tion in the length of the frond, stipe, lamina and basal pinna (Table 
2). This may be ascribed to the diverse altitudes, climates and 
vegetation types it occupies throughout its broad range. The species, 
however, shows little variation in pinnule outline and palea struc- 
ture, distribution and density. Stipe-base paleae are mostly 
ferrugineous, but in rare cases the larger paleae are densely 
impregnated with secondary compounds giving them a dark brown 
colour. The indusium also shows significant variation in size, shape 
and the presence or absence of central processes. Basal pinnae may 
be deflexed or not. Possible causes of these variations in plants 
occurring in close proximity under similar growing conditions 
remain unknown. 



Table 2 Variation in frond, stipe lamina and basal pinna length in 
Polystichum transvaalense N.C. Anthony. 



Ramge (mm) 



x (S.D.) 



Frond 


212-1045 


713.9(204.1) 


41 


Stipe 


77-535 


283.3(101.6) 


42 


Lamina 


135-670 


422(116.1) 


52 


Pinna 


25-140 


86.3 (27.2) 


52 



DISTRIBUTION AND ECOLOGY. Polystichum transvaalense is widely 
distributed in temperate and tropical Africa. The distribution largely 
follows the escarpment and mountain ranges on the eastern parts of 
the continent. In South Africa it occurs from the Drakensberg 
foothills in the Eastern Cape along the KwaZulu-Natal Drakensberg 
escarpment, the Eastern Cape and southern KwaZulu-Natal mid- 
lands, the Free State-KwaZulu-Natal and Mpumalanga-Northern 
Province escarpments to the Soutpansberg in the Northern Province. 
A single collection is also known from the foothills of the 
Riviersonderend Mountains in the southern Cape. In Zimbabwe it is 
found in the Chimanimani and Vumba Mountains extending to the 
Zomba Plateau, the Kirk Mountains and the Nyika Plateau in 
Malawi and Zambia. In Tanzania it occupies the Uluguru- and 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



53 



Usambara Mountains, Mt. Meru and Mt. Kilimanjaro. Further north 
it occurs in the mountainous areas of Kenya, Uganda, Ethiopia, the 
Imatong Mountains in Sudan and the Kivu Ridge in the Democratic 
Republic of Congo. A disjunction in this pattern arise in that P. 
transvaalense is also found on Mt. Cameroon and Bioko in the Gulf 
of Guinea. 

The lithology, climate and vegetation associated with Polystichum 
transvaalense varies considerably throughout its range. In the south- 
ern Cape it occupies isolated forest patches in Mesic Mountain 
Fynbos (Moll et al., 1984) at c. 365 m in acidic sandy soils and on 
rocks of the Table Mountain Sandstone formation. Polystichum 
transvaalense is an exclusively forest growing species, generally 
growing as individuals on streambanks or on rocks along streams 
but rarely also as low-level epiphytes. Polystichum transvaalense 
occurs in several forest types as defined by Acocks (1988). In the 
Eastern Cape it is found in Pondoland Coastal Plateau Sourveld 
forests and rarely in Typical Coast-belt forests. In the Natal mid- 
lands and Drakensberg escarpment where it may appear at elevations 
up to 1840 m it occurs in 'Ngongoni Veld forests and Highland 
Sourveld forests. In Mpumalanga these forests are replaced by 
Northeastern Mountain Sourveld forests that extend to the 
Soutpansberg. North of the Limpopo the species is found at eleva- 
tions ranging between 1300 and 2825 m in largely Undifferentiated 
Afromontane forests, as defined by White (1983), but often also in 
single-dominant Afromontane forests such as Juniperus procera 
forests at 2400 m in Ethiopia and in Afromontane bamboo at 1 828 m 
on Mt. Malati in Tanzania and at 2745 m at Kinangop in Kenya. 

8. Polystichum wilsonii H. Christ in Bot. Gaz. 51: 353 (1911). 

Type: China, Szechuan Province, Mupin, woodlands, 4000- 

6000 ft, Wilson 2614 (BM!-holotype). 
Fig. 9. 

Polystichum lobatum var. ruwensoriense Pirotta in L.A. di Savoia, // 
Ruwensori I: 478 (1909). Type: Ruwenzori, nella foresta 
scendendo da Kichuchu a Nakitava, Roccati et Cavalli-Molinelli 
s.n. (TO-holotype). 

Polystichum aculeatum var. mildbraedii Brause in Bot. Jahrb. Syst. 
53: 379 (1915). Type: Fernando Po (Bioko), Nordseite des Pics 
Sta. Isabel oberhalb Basile, Grasflur-Region des Gipfels mit viel 
Ericinella, zwischen Gras, c. 2700 m, Mildbraed 7180 (B!- 
holotype). 

Polystichum aculeatum var. rubescens Bonap., Notes Pterid. 14: 

214 (1923). Type: Tanzania, Kilimanjaro, zone super des forets, 
2760 m, Alluaud 48 (P!-holotype). 

Polystichum aculeatum var. stenophyllon Bonap., Notes Pterid. 14: 

215 (1923). Type: Kenya, Mont Kenya, versant ouest, foret 
interfere, 2400 m, Alluaud 241 (P!-holotype). 

Polystichum fuscopaleaceum Alston in Bol. Soc. Brot. ser. 2, 30: 22 
(1956). Type: Cameroon, Victoria District, Cameroon Mountain, 
SW of hut 2, in gully woodland, 9100 ft, Keay FHI 28602 (BM!- 
holotype). 

Polystichum setiferum var. fuscopaleaceum (Alston) Schelpe in Bol. 
Soc. Brot. ser. 2, 41: 216 (1967). 

Polystichum fuscopaleaceum var. ruwensoriense (Pirotta) Pic.Serm. 
inWebbia32:90(1977). 

Polystichum alticola Schelpe & N.C. Anthony in Contr. Bolus Herb. 
10: 144 (1982). Type: South Africa, Ladismith, Swartberg, 
Toverkop, 2160 m, Esterhuysen 26699 (BOL!-holotype; NBG!, 
PRE!, isotypes). 

Plants terrestrial or epilithic. Rhizome short, to 130 mm long, erect 
to suberect, to 10 mm in diameter, rarely branched, set with roots, 
crowded, persistent stipe bases, and paleae; paleae broadly attached, 



castaneous, chartaceous, narrowly linear, the margins with small, 
widely spaced cellular outgrowths, the apex mostly terminating in 
an acicular cell, rarely in a small thin-walled cell, to 12 x 1 mm. 
Fronds caespitose, 8-12 per plant, suberect to arching, to 1.05 m 
long; stipe proximally castaneous, stramineous distally, adaxially 
sulcate, to 450 mm long x 5 mm in diameter, sparsely to densely 
paleated; larger paleae broadly attached, often slightly bullate, 
spreading, extending to the rachis, concolorous or bicolorous, 
chartaceous to crustaceous, broadly ovate-acuminate to ovate-acu- 
minate, cordate to cordate-imbricate, the margins widely to closely 
fimbriate, fimbriae generally straight, the apex entire, terminating in 
an acicular cell, to 23 x 9 mm; smaller paleae apically or basally 
directed, stramineous, chartaceous, narrowly triangular to subulate, 
short-stalked, often somewhat auricled, the margins proximally 
with long straight, angular or curved outgrowths, distally with few 
widely spaced, short or long marginal outgrowths, the apex entire, 
terminating in an acicular cell, to 13 x 7 mm: lamina 2-pinnate, with 
up to 29 free pinna pairs, herbaceous to firmly herbaceous, pale to 
dark green adaxially, paler abaxially, narrowly elliptic, to 625 mm 
long, the proximal pinnae reduced, deflexed: rachis stramineous, 
adaxially sulcate, densely set with paleae similar to but smaller than 
those on the stipe, paleae restricted to the abaxial surface, to 9 x 3 
mm: pinnae 1 -pinnate, with up to 12 free pinnule pairs, proximally 
widely spaced, distally closely spaced and somewhat overlapping, 
folded ventrally along the rachis (conduplicate), narrowly triangular 
to oblong-attenuate, the proximal pinnae to 88 mm long x 20 mm 
wide: pinna-rachis stramineous, adaxially sulcate, densely paleated; 
paleae short- stalked, narrowly ovate to narrowly triangular, the 
margins proximally with long straight or angular outgrowths, apically 
with few widely spaced short or long outgrowths, the apex entire, 
terminating in an acicular cell: pinnules asymmetric, acroscopically 
auriculate, narrowly trullate to trullate, to 12 mm long, serrate, long- 
aristate; adaxially with straight or slightly twisted filiform paleae, 
simple or proximally with short straight or curved marginal 
outgrowths, the apex terminating in an acicular cell; abaxially with 
straight or proximally somewhat twisted, subulate-hastate paleae, 
the margins with short straight or angular outgrowths at the base, the 
apex entire, terminating in an acicular cell. Venation raised. Sori 
circular, c. 1 mm in diameter, terminal or near terminal on abbrevi- 
ated vein branches: sporangium with 1 1-(15)-24 indurated annulus 
cells; stalk eglandular: indusium stramineous, peltate, circular or 
reniform, repand to erase, often with small central processes, per- 
sistent, the maximum radius 0.51-(0.75)-1.09 mm. Spores 64 per 
sporangium, brown, the perispore smooth or tuberculate, spiculate, 
closely perforated, the exospore 32-(41.74)-52 x 24-{30.16)-40 
um. Chromosome number 2n=164. 

MATERIAL EXAMINED 

BIOKO: Fernando Po, Mann s.n. (K); cratera del pico Basile, km 23, junto 
a la cumbre, 3000 m, Carvalho 3652 (B, BR). 

CAMEROON: Mt. Cameroon, 3700 m, Breteleret al. 69 (K, P, WAG); 
Mt. Cameroon, 1950 m, Breteleret al. 75 (K, P, WAG); Bambutos, 2600 m, 
Felix 5430 (P); Mt. Cameroon, 3600 m, Annet 126 (P); Cameroon, mont 
versant, 3000 ft, Meurillon 1 158 (BR, K, P); piste du village d'Okon au mert 
Okon, 3008 m (45 km SSO de Nkambe), Letouzey 8940 (K, P); Mt. Cameroon, 
haut plateau, 3600 m, Annet 128 (P); Bambutos, station mi-ombragee, vers 
2300 m, sine coll. 30 (P); Mt. Cameroon, 7-10000 ft, Mann 1376 (K); 
Cameroon Mountain, above 2nd hut, 12000 ft, Hutchinson & Metcalfe 48 
(K); Mt. Cameroon, 11000 ft, Steele 22, 27 (K); Mt. Cameroon, 11000 ft, 
Migeod 190 (K); Mt. Cameroon, oberhalb Buea, 2800 m, Mildbraed 10883 
(B, K); Buea, 3000 m, Preuss 787 (B), 788 (B, S); Kamerun-Berg, standort 
iiber Buea, unteren Fako Plateau, 2800 m, Mildbraed 3377 (B); 
Kamerungebirge, Buea, Deistel s.n. (B); Kamerun-pitz, 3500-3600 m, 
Bornmuller 26 (B); Mt. Cameroon, 3800 m, Hintz 29 (B); Buea, Wonjombia 
faco, Reder 1026 (B). 



54 



J.P. ROUX 




Fig. 9 Polystichum wilsonii. A, proximal part of lamina; B, rhizome; C, adaxial surface of pinnule; D, abaxial surface of fertile pinnule. All drawn from 
Roux 2529 (NBG). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



55 



DEMOCRATIC REPUBLIC OF CONGO: Kivu District, Virunga 
west, Nyamuragira, Stauffler 178 (BR, PRE); P.N.A. Kabara, flanc N. du 
volcan Karisimbi, 3000 m, Jean Louis 5301 (BR, K, P); Volcan Karisimbi (au 
NE du lac Kivu), 3500 m, Humbert 8563 (BR, P); Kivu, Volcan Mikeno, 
2500-3400 m, Humbert 8010 (BR, P); Mt. Kinangop, 2800 m, Alluaud 262 
(P, S); P.N.A. Nyamlagira, 3000 m, Germain 3476 (BR); Kivu District, SW 
side of Mt. Mikeno, 10500 ft, Chaplin 373 (BR); 129'S, 2926'E, Goma 
territory, versant ouest du Karisimbi, 3360 m, Bamps 2984, 2995 (BR); Pare 
National Albert, ruisseau affl. de la Nososa, (a Test de Mahungu), 3180 m, 
Fredericq 9152 (BR); Viroenga Park, Karisimbi-massif, 3465 m, Van der 
Veken 9130, 9135 (BR); Kivu, upper Ruamoli Valley, 1 180 ft, /to 77 8 (BR); 
Viroenga Park, Karisimbi-massif, 3 km van de gite Rukumi, 3330 m, Van 
der Veken9\40 (BR); massif du Karisimbi, a 500 m env. du gite de Rukumi, 
3500 m,Auquier 2290 (BR); Kivu Province, upper Ruamoli Valley, 12 500 ft, 
Ross 743 (BR); Pare National Albert, selle de Kabora entre le Karisimbi et le 
Mikeno, 3000-3100 m, Lebrun 7344 (BR). 

ETHIOPIA: Bale Region, 45 km N. of Goba, Sannetti Plateau, 3900 m, 
Tadesse 5545 (ETH); SE of Dinsho on road from Goba to Shashemene near 
proposed HQ of Bale National Park, 10400 ft, Gilbert 1812 (ETH, K); Bale 
Province, Bale Mountain National Park, E. of Garba Goracha camp, 4070 m, 
Hedberg 5649 (ETH); pass just N. of the summit of Cara Mulatta Mountain, 
c. 10200 ft, Burger 1480 (ETH); Choke Mountain, Gojjam, vicinity of the 
upper Ghiedeb Valley, Flenley & Evans 327 (ETH, K), Begemdir Province 
Simien, Buahit, 3870 m, Hedberg & Aweke 5461 (ETH); Bale Mountains, 
Finchaya Habera, 3510 m, Miehe 266, 335 (ETH); Bale Mountains, above 
Goba, 3500 m, Miehe 3086 (ETH); Bale Mountains, above Rira, 3530 m, 
Miehe 2343 (ETH); Shoa Province, Arussi Mountains, 35 km S. of Mount 
Chillalo 32 km on track to Ticcio via Robie turnoff, 35 km S. of Asella, 3275 
m, Ash 2330 (ETH); Bale region, Mendeyou Auraja, c. 5-7 km on Fincha 
Haberra-Soddota track, 3500-3580 m, Tadesse 7813 (ETH); Shoa, Lake 
Wonchi, outer rim of caldeira, Gilbert & Tewolde 3279 (ETH, K); Bale 
region, Mendeyou Awraja Fincha Heberra, 3490-3510 m, Tadesse 7713 
(ETH); in rupibus umbrosis Demeski, 10500 ft, Schimper 244 (P); Matssehe 
Dedschem, 12000 ft, Schimper 1398 (P); Arussi Prov., Chillalo Awraja, 
Galama Mountains (30 km ESE of Asella) c. 3 km E. of Boraluco, 3750 m, 
Hedberg 4233 (K, PIC.SERM.); Bale region, Dello Awraja, c. 3.7 km N. of 
Kecha towards Rira, 2620 m, Tadesse 5143 (ETH); Shewa region, Wonchi 
Mountains, edge of volcanic crater 20 km to SSE of Ambo, Pavlov & Petelin 
138 (ETH); in regio media montis, Schimper 180 (B, K, P, S); Begemder 
Province, Semian Mountains, De Wilde 175 (BR, WAG); Mt. Borullccu, 
along road to Ticco c. 30 km SE of Asella, c. 4000 m, De Wilde 9039 (WAG); 
c. 25 km SE of Asella, W. slope of Mt. Boruluccu, c. 3800 m, De Wilde 8089 
(WAG); c. 3 km E. of Asella, c. 175 km SSE of Addis Ababa, W. slope of Mt. 
Cilalo, c. 2700 m, De Wilde 6623 (BR, WAG); Shoa Province, Menagesha 
State Forest on the W. slope of Mt. Wuchacha, 2600 m, Friis 1 209 (K); c. 1 75 
km from Addis Abeba on Dessie road, 10500 ft, Gilbert 454 (K); Bale region, 
10-15 km SE of Goba on road towards Masslo, 3200-3400 m, Thulin 3678 
(K); Gara Mullato Mountains, 10800 ft, Burger 1907 (K); Scioa, Monte 
Wochacha, 3250-3300 m, Pichi Sermolli 6740 (B, BR, K); Mussolini Pass, 
between Dera Sina and Debra Berhan, c. 3000 m, De Wilde 9654 (WAG); 
Ethiopia, Schimper 1398 (B); ad rupes locis humidis umbrosis pr. Demerts, 
10500 ft, Schimper 244 (B); Arussi, Catena dei Monti Galamo-Sagatu, c. 
3100 m, Pichi Sermolli 6828 (BR). 

KENYA: Mt. Kenya, forest end, 9600 ft, Mclaughlin 676 (BOL, PRE); 
Mt. Elgon, versant est, Arambourg et al. 134, pro parte (P); W. slopes of Mt. 
Kenya, along the trail from West Kenya Forest Station to summit, c. 3630 m, 
Mearns 1421, 1502, (P); W. slopes of Mount Kenya, along trail from West 
Kenya Forest Station to summit, c, 3000 m, Mearns 1702 (B, P, S); Mt. 
Kenya, c. 7500 ft, Schippers K164 (WAG); Aberdares National Park, c. 
10500 ft, Schippers K110 (WAG); Aberdares National Park, c. 10300 ft, 
Schippers K78 (WAG); Shira Plateau, W. Kilimanjaro, 3200 m, Schippers 
T1052 (WAG); Mt. Meru, c. 2000 m, Schippers T777 (WAG); Mt. Kenya, 
Kinangop, Aberdare, 8800-8900 ft, Chandler 2266 (K); Aberdare Moun- 
tains, James s.n. (K); Aberdare Mountains, Ramsden s.n. (K); Aberdare 
range, near W. part of the Nyeri track, 3 1 00 m, Hedberg 1533 (K); near Molo, 
Mau Forest, 8000 ft, Gardner 975 (K); Narok District, 20 miles from 
Olokurto on road to Elburgon, c. 9600 ft, Glover et al. 1096 (K); SE 
Aberdares, Kitikuya, 8500 ft, Gardner s.n. (K); Rift Valley, Nakuru District, 
E. Mau Forest Reserve, 2750 m, Geesteranus 5908 (BR, K, L, PRE, S); Mt. 



Elgon, E. slope above Tweedie's saw-mill, 2550 m, Hedberg 68 (K, S); North 
Forest, 9200 ft, Schippers K364 (WAG); Mt. Aberdare, c. 3200 m, Fries 2644 
(B); Mt. Elgon, 2700 m, Gravik s.n. (S); Mt. Elgon, 3800 m, Gravik s.n. (S); 
Mt. Kenya, Sagana Valley, 10500 ft, Schelpe 2713 (BR); Mt. Aberdare, pr. 
'West Kenya Forest Station', 2350 m, Fries 775 (BR); Mt. Kenya, 10000 ft, 
Meyerscough K2, K3, K16, K18, K22, K26, K30, K31 (BOL). 

LESOTHO. 2828 (Bethlehem): Butha Buthe District, Khatibe B camp, 
9500 ft (DC), Troughton B26 (GRA); Leribe, Dieterlen 167 (BOL, P, pro 
parte); 1 km from Moteng store, Roux 1 294 (NBG). 2927 (Maseru): between 
Blue Mountain Pass and Likholaneng, 8700 ft (BD), Schmitz 7266 (PRE); 
Morija, Dieterlen 1309 B-only (PRE). 2928 (Marakabei): Mamalapi, 8000 
ft (AC), Jacot-Guillarmod 690 (PRE); Mamalapi, 9000 ft, Compton 21331, 
21334, 21339 (NBG); hill at Bushmen Pass, beyond little Bokong, 9000 ft, 
Bevis 102 (PRE); Blue Mountain Pass, Roux 2227 (NBG); mountain road, 60 
miles from Maseru, 8000 ft, Bowmaker 23 (BOL); mountain road, 38 miles 
from Maseru, 7500 ft, Bowmaker 25 (BOL); Lehaha-la-Sekhomgana, 9 1 00 ft 
(AD), Jacot-Guillarmod 206 (PRE); Semonkong, waterfall gorge, c. 7000 ft 
(CC), Davidson 3023 (PRE); Semonkong, at Le Bihan Waterfall, Roux 1493 
(NBG). 2929 (Underberg): Between Mokhotlong and Sani top, 15 km 
from Mokhotlong, 2200 m (AC), Matthews 887 (NBG, PRE); 15 km past 
Thaba-Tseka turnoff on Sani road, Roux 1 344 (NBG); Sehlabathebe National 
Park (CC), Schmitz 7122 (PRE); Sehlabathebe National Park, Matthews 987 
(NBG); Sehlabathebe area, on way to Devils Knuckles, 9500 ft, Davis 181 
(NU); Sehlabathebe area, Devils Knuckles, c. 9000 ft, Davis 176 (NU); 
Sehlabathebe National Park, 2250 m, Hoener 1658 (BOL). 

SOUTH AFRICA. 2730 (Vryheid): Wakkerstroom, Oshoek, 6400 ft, 
(AD), Devenish 195, 638 (PRE). 2731 (Louwsburg): Nongoma, c. 1000 ft 
(DC), Tosh s.n. (NU). 2828 (Bethlehem): Clarence (CB), Van Hoepen s.n. 
TM 18230 (PRE); gully between the Witches and the Sentinel (DB), Roux 
1906, 2529 (NBG); versant N. du Mont-aux-Sources, region de Witzies 
Hoek, c. 1800 m, Junod 14 (P); Royal Natal National Park (DB), Hafstrom & 
Acocks 1699 (PRE); Royal Natal National Park, Gudu Forest, Roux 2510a, 
2511 (NBG); Mont-aux-Sources, 8000 ft (DD), Dyke 5489a (NBG); Royal 
Natal National Park, Plowmans Kop, Aerck 1966 (S); Mont-aux-Sources, 
10000 ft, Sim s.n. TM521c (PRE); Mont-aux-Sources, Mogg 4222 (PRE); 
Mont-aux-Sources, 3100 ft, Marloth 2862 (BOL). 2829 (Harrismith): 
Harrismith, Platberg, Zig-Zag Pass, 1800 m (AC), Jacobsz 4715 (PRE); 
Harrismith, Platberg, Donkie Pass, 1850 m, Jacobsz 4729, 4730 (PRE); 
Platberg, 6800 ft, Roux 782 (NBG); Harrismith, Platberg, Roux 2521, 2524, 
2526 (NBG); Harrismith, farm Bosch Hoek (AD), Roux 892 (NBG); 
Harrismith District, farm Klavervlei (CA), Roux 876 (NBG); Oliviershoek 
Pass, S. of Seheletwane,/tou*2516, 2517 (NBG); MnWeni Pass, 8000-9000 
ft (CB), Esterhuysen 27838 (BOL); MnWeni area, Pinnacles Gully, 9000 ft, 
Esterhuysen 29595 (BOL); MnWeni area, Mbunduni scree, c. 6000 ft, 
Esterhuysen 27816 (BOL); Drakensberg, Injasuti area, 6500-8500 ft (CC), 
Esterhuysen 26045 (BOL, K, NBG, PRE); along Cathedral Peak path, 1550 
ft, Goetghebeur457l (BR, PRE); Cathedral Peak, Ruch 2030, 2300 (PRE); 
Cathedral Peak area, 5000 ft, Harding 38 (NU); Cathedral Peak Forest, 
Killick 981 (NU). 2929 (Underberg): Cathedral Peak Forest Research 
Station, 6100 ft (AB), Killick 981 (PRE); summit of Cathedral Peak, 7700 ft, 
Schelpe p.30 (NU); Cathedral Peak area, Cleft Peak path, 8000 ft, Schelpe 
557 (NU); upper Tsanatalana Valley, near Cleft Peak, 9800 ft, Schelpe 7227 
(BOL); Champagne Castle, Bayer 1443 (PRE), 1445 (NU, PRE); Giants 
Castle (AD), Symons 134 (PRE); Mpendhle Distr., Mulangane Ridge, above 
Carter's Nek, 7000-7300 ft (BC), Milliard & Burtt 1 695 1 (BOL, NU), 1 6969 
(BOL, NU, PRE); Mpendhle District, Highmoor Forest Reserve, ridge SE of 
Giants Castle, headwaters of Elandshoek River, c. 8100 ft, Milliard & Burtt 
1 6 1 92 (BOL, NU); near Rosetta, 5000 ft (BD), Thode s.n. (NBG); Drakensberg 
Garden State Forest Reserve, 9500 ft (CA), Van Jaarsveld 6531 (NBG); 
Garden Castle Forest Reserve, Mlambonya Valley, 6200 ft, Hilliard & Burtt 
14972 (BOL, NU); upper tributaries S. of Mkomazi River (CB), Hilliard & 
Burtt 15853 (NU, PRE); Bamboo Mountain, McClean 684 (PRE); Sani Pass, 
wet slope below waterfall, 6900 ft, Hilliard & Burtt 1 7976 (NU, PRE), 1 7983 
(BOL, NU, PRE); Sani escarpment, c. 9000 ft, Marker s.n. (GRA); Underberg 
District, 5-7 miles NNW of Castle View farm, headwaters of Mlahlangubo 
River, 8500 ft, Hilliard & Burtt 15331 (BOL, K, NU); headwaters of 
Mlahlangubo River, c. 7800 ft, Milliard & Burtt 13714 (NU); Underberg 
District, Gxalingenwa Valley between Sani Pass and Polela Valley, 7400 ft, 
Milliard & Burtt 1 7 1 99 (BOL, NU); Ndumeni area (CC), Everson s.n. (BOL); 



56 

Bulwer (DD), Allsopp 850, A-only (NU); Bulwer, Henkel s.n., A-only (NU); 
Xumeni Forest, Rycroft 519 (NU). 2930 (Pietermaritzburg): Nottingham 
Road District, 'Drayton', 5400 ft (AC), Smith 147 (NU); York, 'Benuie', c. 
4000 ft (AD), Fisher 1040 (NU); Impendhle, Boston, 4500 ft (CA), Beanie 
77 (NU); Pietermaritzburg, Zwaartkop (CB), Sim s.n. (NU, PRE). 3027 
(Lady Grey): Lady Grey, mountain left of summit of Jouberts Pass on road 
to Barkley East (CA), Roux 1 136 (NBG); Wittebergen, Ben McDhui, 9550 ft 
(DB), Galpin 6934 (BOL, GRA. PRE), 6935, 6939 (BOL, PRE); road 
between Naude's Nek and Ben McDhui, Roux 1180 (NBG); Barkley East 
District, Ben McDhui, Bell River Gorge, c. 8000 ft, Milliard & Burn 16526 
(BOL, K, NU); zwischen Passtrasse Maclear und Naude's Nek, Werdermann 
& Oberdieck 1118 (B); Barkley East District, Ben McDhui, 9550 ft, Galpin 
6939 (B); Ben McDhui, c. 9000 ft, Milliard & Burn 16406 (BOL, NU); Ben 
McDhui, 8900 ft, Milliard & Burn 16495 (NU). 3028 (Matatiele): near 
summit of Ongeluks Nek Pass (AD), Roux 1383 (NBG); Rhodes, Naude's 
Nek Pass (CA), Roux 2475, 2477 (NBG). 3029 (Kokstad): upper slopes of 
Inungi Range, Matatiele, c. 5500 ft (CA), Acocks 12207 (PRE); Kokstad 
(CB), McLoughlin S38 (PRE); Mt Currie Nature Reserve, Kokstad, Crouch 
511 (NU); Kokstad, McLoughlin 746, 753 (BOL). 3030 (Port Shepstone): 
Oribi Gorge (CB), Slinger59 (NU). 3127(Lady Frere): Barkley Pass between 
Elliot and Barkly East (BB), Roux 2469 (NBG); Bastervoetpad, between 
Ugie and Barkley Pass, Roux 247 1 , 2474 (NBG). 3128 (Umtata): summit of 
Biziya Mountain, 1250 m (AD), Stever 898 (PRE). 3225 (Somerset East): 
near Somerset East (DA), MacOwen s.n. (P). 3226 (Fort Beaufort): Upper 
Zwart Kei, Mount Hope farm, 5300 ft (BC), Galpin 5621 (GRA, PRE); 
Katberg Pass summit, farm Pleasant View, Roux 2698 (NBG). 3319 (Worces- 
ter): Hex River mountains, shale band between Buffels Dome and Milner 
Peak, 5000 ft (AD), Esterhuysen 28708 (BOL, NU, PRE); Roodeberg 
(Matroosberg group), 6000 ft (BC), Esterhuysen 27695a (BOL); Worcester 
Division, shale band below Milner Peak, 5000 ft (CB), Esterhuysen 14885 
(PRE); Hex River Mountains, Moraine kloof, 4000 ft (DD), Esterhuysen 
28075 (BOL). 3321 (Ladismith): Swartberg near Ladismith, Toverkop 
(AD), Esterhuysen 28241 (BOL). 

TANZANIA: Mt Meru, Arumeru District, Gereau 1623 (PRE); 
Kilimandjaro-Siid, Korongo, c. 3000 m, Schlieben 4869 (B, K, PRE, SRGH); 
Kilimanjaro, tra la Peters Hut a la Bismarks Hut, c. 2900 m, Pichi Sermolli 
5 1 36 (BR, K, P); Ob. Urwald iiber Kibosho, c. 2800 m, Uhlig 1 85 (B, K); Mt. 
Meru, W. slopes above Olkakola Estate, 3300 m, Medberg 2306 (K, S); 
Kilimanjaro, above Marungu, c. 2 km from Peter's Hut, 3700 m, Hedberg 
1288 (BR, K, S); Mbeya, Kilando, 8000 ft, Herb. I.R.L.C.S. 6700 (K); 
Kilimanjaro, Petershutte, 4100 m, Peter 1212 (B); Kilimanjaro, 2000-3000 
m, Meyre s.n. (B); Kilimanjaro, Volkens 1 155 (B); Usambara, Hoist 3824 (B); 
Kissenye, Ninagongo, 3000 m, Mildbraed 1372 (B); NO Kivu, W. Kalago, c. 
2300 m, Mildbraed 1651 (B); Arusha National Park, crater of Mt. Meru, 
below Njeku Hut, 2560 m, Pocs & Kornas 652 I/A (BR). 

UGANDA: Mt. Elgon, 9000 ft, Dummer3560 (BOL, K, NBG); Ruwenzori 
Mountains, Nyamagasani Valley, 12500 ft, Loveridge 197 (K, SRGH); 
Ruwenzori, Lanuri c. 3500 m, Bequaert 4544 (P); Ruwenzori, le vallee du 
Mobuku, Val de Kabuamba, 3500 m, Alluaud 274 (P); Ruwenzori, vail, du 
Mobuku, abri sous roche de Buamba, 3500 m, Alluaud 275, (K, P); Ruwenzori 
(Est), vallee du Mobuku, rocher de Kichuchu, 3000-3200 m, Alluaud 310 
(P); Ruwenzori (Est), vallee du Mobuku, abri sous roche de Buamba, 3500 m, 
Alluaud 276 (P); Toro District, Ruwenzori, Bigo, 3350 m, Osmaston 3921 
(K); Ruwenzori, Nyamudamba, c. 10000 ft, Scott Elliot 8094 (K); NE Elgon, 
Tweedie 2745 (K); Ruwenzori, Mijusi Valley, 3500 m, Medberg 613 (K, S); 
Mt. Elgon, c. 1 1 000 ft, Allen 3676 (K); on Elgon at Benet, 9100 ft, Eggeling 
2454 (K); Mt. Elgon, Rose 10267 (K); Western Province, Bigo, R. Bujuku 
Valley, 3550 m, Osmaston 1738 (BR); Ruwenzori, c. 3500 m, Bequaert 4544 
(BR). 

ZIMBABWE: Inyanga District, 6500 ft, Chase 5100 (NU); Inyanga, 
7000 ft, Patterson 29 (GRA); Vumba Mountains, Umtali District, Eagle 
School road, Jackson 29 (GRA). 

WITHOUT EXACT LOCALITY: Natalia, Buchanan 83 (B); loco 
incerto, ex Herbario Natalensi, sine coll. s.n. (S); loco incerto, Buchanan s.n. 
TM522c (PRE); Natal, Medley-Wood s.n. TM520c (PRE); loco incerto, 
Dinter 575, A-only (B); Muhonora, 3500 m, De Wine 1962 (BR); Karisimbi 
(versant sud) nr. Biuri, c. 3000 m, De Wine 1246 (BR); South Africa, ?Rivier, 
Lincke 57 (BR); Basutuland, Koopoeitz s.n. (GRA); loco incerto, sine coll. 
s.n. NH-9784 (NH). 



J. P. ROUX 

Sledge (1973) cited Polystichum fuscopaleaceum as synonymous 
with P. setiferum var. nigropaleaceum (H. Christ) Sledge [= P. 
nigropaleaceum (H. Christ) Diels]. Christ (1893) described this 
variety from a single specimen collected by H.F. Blanford at 4000 ft 
in the Jumna valley between Mussoorie and Lokwah, western 
Himalayas. Sledge did not examine the type of this variety as he was 
unable to locate it. Fraser-Jenkins (Fraser-Jenkins & Khullar, 1985) 
reported he had studied the type in the Manchester Herbarium 
(MANCH). A Blanford specimen from the same locality has since 
been located in the Paris Herbarium (P!) and may serve as an 
isotype. This plant shows no clear affinity with either R 
fuscopaleaceum or P. setiferum, but rather to the P. luctuosum group 
(section Xiphopolystichum) as was suggested by Christ. Fraser- 
Jenkins & Khullar (1985) consider it synonymous with P. discretion 
(D. Don) J. Sm. 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum wilsonii 
and P. transvaalense occur sympatrically and often grow side by 
side. This has led to a great deal of confusion in separating the two 
taxa from one another. Pichi Sermolli (1977) referred to them as 
altitudinal vicariants, with P. wilsonii (as P. fuscopaleaceum var. 
ruwensoriense) occurring at higher elevations, having narrower 
blades, and a denser covering of wider, approximately rounded, 
acuminate paleae with those on the stipe being pale. In 1985 Pichi 
Sermolli added further observations as to how the two taxa differ. In 
var. ruwensoriense (= P. wilsonii) he found the apical part of the 
pinnae to be acute, moderately incised and usually provided with 
sori, whereas in var. fuscopaleaceum (= P. transvaalense) it is 
acuminate, deeply incised and devoid of sori. 

Jacobsen (1978), who refers to P. wilsonii as the 'Drakensberg 
Form' of P. setiferum var. fuscopaleaceum, provides some charac- 
teristics of the species and ascribes several collections in the National 
Herbarium, Pretoria (PRE) to it. Many of these collections, how- 
ever, belong to other species. 

Polystichum wilsonii is separated from P. transvaalense by the 
slightly shorter and narrower fronds, shorter stipe, a more pro- 
nounced reduction and deflexing of the basal pinnae, and in the 
paleae. Larger paleae in P. wilsonii extend from the stipe to the 
rachis, are generally complanate at maturity and somewhat pol- 
ished. The smaller paleae are more rigid with shorter and less 
divided marginal outgrowths. In P. transvaalense the larger paleae 
are mostly restricted to the stipe and proximal part of the rachis and 
become somewhat shrivelled at maturity. The marginal outgrowths 
on the proximal part of the smaller stipe paleae are also longer, more 
divided, and more twisted. Polystichum wilsonii, although often 
present in forests at lower elevations with P. transvaalense, is 
predominantly a high altitude species occurring in exposed conditi- 
ons. A natural hybrid between the putative species was described as 
P. x saltum (Roux, 1997a). 

Polystichum wilsonii forms part of the section Lasiopolystichum 
Daigobo assemblage of species. More recently (Kung & Zhang, 
1998) P. wilsonii has been placed as a synonym of P. sinense H. 
Christ. I choose to maintain the two as distinct species pending 
critical study. 

VARIATION. Morphological variation in Polystichum wilsonii is 
mostly restricted to the larger paleae present on the stipe and 
abaxially on the rachis. Variation is most apparent in the size, 
density and colour of the paleae. Larger paleae are broad in plants 
growing in more exposed habitats; they are more densely set. In 
plants from deeply shaded forest habitats, however, the larger paleae 
cannot be readily separated from the smaller paleae, especially on 
the rachis. Palea size and density thus appear to be environmentally 
influenced. No correlation could be drawn between habitat and 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



57 



palea colour, consistent with Schelpe's (1967) remark that forms 
with dark paleae intergrade with forms with pale stipe paleae. The 
large paleae are generally stramineous and concolorous. In some 
plants, however, the larger stipe paleae are bicolorous with the 
proximal central part of the paleae being densely impregnated with 
phenolic substances and castaneous. In some plants these bicolorous 
paleae are restricted to the proximal part of the stipe, whereas in 
others they may extend to the basal pinnae. More rarely the larger 
paleae on the rachis are nitid, densely impregnated throughout, 
almost black, and extend to the lower half of the rachis. 

Schelpe & Anthony (1986), in their key to the South African 
Polystichum species, used the direction and length of the pinnule 
aristae to distinguish between taxa. In some plants the basal basiscopic 
aristae of each pinnule may fold over the adaxial surface of the 
pinnule lamina, but in others they may not. Arista length also varies 
considerably. In some plants it may be relatively short but in others 
unusually long. In some plants the basal basiscopic aristae curve 
away from the pinnule lamina. Indusium size shows some variation 
with the margins ranging from repand to erose. 

DISTRIBUTION AND ECOLOGY. Polystichum wilsonii has a wide 
distribution ranging from Africa to the Uttar Pradesh mountains in 
northern India, and to Bhutan, China (Szechuan) and Taiwan (Ilan, 
Taichung, Hsinchu). In the study area P. wilsonii has a disjunct 
distribution. In South Africa it occurs on the southern Cape moun- 
tains, along the KwaZulu-Natal Drakensberg and into Lesotho, 
extending along the Free State-KwaZulu-Natal escarpment as far 
north as the Vryheid District. To the north it occurs in the mountain- 
ous areas of Zimbabwe, Tanzania, Kenya, Uganda, Ethiopia and the 
Kivu Ridge in the Democratic Republic of Congo. The species is 
also known from Mt. Cameroon and the island of Bioko, 32 km from 
the mainland in the Gulf of Guinea. It has furthermore been recorded 
from Grande Comore c. 300 km from the mainland in the Mozam- 
bique channel. Although the higher ground of the Zambezian Region, 
which includes Zimbabwe and Malawi, supports Afromontane plant 
communities (White, 1983), it is rare in this region with only one 
collection known from Zimbabwe. 

The lithology, climate and vegetation associated with Polystichum 
wilsonii vary considerably through its range. In the southern Cape P. 
wilsonii occurs at 1500-2000 m in acidic sandy soils derived from 
sediments of the Cape Supergroup. These soils support the unique 
Mesic Mountain Fynbos (Moll et al., 1984). This area is the only part 
of the distribution range of the species that experiences winter 
rainfall (April-September). 

In the Drakensberg Polystichum wilsonii is associated with the 
Clarens Sandstone formation, the Drakensberg Basalt Formation 
and the intrusive Karoo dolerites. At lower elevations in the 
Drakensberg (1250-1800 m) the species commonly occurs along 
streambanks or on rocks in Undifferentiated Afromontane forests 
and scrub forests confined to sheltered ravines and mountain slopes. 
These forests are largely associated with the Clarens Sandstone 
Formation. At higher elevations (> 1600- 1800 m) the Drakensberg 
Basalt formation and the intrusive Karoo dolerites are prevalent. 
These formations support the Themeda-Festuca Alpine veld (Acocks, 
1988). Here P. wilsonii occurs among boulders along streams, in dry 
exposed rock crevices or in wet and shaded rock overhangs. 

The Ethiopian and Kenyan highlands, Mt. Elgon, Mt. Meru, Mt. 
Kilimanjaro, Mt. Cameroon, Bioko and the Comoro Islands are all 
of volcanic origin or consist in part of volcanic deposits. Many of the 
isolated mountains are still volcanically active today. Also the Kivu 
Ridge, which is largely composed of Precambrian rocks, has local 
exposures of volcanic deposits. On all these mountains the vegeta- 
tion diminishes in structure from the lower slopes to the summit. 



Local features such as aspect, exposure incidence of frost, depth of 
soil and overall patterns of climate contribute to modify the vegeta- 
tion (White, 1983). At these elevations the plants become smaller 
and the apical pinnae more pronouncedly conduplicate along the 
rachis. 

In tropical Africa Polystichum wilsonii occurs in a wide range of 
vegetation types. At lower elevations on Mt. Elgon (2550 m) and the 
Ethiopian highlands (2700 m) it occurs in Undifferentiated 
Afromontane forests. On the Ethiopian highlands the species also 
occurs in single-dominant Afromontane forests such as Juniperus 
procera forests on Mt. Wuchada (2600 m) and Hagenia abyssinica 
forests in the Bale Mountains. On Mt. Kenya it has been recorded 
from the Afromontane bamboo zone. On Mt. Kenya (3200 m), Mt. 
Meru (3300 m), Mt. Elgon (3500 m) and the Bale Mountains (3500 
m) it occurs in Afromontane bushland and thicket. Again on the Bale 
Mountains (3500 m) and on the rim of the caldeira round Lake 
Wanchi (3650 m) it occurs in Afromontane and Afroalpine shrubland. 
With an increase in elevation the latter vegetation type is replaced by 
Afromontane and Afroalpine grassland. Polystichum wilsonii has 
been recorded from this vegetation type on Mt. Kilimanjaro (3000 
m) and the Ethiopian highlands (3900 m). On Mt. Cameroon P. 
wilsonii has been recorded from 1950 m to 3800 m and on the island 
of Grande Comore from 1000 m to 1400 m. In both cases the plants 
were associated with lava flows. 

Growth in Polystichum wilsonii shows a degree of seasonality. In 
the Drakensberg several new fronds are produced almost simultane- 
ously at the onset of the rainy season in November. This pattern is 
retained in cultivated plants. Several Afroalpine vegetation types are 
subject to periodic burning. Fires, however, appear to have little or 
no damaging effect on the rhizomes. 

9. Polystichum x saltum J.P. Roux in Bot. J. Linn. Soc. 124: 376, 
fig. 1 (1997). Type: South Africa, KwaZulu-Natal. 2828 (Bethle- 
hem): Royal Natal National Park, Gudu Forest, near Gudu 
Waterfall, c. 1800 m (DB), Roux 2510b (NBG!-holotype). 

Plants terrestrial or epilithic. Rhizome erect to suberect, to 20 mm in 
diameter, densely set with roots, closely set persistent stipe bases, 
and brown to ferrugineous paleae. Fronds caespitose, to 19 per 
plant, erect to arching, to 400 mm long: stipe proximally stramineous, 
greenish distally, shallowly sulcate adaxially, to 1 10 mm long, to 4 
mm in diameter, densely paleated, the paleae of two types; larger 
paleae broadly attached, brown to ferrugineous, chartaceous, lan- 
ceolate to narrowly lanceolate, cordate, often slightly auriculate, the 
margins closely to widely set with short and long, straight or curved, 
often forked projections, the apex always terminating in a long 
acicular cell, to 1 1 x 3.5 mm; smaller paleae brown to ferrugineous, 
chartaceous, short- or long-stalked, narrowly triangular, cordate to 
cordate-imbricate, the proximal margins closely set with short and 
long, straight or angular, simple or branched projections, the number 
and size of the projections reduced distally, the apex usually simple, 
terminating in a long acicular cell, to 6 x 1 mm: lamina 2-pinnate, 
narrowly ovate, to 300 mm long, with up to 17 free pinna pairs: 
rachis greenish throughout, adaxially shallowly sulcate, densely 
paleated, the proximal paleae of two types; larger paleae similar to 
those on the stipe and reduced in size towards the middle of the 
lamina; smaller paleae short- or long-stalked, ferrugineous, 
chartaceous, narrowly lanceolate to narrowly triangular, slightly 
cordate to cordate-imbricate, often slightly auriculate, the margins 
proximally with short and long, straight or curved, often branched 
projections that are reduced in size and frequency distally, the apex 
usually simple, terminating in a long acicular cell, to 6 x 1 mm: 
pinnae 1 -pinnate, long-stalked, proximally widely spaced, slightly 



58 



J.P. ROUX 



reduced, deflexed, with up to 10 free pinnule pairs, slightly overlap- 
ping distally, narrowly ovate to oblong-attenuate, to 75 x 24 mm: 
pinna-rachis greenish, adaxially shallowly sulcate, sparsely paleated; 
paleae ferrugineous, chartaceous, long-stalked, narrowly triangular 
to narrowly oblong, cordate to cordate-imbricate, the margins proxi- 
mally with long, straight or twisted, simple or forked projections 
reduced in size and frequency distally, the apex usually simple, 
terminating in a long acicular cell, to 3.5 x 0.5 mm: pinnules 
opposite to alternate, firmly herbaceous, pale- to olive-green 
adaxially, slightly paler abaxially, the proximal acroscopic pinnule 
usually slightly longer than the next, asymmetric, ovate to ovate- 
rhomboid, basiscopically narrowly cuneate, acroscopically broadly 
cuneate and auriculate, the auricle often incised midway to costa, 
serrate to doubly serrate, long-aristate, to 18 mm long; adaxially 
sparsely set with a few twisted paleae chiefly along proximal part of 
costa, stramineous, chartaceous, filiform or with a few short mar- 
ginal projections near the base, the apex always terminating in a long 
acicular cell, to 2 mm long; abaxially sparsely paleated, stramineous, 
chartaceous, long-stalked, narrowly deltate to filiform, the margins 
proximally with long, curved or angular, simple or branched projec- 
tions, the apex simple, always terminating in a long acicular cell, to 
3 mm long. Venation raised abaxially. Sori circular, c. 1.2 mm in 
diameter, terminal or near terminal on abbreviated vein branches, 
essentially uniseriate, discrete: sporangium with 13-(14)-17 
indurated annulus cells; stalk eglandular: indusium brown, 
chartaceous, persistent, peltate, circular, frequently with long cen- 
tral processes, fimbriate, the maximum radius 0.8-(0.9)-1.02 mm. 
Spores aborted, the perispore closely perforated. Chromosome 
number 2n=164, meiosis yielding univalents and bivalents (Roux, 
1997a). 

DIAGNOSTIC FEATURES. Polystichum x saltum closely resembles P. 
wilsonii in size, frond and pinnule morphology, and to a certain 
degree in the characteristics of the paleae. The erose to fimbriate 
indusium, however, is more characteristic of P. transvaalense. The 
mean guard cell length, the adaxial epidermal cell length, and the 
mean maximum radius of the indusium are anomalous - being larger 
than that of either progenitor (Roux, 1997a). Perhaps the most 
distinctive diagnostic feature of the taxon is the varying number of 
aborted spores borne in the sporangia. 

DISTRIBUTION AND ECOLOGY. Polystichum x saltum is currently 
known from only one forest in the foothills of the KwaZulu-Natal 
Drakensberg. This forest fragment forms part of the Highland 
Sourveld vegetation type (Acocks, 1988) and is nestled in a shel- 
tered ravine on a steep mountain slope. Forests of this type, situated 
at 1500 to 1700 m, are mostly cool and moist throughout the year, 
even though most of the rainfall occurs during the summer (Novem- 
ber-March). Like its putative parents, P. x saltum also occurs on 
moist moss-covered boulders along streams or on the forest floor in 
permanently moist conditions. Polystichum wilsonii, a taxon mostly 
associated with higher elevations where it occurs in more exposed 
habitats, frequently grows sympatrically with P. transvaalense in 
forests along the Drakensberg. 

lO.Polystichum marionense Alston & Schelpe in J. S. African Bot. 

23: 106, fig. la, t. 34 (1957). Type: Marion Island, Moseley s.n. 

(BM!-holotype). 
Fig. 10. 

Plants terrestrial or epilithic. Rhizome short, decumbent, branched, 
stoloniferous, to 5 mm in diameter, set with roots, closely spaced 
persistent stipe bases, and paleae; paleae sessile, ferrugineous to 
castaneous, scarious. Fronds crowded, to 8 per plant, erect, to 940 



mm long: stipe proximally castaneous, distally stramineous, adaxially 
shallowly sulcate, to 290 mm long x 4 mm in diameter, proximally 
close-set with unicellular pyriform glands, also sparsely to densely 
paleated; paleae sessile, ferrugineous to castaneous, scarious, lan- 
ceolate to broadly ovate, cordate to cordate-imbricate, the margins 
with irregularly spaced, unicellular pyriform glands (which also 
occur superficially) and short or long flagelliform outgrowths ter- 
minating in either a long filiform cell, a long filiform thin-walled 
cell, or rarely in a pyriform glandular cell, the apex terminating in an 
acicular or small thin- walled cell, to 1 1 x 3 mm: lamina 1-pinnate- 
pinnatifid to 2-pinnate, with up to 16 free pinna pairs, narrowly 
ovate to oblong-acute, to 285 mm long, the pinnae proximally wide- 
spaced, imbricate towards the apex, the most proximal pinna pair 
slightly to strongly reduced: rachis stramineous, shallowly sulcate 
adaxially, moderately set with unicellular pyriform glands and 
sparsely to densely paleated; paleae sessile, ferrugineous to 
stramineous, scarious, lanceolate to narrowly ovate, cordate to 
cordate-imbricate, the margins irregularly set with unicellular pyri- 
form glands (which also occur superficially), short 
cuneate-emarginate outgrowths often terminating in a unicellular 
glandular cell, and short or long flagelliform outgrowths (which also 
occur superficially) terminating in either a long filiform cell, a long 
filiform thin-walled cell, or rarely in a pyriform glandular cell, the 
apex teminating in an acicular or thin-walled cell, to 7 x 2 mm: 
pinnae pinnatifid to 1 -pinnate, with up to 5 free pinnule pairs, short- 
stalked, triangular, ovate, deltoid or oblong, to 36 x 18 mm: pinnules 
opposite to alternate, proximally short-stalked and widely spaced, 
sessile and imbricate towards the apex, firmly herbaceous to 
coriaceous, dark green adaxially, slightly paler abaxially, broadly 
ovate to circular, broadly cuneate, the margins shallowly crenate to 
dentate, revolute in plants from exposed habitats, to 1 1 mm long; 
adaxially sparsely set with a few twisted, cartilaginous, castaneous 
paleae chiefly along the pinna-rachis or costa; paleae short-stalked, 
linear to oblong, the margins subentire, with a few short cuneate- 
emarginate outgrowths or rarely with a few unicellular pyriform 
glandular cells and/or flagelliform outgrowths, the apex terminating 
in an acicular cell, to 3 mm long; abaxially sparsely to moderately 
set with hairs and scarious, stramineous to ferrugineous paleae, the 
paleae sessile, narrowly lanceolate to narrowly ovate, cordate, the 
margins (and often superficially) with unicellular, pyriform glandu- 
lar cells, short cuneate outgrowths that often terminate in a unicellular 
glandular cell, and often with a few flagelliform outgrowths termin- 
ating in an acicular or thin-walled cell, to 5 mm long. Venation 
raised. Sori circular, to 1 .5 mm in diameter, medial to inframedial, 
uniseriate, discrete but slightly confluent in depauperate plants; 
exindusiate: sporangium with 11-(14)-20 indurated annulus cells. 
Spores castaneous, the perispore folded to form closely set low 
tubercles, verruculate to echinulate, the exospore 30-(37.78)-78 x 
24-(28.62)-36 |jm. Chromosome number unknown. 

MATERIAL EXAMINED 

MARION ISLAND (4654'S, 3745'E): Black Hagless River near Kildalkey 
Bay, Gremmen s.n. (WAG); Macaroni Bay en route to Stony Ridge, Rand 
3270 (BOL, PRE); grey lava cliffs near Duikers Point, 10 m, Huntley 466 
(NBG-2 sheets, PRE-2 sheets); Marion Island, Mostert 15 (NBG, PRE); 
valley in cliffs above Prinsloo Lake, 25 m, Huntley 788 (NBG-2 sheets); 
Nellie humps, 40 m, Huntley 137 (BOL, NBG); stream adjacent Kildalkey 
hut, 100 m, O'Connor 1003 (BOL, NBG); cliffs at Goodhope Bay, Rand 
3653 (BOL); between station and Skua Ridge, Rand 3766 (BOL); Rand 3 1 92 
(BM, BOL), 3271 (BM, BOL), 3690 (BM, BOL). 

PRINCE EDWARD ISLAND (4638'S, 3757'E): cliffs S. of cave on E. 
coast, 25 m, Huntley 657 (BOL, NBG). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum 
marionense differs from any other taxon in the study area in having 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



59 




1 mm 



Fig. 10 Polystichum marionense. A, habit; B, abaxial surface of fertile pinna. All drawn from Huntley 466 (NBG). 



60 



J.P. ROUX 



a thin, decumbent and branched rhizome, pinnae that are not 
acroscopically developed, raised veins with few dichotomies, 
exindusiate sori situated along the veins and not at the vein endings, 
and paleae with unicellular glandular cells and flagelliform 
outgrowths along the margins and frequently also superficially. 

Alston & Schelpe (1957) considered the species to belong to the 
Polystichum mohrioides (Bory) C. Presl group. In this group they 
included P. plicatum (Poepp.) Hicken from the Andes and South 
Georgia, P. elegans J. Remy from the Andes, P. scopulinum (R.J. 
Eaton) Maxon and P. lemmonii Underw. from the western United 
States, and P. cystostegia (Hook.) J.B. Armstr. from New Zealand. 
Polystichum plicatum and P. elegans are sometimes considered to be 
varieties of P. mohrioides, but I found P. mohrioides to be extremely 
variable and could not on the grounds of the palea, indusium, 
sporangium and spore morphology distinguish between these taxa. 
I furthermore do not consider them related to P. marionense, as in 
this species the paleae bear long flagelliform outgrowths along the 
margins which often terminate in a glandular cell and are exindusiate. 
In P. mohrioides the paleae margins are subentire, but mostly bear a 
few short angular outgrowths. The palea apex always terminates in 
a short acicular cell. The indusia are large and mostly bear a variable 
number of pyriform cells along the margin and often also on the 
adaxial and abaxial surfaces. Polystichum cystostegia and P. 
mohrioides are clearly related as both are characterized by similar 
paleae and indusia. Polystichum scopulinum and P. lemmonii are 
related as can be judged from the paleae with short angular marginal 
outgrowths and apices that terminate in either an acicular cell or a 
small thin-walled cell. This is supported by the findings of Wagner 
(1979). Polystichum scopulinum and P. lemmonii are not considered 
to be related to either P. marionense or P. mohrioides. The affinity of 
P. marionense remains obscure. 

VARIATION. Variation in Polystichum marionense on Marion and 
Prince Edward Islands can be ascribed to environmental influences. 
Plants from well protected sites are large, the pinnae widely spaced, 
and the stipe sparsely paleated. Plants from more exposed sites are 
generally depauperate, densely paleated, and the pinnae coriaceous 
and closely imbricate with the pinnule margins strongly revolute. 
Pinnae of depauperate forms are often arranged perpendicular to the 
lamina axes. Minor variations also occur in the paleae. In some 
collections the flagelliform marginal outgrowths are extremely long 
whereas in others they are short. Also the occurrence of such 
outgrowths from the surface of the palea varies from collection to 
collection. The limitation of unicellular glands to the proximal part 
of the palea surface seems to be fairly constant. Unicellular pyriform 
glandular cells on the adaxial and abaxial surfaces of the lamina 
have only been observed in Huntley 788 (NBG). 

DISTRIBUTION AND ECOLOGY. Polystichum marionense is known 
only from Marion Island, Prince Edward Island and Possetion Island 
of the Crozet group in the Southern Ocean (Alston & Schelpe, 1957; 
Gremmen, 1982). Since the floras of the subantarctic islands are 
poorly known the species may have a wider distribution than is 
currently known. On Marion and Prince Edward Islands the species 
is only known from low-lying areas, with most collections having 
been made at elevations between 10 and 100 m above sea-level. The 
plants form large clumps in basalt rock crevices and at boulder and 
cliff bases. Huntley ( 1 97 1 ) reported the plant to always occur in sites 
protected from the predominantly westerly and north-westerly winds. 

1 1. Polystichum transkeiense W. Jacobsen in /. 5. African Bot. 44: 
169 (1978). Type: South Africa, Transkei, Port St Johns, near 
road to Second Beach, deep shade in forest, 67 m, W.B.G. 
Jacobsen 4301 (PRE!-holotype). 

Fig. 11. 



Plants terrestrial or epilithic. Rhizome prostrate, widely creeping, 
branched, to 10 mm in diameter, set with roots, closely to widely 
spaced persistent stipe bases, and paleae (which are restricted to 
apical region); paleae broadly attached, stramineous to castaneous, 
chartaceous, narrowly lanceolate, cordate to cordate-imbricate, the 
margins repand to erose, generally without thin-walled hair-like 
cells, the apex often flagelliform, mostly terminating in a thin- 
walled cell, to 8.5 x 1 .5 mm. Fronds usually widely spaced, 4-6 per 
plant, arching, to 1.34 m long: stipe firm, adaxially sulcate, proxi- 
mally castaneous, stramineous distally, to 710 mm long x 4 mm in 
diameter, proximally densely paleated; paleae broadly attached, 
castaneous to stramineous, chartaceous, narrowly to broadly ovate, 
cordate to cordate-imbricate, the margins repand, erose to fimbriate, 
with or without thin-walled cells, the apex often flagelliform, ter- 
minating in a thin-walled cell, to 7 x 2.5 mm; distally sparsely 
paleated, becoming glabrous with age: lamina 2- or 3-pinnate, with 
up to 22 pairs of free pinnae, firmly herbaceous, adaxially dark 
green, somewhat paler abaxially, ovate to broadly ovate, to 655 mm 
long, the proximal pinna pair reduced in size: rachis stramineous to 
greenish, adaxially sulcate, sparsely paleated; paleae short-stalked, 
stramineous, chartaceous to membranous, narrowly oblong to nar- 
rowly ovate, cordate to hastate, the margins proximally repand, 
erose, or set with short and/or long irregular outgrowths, often with 
filiform outgrowths terminating in a thin- walled cell, distally repand 
to entire, flagelliform, terminating in a filiform cell or a thin- walled 
cell, to 6 x 1 mm: pinnae 1 -pinnate or 2-pinnate, with up to 20 pairs 
of free pinnules, proximally widely spaced, mostly not overlapping, 
distally frequently overlapping; proximal pinnae narrowly ovate to 
narrowly oblong-attenuate, those towards the middle of the lamina 
ovate, narrowly oblong to oblong-attenuate, to 240 x 75 mm: pinna- 
rachis stramineous, adaxially sulcate, sparsely to densely paleated; 
paleae short-stalked, stramineous, chartaceous to membranous, lin- 
ear, narrowly triangular to narrowly ovate, cordate to hastate, the 
margins proximally with short and/or long irregular outgrowths 
often terminating in a thin-walled cell, distally entire, twisted, the 
apex terminating in a filiform or thin-walled cell: pinnules short- 
stalked, opposite to alternate, widely spaced to overlapping, the 
proximal acroscopic pinnule the largest, the proximal basiscopic 
pinnule on basal pinna pair generally significantly smaller than the 
next basiscopic pinnule, inaequilateral, ovate, ovate-oblong to ovate- 
rhomboid, acuminate to obtuse, acroscopically auricled, shallowly 
to deeply incised, lobate-serrate, the lobes oblong, the proximal 
acroscopic auricle obovate, sharp-tipped to aristate, the costa 
adaxially proximally sulcate, sparsely paleated; paleae stramineous, 
membranous, twisted, simple or proximally with short or long 
angular outgrowths, the apex terminating in a filiform or a thin- 
walled cell, to 2.6 mm long, abaxially sparsely to moderately 
paleated; paleae stramineous, membranous, narrowly triangular to 
narrowly ovate, short-stalked, cordate to cordate-imbricate, the 
margins proximally erose or with short and/or long angular 
outgrowths or with long filiform outgrowths terminating in a thin- 
walled cell, distally entire, flagelliform, twisted, the apex terminating 
in a filiform or thin- walled cell, to 1 .5 mm long. Venation raised. 
Sori circular, c. 1 mm in diameter, near or at the apex of abbreviated 
veins, discrete at maturity: sporangium with 10-(13)-19 indurated 
annulus cells; stalk eglandular: indusium absent. Spores 64 per 
sporangium, brown, the perispore folded to form a reticulum of 
inflated ridges, the ridges with a high crest, variously but mostly 
sparsely echinulate, minutely perforated, the exospore 32-(38.8)- 
46 x 22-(28.4)-36 urn. Chromosome number 2n=164. 

MATERIAL EXAMINED 

SOUTH AFRICA. 2330 (Tzaneen): Woodbush (CC), Jenkins 919 (PRE). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



61 




10 mm 



Fig. 11 Polystichum transkeiense. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A & C, drawn from Roux 2541 (NBG); B, 
drawn from Roux 2539 (NBG). 



62 



J.P.ROUX 



2430 (Pilgrim's Rest): Mariepskop (DB), Burrows 3150 (PRE). 2531 
(Komatipoort): Barberton, Ida Doyer Nature Reserve, 1 100 m (CC), Mutter 
2107 (PRE); Barberton, Maid of the Mist, Thorncroft 40 (PRE). 2730 
(Vryheid): Pongola Bush Reserve, Stinkwood Falls, 1550 m (BC), Glen 
2436 (PRE). 2731 (Louwsburg): Ngome Forest Reserve (CD), Roux 2535, 
2536, 2537, 2538, 2539, 2540, 2541 (NBG); Ngome Forest, Reid 68 (PRE); 
Ngome Forest, 4100 ft, Schelpe 6223 (BOL); Ngome Forest, Strey 8378, 
9381 (BOL, NH), 10487 (NH); Ngome Forest, c. 1000 ft, Schelpe 6244 
(BOL); Ngome Forest, along waterfall path, 1000 m, Glen 93, 97 (PRE). 
2830 (Dundee): Eshowe, Hospital Wood (CD), Lawn 63 (NH); Qudeni 
Forest, 5000 ft (DB), Fisher 802 (NH, NU, PRE), 877 (NU), Qudeni Forest, 
Jordaan 698 (NH, PRE); Qudeni Forest, 5500 ft, Schelpe 6267, 6268 (BOL); 
Qudeni Forest, 5000 ft, Clarkson 87 (NU), 130 (BOL, NU); Qudeni Forest, 
c. 5000 ft, Allsopp 743 (NU); Qudeni, 5000 ft, Fisher 83 1 (NH, NU); Qudeni, 
Ekombe Forest, 5000 ft, Fisher 8 1 7 (NU); Qudeni Forest, 1 1 00 m, MacDevette 
702 (PRE); Qudeni Forest Reserve, Van Wyk 7306 (NH); Qudeni Forest, 5000 
ft, Fisher & Schweickerdt 109 (NH). 2831 (Nkandla): Nkandla Forest (CA), 
Roux 1932 (NBG); Eshowe to Nkandla, 1080 m, Goetghebeur 4443 (BR, 
PRE); Nkandla Forest, Schelpe 1701 (BOL); Nkandla, 3000 ft, Meebold 
12614(M); Nkandla, Schelpe 1701 (NU); Nkandla, Mjton s.n. (NU); Nkandla, 
Lawn 2001 (NH). 2930 (Pietermaritzburg): Karkloof, farm Shawswood 
(AC), Roux 1915 (NBG); Karkloof, Van Jaarsveld 5026 (BOL, NBG); 
Karkloof, farm Ehlateni, Roux 1006, 1007, 1008, 1009, 1010, (NBG), 1011 
(NBG, PRE); Karkloof, 'Braco', 4300 ft, Schelpe 5115 (BOL); Balgowan, 
'Boschfontein', 4000 ft, Schelpe 606, 610 (NU); Balgowan, 4000 ft, Lindahl 
102 (NU); Balgowan, 'Boschfontein', 4000 ft, Fisher 630 (NH, NU); 
Karkloof, 'Elderslie', Rycroft s.n. (NU); Balgowan, Graham 107 (NU); 
Karkloof Forest, Ehlatine, Rycroft 89 (NU); Karkloof Forest, Wirminghaus 
610 (NU); Karkloof Forest, bank of Mshwati River, Wirminghaus 902 (NU); 
Karkloof, Colsbourne farm, Vos & McGregor s.n. (NU); Balgowan, Thomas 
67 (NU); Ahrens, 'Mowbray', c. 5000 ft (BB), Fisher 993 (NU); Dargle, 
Kilgoblin (CA), Smook 579 (NU), 566 (BOL, NU); Dargle, Esterhuysen 
26200 (BOL); Lions River District, Lions Bush, Moll 832, 833 (NU); Dargle, 
Kilgoblin, Smook 661 (NU); Pietermaritzburg, 2500 ft, (CB), Sanderson s.n. 
(PRE); Pietermaritzburg, Winters Kloof, Doige s.n. (PRE); Pietermaritzburg, 
Worlds View, Venter 736 (PRE); Pietermaritzburg, Ferncliff, Schelpe s.n. 
(BOL); Zwaartkop, Sim s.n. (BOL); Pietermaritzburg, Ferncliff Nature Re- 
serve, c. 2500 ft, Cowan 120 (BOL); Pietermaritzburg, Town Bush Valley, 
2500 ft, Tosh et al. (K); Swartkop, Hillary 69 (NU); Pietermaritzburg, 
Swartkop, Duncan-Vale 19 (NU); Sweetwaters, Stinger 58 (NU); 
Pietermatitzburg, Swartkop, 4000 ft, Fisher 123 (NH, NU); Swartkop, 4500 
ft, Nixon 36 (NU); Cascades, Town Bush, 2900 ft, Sidly 49 (NU); 
Pietermaritzburg, Claridge, Carnegie s.n. (NU); Hilton Road, Devlin 43 
(NU); Town Hill, Carnegie 706 (NU); Sweetwaters, Uhjati 52 (NU); 
Pietermaritzburg, Town Bush Valley, Fisher 693 (NU); upper Town Bush 
Valley, 3300 ft, Wand 28 (NU); Town Bush Valley, Tosh Robinson & De 
Villiers 1 (NU); Town Bush Valley, c. 2250 m, Doni 72 (NU); Pietermaritzburg, 
Ferncliff Nature Reserve, 1000 m, Crouch 556, 570, 598 (NU); Swartkop, 
Clarkson 19 (NU); Town Bush Valley, Devlin 34 (NU); Town Bush Valley, 
3000 ft, Fisher 667 (NU); Ferncliff Nature Reserve, 2500 ft, Cowan 155 
(NU); Town Bush Valley, 3000 ft, Nieuwoudt 56 (NU); Winters Kloof, 
Doidge P54 (PRE); Cottingham, farm Keerom, 4500 ft (CC), Strey 8429 
(BOL, NH); Richmond, Enon Forest (CD), Van Jaarsveld 5044 (PRE); 
Inanda (DB), Wood s.n. (B, PRE); Camperdown, Nagle Dam, 3000 ft (DD), 
Wells 1551 (NU). 2931 (Stanger): 10 km from Kwasizabantu towards 
Mapumulo (AA), Van Jaarsveld & Lang 5096, 5098 (BOL, NBG), Van 
Jaarsveld & Jacobs 5851 (NBG); Alexandra District, Moyeni, 750 m (BA), 
Rudatis 1 100 (B, K, NBG, P); Richmond, Enon Forest (CD), Van Jaarsveld 
5044 (BOL, NBG). 3029 (Kokstad): 22 miles E. of Kokstad, 4850 ft (CB), 
Schelpe 4417, 4418 (BOL); Tabankulu Forest Reserve (CD), Wilkins 40 
(PRE); Ingeli Bush (DA), Taylor 5227 (NBG, PRE); Mpetsheni Forest, 
Weza, Roux 1960, 1961 (NBG); Weza Forest, Roux 2493, 2494, 2495, 2497, 
2498 (NBG); Mpetsheni Forest, Weza, c. 1000 m, Nicholas & Marais 1675 
(PRE); Weza Forest, Roux 623 (BOL); Weza Forest, Bangeni Forest, 1 200 m, 
MacDevette 1 534 (NH). 3030 (Port Shepstone): Burntwood, Paddock (CC), 
Strey 5994 (BR, K, NU); Umtamvuna Nature Reserve, Long Kloof, 360 m, 
Abbott 1818 (NH); Umtamvuna Nature Reserve, Gogosa Kloof, Abbott 2101 
(NH); Umtamvuna Nature Reserve, Verassend Kloof, 360 m, Abbott 1821 
(NH). 3128 (Umtata): Tsolo, Nqadu Ridge, c. 1 1 00 m, (BC) Keeler & Cloete 



449 (NH). 3129 (Port St Johns): Port St Johns, Egossa Forest (BC), Strey 
8869 (BOL, NH, NU, PRE); Lusikisiki, Magwa Falls, Strey 67 1 8 (NH, PRE); 
Egossa Forest above Magwa Falls, 1300 m, Venter & Vorsterll (BR, PRE); 
Port St Johns, stream at S. end of airstrip (DA), Roux 582 (BOL, NBG); Port 
St Johns, Hardcastle s.n. (NBG); woods at Port St Johns, Flanagan 2973 
(PRE); Port St Johns, Hardcastle 28 1/283 (PRE); Port St Johns, McLoughlin 
788 (BOL, PRE); Port St Johns, edge of plateau, 1200 ft, Hardcastle 285 
(PRE); Port St Johns, McLoughlin S36 (PRE); Port St Johns, Agate Terrace, 
McLoughlin 780 (BOL); Port St Johns, Isaac s.n. (BOL); Port St Johns, 
Moffets Glen, Roux 589 (BOL); Port St Johns, Schelpe 357, 358 (NU); Port 
St Johns, Flanagan 2473 (PRE). 

SWAZILAND. 2531 (Komatipoort): Piggs Peak, Kings Forest (CD), 
Compton 27831 (NBG, PRE); Havelock, Kings Forest, 5000 ft, Schelpe 
6163, 6169 (BOL). 

WITHOUT EXACT LOCALITY: Port Natal, Krauss 258 (BM); loco 
incerto, Hill 36 (PRE); loco incerto BOL 5771 3 (BOL); Natal, Buchanan s.n. 
(BOL); Natalia, in sylvis montis humidis, sine coll. s.n. (P); Pondoland, 
Buchanan 30 (P); Hlokozi, Alexandra City, 2700 ft, Rudatis 2309 (NBG); in 
sylvis montanis umbrosis humidis, Guienzius 28 (B); Natalia, in sylvis 
humidis, sine coll. B-97066 (B); Natal, McKen s.n. (B); Natal, Buchanan 75 
(B); Pondoland, Buchanan 30 (B); zwischen dem grossen Wasserfall und 
Omsamcaba, Drege s.n. (B); loco incerto, Drege s.n. B-97092 (B); Natal, 
Guienzius s.n. B-97064 (B); loco incerto, sine coll. s.n. NH-26388 (NH); 
Prom. b. spei, Guienzius s.n. (BR); Natal, sine coll. s.n. (BR); Zululand, 
Haygarth s.n. (NH); loco incerto, sine coll. s.n. NH-26465 & 26466 (NH); 
Inanda, Great Noodsberg, Town Hill, P.M.B., sine coll. s.n. NH-26791 (NH); 
loco incerto, sine coll. s.n. NH-26387 A-only (NH). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. The thin, widely creep- 
ing rhizome, the thin-walled cells on the paleae, and the exindusiate 
sori are the most diagnostic features of the species. The affinity of 
Polystichum transkeiense is yet to be determined. 

VARIATION. Polystichum transkeiense shows extreme variation in 
frond morphology, even within a population. Lamina dissection 
varies between 2-pinnate and 3-pinnate with the pinnules showing 
various degrees of dissection. Pinnule lobes may vary between 
broadly elliptic to obovate and narrowly obovate to narrowly ob- 
long. Palea variation is not as significant as that of the lamina. Paleae 
do, however, vary in outline, the degree to which the margins are 
sculptured, the absence or presence of thin-walled cells along the 
margins, and in the apex terminating in a thin-walled cell or a 
filiform cell. Both conditions are usually present in the same plant. 

DISTRIBUTION AND ECOLOGY. Polystichum transkeiense is con- 
fined to the eastern parts of South Africa and northern Swaziland. 
This region receives its rain largely during the summer months 
(September-March). It is an exclusively forest-dwelling species, 
often growing in very wet conditions. In the southern limits of its 
distribution at Port St Johns, P. transkeiense grows in Typical Coast 
Belt Forest that occurs from near sea-level to approximately the 450 
m contour. This region receives 900-1 500 mm of rain per annum. To 
the north and somewhat inland it occurs in forests of the Pondoland 
Coastal Plateau Sourveld where it occupies a plateau to 450 m above 
the sea. The forests are mainly found in protected places along the 
escarpment such as gorges and valleys below cliffs. Rainfall in this 
region is high; 1150-1300 mm of precipitation is measured per 
annum. In the KwaZulu-Natal midlands P. transkeiense occurs in 
forests of the 'Ngongoni Veld, extending between 450 and 900 m 
above the sea and receives on average 750-1300 mm of rainfall per 
annum. Nkandla, Qudeni and Weza are among the most notable 
forests occurring in this vegetation type. In northern KwaZulu-Natal 
Afromontane forests with slightly more tropical affinity occur on the 
inland mountains. The extensive Ngome Forest, where P. transkeiense 
is common, is an example of this forest type. Forests of this type 
occur northwards to the mountains south and west of Barberton. 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 

Rainfall in this region is high, ranging between 900-1950 mm per 
annum (Acocks, 1988). 

Jacobsen ( 1 978) considered 'its tendency to grow isolated in deep 
shade and not in large clusters. . .' as a diagnostic feature of 
Polystichum transkeiense. My observations, however, do not con- 
form with this statement. Polystichum transkeiense does grow as 
isolated plants, but generally forms continuous, dominant stands, 
especially in the Weza, Karkloof, Nkandla and Ngoye forests. 

12. Polystichum magnificum F. Ballard in Kew Bull. 12: 48, f. 1 
(1957). Type: Uganda, Mount Elgon, in the crater (alpine region) 
in a small sheltered cleft on the ridge north of Maji ya moto, 3750 
m, Hedberg 965 (K!-holotype; K!-isotype). 

Fig. 12. 

Plants terrestrial. Rhizome short-decumbent, branched, to 12 mm in 
diameter, set with roots, closely spaced persistent stipe bases, and 
paleae; paleae broadly attached, ferrugineous, chartaceous, linear, 
cordate, the margins subentire or with small, widely spaced, straight 
or curved outgrowths, the apex terminating in an acicular cell, to 30 
x 2 mm long. Fronds 8-12 per plant, erect to suberect, to 1.13 m 
long: stipe proximally castaneous, stramineous distally, adaxially 
sulcate, to 470 mm long x 9 mm in diameter, densely paleated; 
paleae broadly attached, ferrugineous, chartaceous, those on the 
proximal part of stipe narrowly ovate, cordate, the margin with small 
widely spaced projections, the apex terminating in an acicular cell, 
to 40 x 2 mm, those on the distal part of stipe more variable in size; 
the larger broadly attached, narrowly ovate, ovate to narrowly 
oblong, cordate, with the margins variously set with short and/or 
long, simple or forked recurved outgrowths, the apex terminating in 
an acicular cell, to 28 x 8 mm; the smaller short-stalked, narrowly 
triangular, cordate, often somewhat auricled, the margins proxi- 
mally with short and/or long angular or curved outgrowths, distally 
entire or with small projections, the apex terminating in an acicular 
cell: lamina 2-pinnate to 2-pinnate-pinnatifid, coriaceous, adaxially 
dark green, abaxially slightly paler, narrowly ovate to narrowly 
oblong, to 660 mm long, reduced towards the base, often with a 
single proliferous bud near the apex, the bud paleae ferrugineous: 
rachis stramineous, adaxially sulcate, densely paleated; paleae 
stramineous to ferrugineous, the larger broadly attached, ovate to 
narrowly ovate, cordate, the margins proximally with long and/or 
short, straight and/or curved outgrowths, distally entire, the apex 
terminating in an acicular cell, the smaller short-stalked, narrowly 
triangular, the margins proximally with curved or angular outgrowths, 
distally entire, the apex terminating in an acicular cell: pinnae 
mostly somewhat overlapping, narrowly ovate to oblong, not sig- 
nificantly developed acroscopically, to 130 mm long: pinna-rachis 
stramineous, adaxially sulcate, densely set with paleae similar to but 
smaller than those on the rachis; pinnules proximally closely spaced, 
distally alternate, mostly somewhat imbricate, inaequilateral, ovate, 
lobate, crenate, the proximal pinnules acroscopically incised to or 
near to the adaxially sulcate costa, the segments unequally rhomboid 
to obovate, adaxially densely paleated; paleae short-stalked, ferrugi- 
neous, chartaceous, subulate, simple or proximally with short 
marginal outgrowths, often twisted, the apex always terminating in 
an acicular cell, to 5 mm long, abaxially densely paleated; paleae 
short-stalked, ferrugineous, subulate, straight or twisted, proximally 
with short marginal outgrowths, the apex terminating in an acicular 
cell, to 4 mm long. Venation raised. Sori circular, to 2.2 mm in 
diameter, uniseriate, discrete at maturity, terminal or near-terminal 
on abbreviated vein branches: sporangium with 1 1-( 13)-17 indurated 
annulus cells; stalk eglandular: indusium peltate, circular, erose, the 
maximum radius 0.63-(0.94)-1.14 mm, persistent, brown. Spores 



63 

dark brown, 64 per sporangium, the perispore relatively smooth, 
echinulate, closely perforated, the exospore 36-(47.78)-58 x 26- 
(33. 37)^0 urn. Chromosome number unknown. 

MATERIAL EXAMINED 

ETHIOPIA: Bale Region, Dello Awraja, in Harrena Forest c. 3.3 km N. of 
Rira, 3040 m, Mesfin 5077 (ETH); Bale Mountains, E. of Kara Deema, 4140 
m, Miehe 1497 (ETH); Bale Mountains, E. of Kara Deema, 4200 m, Miehe 
1541 (ETH); Bale Mountains, Mendoyn Anraja, in Harrena Forest, c. 1-2 km 
S. of Riva village, 2780-2850 m, Mesfin 5355 (ETH); Bale Region, Dello 
Awraja, c. 3.4 km N. of Rira village, 3120 m, Mesfin 5332 (ETH); Arussi, 
Juniper forest, 9000 ft, Thomerson 550 (ETH, K); Darra, bamboo forest, 
9000 ft, Mulvany 48 (K); Bale Province, Rira, 20 miles SW of Goba, 10 800 
ft, Mooney 7192 (K); Mount Tola, Gamu Highlands, 13 000 ft, Mulvany 1 
(K); Bale Province, c. 30 miles S. of Goba, Saneti Plateau, 2720 m, Ash 3567 
(BR). 

KENYA: Mount Elgon, 12 500 ft, Tweedie s.n. (K). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Most striking is the 
large stature of the plants occurring at these high elevations. Also 
diagnostic are the densely paleated fronds, rounded lobes, and the 
proliferous bud borne on the rachis near the frond apex. 

The affinity of Polystichum magnificum is uncertain. Super- 
ficially it appears to belong to section Lasiopolystichum but the 
palea structure and the short, branched decumbent rhizome does not 
support such an affinity. 

DISTRIBUTION AND ECOLOGY. Polystichum magnificum is only 
known from Mount Elgon, on the border between Uganda and 
Kenya, and the Bale, Arussi, and Gamu Gofa regions in southern 
Ethiopia. 

On Mount Elgon Polystichum magnificum occurs at elevations 
ranging from 3700 to 3800 m. In Ethiopia it occurs from 2700 to 
4200 m and grows in a wide range of afromontane vegetation 
communities. These include afromontane forests of the Hagenia 
abyssinica (Bruce) J.F. Gmel. and Juniperus procera Hochst. ex 
Endl. types, afromontane bamboo zone, afromontane scrubland and 
afromontane grassland types as defined by White (1983). 

The species has been reported to form compact patches up to 1.8 
m in diameter at 3000 m on Mount Tola and Mount Gughe in 
Ethiopia. At higher elevations Polystichum magnificum tends to be 
restricted to rock crevices where it is protected from wind and fire. 

13. Polystichum zambesiacum Schelpe in Bol. Soc. Brot. ser. 2, 41: 
215(1 967). Type: Rhodesia (Zimbabwe), Umtali District, Henkels 
Nek, Stapleford, Schelpe 5751 (BOL!-holotype; BOL!-isotype). 

Fig. 13. 

Plants terrestrial or epilithic. Rhizome short-decumbent to suberect, 
short-branched, to 25 mm in diameter, set with roots and closely 
spaced persistent stipe bases, the older parts nude, the apical part 
densely paleated; paleae broadly attached, rugose, ferrugineous, 
linear, truncate to cordate, the margins variously fimbriate, often 
also with a few long, straight, recurved, filiform outgrowths, to 30 x 
2 mm. Fronds caespitose, 5-8 per plant, arching, to 1.8m long: stipe 
proximally castaneous, stramineous distally, adaxially sulcate, to 
840 mm long x 9 mm in diameter, proximally densely paleated; 
larger paleae more widely spaced and becoming smaller distally, 
membranous, ferrugineous throughout or with a narrow membra- 
nous margin and a dark, nitid centre, ovate, cordate, the margins 
fimbriate, the apex cuspidate or flagelliform, terminating in a thin- 
walled cell or a filiform cell, to 16 x 10 mm; smaller paleae 
short-stalked, narrowly ovate to narrowly triangular, cordate to 
cordate-imbricate, the margins proximally with irregular angular 
outgrowths, often also with long, twisted, filiform outgrowths often 
terminating in a small thin-walled cell, sparsely fimbriate distally, 



64 



J.P. ROUX 




2 mm 



5 mm 



Fig. 12 Polystichum magnificum. A, middle pinnae of lamina; B, adaxial surface of pinnule; C, abaxial surface of fertile pinnule; D, section of abaxial 
surface of rachis. All drawn from Hedberg 965 (K). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



65 




Fig. 13 Polystichum zambesiacum. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A & C, drawn from Schelpe 5751 (BOL); 
B, drawn from Burrows 2935 (PRE). 



66 



J.P. ROUX 



the apex flagelliform, terminating in a small thin-walled cell or a 
filiform cell; lamina 2-pinnate to 3-pinnate, with up to 30 free pinna 
pairs, firmly herbaceous, narrowly ovate to ovate, up to 890 mm 
long, the proximal pinna pair often slightly reduced and often 
somewhat deflexed: rachis stramineous, adaxially sulcate, often 
somewhat flexuous towards the apex, sparsely paleated; paleae 
short-stalked, chartaceous, stramineous to ferrugineous, narrowly 
linear, narrowly ovate to narrowly triangular, cordate to cordate- 
imbricate, the margins proximally with straight or angular outgrowths 
or with long, twisted, filiform outgrowths, distally entire or sparsely 
fimbriate, the apex terminating in a small thin-walled cell or a 
filiform cell, to 1 .8 mm long: pinnae 1 -pinnate to 2-pinnate, with up 
to 27 free pinnule pairs, generally widely spaced but often imbricate, 
lanceolate, narrowly ovate to oblong-attenuate, the proximal pinnae 
to 268 mm long: pinna-rachis stramineous, adaxially sulcate, set 
with paleae similar to but smaller than those on the rachis: pinnules 
opposite to alternate, widely spaced or imbricate, asymmetric, 
ovate-oblong, narrowly ovate to narrowly triangular, acroscopically 
auricled, usually somewhat reclinate, lobate-serrate, obtuse, the 
proximal acroscopic pinnules on basal pinnae often reduced, be- 
coming larger towards middle of lamina, the proximal pinnules 
simple or incised to or near to costa, to 45 mm long; costa proximally 
adaxially sulcate, glabrous or with a few short-stalked, stramineous, 
twisted paleae; paleae filiform, simple or proximally with a few 
twisted outgrowths, the apex terminating in a small thin-walled cell 
or a filiform cell, to 1 .2 mm long, adaxially sparsely paleated; paleae 
short-stalked, narrowly triangular to narrowly ovate, cordate, the 
margins proximally with short or long angular, often filiform 
outgrowths, the apex entire or sparsely fimbriate, terminating in a 
small thin-walled cell or a filiform cell, to 0.8 mm long. Venation 
raised or obscure. Sort circular, terminal, medial or inframedial on 
unabbreviated or abbreviated vein branches, discrete at maturity, c. 
1 mm in diameter: sporangium with 9-(14)-20 indurated annulus 
cells; stalk eglandular; indusium peltate, circular to reniform, repand 
to erose, the maximum radius 0.19-(0.33)-0.5 mm, persistent, 
brown. Spores 64 per sporangium, brown, the perispore folded to 
form inflated reticulate ridges and tubercles, the ridges and areas 
between variously perforated, echinate to echinulate, the exospore 
30-(37.77)-54 x 22-(27.49)-36 urn. Chromosome number un- 
known. 

MATERIAL EXAMINED 

MALAWI: Nyika Plateau, 2250 m, Brass 17255 (K, PRE, SRGH); Mount 
Mulanje, Lichenya Plateau, Nessa Path Forest, 1800 m, Chapman 8262 
(MAL, PRE); Zomba Plateau, W.-facing cliff edge below Malumbe Peak, 
1900 m, Berrie 602 (MAL); Zomba Plateau, Malumbe Peak, Roux 2928 
(NBG); Mulanje Mountains, Lichenya Plateau, Pawek 3815 (K); Mount 
Mulanje, Lichenya Plateau, 1 820 m, Brass 1 6566 (K, SRGH); Nyika Plateau, 
Kasaramba Peak, 8400 ft, Simon et al. 1730 (K, SRGH); Mount Mulanje, 
Lichenya Plateau, 1890m,fira 16820 (K, SRGH); Mount Mulanje, Lichenya 
Plateau, 1950 m, Richards 16556 (K); Mount Mulanje, Nayawani Forest, 
6400 ft, Newman & Whitmore 547 (BR, SRGH); Mt. Mulanje, L. Ruo 
Plateau, 6400 ft, Newman & Whitmore 399 (BR). 

MOZAMBIQUE: Manica & Sofala, Penhalonga, 4500 ft, Schelpe 5325 
(BOL); Manica & Sofala, Gorongosa Mountains, Gogogo Peak, 5000 ft, 
Schelpe 5518 (BOL); Manica & Sofala, Gorongosa, Serra da Gorongosa, 
vertente do monte Nhandare, 1750 m, Torre & Pereira 12515 (BOL, BR, 
SRGH); Namuli, Makua country, Last s.n. (K). 

TANZANIA: Uluguru Mountains, S. of Bunduki, NEedge of Lukwangule 
Plateau, on rocky outcrops of Muisile Hill, 2400-2450 m, Pocs et al. 86141/ 
A (WAG); W. Usambaras, towards Mount Kwashenhambu, 1 850 m, Schippers 
T 1578 A (WAG); W. Usambara Mountains, N. slopes of Mount Shegein, 
Shasayo Forest, 1850 m, Schippers T1604A (WAG); Morogoro, Ukaguru 
Mountains in Kilosa District, W. Mamiwa Ridge, 2100-2200 m, Pocs et al. 
86100/B (WAG); Tanga Region, Lushoto District, W. Usambara Mountains, 



Shagayu Forest Reserve, NW slope of the summit 2.5 km ENE of Shagayu 
sawmill, 1850-1950 m, Borhidi et al. 84847 (ETH); Morogoro District, 
Uluguru Mountains, W. part of Lukwangule Plateau, 2400-2500 m, Harris et 
al. 3726 (K); Usagara: Itumba, Wood s.n. (K); Uluguru Mountains, above 
Morogoro, NE ridge of Bondwa between Morningside and Mwere Valley, 
1050 m, P6cs 6537/C (PIC.SERM.); Uluguru Mountains, NW slope of 
Bondwa, along road to Morningside, Faden et al. 70/654 (BM, BOL); 
Uluguru Mountains, Morningside to Bondwa, Faden 70/316 (BOL, K); 
mainland W. of Zanzibar, Last s.n. (K); N. Uluguru Forest Reserve, Lupunga 
Peak, W. side, 2000 m, Hall s.n. (K); Morogoro: Uluguru gebirge, Lupanga, 
2100 m, Schlieben 2977 (B, BR). 

ZIMBABWE: Inyanga, Pungwe Gorge, 6000 ft, Schelpe 5699 (BOL); 
Inyanga, circular drive on N. rim of Pungwe Gorge, 7000 ft, Mitchell 148 
(BOL); Umtali, Stapleford Forest Reserve, W. of Rupere Peak, 5500 ft, 
Chase 7429 (BOL, K); Inyanga, Mitchell 1082 (BOL, K, NU, SRGH); 
Umtali, 'Cloudlands', Vumba, 5200 ft, Schelpe 5365 (BOL); Inyanga, ad 
dejectum fluminis Pungwe, c. 1700 m, Fries et al. 3795 (BOL, K); Umtali 
District, Vumba Mountains, 'Cloudlands', forest E. of Cripps Grid, Chase 
8345 (BOL, K); Umtali District, on Lords Head property, Vumba Mountains, 
5300 ft, Chase 8343, 8344 (BOL, K, SRGH); Melsetter, forest in gully on W. 
side of N. end of Bundi Valley, Mitchell 514 (BOL, K, SRGH); Umtali 
District, Stapleford Forest Reserve, 5600 ft, Chase 8373 (BOL, K, SRGH); 
Stapleford Forest Reserve, lower part of road to Henkels Nek, 5600 ft, Chase 
8371 (BOL, SRGH); Umtali District, Vumba Mountains, Williams VUM18 
(BOL); Vumba Mountains, Williams VUM1, VUM10, VUM11, VUM12, 
VUM13 (BOL); Chimanimani Mountains, Gwasha, Williams STP1, STP2 
(BOL); Inyanga, 6500 ft, Chase 5100 (BOL); Inyanga, Pungwe Gorge, 
Inyangani Mountains, 6500 ft, Chase 5240 (BOL); Umtali, Vumba Moun- 
tains below Castle Beacon, 5600 ft, Chase 7489 (BOL); Inyanga, circular 
drive below Inyangani Mountain, 7500 ft, Mitchell 135 A (BOL); Umtali, 
Imbeza Forest Estate, Zuwanne indigenous forest, 5150 ft, Jacobsen 3838 
(BOL, SRGH); Stapleford Forest Station, 6000 ft, Taylor 3234 (BOL, 
SRGH); Vumba Mountains, SE slope of Castle Beacon, 1675 m, Burrows 
2944, 2945 (PRE); Inyanga, S. tip of Mount Inyangani, 2040 m, Burrows 
2935 (PRE); Pungwe Gorge, in drier part of ravine, Schweikerdt 2412 (M, 
PRE); Umtali, Vumba, Castle Beacon, 6000 ft, Fisher 1638 (NU, PRE); 
Vumba Mountains, Chase 1 102 (PRE); W. slope of S. tip of Mount Inyangani, 
2000 m, Burrows 2940 (PRE); Inyanga, new beacon on Mount Inyangani, 
2540 m, Burrows 2828 (PRE); Inyanga, montane forest, 6000 ft, Chase 4024 
(PRE); Vumba Mountains, 5400 ft, Chase 8347 (K); Vumba Mountains, 
Bunga Forest, Jacobsen 3037 (SRGH); Inyanga, lower E. slope of Inyangani, 
c. 1480 m,Miiller32\4 (SRGH); lower SE slope of Mt.Pene, \350m,Muller 
2798 (SRGH); Inyanga, top of escarpment 2 km N. of Honde View, c. 1 840 m, 
Muller 3243 (SRGH); Melsetter, Chimanimani, Bundi River, 5500 ft, Whellan 
2184 (SRGH); Vumba Mountains, forest on E. slope of Castle Mountain, 
Jacobsen 3033 (SRGH); Stapleford Forest Reserve, Ruperi Peak, 6100 ft, 
Chase 4656 (SRGH); Stapleford Estate, montane forest, Roux 2828 (NBG); 
SE edge of Vumba Hotel forest, Chase 4022 (SRGH); Stapleford Forest 
Reserve, Chase 8372 (SRGH); Melsetter District, Orange Grove, Chase 
3088 (BR, NU, S, SRGH); Inyanga District, source of Inyahupina River, 
Romneydale, 6100 ft, Chase 2089 (NU, SRGH); 30 miles S. of Umtali, 6000 
ft, Grout 33 (NU); Umtali, Pioneer farm, Fisher & Schweickerdt 306 (NU); 
Umtali District, Vumba Mountains, Natseland, Chase 3428 (NU); Umtali 
District, Nyagari farm, N. of Zwitembo, Chase 3156 (NU); Inyanga District, 
Chase 3206 (NU); Inyanga District, Pungwe Falls, 6000 ft, Chase 3197 
(NU); Umtali District, Stapleford Forest Reserve, Chase 4520 (NU); Umtali 
District, Penhalonga, Chase 3132 (NU); Inyanga, Pungwe view point, Chase 
3206 (NU); Umtali, Chase s.n. (NU); a 15 km a 1'E. d'Inyanga, sommet du 
mont Mimunzi, 1814'S, 3253'E, 1950 m, Bamps et al. 272 (BR); Vumba 
Mountains, Umtali District, Cloudlands, Jackson 8 (GRA). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum 
zambesiacum can be easily recognized among African members of 
the genus by its long and narrow rugose rhizome paleae, long- 
attenuate pinnae and (although variable) obtuse pinnule lobes, 
conspicuously small indusia, and the palea morphology. 

Polystichum zambesiacum is not considered to be related to any 
other taxon in the study area. 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



67 



VARIATION. Schippers (1993) suggested that the Tanzanian plants 
may be a different but related species to Polystichum zambesiacum. 
He noted the Usambara Mountain plant to differ from that of the 
southern highlands in the more shiny fronds, the more deeply 
divided pinnae, and the segment margins having more clearly 
aristate teeth. Polystichum zambesiacum is clearly a very variable 
taxon with lamina division ranging between 2-pinnate and 3-pin- 
nate. Most apparent is the degree to which the size, shape and 
incision of the pinnules vary. This, however, cannot be linked to 
distribution as 3-pinnate plants have been recorded from the Pungwe 
Gorge in Zimbabwe [Chase 5240 (BOL)] and the Ukagura Moun- 
tains in Tanzania [Pocs et al. 86100/B (WAG)]. In the Uluguru 
Mountains, like in the rest of its distribution, the pinnules are mostly 
variously incised and lobate-dentate to ovate-serrate. One collection 
from this region [Faden et al. 70/316 (BOL, K)] shows hardly any 
incision of the pinnules which are poorly lobate. Lamina division 
can thus not be considered in subdividing the taxon. In some plants 
the proximal pinnule of the basal pinnae is reduced in size and thus 
not auriculate as the distal pinnae. The basiscopic pinnules of these 
pinnae are often more significantly reduced than the acroscopic 
pinnules. In others, however, the pinnules show no reduction of size 
on the lower pinnae. 

Stipe paleae also show variation. In some plants the larger paleae 
are variously impregnated with the central part of the paleae often 
dark brown to black and nitid with a narrow membranous margin. 
These paleae are generally persistent for a long time, resulting in the 
non-impregnated margin being worn away. In others the paleae are 
reddish brown and remain membranous. 

DISTRIBUTION AND ECOLOGY. Polystichum zambesiacum ranges 
from the eastern highlands of Zimbabwe and adjacent Mozambique 
to Mount Mulanje and the Nyika Plateau in Malawi and the Uluguru, 
Usambara and Pare Mountains in Tanzania. 

Polystichum zambesiacum is a terrestrial or epilithic species 
occurring in high-altitude, evergreen, montane mist forests, along 
forest margins and streambanks in forests. It has been reported on 
Mount Inyangani, Zimbabwe, from open montane grassland, and in 
the Uluguru Mountains from rock outcrops. 

Being a high altitude species, Polystichum zambesiacum is re- 
stricted to the often isolated high mountains of east Africa. In 
Zimbabwe and Mozambique it is confined to the Chimanimani and 
Nyanga Mountain ranges occurring between 1370 and 2540 m. In 
Malawi the fern is only known from Mount Mulanje in the south 
(1800-1950 m) and the Nyika Plateau in the north where it occurs 
between 1800 and 2550 m. In the Uluguru Mountains, Tanzania, it 
occurs at elevations ranging between 1050 and 2500 m. Further 
north, in the Usambara and Pare Mountains it occurs at 1 850 to 1950 
m, but is reported to be rare (Schippers, 1993). 

H.Polystichum monticola N.C. Anthony & Schelpe in Bothalia 
15: 554 (1985); Fl. Sthn. Afr., Pterid.: 257 (1986); Burrows, Sthn. 
Afr. ferns and fern allies: 3 14, f. 75, t. 320a-c (1990). Type: Cape 
Peninsula, Table Mountain, Dark Gorge, below saddle SE side, 
sheltered gully, dry in summer, on steep rocky slopes, Esterhuysen 
26685 (BOL!-holotype; B, C, CHR, G, GH, K, M, MO, NBG!, 
NU, P, PR, PRE!-isotypes). 

Fig. 14. 

Plants terrestrial or epilithic. Rhizome short-decumbent, closely 
branched, to 14 mm in diameter, set with roots, crowded persistent 
stipe bases, and paleae; paleae broadly attached, ferrugineous, 
membranous, narrowly oblong to narrowly linear, cordate, the 
margins entire or with widely spaced, short, apically or basally 
directed outgrowths, the apex acicular or with a thin-walled apical 



cell, to 26 x 1 .8 mm. Fronds closely spaced, 5-8 per plant, suberect 
to arching, to 840 mm long: stipe proximally castaneous, stramineous 
distally, adaxially sulcate, to 385 mm long, to 5 mm in diameter, 
densely paleated; larger paleae broadly attached, concolorous or 
bicolorous, stramineous or with a castaneous central section, these 
occurring on the proximal third of the stipe, lanceolate to broadly 
ovate, cordate to cordate-imbricate, the margins with closely to 
widely spaced, short, straight or curved, often branched fimbriae, 
the apex acicular or with a thin-walled apical cell, to 20 x 7 mm; 
smaller paleae short-stalked, narrowly lanceolate to narrowly trian- 
gular, cordate to cordate-imbricate, the margins proximally with 
closely spaced, short and long, often branched fimbriae, the apex 
entire or with short, firm, widely spaced fimbriae terminating in an 
acicular cell or with a thin-walled cell: lamina 2-pinnate to 2- 
pinnate-pinnatifid, firmly herbaceous, dark green adaxially, paler 
abaxially, narrowly ovate to elliptic, to 510 mm long, proximally 
somewhat reduced, the proximal pinnae usually somewhat deflexed: 
rachis stramineous, adaxially sulcate, densely paleated; paleae short- 
stalked, ferrugineous, chartaceous, narrowly lanceolate, narrowly 
ovate or narrowly triangular, cordate to cordate-imbricate, the mar- 
gins proximally with long and short, straight or twisted marginal 
outgrowths becoming simple towards the apex, the apex usually 
acicular, rarely terminating in a thin-walled apical cell, to 1 1 .5 x 2.5 
mm: pinnae often widely spaced proximally, somewhat overlapping 
distally, oblong-attenuate to narrowly oblong-attenuate, to 170 x 30 
mm: pinna-rachis stramineous, adaxially sulcate, densely set with 
paleae similar to but smaller than those on the rachis, to 7 x 1 .5 mm: 
pinnules opposite to alternate, closely spaced, often imbricate, 
asymmetric, narrowly ovate to ovate, acroscopically auricled, ser- 
rate to lobate-serrate, sharp-tipped to aristate, the proximal acroscopic 
pinnule the largest, the proximal pinnules often acroscopically 
incised to or near to costa, to 22 mm long; adaxially with a few 
simple and filiform or proximally hastate paleae along the proximal 
part of the costa, the apical cell always acicular, to 4.5 mm long; 
abaxially sparsely set with ferrugineous, short-stalked, narrowly 
triangular to subulate paleae, proximally usually with a few short or 
long, straight or curved outgrowths, the apical cell always acicular, 
to 4 x 0.9 mm long. Venation immersed. Sori circular, c. 1 .5 mm in 
diameter, essentially uniseriate, terminal or near terminal on abbre- 
viated vein branches: sporangium with 7-( 1 2)-2 1 indurated annulus 
cells; stalk eglandular: indusium stramineous to castaneous, often 
black-centred, peltate, circular, entire to repand, the maximum 
radius 0.31-(0.61)-1.04 mm, persistent. Spores brown, 32 per spo- 
rangium, the perispore variable, folded to form a reticulum of 
inflated or compressed ridges and/or tubercles, the crests and areas 
between echinulate to spiculate, the areas between ridges perfo- 
rated, the exospore 38-(59.07)-74 x 22-(43.76)-60 Jim. 
Chromosome number 2n=246, apogamous. 

MATERIAL EXAMINED 

LESOTHO. 2828 (Bethlehem): Leribe (CC), Dieterlen 167, in part (B); 
Mafeteng, Mont Ka-majapela, Dieterlen s.n. (BR); Leribe, 5000-6000 ft, 
Dieterlen 695 (NBG, PRE, SAM). 2927 (Maseru): Mamathes District, 5850 
ft (BB), Guillarmod 835 (PRE); Roma, 5550 ft (BC), Ruch 1909 B-only 
(PRE); Roma, W. ravine, Schmitz 399 (PRE); Roma, SE-facing slope, 
Schmitz 402 (PRE), Roma, Schmitz 6888 (PRE); Morija (DA), Dieterlen 
1309 A-only (PRE). 2929 (Underberg): Sehlabathebe National Park, 
Mofoqoi, 2350 m, Hoener 1479 (BOL). 

SOUTH AFRICA. 2730 (Vryheid): Hlobane, Johnstone 295 (NH). 2828 
(Bethlehem): Clarens, mountain N. of Leibrandt Kloof (CB), Roux 937 
(NBG); Golden Gate National Park, NE of Glen Reenen Camp, 6200 ft (DA), 
Liebenberg 7498 (PRE); Golden Gate National Park, Wonderhoek, 
Gertenbach & Groenewald9\ 67 (PRE): Golden Gate National Park, Roberts 
3235 (PRE); Golden Gate National Park, Rossouw 406 (BOL); N. of Mont- 
aux-Sources, Witsieshoek area, 1800 m (DB), Junod 14 (P); Royal Natal 



68 



J.P. ROUX 




10 mm 



10 mm 



Fig. 14 Polystichum monticola. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. A, B, drawn from Roux 2581 (NBG); C, 
drawn from Van Jaarsveld 6494 (NBG). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



69 



National Park, Gudu Forest, Edwards 325 (NU, PRE); Royal Natal National 
Park, Gudu Forest, Roux 25 1 3 (NBG); shady gully on the Lion facing the 
Tugelana Valley, c. 6300 ft, Esterhuysen 35645 (B, BOL, NBG, NU, PRE); 
Witsieshoek, 6000-7000 ft, Thode s.n. (NBG); Royal Natal National Park, 
Gudu Forest, 5000 ft, Schelpe 1455 (NU); Mont-aux-Sources, Bottomley s.n. 
(PRE); Mont-aux-Sources, Tugela River, near gorge, Hutchinson 41 (PRE); 
Mont-aux-Sources, Schweickerdt 760B (PRE); Royal Natal National Park, 
West 1 284 (PRE); Quaqua mountains, Lefika, Rustenberg s.n. (P); versant N. 
du Mont-aux-Sources, region de Witsies Hoek, c. 1 800 m, Junod 1 4 (P). 2829 
(Harrismith): Swinburne, Boschhoek,E. slopeof Manyenyeza(AC),Jacofaz 
4711 (PRE); Harrismith, Platberg, valley below Monkey Point, Roux 2520 
(NBG); Platberg, 1600 m, NBG Exped. s.n. (BOL); Harrismith, Platberg, 
1600 m, sine coll. 151191 (NBG); Harrismith, Platberg, dolerite cliffs left of 
summit of Donkey's Pass, Roux 2528 (NBG); Platberg, in shade of concrete 
furrow leading to Hawkins Dam, 2220 m, Roux 2527 (NBG); Harrismith, 
Platberg, Roux 2523 (NBG); Harrismith, Kerkenberg, Schelpe 7278 (BOL); 
Cathedral Peak area, nKdhlanhla Forest, Schelpe p.47 (NU); Cathedral Peak 
area, 5000 ft, Brass 76 (NU); Cathedral Peak area, Indumeni Forest, 5 1 00 ft, 
Schelpe 737 (NU); Van Reenen, 6800 ft (AD), Schlechter 6932 (B, NBG, 
PRE); Oliviershoek Pass, forests S. of Seheletwane (CA), Roux 25 1 8 (NBG); 
Oliviershoek, 6000-7000 ft, Thode s.n. (NBG); Cathedral Peak Hotel, along 
Cathedral Peak path, 1500 m (CC), Goetghebeur4563 (BR, PRE); Cathedral 
Peak area, Brass 67, 79, 82, 96, 98 (NU); Cathedral Peak Forest Station, 6050 
ft, Killick 1 134 (NU); Cathedral Peak area, 2000-3000 ft, Wilker 69 (NU); 
Drakensberg, MnWeni Pass, c. 8500-9000 ft, Esterhuysen 27830 (PRE). 
2831 (Nkanhla): Ngoye, 2-3000 ft (DC), Medley-Wood 10886 (PRE). 2926 
(Bloemfontein): Thaba 'Nchu Mountain, 6500 ft (BB), Roberts 2998 (PRE). 
2929 (Underberg): Giants Castle Nature Reserve, forests above Hillside 
campsite, c. 1600 m (AB), Roux 2499, 2501, 2506, 2507 (NBG); Injasuti 
area, 5000 ft, Esterhuysen 26041 (BOL); Kamberg area, Storms Heights, c. 
7000 ft (BC), Milliard & Burn 11795 (NU, PRE); Mpendhle District, 
Mulangane Ridge, above Carter's Nek, 7000-7300 ft, Milliard & Burn 1 695 1 
(BOL); 18406 (NBG, PRE); Kamberg, 'Game Pass', 6100 ft, Gordon-Gray 
85 (NU); Drakensberg Garden State Forest Reserve, near Mlambonja River 
(CA), Van Jaarsveld 6492, 6494 (NBG); Drakensberg Gardens, 6000 ft, 
Bronhead 59 (NU); Drakensberg Gardens, Dyer 73 (NU); Drakensberg 
Garden area, 6000 ft, Schelpe p54 (NU); Drakensberg Garden Hotel, up to 
Rhino Peak, along Mlambonja River, 2000 m, Goetghebeur45 1 9 (BR, PRE); 
Gxalingenwa Valley between Sani Pass and Polela Valley, c. 6700 ft (CB), 
Milliard & Burn 17076 (NU, PRE); Cobham Forest Station, Ndlovini, 
Troutbeck, c. 6000 ft, Milliard & Burn 13311 (NU), 13329 (BOL, NU); 
Cobham Forest Reserve, Sipongweni, 6500 ft, Milliard & Burn 141 35 (BOL, 
NU, PRE); Cobham Forest Reserve, 'Lakes' Cave area, c. 7800 ft, Manning 
et al. 1 59 1 8 (BOL, NU); Garden Castle area, 9000 ft (CD), Crooked 62 (NU); 
Garden Castle area, Elliott 31 (NH), 37 (NU); Boston, Impendhle, 5000 ft 
(DB), Randies 1 85 (NU); Impendhle, 5 200 ft, Clarkson 1 33 (NU); Impendhle, 
c. 5000 ft, Huntley 167 (NU); Bulwer, Marwaga Mountain, farm Sunset, 
1810 m (DC), Roux 2309 (NBG); Farm Sunset, 5800 ft, Rennie 1441 (NU, 
PRE); Himeville District, 5000 ft, Webb 101 (NU); Farm Sunset, gully above 
dams, 6000 ft, Rennie 1055 (NU); Polela District, near Bulwer (DD), Henkel 
s.n. (NU); Bulwer, 5100 ft, Clarkson 182 (NH, NU, PRE); Bulwer, Allsopp 
843 (NU). 2930 (Pietermaritzburg): Zwaartkop (CB), Sim s.n. (NU, PRE). 
3018 (Kamiesberg): Kamiesberg, Rooiberg, 5500 ft (AC), Rourke 1684 
(BOL, NBG, PRE); Rooiberg, farm Damsland, Roux 2453 (NBG). 3027 
(Lady Grey): Herschel District, Majuba Nek, Hepburn 262, B-only (GRA). 
3028 (Matatiele): Ongeluks Nek, c. 4 km from Lesotho border post, 2250 m 
(AB), Matthews 916 (NBG). 3029 (Kokstad): Mt. Currie, farm Highland 
Home (AB), Roux 2488, 2490 (NBG); Mt. Currie, Edwards 214a (NU); Mt. 
Currie, 5200 ft, Edwards 44 (NU); Kokstad (CB), Mogg 1927 (PRE). 3030 
(Port Shepstone): Ixopo (AA), Hancock s.n. (NU). 3124 (Hanover): 
Compassberg, near top, c. 2440 m. (DC), Trollip s.n. (PRE); top of 
Compassberg, Coetzee s.n. (PRE); Compassberg, farm Grootkop, 6300 ft, 
Acocks 23447 (PRE). 3126 (Queenstown): Broughton, near Molteno (AD), 
Flanagan s.n. (SAM); Broughton near Molteno, Flanagan 1681 (PRE); 
Broughton, Molteno, 6300 ft, Flanagan 527 (PRE); Queenstown, Hangklip 
Mountain, 6600 ft (DD), Roberts 20 1 2 (PRE). 3128 (Umtata): Maclear, farm 
Woodcliffs (AB), Roux 2481, 2484, 2485 (NBG). 3218 (Clanwilliam): 
Clanwilliam Division, between Tafelberg and Spout, 6000 ft (BB), Schelpe 
1960 (BOL, K). 3219 (Wuppertal): Gideon's Kop, 1500 m (CB), Burrows 



1235 (BOL). 3224 (Graaff-Reinet): Mount Oudeberg near Graaff-Reinet 
(DD), sine coll. 96897 (B); in fissuris rupium in monte Oudeberg prope 
Graaff-Reinet, 4800 ft, Bolus 1 736 (BOL, K). 3225 (Somerset East): Montis 
Boschberg (DA), MacOwen 870 (BR). 3226 (Fort Beaufort): Katberg, 
3500-4000 ft (BC), Baur 865 (B, GRA); Katberg Pass summit, farm Pleasant 
View, Roux 2697 (NBG); Hogsback, Elandsberg summit (DB); Roux 2688 
(NBG); Hogsback, Zincucha Forest, Roux 2683 (NBG). 3318 (Cape Town): 
in numerosis umbrosis montis Tafelberg, Paradys (CD), MacOwen s.n. (P); 
Table Mountain, Dark Gorge, below Saddle, SE side, Esterhuysen 26563 (B, 
BOL, PRE); Stellenbosch, Jonkershoek, Langrivierkloof (DD), Roux 2580, 
2581, 2582, 2583 (NBG); Stellenbosch, Simonsberg, 3000 ft, Esterhuysen 
25453 (BOL); Stellenbosch, Helderberg, Disa Gorge, Esterhuysen 28475 
(BOL). 3319 (Worcester): Great Winterhoek Mountains, 4000-5000 ft 
(AA), Esterhuysen 26982 (B, BOL, NBG, PRE); Tulbagh, W. slopes of 
Swartgat Peak, Witzenberge, 4000 ft (AC), Esterhuysen 1 69 14 (BOL, NBG); 
Tulbagh, Great Winterhoek Mountains, 5500 ft, Esterhuysen 19787 (BOL, 
NBG); Ceres, Baviaansberg (BA), Stokoe s.n. (NBG, SAM); shale band 
below Milner Peak, Hex River Mountains, 5000 ft (AD), Esterhuysen 14264 
(BOL), 14885 (BOL, NBG); Milner Ridge Peak and Buffels Dome, 5000 ft, 
Esterhuysen 28708B (BOL); Ceres, Slab Peak, 1310m, Winter 431 (NBG); 
Hex River Mountains, Moraine Kloof, c. 4000 ft, Ashton 352 (BOL); 
Goudini, Waaihoek Mountains, 4000 ft (CB), Barnard s.n. (SAM). 3320 
(Montagu): Boesmansbos (DD), Adamson s.n. (BOL). 3321 (Ladismith): 
Swartberg, near Ladismith, Toverkop, 6500 ft (AC), Esterhuysen 26698 (B, 
BOL, NBG, NU, PRE); below Toverkop on S. slope of Swartberg, 5000- 
6000 ft, Esterhuysen 14013 (BOL, PRE); cliffs at base of Toverkop, 6500 ft, 
Esterhuysen 18511 (BOL). 3322 (Oudtshoorn): Prince Albert Division, 
Swartberg Mountains (AC), Stokoe 9410 (NBG, SAM); Prince Albert, Popta 
s.n. (L); Blesberg, 6000 ft (BC), Esterhuysen 24920 (BOL); Montagu Pass 
(CD), Schweickerdt 4705 (PRE); Kammanassie Mountains, Mannetjiesberg, 
4200 ft (DB), Matthews 1023 (NBG); S. slope of Mannetjiesberg, 
Kammanassie Mts, 5000 ft, Esterhuysen 1 8396 (BOL). 3323 (Willowmore): 
Hoopsberg, S. slope, 5000 ft (CB), Esterhuysen 6557 (BOL). 3324 
(Steytlerville): S. slopes of Kouga Peak near Joubertina (CA), Esterhuysen 
16280 (BOL, NBG); Uitenhage, Cockscomb, Great Winterhoek mountains, 
4800 ft (DB), Esterhuysen 27090 (BOL, PRE). 

ZIMBABWE: Victoria Falls, Sim s.n. (PRE). 

WITHOUT EXACT LOCALITY: Orange Free State, Wittebergen, ad 
Caledonrivier, Rehmann 3938, 3978 (B); Natal, sine coll. s.n. (NBG); 
Blinkwater Bush, Graham 84 (NU); Transvaal, Repton 5B (PRE); in summo 
monte Koudveld, 6500 ft, Tyson 140 (PRE); in sylvis umbrosis faecium 
montium Hott. Holland., sine coll. 341 (S); Malappa's Place, Rustenberg s.n. 
(P); Natal, sine coll. s.n. (BR); Basotholand, Koopowitz s.n. (GRA); Bushmans 
Cave, Lubke s.n. (GRA). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum monticola 
is characterized by its largely montane habitat, the usually large 
stands it forms, the short decumbent rhizome clothed by long, 
narrowly linear to cuspidate paleae, the closely spaced, firmly 
herbaceous fronds, smallish pinnules, bicolorous larger paleae on 
the proximal part of the stipe, short-stalked paleae with proximal 
margins that bear stiff, often branched outgrowths, and with an 
apical cell that is not thin-walled, the relatively large repand to 
entire, persistent indusia, the 32 spores borne by each sporangium, 
and the somatic chromosome number of 2n=246. 

In palea morphology Polystichum monticola exhibits features of 
sections Lasiopolystichum and Metapolystichum. Both of these 
sections are characterized by short, unbranched, suberect to erect 
rhizomes resulting in the plants growing as individuals. In P. 
monticola, however, the rhizome is decumbent and mostly repeat- 
edly branched: as a result plants form large clonal stands. In spore 
morphology it is closer to section Metapolystichum than to section 
Lasiopolystichum . 

VARIATION. Depending on habitat, Polystichum monticola shows 
considerable variation in frond size, lamina texture, pinnae arrange- 
ment and pinnule size. Plants from xeric habitats have short fronds 
with closely spaced pinnae and small imbricate pinnules that are 



70 



J.P. ROUX 



firm-herbaceous in texture. Some collections proved to have unusu- 
ally large indusia, with a maximum radius up to 1.04 mm having 
been measured on a plant from Milner Peak in the Hex River 
Mountains [Esterhuysen 14885 (NBG, BOL)]. Indusia of the type 
collection are unusually small with the mean radius being merely 
0.36 mm (n=6). Collections with these unusually large indusia occur 
at random throughout the distribution of the species and cannot be 
ascribed to any environmental condition. The same pattern applies 
to the often black-centred indusia. Although the number of indurated 
annulus cells per sporangium is relatively uniform in the species, 
unusually low and high numbers have been recorded. A plant from 
Leribe, Lesotho [Dieterlen 695 (SAM)] has sporangia with the 
number of indurated annulus cells varying between 7 and 13 
(x=10.66; n=50), whereas a plant from Platberg, Harrismith [Roux 
2523 (NBG)] has sporangia with the number of indurated annulus 
cells varying between 13 and 17 (x=14.7; n=50). Plants from forest 
habitats generally have larger fronds that are softer in texture to 
those occurring in more exposed habitats where the fronds are 
smaller and firm-herbaceous to subcoriaceous. 

DISTRIBUTION AND ECOLOGY. Polystichum monticola is confined 
to southern Africa where it has been recorded from the Northern 
Cape, Western Cape, Eastern Cape, KwaZulu-Natal and Free State, 
as well as in Lesotho, with one isolated record from the Victoria 
Falls in Zimbabwe that needs to be confirmed. The species is largely 
confined to the mountains ranging from the Cape Peninsula, along 
the southern Cape mountains to the Drakensberg Escarpment as far 
north as Platberg in the northeastern Free State. Plants have also 
been recorded from outlying locations such as the Kamiesberg in the 
Northern Cape Province and from Thaba 'Nchu Mountain in the 
eastern Free State. This apomictic taxon occurs at elevations ranging 
from 600 to 2740 m in often xeric environments. The habitat 
includes rock crevices in screes, cliff bases, streambanks, forest 
margins and forest floors. Often growing in exposed habitats where 
it forms large masses, the species is frequently exposed to veld fires, 
from which it soon recovers. 

15. Polystichum dracomontanum Schelpe & N.C. Anthony in 
Contr. Bolus Herb. 10: 145 (1982). Type: Natal, Bergville Divi- 
sion, on banks above stream in side kloof west of main kloof, 
shortly above the Singati Cave (E. of Mont-aux-Sources), in 
unburnt sparse bush or small trees or in the open, c. 6000 ft, 
Esterhuysen 35646 (BOL!-holotype; B!, BOL!, C, GH, K, M, 
MO, NU!, P, PRE!, S-isotypes). 

Fig. 15. 

Plants terrestrial or epilithic. Rhizome decumbent, stoloniferous, to 
10 mm in diameter, densely set with roots, persistent stipe bases, and 
paleae; paleae ferrugineous, broadly attached, chartaceous, linear- 
attenuate, entire, the apex terminating in a small thin-walled cell, to 
28 x 2 mm. Fronds closely spaced, 5-7 per plant, erect or arching, to 
1.15 m long: stipe proximally castaneous, stramineous distally, 
adaxially sulcate, to 540 mm long x 6 mm in diameter, initially 
densely paleated, becoming glabrous with age, proximally with 
paleae similar to those on the rhizome, the paleae distally of two 
types; larger paleae broadly attached, ferrugineous to castaneous 
throughout or stramineous to ferrugineous with a castaneous central 
region or apex, chartaceous, narrowly ovate to ovate, cordate, the 
margins with short, straight or curved projections, the apex often 
flagelliform, terminating in a thin-walled cell, to 25 x 6 mm; smaller 
paleae ferrugineous to stramineous, membranous, narrowly ovate to 
lanceolate, cordate to cordate-imbricate, the margins closely set 
with short and/or long, straight or curved, often branched outgrowths, 
the apex flagelliform, acicular or terminating in an oblong to clavate 



thin-walled cell, to 1 1 .5 x 1.8 mm: lamina 2-pinnate to 2-pinnate- 
pinnatifid, with up to 24 free pinna pairs, ovate, to 610 mm long: 
rachis stramineous to greenish, adaxialy sulcate, initially densely 
paleated; paleae short-stalked, membranous, narrowly lanceolate to 
narrowly ovate, cordate to cordate-imbricate, the margins proxi- 
mally with long and/or short, straight or curved, often branched 
outgrowths, the apex acicular or flagelliform and terminating in a 
thin-walled cell, to 7.3 x 1.6 mm: pinnae proximally short-stalked, 
1 -pinnate to 1-pinnate-pinnatifid, with up to 18 free pinnule pairs, 
proximally widely spaced, often slightly reduced, distally often 
overlapping, ovate to narrowly ovate, to 105 mm long: pinna-rachis 
stramineous, adaxially sulcate, initially densely paleated; paleae 
stramineous, membranous, narrowly oblong to narrowly triangular, 
cordate to cordate-imbricate, the margins proximally with long and/ 
or short, curved, often branched outgrowths, the apex acicular, to 
9.75 x 2 mm: pinnules opposite to alternate, firmly herbaceous to 
coriaceous, dark green adaxially, paler abaxially, proximally widely 
spaced to imbricate, asymmetric, ovate to ovate-rhomboid, often 
somewhat falcate, acroscopically auricled, serrate to doubly-serrate, 
sharp-tipped to strongly aristate, the margins somewhat revolute, 
the proximal acroscopic pinnule usually slightly longer than the 
next, to 22 mm long; adaxially glabrous or with a few stramineous, 
membranous, filiform, acicular paleae along proximal part of the 
costa, to 9.5 mm long, abaxially sparsely paleated; paleae short- 
stalked, stramineous, membranous, filiform, narrowly oblong to 
narrowly triangular, cordate, the margins proximally with short and/ 
or long, straight or curved outgrowths, the apex terminating in an 
acicular cell or with a thin-walled cell, to 8 mm long. Venation 
immersed or raised. Son circular, c. 2 mm in diameter, terminal or 
near-terminal on abbreviated vein branches, uniseriate, discrete to 
confluent at maturity: sporangium with 10-(13)-21 indurated annu- 
lus cells; stalk eglandular: indusium ferrugineous to stramineous, 
chartaceous, persistent, peltate, circular, with or without central 
processes, repand to weakly erose, the maximum radius 0.63- 
(0.81)-1.09 mm. Spores 64 per sporangium, brown, the perispore 
globose or folded to form inflated or narrow reticulate ridges, 
minutely and sparsely perforated, the ridges and areas between 
echinate to echinulate, the exospore 25-(54.24)-70 x 28-(41.04)- 
58 (im. Chromosome number unknown. 



MATERIAL EXAMINED 

LESOTHO. 2927 (Maseru): pente de la montagne de Mathatha district de 
Mafeteng (CC), Dieterlen s.n. (P). 

SOUTH AFRICA. 2828 (Bethlehem): Tugela Gorge above chain ladder, 
6200 ft (DB), Milliard & Burtt 15445 (BOL, NU); Royal Natal National Park, 
Tugela Gorge, scrub above chain ladder, Roux 27 1 5 (NBG); tributary flowing 
into the Singati, E. of Mont-aux-Sources (DD), Esterhuysen 35644 (B, BOL, 
NBG, PRE); Bergville, Mont-aux-Sources, Schweickerdt 760 (PRE); 
Mbunduni (MnWeni area), 9000 ft, Esterhuysen 27811 (BOL); MnWeni 
Pass, 8000 ft, Esterhuysen 27839 (BOL, PRE). 2829 (Harrismith): Farm 
Bosch Hoek (AD), Roux 896, 897, 898 (NBG); MnWeni area, foot of Rockies 
Pass, 5500 ft, (CB), Esterhuysen 21656 (BOL); Rockies Pass, 8000 ft, 
Edwards 2145 (NU); Cathedral Peak area, shady side of kloof (CC), 
Esterhuysen 15486 (BOL, NBG); Cathedral Peak, sheltered slopes below 
Cleft Peak, 8000 ft, Esterhuysen 101 99a (BOL, PRE); between Cathedral 
Peak and Royal Natal National Park, MnWeni Pass, 9000 ft, Edwards 85 1 
(NU, PRE); SE slope of The Camel, 8700 ft, Everson 73, 74 (BOL); NE 
facing slope of The Camel, 8700 ft, Everson 75, 76 (BOL); Bergville, Orange 
Peel Gap, 7200 ft, Everson 71, 72 (BOL); NE slope of The Camel, 7000 ft, 
Schelpe 756 (NU); Orange Peel Gap, 2420 m, Crouch 647 (NU); Cathedral 
Peak, Nixon s.n. (NU); Cathedral Peak area, Umbonbonja River, 6000 ft, 
Schelpe 1096 (NU). 2929 (Underberg): Injasuti Valley, Solitude (AB), 
Malan 1 (BOL, NBG); Injasuti Nature Reserve, Leucosidea scrub at camp- 
site, Roux 2721 (NBG); Ndedema Valley, 6000 ft, Noel 1 28 1 (GRA); Injasuti 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



71 




10 mm 



Fig. 15 Polystichum dracomontanum. A, proximal part of lamina; B, rhizome; C, fertile pinnule. All drawn from Esterhuysen 35646 (BOL). 



72 



J.P. ROUX 



area, 6500 ft, Esterhuysen 35219 (BOL); Injasuti area, 6500-8000 ft, 
Esterhuysen 26039 (BOL); Injasuti area, 7000 ft, Esterhuysen 26050 (K); 
Tabamhlope Mountain, 6000 ft (BA), West 184 (PRE); Mulangane Ridge, 
above Carter's Nek, 7000-7300 ft (BC), Milliard & Burtt 17032, 17529 
(BOL, NU, PRE); 5-7 miles NNW of farm Castle View, headwaters of 
Mlahlangubo River, 6700 ft (CB), Milliard & Burtt 15188 (BOL, K, NU); 
Cathedral Peak, Ndumeni Valley, 1950 m (CC), Farrell2\ (NH). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum 
dracomontanum may be confused with P. monticola, which occurs 
in the same region and often in similar habitats. It can, however, be 
separated from it in the decumbent rhizome that produces slender 
stoloniferous branches, in the stipe that becomes near glabrous with 
age, and in the ovate lamina. Other diagnostic features are the entire 
rhizome paleae that evidently always terminate in a short thin- 
walled cell, in contrast to the often acicular apical cell in P. monticola. 
Paleae from the stipe, rachis, pinna-rachis and abaxial lamina sur- 
face appear to terminate more often in a thin- walled cell in contrast 
to those of P. monticola, where the apical cell appears to be largely 
acicular. The adaxial surface of the pinnules is largely glabrous but 
a few filiform paleae may occur proximally along the costa. These 
paleae always terminate in an acicular apical cell. Paleae from the 
abaxial surface of the pinnules largely terminate in an acicular apex, 
but paleae terminating in a short thin-walled cell are not unknown. 
The smaller paleae in P. dracomontanum are stramineous and 
membranous. The coriaceous lamina and somewhat revolute pinnule 
margins are also diagnostic. Micromorphological characters sepa- 
rating P. dracomontanum from other taxa are the small, almost 
square, adaxial epidermal cells with almost straight anticlinal walls 
and the almost circular stomata that are visible at a x 1 2 magnifica- 
tion. Considering the rhizome and palea morphology, P. 
dracomontanum is allied to the P. pungens group of species. 

VARIATION. Relatively little variation occurs within the species. 
Variation was observed in the colour of the larger stipe paleae. In 
most cases they are stramineous to ferrugineous throughout, but in a 
few collections they are centrally castaneous with a narrow 
stramineous margin. The paleae also show some variation in that 
some terminate in a long acicular cell whereas others may terminate 
in a short, thin-walled cell. Pinnules vary in size and in the margins 
that may be sharp-tipped to aristate. 

DISTRIBUTION AND ECOLOGY. Polystichum dracomontanum is 
largely confined to the northern Drakensberg Escarpment between 
Lesotho and KwaZulu-Natal where it occurs on both the lower 
Clarens Sandstone and the upper basalt formations. The species also 
occurs further northwards along the escarpment between the Free 
State and KwaZulu-Natal. Isolated populations have also been 
reported from the Mafeteng District in southeastern Lesotho. It 
occurs at elevations ranging from 1675 to 2745 m. Within this 
distribution the species is restricted to two vegetation types as 
defined by Acocks (1988). Along the high Drakensberg escarpment 
it occurs in Themeda-Festuca Alpine Veld that receives an annual 
precipitation of 600-1 900 mm. Polystichum dracomontanum occurs 
in grasslands and scrub forests associated with this vegetation type. 
To the north, along the escarpment between the Free State and 
KwaZulu-Natal, dominated by the Clarens Sandstone formation, the 
species occurs in sheltered forests of the Highland Sourveld type. 
Precipitation in this region measures between 750 and 1500 mm. 
The habitat includes streambanks, boulder bases, screes and scrub, 
and rarely also forests. Polystichum dracomontanum prefers moist 
cool slopes in shaded gullies and kloofs where it often forms large 
stands. At certain sites the species is subjected to regular veld fires, 
but this appears to have no adverse effect on plants. 



1 6. Polystichum incongruum J.P. Roux in Bot. J. Linn. Soc. 125: 36 
(1997). Type: South Africa, 3320 (Montagu): Swellendam, 
Marloth Nature Reserve, Koloniesbos, in scree on dry E.-facing 
slope (CD), Roux 2377 (NBG!-holotype). 

Fig. 16. 

Plants terrestrial. Rhizome decumbent, stout, to 16 mm in diameter, 
sparsely branched, set with roots, crowded stipe bases, and castaneous 
to ferrugineous, chartaceous paleae. Fronds crowded, to 8 per plant, 
suberect to arching, to 1.8 m long: stipe proximally castaneous, 
stramineous for most of its length, adaxially sulcate, to 930 mm long 
x 7 mm in diameter, moderately to densely paleated; paleae at stipe 
base of two types, the larger broadly attached, ferrugineous to 
stramineous, membranous, narrowly to broadly ovate-acuminate, 
rarely with unicellular clavate cells on the paleae surface, cordate to 
cordate-imbricate, the margins with short close-set outgrowths, the 
apex often flagelliform, terminating in an acicular cell or an oblong 
thin-walled cell, to 20 x 6 mm, the smaller sessile to short-stalked, 
stramineous, membranous, narrowly ovate-acuminate to narrowly 
triangular, truncate to cordate-imbricate, with short and/or long, 
straight or curved, somewhat lacerate outgrowths proximally, the 
apex flagelliform, entire and twisted, mostly terminating in an 
acicular cell but often also in a thin- walled cell: lamina 2-pinnate to 
3-pinnate, with up to 27 free pinna pairs, ovate to broadly ovate, to 
870 mm long, the pinnae more widely spaced proximally, the distal 
pinnae often imbricate, the proximal pinnae not to strongly reduced, 
not or slightly deflexed: rachis stramineous, adaxially sulcate, 
moderately to densely paleated; paleae sessile to short-stalked, 
ferrugineous to stramineous, membranous, narrowly ovate-acumi- 
nate to narrowly triangular-acuminate, truncate, cordate or 
cordate-imbricate, the margins proximally with short and/or long, 
curved, often branched outgrowths reduced in size and number 
towards the apex, the apex often flagelliform, twisted, terminating in 
an acicular cell or an oblong thin-walled cell, to 12x3 mm: pinnae 
1 -pinnate to 2-pinnate, with up to 21 free pinnule pairs, narrowly 
oblong-attenuate to narrowly ovate-attenuate, themiddle pinnae to 
265 mm long, to 95 mm wide: pinna-rachis stramineous, adaxially 
sulcate, moderately to densely paleated; paleae sessile to short- 
stalked, ferrugineous to stramineous, membranous, narrowly 
ovate-acuminate, narrowly triangular-acuminate to subulate, cor- 
date to cordate-imbricate, the proximal margins with short and/or 
long, often branched outgrowths reduced in size and number to- 
wards the apex, the apex often flagelliform, twisted, largely 
terminating in an acicular cell but often also in an oblong thin- walled 
cell, to 7 x 2 mm: pinnules opposite to alternate, closely to widely 
spaced, often slightly imbricate, firmly herbaceous to coriaceous, 
dark green adaxially, slightly paler abaxially, inaequilateral, ovate to 
narrowly trullate, often somewhat falcate, basiscopically cuneate, 
acroscopically cuneate to truncate and auriculate, the larger pinnules 
commonly deeply incised to form free or nearly free, narrowly 
ovate, narrowly elliptic to narrowly obovate ultimate segments, the 
margins serrate to doubly serrate, rarely aristate, the proximal 
acroscopic pinnule to 60 mm long, to 13 mm wide, often overlap- 
ping the pinna-rachis above; adaxially sparsely set with stramineous, 
membranous, twisted paleae chiefly along the costa, these filiform, 
narrowly linear to narrowly linear-hastate, simple or with few short 
marginal outgrowths proximally, the apex usually terminating in an 
acicular cell but often also in an oblong thin-walled cell, to 4.5 mm 
long, abaxially moderately paleated; paleae stramineous, membra- 
nous, subulate to narrowly triangular, twisted, short-stalked, truncate 
to cordate, proximally with short and/or long, often branched, 
somewhat lacerate marginal outgrowths, the apex usually termin- 
ating in an acicular cell, up to 3.5 mm long. Venation adaxially 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



73 




10 mm 



Fig. 16 Polystichum incongruum. A, middle pinnae of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2377 (NBG). 



74 

obscure, raised abaxially. Sori circular, up to 1 .5 mm in diameter, 
terminal or near terminal on abbreviated vein branches, essentially 
uniseriate, discrete: sporangium with 1 1-(13)-21 indurated annulus 
cells; stalk glandular or eglandular: indusium stramineous or 
stramineous and black-centred, persistent, circular, simple or with 
central processes, the margins entire, minutely undulate to erose, 
often bearing clavate unicellular glands, the maximum radius 0.22- 
(0.46)-0.82 mm. Spores stramineous, the perispore folded to form 
narrow and broad reticulate ridges, the ridges crested, the crests and 
areas between minutely echinulate, porate, the exospore 30-(43.84)- 
72 x 22-(33.22)-52 urn. Chromosome number 2n=164, n=82. 

MATERIAL EXAMINED 

SOUTH AFRICA. 3226 (Fort Beaufort): summit of Katberg Pass (BC), 
flow* 2426 (NBG); Katberg forests, 3000 ft, Adams 113, 118, 134, 147(NU); 
Hogsback, Zingcuka Forest (DB), Roux 2409, 2412 (NBG); Hogsback, 
Elandsberg, SE slope above scree, 4900 ft, Furness & Phillipson 49 (PRE); 
Hogsback, Auckland Forest Station, Roux 2418, 2419, 2420, 2421 (NBG); 
Hogsback Forest Reserve, 750 m, Dahlstrand 2926 (NBG); Hogsback Forest 
Reserve 'Fern Walk', 800 m, Dahlstrand 1550 (PRE); Auckland Forest, 
Hogsback, 750 m, Dahlstrand 28 1 9 (NBG); Hogsback, 3900 ft, Griffen 1 852 
(PRE); Hogsback, Buckner s.n. (NU); Hogsback, Dahlstrand 1853 (NU); 
Hogsback, 4000 ft, Hoal 36 (NU); Middle Drift District, Hogsback, 3900 ft, 
Schelpe 6556 (B, PRE). 3227 (Stutterheim): Keiskamma Hoek, Gxulu 
Mountain, 5500 ft (BA), Story 3507 (PRE); Keiskammahoek, Gxulu Kop, 
4500 ft (CA), Wells 3343 (PRE); Perie Forest (CB), Sim s.n. (PRE); 
Stutterheim, Evelyn Valley, Taylor 4203 (NBG); Mount Kemp, 5000 ft, Sim 
s.n. (PRE); Perie Forest, 4000 ft, Sim s.n. (PRE); Amabele (DA), Hardcastle 
293 (NBG, PRE); 1 km past Amabele station on road to Stutterheim, Roux 
2678 (NBG). 3320 (Montagu): Swellendam, Koloniesbos (CD), Roux 2373, 
2377, 2589, 2590, 2591 (NBG); Swellendam, Duiwelsbos, along trail lead- 
ing to Die Plaat, Roux 2592, 2593, 2595 (NBG); Swellendam, Wamakersbos, 
Roux 2594 (NBG); Heidelberg, Grootvadersbos, Safraandraai (DD), Roux 
2598 (NBG); Grootvadersbosch West, 1200 ft, Kruger 1322 (NBG, PRE); 
Grootvadersbos, Plantkunde Dept. Univ. van Stellenbosch s.n. (NBG); 
Swellendam, Strawberry Hill, Esterhuysen 10371 (BOL). 3322 
(Oudtshoorn): Mossel Bay, Robinson Pass, Ruitersbos, Boesmansrivier, 
1200 m (CC), Roux 2603, 2604, 2605, 2606 (NBG); Mossel Bay, Ruitersbos 
Forest Station, Roux 2381 (NBG); Groot Brakrivier, Jonkersberg Forest 
Station, Langbos, 380 m, Roux 2607, 2608, 2609, 2610, 2611, 2612 (NBG); 
Robinson Pass, 1700 ft, Schelpe 4989 (BOL); George, 1 km from turnoff to 
Witklippen Forest on old George-Knysna road (CD), Roux 2391, 2392 
(NBG); George, lower circular drive, 275 m, Cameron 90 (PIC-SERM); 
George, Paterson 1239 (BOL); George, Schlechter 525 (PRE); George 
District, 200 m, Humbert 9834 (PRE); George, Montagu Pass, forest at 
summit of pass, Roux 2613 (NBG); Montagu Pass, Rehmann 118 (B); 
Woodville Forest, 440 m (DA), Roux 2622, 2623, 2624, 2625 (NBG); 
George, on road to Bergplaas Forest Station, 1 km past turnoff to Woodville 
hiking trail (DC), Roux 2395, 2396, 2397, 2398 (NBG); George, Saasveld 
Forest Station, Groenkop Research Area, Roux 2434 (NBG); George, 
Groenkop, Swartrivier, 300 m, Van Daalen 167 (BOL); old road between 
George and Knysna, above Touws River, Knysna side, 260 m, Roux 2620, 
2621 (NBG); George, Groenkop Forest, Geldenhuys 394 (BOL); Wilderness, 
Levyns s.n. (BOL); George, Wilderness, Mogg 11656 (PRE); George, 
Schlechter s.n. TM525 (PRE); in silvis pr. George, 300 m, Schlechter 2441 
(B); George, Wilderness, Jacobsen 2292 (PRE); George, Saasveld, forest 
above the reservoir, Roux 2384, 2385 (NBG); George, 1 .2 km from Saasveld 
turnoff on road to George, Roux 2437, 2438, 2439, 2440, 2441, 2614, 2615, 
2616, 2617, 2618 (NBG); old road between George and Knysna, above 
Touwsriver, George side, 210 m, Roux 2619 (NBG); Goudveld Forest 
Reserve, Jubilee Creek forest walk, 340 m (DD), Roux 2627, 2629 (NBG); 
Farleigh Forest Station, forest above Platbos hut, Roux 2401 (NBG). 3323 
(Willowmore): Concordia Forest Station, near Witklipdraai (CC), Roux 
2637 (NBG); Knysna, Kom-se-Pad, Gouna Forest, Grootdraai picnic site, 
Roux 2638 (NBG); Knysna, Kom-se-Pad, 2.2 km from T-junction to 
Diepwalle, Roux 2405 (NBG); Knysna, Kleinbos, Buffelsnek, Schelpe 4312 
(BOL); Knysna, Paardekop, Steyn 720 (NBG); Knysna, Deepwalls, Schonau 
318 (BOL); Knysna, Gouna, Schelpe s.n. (BOL); Keurbooms River Forest 



J.P. ROUX 

Reserve, 200 m (CD), Dahlstrand 1355 (NBG); Bloukrans Pass, Varkrivier 
(DC), Roux 2649, 2650, 2651 (NBG); Bloukrans Forest Station, Platbos, 
along hiking trail, 260 m, Roux 2645 (NBG); Bloukrans Pass, 300 ft, Schelpe 
4342 (BOL); Bloukrans Pass, Acocks 21298 (PRE); Tsitsikama Forest Re- 
serve (DD), Roux 2647 (NBG); Storms River Forest Reserve, 100 m, 
Dahlstrand 1693 (PRE); Stormsriver, 250 ft, Schlechter 5963 (PRE); 
Stormsriver Gorge, 400 ft, Jacobsen 2331 (PRE); in umbr. pr. Storms River, 
80 m, Schlechter 5963 (B); Tsitsikama Forest Reserve, 260 m, Roux 2648 
(NBG). 3423 (Knysna): Knysna, Kaffirkop Forest, 420 m (AA), Roux 2640, 
264 1 , 2642 (NBG), near Knynsna, Mitchell s.n. (M, PRE); Knysna, Kaffirkop 
Forest, Roux 1994, 1995, 1996 (NBG); Knysna, Marloth 5710, 5711 (PRE); 
Knysna, Rex s.n. (PRE); Knysna, Mitchell s.n. (BOL); Tzitzikamma forest, 1 
mile E. of Storms River village, 850 ft (BB), Schelpe 4372 (BOL); Storms 
River mouth, Maguire 507 (NBG). 3424 (Humansdorp): Hofman's Bosch 
(BB), Britten s.n. (PRE). 

WITHOUT PRECISE LOCALITY. Puspasvlei, Voormansbosch, 
Duivelsbosch and Keurboom, 1000-4000 m, Zeyher 4610 (PRE); George to 
Wilderness, Moss 6280 (PRE); Zuurberg, Rogers s.n. (PRE); loco incerto, 
Dahlstrand 1303 (NU); loco incerto, Zeyher s.n. B-97089 (B); loco incerto, 
Burchell 5200 (B); Kaffrarian forests, sine coll. 96893 (B); loco incerto, sine 
coll. B-96855 (B); Pr. b. sp, Zeyher 4610 (B); Cap. b. Sp., Krebs 360 (B); 
Prom. bon. spei, During s.n. (B); Cap. b. sp., Drege s.n. (B); Pr. b. sp., Ecklon 
& Zeyher 63 (B); Cap Colonie, Breutel s.n. (B); loco incerto, Herb. Lipzig, A 
& B only B-97050 (B); loco incerto, Herb. Lipzig 97051 (B); loco incerto, 
Braun s.n. (B); loco incerto, Gueinzius s.n. B-97069 (B); between Plettenberg 
Bay and Humansdorp, Rodin 1191 (BOL, PRE, S); loco incerto, d'Urban s.n. 
(B); loco incerto, Hort. bot. Berol. B-97049 (B); loco incerto, Hort. bot. 
Berol. B-97047 (B); Pr. b. spei, sine coll. B-97048 (B). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Differentiation in the 
field between this species and Polystichum pungens is not always 
easy because of their sympatric distribution, the variation within and 
among populations, and the absence of stable macromorphological 
characters. However, P. incongruum can be separated from P. pungens 
by its thicker, more stout rhizome. The pinnules in the former 
species are narrower, more slender and acuminate (often also slightly 
falcate), compared to the somewhat ovate to ovate-oblong, almost 
obtuse pinnules of P. pungens. A further character separating P. 
incongruum from P. pungens is the frequent occurrence of clavate 
unicellular glands along the sporangium stalk in the former species. 
Clavate unicellular glands occurring along the indusium margin 
have also been observed in some populations in the southern Cape, 
a feature never occurring in P. pungens. The former species is 
furthermore a sexual tetraploid (2n=164), whereas P. pungens is a 
sexual octoploid (2n=328). 

VARIATION. Variation in lamina morphology is perhaps the most 
apparent, hence the specific epithet (incongruens = inconsistent). 
Lamina division may vary between 2-pinnate to 3-pinnate, the 
pinnae being narrowly oblong-attenuate and widely spaced to nar- 
rowly ovate-attenuate and imbricate, with extreme variations often 
occurring within populations. Proximal pinnae may be reduced or 
not with the length ratio between these and the middle pinnae 
ranging between 1 : 1 and 1 :0.42. The most proximal pinna pair is 
often deflexed. Pinnules also show a large degree of variation in size 
and incision as illustrated by Roux (19976). Proximal acroscopic 
pinnules also show a large degree of variation in length. In some 
plants these pinnules are only slightly longer than the next pinnule, 
but in others the pinnules may extend beyond the pinna-rachis of the 
pinna above. These variations do not appear to be environmentally 
induced, but the variation in frond size and lamina texture is clearly 
influenced by the environment. Plants occurring in exposed habitats 
in the Amatola Mountains have short erect fronds, a coriaceous 
lamina, and sprout from a stout rhizome, whereas plants growing in 
shaded habitats have long and slender arching fronds with a herba- 
ceous lamina, and the rhizome is slender and branches freely. Palea 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 

morphology shows minor variations between plants from the south- 
ern and eastern part of the distribution. In the southern part of the 
distribution the apices of the generally long marginal outgrowths of 
the paleae tend to split leaving a somewhat lacerated appearance, 
whereas in the eastern part of the distribution the outgrowths are 
short and tend not to split at the apices. Indusia vary in size, shape, 
the absence or presence of central processes, and in the occurrence 
of clavate unicellular glands along the margin. When glandular cells 
are present along the indusium margin of a specific collection not all 
indusia will bear them. Glandular cells along the sporangium stalk 
may be present or absent. Since these variations occur randomly 
they are not considered to be environmentally induced. 

DISTRIBUTION AND ECOLOGY. Polystichum incongruum is confined 
to the Western and Eastern Cape Provinces of South Africa. The 
species has a somewhat disjunct distribution, with a southern centre 
ranging from Swellendam to Hofmans Bosch and an eastern region 
centred in the Amatola Mountains. In the Swellendam region the 
species occurs in forests of the 'Ngongoni veld type (Acocks, 1988). 
These forests are small, isolated, and confined to deep sheltered 
ravines and steep slopes below the south-facing cliffs. In the south- 
ern Cape the species is confined to the Knysna forest type, which is 
more extensive. This region receives a well-distributed rainfall that 
ranges between 460-1250 mm per annum. Soil in this region is 
sandy and is largely derived from weathered Table Mountain Sand- 
stone. In the eastern part of its distribution the species occurs at 
elevations ranging between 600 and 1350 m where forests of the 
Dohne Sourveld type are predominant. In this region, however, the 
species is not confined to forested habitats but also occurs above the 
tree-line. The region receives an annual rainfall of 600-1000 mm 
with regular snowfalls during winter and the soil is predominantly of 
doleritic derivation. 

In the southern part of its distribution the species is confined to 
forests where it forms small or large clonal stands and is especially 
common in light shade along streambanks, road cuttings and forest 
clearings. It may occur in dryish or very wet conditions. In the 
eastern part of its distribution the species occurs in more varied 
habitats ranging from natural forests to pine plantations but also 
occurs above the tree line forming large stands along streams, on 
screes and at boulder and cliff bases. Plants in the latter habitats are 
exposed and generally stunted with short erect fronds. In this region 
the plants are frequently subjected to veld fires but this appears not 
to adversely affect them. 

IT.Polystichum pungens (Kaulf.) C. Presl, Tent, pterid.: 83 (1836); 

Schelpe & Anthony, Fl. Sthn. Afr, Pterid. : 254 ( 1 986), pro parte; 

Burrows, Sthn. Afr. ferns and fern allies: 312 (1990), pro parte. 

Type as for Aspidium pungens Kaulf. 
Fig. 17. 

Aspidium pungens Kaulf, Enum.fil: 242 (1824). Type: Habitat in 
Promentorio b. spei, Chamisso s.n. (LE-holotype; BOL!-photo- 
graph). 

Dryopteris pungens (Kaulf.) Kuntze, Rev. gen. pi. 2: 813 (1891). 

Plants terrestrial or epilithic. Rhizome decumbent, sparsely branched, 
to 370 mm long, to 20 mm in diameter, set with roots and closely to 
widely spaced persistent stipe bases, the older parts nude, the apical 
part densely paleated; paleae broadly attached, castaneous to ferru- 
gineous, chartaceous, narrowly lanceolate to narrowly ovate, truncate 
to cordate, the margins subentire to erose, the apex flagelliform, 
generally terminating in a small thin- walled cell, to 17 x 3 mm. 
Fronds 5-6 per plant, suberect to arching, to 1.4 m long: stipe 
proximally castaneous, stramineous distally, adaxially sulcate, to 



75 

685 mm long x 7 mm in diameter, sparsely to densely set with 
conspicuous larger and smaller paleae; larger paleae more frequent 
proximally, widely spaced and smaller distally, proximally 
castaneous, broadly attached, distally ferrugineous, chartaceous, 
narrowly to broadly ovate, often oblique, cordate to cordate-imbri- 
cate, the margins minutely erose to short-fimbriate, the apex 
flagelliform, terminating in a long acicular cell or a small oblong 
thin- walled cell, to 2 1 x 6 mm; smaller paleae short-stalked, ferrugi- 
neous to stramineous, chartaceous to membranous, narrowly 
triangular, narrowly lanceolate to narrowly ovate, cordate to cor- 
date-imbricate, the margins proximally erose or with short and/or 
long, straight or curved outgrowths, the apex entire, flagelliform, 
terminating in a long acicular cell or a small oblong thin-walled cell: 
lamina 2-pinnate to 2-pinnate-pinnatifid, with up to 21 free pinna 
pairs, firmly herbaceous, adaxially dark green, abaxially slightly 
paler, ovate to broadly ovate, to 704 mm long, the pinnae often 
slightly imbricate distally, more widely spaced proximally, the most 
proximal pinna pair slightly reduced, often somewhat deflexed: 
rachis stramineous to greenish, adaxially sulcate, sparsely to densely 
paleated; paleae short-stalked, ferrugineous to stramineous, mem- 
branous, ovate, narrowly ovate to narrowly triangular, sessile or 
short-stalked, cordate to cordate-imbricate, the proximal margins 
erose to sparsely fimbriate or with short and/or long, curved or 
angular, often branched outgrowths that reduce in size and number 
towards the apex, the apex flagelliform, terminating in a long 
acicular cell or a small thin-walled cell: pinnae 1 -pinnate to 1- 
pinnate-pinnatifid, with up to 24 free pinnule pairs, narrowly 
ovate-attenuate to narrowly oblong-attenuate, to 272 x 48 mm; 
pinna-rachis stramineous, adaxially sulcate, moderately to densely 
paleate; paleae ferrugineous to stramineous, membranous, narrowly 
triangular, short-stalked, cordate-imbricate, the proximal margins 
with short and/or long, often branched outgrowths reduced in size 
and number towards the apex, the apex flagelliform, twisted, termin- 
ating in an acicular cell or a small thin-walled cell, to 4.5 mm long, 
each pinna often subtended by one or more large, often bullate, 
broadly ovate, cordate, minutely erose to fimbriate paleae: pinnules 
opposite to alternate, widely spaced to slightly imbricate, the proxi- 
mal acroscopic pinnule the largest, often significantly longer than 
the next, especially towards the middle of the lamina, each sub- 
tended by one or more large, often bullate, broadly ovate paleae, 
similar to but smaller than those on the rachis, inaequilateral, ovate, 
ovate-oblong to ovate-rhomboid or trullate, often somewhat falcate, 
basiscopically cuneate, acroscopically cuneate to truncate and au- 
riculate, the larger pinnules commonly deeply incised near to the 
costa forming a nearly free auricle acroscopically, the margins 
serrate to lobate-serrate, sharp-tipped, rarely aristate, the costa 
proximally adaxially sulcate, most proximal acroscopic pinnule to 
50 mm long, to 19 mm wide, often reaching beyond pinna-rachis 
above; adaxially sparsely set with paleae chiefly along costa, fili- 
form to taeniform, the margins entire or proximally with a few short 
curved or long angular outgrowths, the apex terminating in a long 
acicular cell or a small thin-walled cell, to 6 mm long; abaxially 
sparsely to moderately paleate, the paleae stramineous, membra- 
nous, short-stalked, subulate, narrowly triangular to narrowly ovate, 
cordate to cordate-imbricate, the margins proximally with short and/ 
or long, angular outgrowths, the apex entire, filiform, terminating in 
a long acicular cell or a small thin-walled cell, to 3.7 mm long. 
Venation adaxially obscure, raised abaxially. Sori circular, c. 1 mm 
in diameter, terminal or near terminal on abbreviated vein branches, 
essentially uniseriate, discrete at maturity: sporangium with 10- 
( 1 2.8)- 1 9 indurated annulus cells; stalk eglandular: indusium peltate, 
stramineous, castaneous or black, nitid, amorphous to circular, 
entire to repand, the maximum radius 0.26-(0.5)-0.8 mm in diameter, 



76 



J.P. ROUX 




10 mm 



Fig. 17 Polystichum pungens. A, proximal part of lamina; B, rhizome; C, abaxial surface of fertile pinnule. All drawn from Roux 2367 (NBG). 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



77 



persistent, brown. Spores pale brown, 64 per sporangium, the 
perispore folded to form inflated reticulate ridges, the ridges crested, 
the ridges and areas between sparsely to densely echinulate, vari- 
ously porate, the exospore 30-(49.31)-62 x 28-(38.08)-56 prn. 
Chromosome number 2n=328. 

MATERIAL EXAMINED 

SOUTH AFRICA. 2330 (Tzaneen): Woodbush (CC), Wager s.n. CH7461 
(PRE); De Hoek, Woodbush, Schweickerdt s.n. TM1852C (PRE). 2430 
(Pilgrim's Rest): Pilgrim's Rest, Mount Sheba Nature Reserve, Waterfall 
trail (DC), Roux 2554 (NBG). 2730 (Vryheid): Hlobane, Mto'.a forest (DB), 
Johnstone 295 (NU). 2828 (Bethlehem): Royal Natal National Park, Gudu 
forest, Gudu waterfall (DB), Roux 2512 (NBG); Royal Natal National Park, 
Gudu forest, Schelpe 1454 (NU). 2829 (Harrismith): Farm Whitestones 
(CA), Roux 1901 (NBG, PRE); Bezuidenhouts Pass, farm Whitestones, Roux 
1684 (NBG, PRE); Oliviershoek Pass, forests S. of Seheletwane, Roux25\9 
(NBG); Cathedral Peak area, Indumeni forest, 5100 ft (CC), Schelpe 781 
(NU). 2831 (Nkandla): Eshowe (CD), Laura s.n. CH6421 (PRE). 2929 
(Underberg): Giants Castle Nature Reserve, forest above Hillside camping 
site (AB), Roux 2500, 2502, 2505 (NBG); Lions River District, Lions Bush, 
5000 ft (BD), Moll 832 (PRE). 2930 (Pietermaritzburg): Buccleuch (AD), 
Sim s.n. CH3641 (PRE); Hilton Road (CB), Wager s.n. B-only (NU); Hilton 
District, Cedara Dam, 3200 ft, Churcher s.n. (NU); Zwaartkop, Sim s.n. PRE- 
11026 (PRE); Sim s.n. CH387 (PRE); Sim s.n. TM1239C (PRE). 2931 
(Stanger): prope Mapumulo (AA), Abraham s.n. (B). 3029 (Kokstad): 
Insizwa forest (CC), Strey 10749 (PRE). 3030 (Port Shepstone): Paddock, 
Burntwood (CC), Strey 5994 (PRE). 3128 (Umtata): Tsolo District, Ngadu, 
3200 ft (BC), Von Breitenbach 1330 (PRE). 3129 (Port St Johns): Port St 
Johns (DA), Wager s.n. CH2996 (PRE). 3225 (Somerset East): Boschberg 
(DA), MacOwen s.n. (P); forest kloofs of the Boschberg mountain, sine coll. 
s.n. (P); in sylvis ad ped. Mont. Boschberg, 3000 ft (DA), MacOwen s.n. (B). 
3226 (Fort Beaufort): Katberg forests (BC), Adams 168, 175 (NU); 
Hogsback, Auckland forest (DB), Roux 510 (NBG); Hogsback forest, 4250 
ft, Schirach 280 (NBG); Hogsback, Stirton 6267 (PRE). 3227 (Stutterheim): 
Fort Cunninghame, 3300 ft (AD), Galpin 2446 (PRE); Isidenge forest (CA), 
Roux 1981 (NBG); Keiskamma Hoek, 2000 ft, Ely s.n. (PRE); Cathcart, Fort 
Cunningham forest Reserve (CB), Roux 2428, 2429, 2432 (NBG); Frankfort, 
Sim s.n. (PRE); Pirie, Sim s.n. (PRE); Dohne Hill, Sim s.n. (PRE); Komgha 
(DB), Flanagan s.n. (PRE). 3318 (Cape Town): Nursery Gorge (CD), 
Schelpe s.n. BOL-35933 (BOL); head of Nursery Gorge, Esterhuysen 2585 1 
(BOL); top of Nursery Gorge, Esterhuysen 15355 (BOL); Devils Peak, Dark 
Gorge, Esterhuysen 26564 (BOL, NBG); Table Mountain, Hiddingh Ravine, 
Esterhuysen 25862 (BOL); Kirstenbosch, sine coll. BOL-55808 (BOL); 
Devils Peak, waterfall, Wolley-Dod 915 (BOL); Table Mountain, Skeleton 
Gorge, Schelpe s.n. (BOL); SE slopes of Devils Peak, Pillans 2694 (BOL); 
Skeleton Gorge, Esterhuysen 26674 (BOL); Skeleton Gorge, Roux91 (BOL); 
mountain woods at back of Newlands, sine coll. s.n. (P); Newlands Ravine 
above contour path, Roux 2370, 2371 (NBG); Lubberts Gift, Roux 2372 
(NBG); Kirstenbosch, contour path, Compton 14629 (NBG); Window Gorge, 
Roux 36 (NBG); Dark Gorge, Roux 2367, 2368a, 2369 (NBG); Window 
Gorge, Wasserfall 84, 156 (NBG); Newlands, Paradise, Rawson s.n. (SAM). 
3320 (Montagu): Heidelberg, Grootvadersbosch (DD), Roux 2596 (NBG); 
Grootvadersbosch, Safraandraai, Roux 2597, 2599 (NBG); Grootvadersbosch, 
Stinkhout hiking trail, Roux 2378, 2379, 2380 (NBG); Grootvadersbosch, 
Taylor 1228 (BOL); Grootvadersbosch, near end of road running past red- 
woods, Roux 2600 (NBG). 3322 (Oudtshoorn): George, Wildernes (DC), 
Compton 14305 (NBG); Goudveld Forest Station, Krisjan se Nek picnic site 
(DD), Roux 2626 (NBG). 3325 (Port Elizabeth): Enon (BC), sine coll. B- 
97063 (B). 3326 (Grahamstown): in sylvis prope Grahamstown (BC), 
MacOwan s.n. (P); kloofs near Grahamstown, Holland s.n. (NBG). 3418 
(Simonstown): Diepgat, kloof below SW Triplets (BB), Esterhuysen 27060 
(BOL); ravines of the Helderberg, Parker 4311 (BOL). 3419 (Caledon): 
Riviersonderend, Oubos (BB), Roux 2586 (NBG). 3423 (Knysna): Knysna 
(AA),Marloth 1901 (L). 

SWAZILAND. 2631 (Mbabane): Millers Falls, 4500 ft (AC), Compton 
25967 (NBG); 5 km NW of Mbabane, 1200 m, Kemp 896 (PRE). 

WITHOUT EXACT LOCALITY: loco incerto, sine coll. BOL-55877 
(BOL); Albany District, Cooper 1415 (P); Africa austral, Drege s.n. (P); Cap. 
b. spei, Bojer s.n. (P); Cap. b. spei, Herb Musei Palat. Vindob. 126 (P); in 



umbrosis montium Hottentots Hollandiae, Zeyher s.n. (SAM); Drakensberg, 
Bottomley s.n. CH5018 (PRE); Fort Beaufort District, Myburg s.n. (NBG); 
Bedford District, Van Rensburg s.n. (NBG); Katberg, Young s.n. (PRE); 
Natal, Pondoland and Zululand midlands, Watt & Brandwyk 336 (PRE); 
district of Albany, Cooper 1415 (PRE); loco incerto, Flanagan s.n. (PRE); 
Kaffirland, St. Augustine, Baur 215, B-only (B); Afr. austr., sine coll. s.n. 
(B); Prom. b. spei, Krebs 360 (B), Natalia, Buchanan 85 (B); Natalia, in sylva 
ad fr. Tugela, Gueinzius s.n. (B). 

Controversy as to the correct name for this taxon has existed for a 
long time. Sim (1892) initially labelled this species as Aspidium 
aculeatum var. pungens, but by 1915 he realized that two entities 
could be recognized, a forest dwelling species that he referred to 
Polystichum aculeatum and a montane form that he referred to P. 
pungens. Becherer (1937), however, proposed the name P. lucidum 
(Burm.f.) Becherer (= Asplenium lucidum Burm.f.) for the forest 
growing species, a name that became well entrenched (Schelpe, 
1969; Roux, 1979; Jacobsen, 1983). Following a reinterpretation of 
the types, Anthony & Schelpe (1985) concluded that Asplenium 
lucidum Burm.f. is synonymous with Asplenium adiantum-nigrum 
L. This largely follows the view of C.V. Morton who distributed 
photographs of what he believed to be the type of the species. A 
review of these anomalous typifications has been provided by Roux 
(1994). Since it was concluded that A. lucidum is synonymous with 
A. adiantum-nigrum, a new name was required for the forest species. 
The next available name for the species is P. pungens (Kaulf.) C. 
Presl (= Aspidium pungens Kaulf.). 

DIAGNOSTIC FEATURES AND RELATIONSHIPS. Polystichum pungens 
forms part of a species group characterized by decumbent rhizomes 
and paleae that usually terminate in a long flagelliform apex. It can, 
however, be separated from other taxa in the group by not having 
glandular cells on the sporangium stalk and by the longer, more 
slender rhizome. Furthermore, P. pungens has a somatic chromo- 
some number of 2n=328, versus 2n=164 in P. incongruum with 
which it may be confused and to which it evidently is related. 

VARIATION. Variation in pinnule size and shape may be influenced 
by numerous environmental factors. Smaller pinnules may be ovate 
in outline and shallowly lobate-dentate. As the pinnules increase in 
size, they become more deeply lobate in the proximal part of the 
pinnule, often extending to the costa, resulting in the proximal 
acroscopic segment being short-stalked. The proximal acroscopic 
pinnule is generally longer than the following pinnule. The length 
ratio of the proximal and the following pinnule taken from the 
central part of the lamina ranges between 1:0.91 [Compton 14629 
(NBG)] and 1:0.56 [Roux236Sa (NBG)]. 

Paleae vary mostly in shape and in the degree to which the margin 
is sculptured. Although most paleae terminate in a long filiform 
apical cell, some do terminate in a short, thin-walled cell. In one 
collection [Esterhuysen 26564 (BOL, NBG)] unicellular glandular 
cells also occur along the palea margin as well as on the surface of 
the larger rhizome paleae. 

Indusium size and outline vary considerably within the species. 
Although indusia are generally peltate, often some are reniform or 
have the flange not fully 360 developed. Both conditions are 
frequent within a single plant. Although the general outline of the 
indusium may be considered circular, it is often irregular with the 
margins varying from subentire to repand. Indusia appear to increase 
in size from the western part of the distributional range to the east. 
The plant with the smallest mean indusium radius was recorded 
from Table Mountain [Compton 14629 (NBG), x=0.3 mm; n=6] and 
the plant with the largest mean radial length is from the George 
region [Roux 2626 (NBG), x=0.7 1 mm; n=6]. The maximum radius 
of the indusia varies between 0.26 and 0.78 mm. Plants as far east as 



78 



J.P.ROUX 



Port Elizabeth usually have stramineous indusia, whereas plants 
ranging from the Boschberg farther north have dark, almost black 
indusia, and are uniform in outline. 

DISTRIBUTION AND ECOLOGY. Polystichum pungens is restricted to 
South Africa and Swaziland. In this region the species occurs from 
Table Mountain on the Cape Peninsula to the Hottentots Holland 
Mountains, along the Riviersonderend, Langeberg, Outeniqua and 
Great Winterhoek Mountains to Port Elizabeth and Grahamstown. 
Inland it occurs from the Boschberg at Somerset East to the Amatola 
Mountains and along the Drakensberg foothills to the Wolkberg in 
the Northern Province. 

In the eastern part of its distribution P. pungens is restricted to 
isolated climax forest patches largely restricted to the southern 
mountain aspects and sheltered ravines. This region, and the more 
extensive Knysna forest complex, is subject to a high, well-distrib- 
uted rainfall and acidic sandy soils. At Grahamstown and on the 
Boschberg the species occurs in temperate scrub forest subject to 
more seasonal precipitation. In the Amatolas the species occurs in 
forests of the Dohne Sourveld type that lie between 600-1350 m 
above sea level. From here the distribution extends to the eastern 
slopes and foothills of the Drakensberg. Forests in this region are of 
the Highland Sourveld type, which is largely confined to the deep 
gorges and protected mountain slopes occurring at elevations rang- 
ing from 1350-2150 m. To the north the species occurs in forests of 
the Northeastern Mountain Sourveld. In the Eastern Cape and 
southern KwaZulu-Natal it has been reported from forests of the 
Pondoland Coastal Plateau Sourveld that are found at an elevation of 
300-450 m. These forests are largely confined to the escarpment, 
gorges and valleys below krantzes. In the KwaZulu-Natal midlands 
it is confined to forests of the Mist Belt 'Ngongoni Veld, whereas in 
northern KwaZulu-Natal it occurs in 'Ngongoni Veld. 

Polystichum pungens is a terrestrial or epilithic species occurring 
as isolated individuals or often also as large clones on dryish or 
moist slopes in partially to deeply shaded conditions. In Newlands 
Ravine on Table Mountain, however, the species forms extensive 
stands on exposed east-facing slopes. 



ACKNOWLEDGEMENTS. I would like to thank Braam van Wyk for supervi- 
sion throughout the course of my PhD study. My appreciation also goes to the 
collections managers who kindly made their material available for study, and 
to the anonymous reviewers for their constructive criticism on an earlier draft 
of this paper. Yvonne Reynolds is thanked for obtaining literature not locally 
available. 



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Meyer, E. 1960. Zur Gattung Polystichum in Mitteleuropa. Willdenowia 2: 336-342. 
Mitui, K. 1965. Chromosome studies on Japanese ferns. I. Journal of Japanese Botany 

40: 117-124. 
1968. Chromosomes and speciation in ferns. The Science report of the Tokyo 

Kyoiku Diagaku Section B. 13: 285-333. 



POLYSTICHUM (DRYOPTERIDACEAE) IN AFRICA 



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Moll, E.J., Campbell, B.M., Cowling, R.M., Boss, L., Jarman, M.L. & Boucher, C. 

1 984. A description of major vegetation categories in and adjacent to the Fynbos 

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1977. Fragmenta pteridologiae VII. Webbia 32: 69-93. 

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Raven, P.M. & Axelrod, D.I. 1974. Angiosperm biogeography and past continental 

movements. Annals of the Missouri Botanic Garden 61: 539-673. 
Roux, J.P. 1979. Cape Peninsula ferns. Kirstenbosch. 
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641-642. 

1997a. The morphology and cytology of a new Polystichum (Pteridophyta: 

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\997b. A new species of Polystichum (Pteridophyta: Dryopteridaceae) from 

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19966. A taxonomic study on the fern genus Polystichum Roth sect. 

Metapolystichum Tagawa from China (III). Acta Phytotaxonomica Sinica 34: 194- 
203. 



SYSTEMATIC INDEX 



Accepted names are in roman and synonyms in italics. 



Acropelta Nakai 35 

Acropelta omeiensis (C. Chr.) Nakai 35 

Aetopteron House 35 

Aspidium aculeatum (L.) Sw. 45 

Aspidium aculeatum subsp. angulare (Kit. ex Willd.) 

Asch. 47 

Aspidium angulare Kit. ex Willd. 47 
Aspidium hastulatum Ten. 47 
Aspidium lobatum (Huds.) Sw. 45 
Aspidium luctuosum Kunze 39 
Aspidium macleae Baker 36 
Aspidium pungens Kaulf. 75 
Aspidium tsus-simense Hook. 39 
Aspidium volkensii Hieron. 41 
Cheilanthes speciosissima Kunze 35 
Dryopteris aculeata (L.) Kuntze 45 
Dryopteris aculeata subsp. angularis (Kit. ex Willd.) 

Schinz & Thell. 47 

Dryopteris pungens (Kaulf.) Kuntze 75 
Dryopteris setifera (Forssk.) Woyn. ex Schinz & 

Thell. 47 
Dryopteris setifera subsp. angularis (Kit. ex Willd.) 

Maire 47 

Dryopteris setifera subsp. lobata (Huds.) Maire 45 
Hemesteum H. Lev. 35 
Hypopeltis Michx. 35 
Hypopeltis lobulata Bory 35 
Papuapteris C. Chr. 35 
Papuapteris linearis C. Chr. 35 
Plecosorus Fee 35 
Plecosorus mexicanus Fee 35 
Polypodium aculeatum L. 45 
Polypodium lobatum Huds. 45 
Polypodium lonchitis L. 35 



Polypodium setiferum Forssk. 47 
Polystichum Roth 35 

Polystichum aculeatum (L.) Roth 35, 36, 45, 46 
Polystichum aculeatum subsp. angulare (Kit. ex 

Willd.) Vollm. 47 

Polystichum aculeatum van mildbraedii Brause 53 
Polystichum aculeatum var. rubescens Bonap. 53 
Polystichum aculeatum var. stenophyllon Bonap. 53 
Polystichum alticola Schelpe & N.C. Anthony 53 
Polystichum angulare (Kit. ex Willd.) C. Presl 47 
Polystichum barbatum C. Chr. 41 
Polystichum x bicknellii (H. Christ) Hahne 47 
Polystichum coursii Tardieu 45 
Polystichum cystostegia (Hook.) J.B. Armstr. 60 
Polystichum discretum (D. Don) J. Sm. 56 
Polystichum dracomontanum Schelpe & N.C. 

Anthony 36,70, 71 
Polystichum elegans J. Remy 60 
Polystichum falcinellum (Sw.) C. Presl 37 
Polystichum fuscopaleaceum Alston 53 
Polystichum fuscopaleaceum var. ruwensoriense 

(Pirotta) Pic.Serm. 53 
Polystichum glaciale H. Christ 35 
Polystichum incongruum J.P. Roux 36, 72, 73 
Polystichum kalambatitrense Tardieu 37 
Polystichum kilimanjaricum Pic.Serm. 36, 42, 44 
Polystichum lemmonii Underw. 60 
Polystichum lineare (C. Chr.) Copel. 35 
Polystichum lobatum (Huds.) Bastard 45 
Polystichum lobatum var. luctuosum (Kunze) H. 

Christ 39 

Polystichum lobatum var. ruwensoriense Pirotta 53 
Polystichum lonchitis (L.) Roth 35 
Polystichum luctuosum (Kunze) T. Moore 36, 37, 40, 42 



Polystichum macleae (Baker) Diels 36, 38 
Polystichum magnificum F. Ballard 36, 63, 64 
Polystichum marionense Alston & Schelpe 36, 58, 59 
Polystichum mohrioides (Bory) C. Presl 60 
Polystichum monticola N.C. Anthony & Schelpe 36, 

67,68 

Polystichum munitum (Kaulf.) C. Presl 37 
Polystichum nigropaleaceum (H. Christ) Diels 56 
Polystichum omeiense C. Chr. 35 
Polystichum pauciaculeatum Bonap. 45 
Polystichum plicatum (Poepp.) Hicken 60 
Polystichum pungens (Kaulf.) C. Presl 36, 75, 76 
Polystichum x saltum J.P. Roux 36, 56, 57 
Polystichum scopulinum (R.J. Eaton) Maxon 60 
Polystichum setiferum (Forssk.) T. Moore ex 

Woyn. 36,47,48 
Polystichum setiferum var. fuscopaleaceum (Alston) 

Schelpe 53 
Polystichum setiferum var. nigropaleaceum (H. Christ) 

Sledge 56 
Polystichum speciosissimum (Kunze) R.M. Tryon & 

A.F. Tryon 35 

Polystichum transkeiense W. Jacobsen 36, 60, 61 
Polystichum transvaalense N.C. Anthony 36, 49, 50 
Polystichum tsaratananense Tardieu 45 
Polystichum tsus-simense (Hook.) J. Sm. 39 
Polystichum volkensii (Hieron.) C. Chr. 36, 41, 43 
Polystichum wilsonii H. Christ 36, 53, 54 
Polystichum zambesiacum Schelpe 36, 63, 65 
section Lasiopolystichum Daigobo 45, 52, 56, 63, 69 
section Metapolystichum Daigobo 47, 69 
section Xiphopolystichum Daigobo 41 
Sorolepidium H. Christ 35 
Sorolepidium glaciale (H. Christ) H. Christ 35 



Bull. nat. Hist. Mus. Lond. (Bot.) 30(2): 81-99 Issued 30 November 2000 

Recent records of pteridophytes for Belize, 
Central America xy O s 

DAVID A. SUTTON 

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD 

ANGELA HUGHES 

International Centre for Underutilized Crops, Institute of Irrigation and Development Studies, University of 
Southampton, Southampton SO 17 1BJ 

BARBARA BULMER-THOMAS 

55 Maze Hill, Greenwich, London, SEW 8XQ 

CONTENTS 

Introduction 82 

Listoftaxa 83 

Aspleniaceae 83 

Blechnaceae 84 

Cyatheaceae 84 

Davalliaceae 84 

Dennstaedtiaceae 85 

Dicksoniaceae 85 

Dryopteridaceae 85 

Gleicheniaceae 86 

Grammitidaceae 86 

Hymenophyllaceae 86 

Isoetaceae 87 

Lomariopsidaceae 87 

Lophosoriaceae 88 

Lycopodiaceae 88 

Marattiaceae 88 

Metaxyaceae 89 

Ophioglossaceae 89 

Polypodiaceae 89 

Psilotaceae 90 

Pteridaceae 90 

Salviniaceae 91 

Schizaeaceae 91 

Selaginellaceae 92 

Tectariaceae 92 

Thelypteridaceae 93 

Vittariaceae 95 

Woodsiaceae 95 

Discussion 96 

References ... 98 



SYNOPSIS. Belize has one of the richest pteridophyte floras in the world, expressed on an area basis. Analysis of recent 
collections and the widely scattered literature for the country reveals that the number of species has risen by approximately 20% 
since the publication of the major regional flora, Flora Mesoamericana, in 1995. Investigation of the dates of discovery for each 
taxon produces a graph that indicates many more taxa are yet to be discovered in the country. A preliminary assessment is made 
of areas in Belize likely to reveal new records. 



The Natural History Museum, 2000 



82 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



INTRODUCTION 



The vascular plants of Belize have never been treated adequately in 
a national flora. Early botanical investigation during the nineteenth 
century was focussed very much on economically important species, 
especially timber, latex and fruit, though the list produced for the 
now defunct Botanic Station by Campbell ( 1 899) included a number 
of native ferns and orchids. Other early accounts of the flora (Blake, 
1917; Record, 1925), based on exploration and forestry, excluded 
pteridophytes. Standley & Record (1936) provided a good floristic 
grounding for the country with the Forests and flora of British 
Honduras, though this lacks some of the elements of a regular flora 



and the content is biased strongly towards trees both in species 
coverage and detail. Herbaceous angiosperms and pteridophytes are 
merely listed, except where they are believed to be new species. 
However, this single publication provided the earliest published 
record for almost 25% of species in the entire flora, far exceeding the 
significance of any other publication for Belize in this respect. A 
number of checklists have appeared subsequently for the country. 
Those by Dwyer and others (Spellman et al., 1975; Dwyer & 
Spellman, 1981) excluded pteridophytes, but some records appear 
in the catalogue by Schipp (1934). The catalogue of the Belize 
National Herbarium (BRH) by Vargas & Shawe (1997) includes a 
number of plant records for the country not published elsewhere. 




Fig. 1 Districts of Belize. Shaded areas represent 300 m and 600 m contours. 



RECENT RECORDS OF PTERIDOPHYTES 



83 



Recent expedition reports and surveys (for example Parker et al., 
1993) include many of the new records of pteridophytes for the 
country. 

The regional Flora of Guatemala expressly included Belize (as 
British Honduras) in its remit (for example, Standley & Steyermark 
1958) though for the majority of pteridophyte taxa listed (Mickel, 
1981; Smith, 19816; Stolze, 1976, 1981, 1983; Stolze & Hickey, 
1983; 011gaard, 1983) there is no explicit mention of the presence in 
or absence from Belize. For most vascular plants, this publication 
still provides the most complete coverage of the flora for Belize to 
date, though for pteridophyes it has been superseded by the regional 
Flora Mesoamericana (Davidse et al., 1994, 1995). This latter 
project will provide a comprehensive coverage of the species of 
Belize on its eventual completion and it has a unique strength in the 
development of a live link to the TROPICOS (Solomon et al., 2000) 
database in 1998, so that the Internet version of the flora is updated 
on a continual basis. 

Since the publication of the volume of Flora Mesoamericana, 
there has been considerable interest in the pteridophytes of Belize 
arising out of a number of projects (for example, Hughes, 1998) and 
recent fieldwork. A list of current names used for Belize, with an 
indication of their status is given below. The report includes an 
analysis of over 2500 pteridophyte specimen records from The 
Natural History Museum in London (BM) and other herbaria at 
AAU, B, BR, BRH, CR, DS, F, G, GH, K, LAGU, MEXU, MICH, 
MO, NY, S, SEL, TEX, UAMIZ, UC, UCWI, US and Z. Full use has 
been made of information sources on the Internet, and the authors 
particularly express their gratitude to Missouri Botanical Garden for 
making TROPICOS available on the World Wide Web. 



LIST OF TAXA 

The following list is arranged alphabetically by family following the 
family delimitation used in Flora Mesoamericana (Davidse et al., 
1 995). Taxa previously recorded in Flora Mesoamericana are shown 
in italic and the superscript reference indicates the account and page 
number. Representative material is shown with one voucher for each 
district (Fig. 1) where the species has been recorded, except where 
the taxon has particular significance as a new record or endemic. 
Preference in citation is given to specimens cited in a monograph, 
revision or a recent regional flora. A superscript reference number or 
numbers following a specimen is to the source included in the 
bibliography where that specimen is listed. For new records not 
included in Flora Mesoamericana, the name is shown in bold and 
additional specimens are listed where known. An indication is given 
of where the taxon was first listed for Belize. 

ASPLENIACEAE 

Asplenium abscissum Willd., Sp. pi. 5: 321 (1810) 2:295 

Toledo, G. Davidse & D.L. Holland 36836 (MO 115 ). 
Asplenium auritum Sw. in J. Bot. (Schrader) 1800(2): 52 (1801) 2:2 - 

Cayo, A. Hughes 64 (BM-000557717! 22 ); Toledo, T.B. Croat 

24260 (MO 2 ). 
Asplenium cirrhatum Rich, ex Willd., Sp. pi. 5: 321 (1810) 2:2< 

Toledo, B.H. Allen 15443 (MO 2 ' 3 ' 115 ). 
Asplenium cristatum Lam., Encycl 2(1): 310 (1786) 2:3( 

Cayo, A. Hughes 98 (BM-000557710! 22 ); Stann Creek, W.A. 

Schipp 50 (BM 2 , BRH 125 , NY); Toledo, B.K. Hoist 4397 (MO 91 ). 
Asplenium delitescens (Maxon) L.D. Gomez in Brenesia 8: 52 

(1976^301 

Cayo, A. Hughes 12 (BM-000531878!, BM-000557708! 22 , 



BRH!); Stann Creek, P.H. Gentle 2714 (F, GH, K 2 , MICH, NY); 
Toledo, P.H. Gentle 6261 (F, G, NY, S, UC). 

Asplenium dentatum L., Sp. pi. 2: 1080-1081 (1753) 

Stann Creek, P.H. Gentle 2712 (NY); Toledo, T.E. Hawkins 1631 
(MO 115 ). 

A new name for Asplenium trichomanes-dentatum L., listed for 
Belize in Flora Mesoamericana (Adams, 19956: 322) based on 
extralimital distribution in Proctor (1985: 375) but without refer- 
ence to a specimen. The name is automatically corrected according 
to the International Code of Botanical Nomenclature (Tokyo 
Code 23.8, ex. 14, Greuter et al., 1998). Vargas & Shawe (1997: 
22) listed a specimen (W.A. Schipp 1066, BRH) under the name 
A. dentatum L., but this material has also been identified as A. 
macilentum by Adams (19956: 309) based on a duplicate at K. 

Asplenium formosum Willd., Sp. pi. 5: 329 (1810) 2:304 

Cayo, G.R. Proctor 29902 (BM 2 ); Stann Creek, R. Rivera et al. 
2576 (BRH 125 ); Toledo, G. Davidse 35782 (MO 115 ). 

Asplenium heterochroum Kunze in Linnaea 9: 67 ( 1 834) 2: 306 
Cayo, A. Hughes 148 (BM-000557731! 22 ); Stann Creek, W.A. 
Schipp 211 (BM 2 , BRH 125 ); Toledo, G. Davidse & A.E. Brant 
32131 (MO 115 ). 

Asplenium juglandifolium Lam., Encycl. 2(1): 307 (1786) 2 3( 
Toledo, B.K. Hoist 4027 (MO 91 ' 115 ). 

Asplenium laetum Sw., Syn.fil: 79, 271 (1806) 2:308 

Stann Creek, WA. Schipp S-277 (B 115 ); Toledo, P.H. Gentle 7378 
(BM 2 ' 115 , F 115 , G 115 , MICH 115 , NY 115 , S" 5 , UC 115 ). 

Asplenium macilentum Kunze ex Klotzsch in Linnaea 20: 351 

(1 8 47)2: 309 

Toledo, W.A. Schipp 1066 (BRH 125 , K 2 , NY). 
Adams (1995: 309-310) considered that material cited as A. 
monodon Liebm. was more closely related to A. macilentum than 
A. auritum, although it was included as a synonym in the latter 
taxon by Stolze ( 198 1 : 60). It is not clear if A. monodon occurs in 
Central America and the name may have been misapplied. 
Asplenium palmeri Maxon in Contr. U.S. Natl. Herb. 13: 39 (1909) 2 

313 

First recorded for Belize in extralimital distribution for the 
Pteridophyte flora of Oaxaca, Mexico (Mickel & Beitel, 1988: 
65) but without explicit reference to specimens. Vargas & Shawe 
(1997: 22) list a specimen collected in 1929 from the Stann Creek 
District ( W.A. Schipp 211, BRH) under this name, but a duplicate 
at BM has been identified as A. heterochroum by Adams (19956: 
306). Material needs to be located to support the record for 
Belize. 

Asplenium pteropus Kaulf., Enum. filic. : 1 70 ( 1 824) 2: 3I4 

Stann Creek, WA. Schipp 365 (MO 2 , NY); Toledo, B.K. Hoist et 
al. 5 195 (MO 115 ). 

Asplenium pumilum Sw., Prodr.: 129 (1788) 2;315 

Cayo, A. Hughes 136 (BM!); Toledo, T. Arnason & J. Lambert 
17182 (MO 2 ), G. Davidse & B.K. Hoist 36181 (MO 115 ). 

Asplenium riparium Liebm. in Kongel. Danske Vidensk. Selsk. Skr., 
Naturvidensk. Math. Afd. ser. 5, 1: 244 (1849) 
Toledo, G. Davidse & D.L Holland 36837 (MO 115 ), B.K. Hoist 
5332 (MO 115 ). 

Not previously recorded from Belize in the Central American 
literature, but material collected from Toledo District in 1 996 and 
1997 is listed in TROPICOS and identified as this taxon by A.R. 
Smith in 1997. 

Asplenium salicifolium L., Sp. pi- 2: 1080 (1753) 2:318 

Asplenium salicifolium var. salicifolium 1 3 
Cayo, T.B. Croat 23773 (MO 2 ). 

Asplenium serra Langsd. & Fisch., PL Voy. Russes monde 1: 16, 1. 19 
(1810) 2:319 



84 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



Cayo, B.H. Allen 15177 (BM-000543291!, MO 2 - 3 ' 115 ). 

Asplenium serratum L., Sp. pi. 2: 1079 (1753) 2:319 

Cayo, C.L. Lundell 6235 (MICH 28 , NY); Stann Creek, R. Rivera 
et al. 2602 (BRH 125 ); Toledo, C. Whitefoord 1625 (BM- 
000543328 ! 2 ). 

Asplenium uniseriale Raddi in Opusc. Sci. 3: 291 (1819) 
Toledo, B.K. Hoist 5321 (MO 115 , UC" 5 ). 
Not listed for Belize in the published version of Flora 
Mesoamericana (Adams, 19956: 323), but included in the Internet 
version based on a specimen collected in 1996 from Toledo 
District. 

BLECHNACEAE 

HI cell n 11 in x ant Ulan u in Proctor in Brit. Fern Gaz. 9: 214 (1965) 
Cayo, G.R. Proctor 29904 (BRH 125 ). 

Interpreted in Flora Mesoamericana (Moran, I995ae: 329) as a 
hybrid between B. meridense and B. glandulosum and expected 
from Mesoamerica. Vargas & Shawe (1997: 31) listed an unpub- 
lished herbarium name with the epithet 'antillense' attributed to 
G.R. Proctor and based on a specimen (cited above, collected in 
1968 from Rio Frio Caves in Cayo District) which apparently 
equates to Proctor's Blechnum x antillanum. However, of the 
putative parent species, only B. meridense has been recorded 
from Belize and the status of this specimen is uncertain. 

Blechnum x caudatum Cav., Descr. pi: 262 (1802) 67:326 
Cayo, A. Hughes 34 (BM-000531849!). 

Blechnum ensiforme (Liebm.) C. Chr., Index filic.: 153 (1905) 
Toledo, G. Davidse & D.L. Holland 36781 (MO 115 , UC" 5 ). 
Not recorded from Belize in the published version of Flora 
Mesoamericana (Moran, \995ae: 327), but included in the 
Internet version based on a specimen (G. Davidse & D.L. Hol- 
land 36781, UC) collected from Toledo District in 1997 and 
identified by A.R. Smith in the same year. 

Blechnum fragile (Liebm.) C.V. Morton & Lellinger in Amer. Fern 
J. 57:68(1967) 67:328 

Cayo, B.H. Allen 15204 (MO 3 - 67 - " 5 ); Toledo, B.K. Hoist et al. 
5303 (MO 115 ). 

Blechnum gracile Kaulf., Enum. filic.: 158 (1824) 67:328 
Toledo, G. Davidse & A.E. Brant 31915 (MO 67 115 ). 

Blechnum meridense Klotzsch in Linnaea 20: 349 (1847) 

Toledo, G. Davidse 36979 (MO 115 , UC), D.L. Holland65 (MO 115 ), 
D.L. Holland & B. Kid 100 (MO 115 ). 

Not recorded from Belize in the published version of Flora 
Mesoamericana (Moran, \995ae: 329), but included in the 
Internet version based on a specimen (G. Davidse 36979, UC) 
collected from Toledo District in 1997 and identified by A.R. 
Smith in the same year. 

Blechnum occidentale L., Sp. pi. 2: 1077 (1753) 67:329 

Cayo, A. Hughes93 (BM-000531830!, BM-000531837!); Stann 
Creek, W.A. Schipp 162 (BRH 125 , NY); Toledo, G. Davidse & 
A.E. Brant 32304 (MO 67 - 115 ). 

Blechnum polypodioides Raddi in Opusc. Sci. 3: 294 (1819) 67:329 
Cayo, H.H. Bartlett 11643 (US). 

Moran (\995af: 329) did not base his record for this taxon on a 
specimen, but on extralimital distribution in Flora of Chiapas 
(Smith, 198 la: 60). The Bartlett specimen cited above is at US, 
and a duplicate is to be expected at MICH. 

Blechnum serrulatum Rich, in Actes Soc. Hist. Nat. Paris 1: 114 

(1 792)67: 330 

Belize, G. Davidse & A.E. Brant 32999 (MO 67 - " 5 ); Orange Walk, 
G.R. Proctor 35794 (BRH 125 ); Stann Creek, P.H. Gentle 8227 
(NY); Toledo, G. Davidse & A.E. Brant 32496 (MO 115 ). 



Salpichlaena volubilis (Kaulf.) J. Sm. in Hook. & Bauer, Gen.fil.: t. 
93(1841) 68:332 
Toledo, B.K. Hoist 4287 (MO 6 "-"- 115 ). 



CYATHEACEAE 

Alsophila firma (Baker) D.S. Conant in J. Arnold Arbor. 64: 372 

(1983) IO.:89 

Cayo, B.H. Allen 15221 (MO 3 - l01 - 115 ); Stann Creek, A.M. Gentry 

7958 (MO 115 ); Toledo, B.K. Hoist et al. 5301 (MO 115 ). 
Alsophila salvinii Hook, in Hook. & Baker, Syn.fil.: 36 (1866) 

Toledo, B.K. Hoist 5854 (MO 115 ). 

This Central American endemic was not recorded for Belize in 

Flora Mesoamericana (Riba, 1995c: 90), though it has been 

recorded from all surrounding countries and is included here 

based on a specimen collected in 1997 from Toledo District. 

Further material is to be expected from the wetter parts of the 

Maya Mountain divide and the Liquidambar forests. 
Cyathea costaricensis (Mett. ex Kuhn) Domin in Acta Bot. Bohem. 

9: 107(1930) 40:96 

Stann Creek, J.D. Dwyer et al. 583 (MO 40 - 115 ). 
Cyathea divergens Kunze in Linnaea 9: 100 (1834) 40197 
Cyathea divergens var. tuerckheimii (Maxon) R.M. Tryon in Contr. 

Gray Herb. 206: 56 (1976) 40:97 

Cayo, B.H. Allen 15224 (MO 3 - 40 - 115 ); Toledo, B.K. Hoist et al. 

5245 (MO 115 ). 
Cyathea microdonta (Desv.) Domin in Pteridophyta: 263 (1929) 40:98 

Stann Creek, J.D. Dwyer et al. 549 (MO 40 - " 5 ); Toledo, P.H. 

Gentle 1933 (NY). 
Cyathea multiflora Sm. in Mem. Acad. Roy. Sci. (Turin) 5(1790- 

1791): 416 (1793) 40:99 

Cayo, J. Meave 1037 (MO 40 - 115 ); Toledo, M.J. Balick et al. 2543 

(BRH 125 , MO 115 , NY). 
Cyathea myosuroides (Liebm.) Domin in Pteridophyta: 263 (1929) 40199 

Cayo,A.//wg/u>sl32a(BM-000531752!,BM-000531908!,BM- 

000531909!); Stann Creek, W.A. Schipp 191 (BRH 125 , NY); 

Toledo, B.K. Hoist 4285 (MO 91 - 115 ). 
Cyathea schiedeana (C. Presl) Domin in Pteridophyta: 263 (1929) 401 101 

Cayo, J.D. Dwyer 11431 (MO 40 - 115 ); Stann Creek, P.H. Gentle 

8257 (NY); Toledo, B.H. Allen 15430 (MO 3 - 115 ). 
Cyathea ursina (Maxon) Lellinger in Amer. FernJ. 77: 101 (1987) 40: 102 

Stann Creek, P.H. Gentle 3 197 (LL 40 , MICH 23 , MO-photograph 40 - 

" 5 , US-holotype 23 - 30 - 120 - 124 ). 

Reported only from the type collected from Antelope Ridge in 

the Stann Creek Valley of Belize, this Central American endemic 

occurs also in Guatemala and from Nicaragua to Panama. 
Sphaeropteris horrida (Liebm.) R.M. Tryon in Contr. Gray Herb. 

200:20(1970) 89:104 

Toledo, B.K. Hoist 4213 (MO 89 - 91 - 115 ). 

DAVALLIACEAE 

Nephrolepis biserrata (Sw.) Schott, Gen.fil.: t. 3 (1834) 80:286 

Cayo, A. Hughes 130 (BM-000531755! 22 , BM-000531910!); 

Stann Creek, WA. Schipp 394 (BRH 125 , NY 80 ); Toledo, G. Davidse 

& A.E. Brant 32193 (MO 115 ). 
Nephrolepis cordifolia (L.) C. Presl, Tent, pterid.: 79 (1836) 80:287 

Cayo, G.R. Praetor 29881 (BRH 125 ); Toledo, D.E. Breedlove & 

D.C. McClintock 23670 (DS 80 ). 
Nephrolepis multiflora (Roxb.) P.M. Jarrett ex C.V. Morton in 

Contr. U.S. Natl. Herb. 38: 309 (1974) 80:287 

Cayo, A. Hughes 67 (BM-000531799!, BM-000557718! 22 ); Stann 



RECENT RECORDS OF PTERIDOPHYTES 

Creek, R. Rivera et al. 2536 (BRH 125 ); Toledo, B.K. Hoist 4507 

(MO 80 ' 91 -" 5 ). 
Nephrolepis pendula (Raddi) J. Sm. in J. Bot. (Hooker) 4: 197 

(1841) 80:288 

Cayo, C.L. Lundell 6305 (MICH 28 , NY 80 ); Toledo, M.E. Peck 538 

(NY). 
Nephrolepis rivularis (Vahl) Mett. ex Krug in Bot. Jahrb. Syst. 24: 

122(1897) 

Toledo, G. Davidse 36876 (MO 115 , UC), T.E. Hawkins 1425 

(MO 115 ). 

Not recorded for Belize in the published version of Flora 

Mesoamericana (Nauman, 1995: 288), although included in the 

Internet version on the basis of material collected in 1997 from 

Toledo District and identified by A.R Smith in 1997 and 1998. 
Nephrolepis undulata (Afzel. ex Sw.) J. Sm. in Bot. Mag. 72(Com- 

panion): 35 bis (1846) 

Cayo, A. Hughes 104 (BM-000532008!). 

Not recorded for Belize in Flora Mesoamericana (Nauman, 

1995: 288), although recorded from all surrounding countries 

and to be expected in Belize. The material listed here from Cayo 

District is not fully fertile and confirmation of the identification 

is required from further collections. 
Oleandra articulata (Sw.) C. Presl, Tent, pterid.: 78, t. 2, f. 12 

(1836) 90:289 

Toledo, T.E. Hawkins 1542 (MO 115 ). 

DENNSTAEDTIACEAE 

Dennstaedtia bipinnata (Cav.) Maxon in Proc. Biol. Soc. Wash. 51: 

39(1938) 47:152 

Toledo, B.K. Hoist 4474 (MO 47 ' 91 ' 115 ). 
Dennstaedtia cicutaria (Sw.) T. Moore, Index fil: xcvii (1857) 47: 152 

Toledo, W.A. Schipp S-802 (GH 47 - 115 ). 
Dennstaedtia dissecta (Sw.) T. Moore, Index fil.: 305 (1861) 47: 152 

Stann Creek, R. Rivero et al. 2587A (BRH 125 ); Toledo, W.A. 

Schipp S-921 (GH 47 ' 115 ). 
Hypolepis repens (L.) C. Presl, Tent, pterid.: 162 (1836) 48: 156 

Stann Creek, R. Rivero et al. 2532 (BRH 125 ); Toledo, W.A. Schipp 

258 (GH 48 ' 115 , NY). 
Lindsaea klotzschiana Moritz in Ettingsh., Farnkr. Jetztw.: 212, t. 

145, f. 1-2(1865) 49:158 

Toledo, B.K. Hoist 4265 (MO 49 ' 91 ' 115 ). 

Lindsaea lancea (L.) Bedd., Suppl. ferns S. Ind.: 6 (1876) 49: 158 
Lindsaea lancea var. lancea 49 159 

Stann Creek, W.A. Schipp 100 (BRH 125 , NY); Toledo, C. 

Whitefoord 1310 (BM, MO 49 - 115 ). 

Lindsaea quadrangularis Raddi in Opusc. Sci. 3: 294 (1819) 49: 159 
Lindsaea quadrangularis subsp. subalata K.U. Kramer in Acta Bot. 

Neerl.6: 190 (1957) 49: 159 

Stann Creek, W.A. Schipp 200 (BRH 125 , F 49 - " 5 , US); Toledo, G. 

Davidse 36247 (MO 115 ). 
Lindsaea stricta (Sw.) Dryand. in Trans. Linn. Soc. London 3: 42 

(1 797)49: 159 

Lindsaea stricta var. stricta 49 159 

Belize, P.H. Gentle 1536 (K, NY); Cayo, G. Davidse & A.E. 

Brant 33024 (MO 49 ' 115 ); Toledo, G. Davidse & A.E. Brant 31958 

(MO 115 ). 
Lonchitis hirsuta L., Sp. pi. 2: 1078 (1753) 

Toledo, G. Davidse & D.L. Holland 36478 (MO 115 ), G. Davidse 

36888 (MO 115 ), G. Davidse 36983 (MO 115 , UC), B.K. Hoist et al. 

55 16 (MO 115 ). 

Not listed for Belize in the published version of Flora 

Mesoamericana Moran ( 1 995n: 1 60), but included in the Internet 



85 

version on the basis of a collection from Toledo District (G. 

Davidse 36983, UC) in 1997 and identified by A.R. Smith in 

December 1997. This is the first record of the genus for Belize. 
Odontosoria schlechtendalii (C. Presl) C. Chr., Index filic. : 209 

(1905) 51:161 

Toledo, G. Davidse & A.E. Brant 31986 (MO 51 - " 5 , US). 
Pteridium caudatum (L.) Maxon, in Proc. U.S. Natl. Mus. 23( 1226): 

631 (1901) 52:162 

Belize, C. Whitefoord 2452 (BM); Cayo, A. Hughes 65 (BM- 

000531794!, BM-000557716!); Stann Creek, W.A. Schipp 190 

(BRH 125 , NY); Toledo, C. Whitefoord 2204 (MO 52 115 ). 
Saccoloma elegans Kaulf. in Berlin. Jahrb. Pharm. Verbundenen 

Wiss. 21:51 (1827) 53:163 
Saccoloma elegans subsp. chartaceum G.B. Nair ex Cremers & 

K.U. Kramer, Bot. Helvet. 99: 47 (1989) 53: 163 

Cayo, T.B. Croat 24577 (MO 53 - 115 ); Stann Creek, W.A. Schipp 90 

(BRH 125 , NY); Toledo, P.H. Gentle 9315 (NY). 
Saccoloma inaequale (Kunze) Mett. in Ann. Sci. Nat., Bot. ser. 4, 15: 

80(1861) 53:163 

Toledo, W.A. Schipp 315 (MO 53 " 5 , US). 

DICKSONIACEAE 

Cibotium regale Verschaff. & Lem. in ///. Hort. 15: t. 548 (1868) 
Cayo, A.K. Monro & S. Cafferty 2639 (BM-000543340!, BM- 
000543341!, BM-000543342!, BM-000543343!, BRH, LAGU, 
MEXU, MO). 

Known formerly from Chiapas (including the type), Guatemala 
and Honduras to El Salvador (Perez-Garcia, I995b: 86-87), this 
Central American endemic is restricted to mixed woodland of 
Pinus, Quercus and Liquidambar. The vegetation class occurs in 
Belize in the southern part of the Chiquibul Forest and probably 
will be found in other little-explored parts of the Maya Mountain 
Divide. Though the presence of Liquidambar styraciflua L. in 
Belize has been reported since the first half of the twentieth 
century (Stevenson, 1928) and Standley & Record (1936: 147) 
considered that it occurred 'in some abundance in the higher 
parts of the Cockscomb Mountains', material was not collected 
until very recently. A specimen can be found in the Belize 
National Herbarium (Belize: Cayo; J.C. Meerman s.n. (BRH), 
cf. Vargas & Shawe, 1997: 67). General collections from this 
class of vegetation were not made until 1998 by Monro and 
Cafferty, who provide the first record of Cibotium regale. 

DRYOPTERIDACEAE 

Arachniodes denticulata (Sw.) Ching in Acta Bot. Sin. 10: 260 

(1962) 59:2M 

Toledo, B.H. Allen 15391 (AAU 59 - 115 , MO 3 ' 115 ). 
Didymochlaena truncatula (Sw.) J. Sm. in J. Bot. (Hooker) 4: 196 

(1841) 60:212 

Stann Creek, W.A. Schipp 406 (BRH 125 , MO 60 115 ); Toledo, B.K. 

Hoist et al. 5160 (BRH 115 , MO 115 , SEL). 
Olfersia cervina (L.) Kunze in Flora 7: 312 (1824) 61:214 

Stann Creek, W.A. Schipp 402 (BRH 125 , F 36 - 61 - 115 , GH 36 , MICH 36 , 

NY 36 , UC 36 , US 36 , Z 36 ); Toledo, G. Davidse & D.L. Holland 

36737 (MO 115 ). 
Polybotrya caudata Kunze in Linnaea 9: 23 (1834) 62 217 

Cayo, C.L. Lundell 6416 (MICH 28 ); Stann Creek, W.A. Schipp 

273 (BRH 125 , F 62 - 115 ). 
Polybotrya osmundacea Humb. & Bonpl. ex Willd., Sp. pi 5: 99 

(1810) 

Toledo, B.K. Hoist 4048 (MO 91 - 115 ), B.K. Hoist 4049 (MO 91 - " 5 ). 

Not recorded for Belize in Flora Mesoamericana (Moran, 1 995z: 



86 

218), although the species was recorded for Toledo District by 
Parker et al. (1993) and material bearing this identification is at 
MO and listed in TROPICOS. 

Polybotrya polybotryoides (Baker) H. Christ in Bull. Herb. Boiss. 
ser. 2, 1:70(1901) 62:218 

Cayo, T.B. Croat 24566 (MO 62 ' ll5 ); Toledo, B.K. Hoist 4047 
(MO 91 '" 5 ). 

Stigmatopteris sordida (Maxon) C. Chr., Index filic., Suppl. 3: 175 
(1934) 

Toledo, B.K. Hoist 5897 (MO 115 , UC). 

Not recorded for Belize in the published version of Flora Meso- 
americana (Moran, \995aa: 226), but included in the Internet 
version based on a specimen from Toldeo District (B.K. Hoist 
5897, UC) collected in 1997 and identified by A.R. Smith in 
December of the same year. This is also the first record of the 
genus from Belize. 

GLEICHENIACEAE 

Dicranopteris flexuosa (Schrad.) Underw. in Bull. Torrey Bot. Club 

34:254(1907) 38:58 

Belize, P.H. Gentle 9558 (MO 38 115 , US). 
Dicranopteris pectinata (Willd.) Underw. in Bull. Torrey Bot. Club 

34:260(1907) 38:58 

Cayo, C.L. Lundell 6603 (MICH 28 , NY, US); Stann Creek, W.A. 

Schipp 350 (BRH 125 , K!, NY, US); Toledo, D.L. Spellman &W.W. 

Newey 1648 (MO 38 ' 115 ). 
Sticherus palmatus (J.H. Schaffn. ex Underw.) Copel., Gen. fil. 

(Ann. Cryptog. Phytopathol. 5): 28 (1947) 39:61 

Toledo, B.K. Hoist 4270 (MO 39 ' 91 - 115 ). 

GRAMMITIDACEAE 

Cochlidium linearifolium (Desv.) Maxon ex C. Chr. in Dansk 

Botanisk Arkiv 6(3): 23 (1929) 6:371 

Cayo, G. Davidse & A.E. Brant 33087 (MO 115 ); Toledo, B.H. 

Allen 1 5304 (MO 3 ' 6 - 115 ). 
Cochlidium serrulatum (Sw.) L.E. Bishop in Amer. Fern J. 68: 80 



Cayo, G. Davidse & A.E. Brant 33088 (MO 115 ); Toledo, W.A. 

Schipp 213 (N\, UC 6 ). 
Enterosora ecostata (Sodiro) L.E. Bishop in Syst. Bot. 17(3): 348, f. 

1A-C(1992) 1I2:373 

Toledo, W.A. Schipp S-801[a] (F 117 , GH 129 ). 
Lellingeria mitchellae (Baker ex Hemsl.) A.R. Sm. & R.C. Moran in 

Amer. Fern J. 81: 85 (1991) 77:378 

Cayo, A. Hughes 48 (BM-000531771 !); Orange Walk, Mitchell 

s.n. (K-holotype 10 - 19 ' 77 ' 106 ); Toledo, G. Davidse 36126 (MO 115 ). 

Based on a Belizean type, this Central American endemic occurs 

from Chiapas to Panama and is unique in having setulose 

sporangia capsules. 
Micropolypodium taenifolium (Jenman) A.R. Sm. in Novon 2: 423 

(1992) 

Toledo, G. Davidse & D.L. Holland 36701 (MO 115 ), 36782 

(MO 115 ), T.E. Hawkins 1526 (MO 115 ), B.K. Hoist et al. 5248 

(MO 115 ). 

Not included in Flora Mesoamericana (Smith, 1995ft: 384) but 

included here on the basis of material collected from Toledo 

District in 1996-1997 and identified by A.R. Smith in 1997- 

1998. This is the first record of the genus from Belize. 
Terpsichore asplenifolia (L.) A.R. Sm. in Novon 3: 485 (1993)" 4: 387 

Cayo, B.H. Allen 15174 (MO 3 ' " 4 - l15 ); Toledo, B.K. Hoist et al. 

5 189 (MO 115 ). 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 
Terpsichore lehmanniana (Hieron.) A.R. Sm. in Novon 3: 487 

(1 993).. 4: 389 

Cayo, B.H. Allen 15215 (MO 3 ' 114 ' 115 ); Toledo, G. Davidse & D.L. 
Holland 36738 (MO 115 ). 

Terpsichore mollissima (Fee) A.R. Sm. in Novon 3: 487 (1993) 114:39 
Toledo, B.K. Hoist 4040 (MO 91 ' 115 ). 

HYMENOPHYLLACEAE 

Hymenophyllum abruptum Hook., Sp.fil. 1: 88, t. 3 IB (1844) 84:64 

Cayo, H.H. Bartlett 11751 (F 84 ' 115 ). 
Hymenophyllum fucoides (Sw.) Sw. in J. Bot. (Schroder) 1800(2): 

99(1801) 84:66 

Toledo, B.H. Allen 15305 (MO 3 ' 84 - 115 ). 
Hymenophyllum hirsutum (L.) Sw. in J. Bot. (Schrader) 1800(2): 99 

(1801 )84:66 

Cayo, B.H. Allen 15203A (MO 3 ' 84 - 115 ); Toledo, B.K. Hoist et al. 
5304 (MO 115 ). 

Hymenophyllum poly anihos (Sw.) Sw. in/. Bot. (Schrader) 1800(2): 
102(1801) 84:68 

Cayo, C. Whitefoord 9\4l (BM); Stann Creek, W.A. Schipp 114 
(F 84 - 115 ); Toledo, G. Davidse 36124 (MO 115 ). 

Hymenophyllum pulchellum Schltdl. & Cham, in Linnaea 5: 618 
(1830) 84:68 
Toledo, B.K. Hoist 4041 (MO 84 ' 91 ' 115 ). 

Hymenophyllum sieberi (C. Presl) Boschin in Ned. Kruidk. Arch. 4: 
414(1858). 

Toledo, B.K. Hoist el al. 5296 (BRH 115 , MO 115 , UC 115 ), B.K. Hoist 
5974 (MO 115 ). 

Not listed for Belize in the published version of Flora 
Mesoamericana (Pacheco, 1995a: 69), H. sieberi is included in 
the Internet version on the basis of a collection from Toledo 
District (B.K. Hoist et al. 5296, UC) made in 1996 and identified 
by A.R. Smith in May 1997. 

Trichomanes ankersii C. Parker ex Hook. & Grev., Icon, filic. 
2(11): t. 201(1831) 

No material has been collected from this far north in Central 
America according to Flora Mesoamericana (Pacheco, 1995ft: 
73), though it is recorded from Nicaragua to Panama. Standley & 
Record (1936: 60) listed the taxon for Belize but without explicit 
reference to a specimen. There may be material at F, though this 
needs further investigation to verify the identification. 

Trichomanes capillaceum L., Sp. pi. 2: 1099 (1753) 85:73 

Trichomanes capillaceum var. capillaceum* 5 74 
Toledo, B.H. Allen 15411 (MO 3 - 85 - 115 ). 

Trichomanes collariatum Bosch in Ned. Kruidk. Arch. 4: 368 (1858) 85:74 
Stann Creek, W.A. Schipp 364 (BM-000543308!, K!, NY); 
Toledo, M.E. Peck 601 (F 85 - " 5 , K!). 

Trichomanes crispum L., Sp. pi. 2: 1097 (1753) 85:74 

Cayo, B.H. Allen 15172 (MO 3 ' l15 ); Toledo, B.K. Hoist 4023 
(MO 85 - 91 ' 115 ). 

Trichomanes curtii Rosenst. in Repert. Spec. Nov. Regni Veg. 
22(606-608): 5 (1925) 85:75 

No specimen was listed for Belize in Flora Mesoamericana 
(Pacheco, 1995ft: 75), but reference is made to Flora of Guate- 
mala (Stolze, 1976: 78). The latter author also did not cite 
material, but made explicit reference to 'British Honduras'. 
There may be supporting material at F. Trichomanes curtii is also 
recorded from Chiapas and Guatemala, extending southeast from 
Nicaragua to Colombia. 

Trichomanes diaphanum Kunth in Humb., Bonpl. & Kunth, Nov. 
Gen.Sp. 1:25(1816) 85;75 
Toledo, B.K. Hoist et al. 5214 (MO 115 ). 



RECENT RECORDS OF PTERIDOPHYTES 



87 



Trichomanes diversifrons (Bory) Mett. ex Sadeb. in Engl. & Prantl, 
Nat. Pflanzenfam. 1(4): 108 (1899) 85:75 

Belize, C. Whitefoord 1210 (BM-000543306!); Stann Creek, 
W.A. Schipp 237 (K!, NY); Toledo, T.B. Croat 24514 (P 5 - " 5 ). 

Trichomanes ekmanii Wess. Boer in Acta Bot. Neerl. 11: 319, f. 33 

(1 962 )85: 75 

Stann Creek, W.A. Schipp 324 (BM-000543311!, BRH 125 , K, 
MO 85 - 115 , NY). 

Trichomanes elegans Rich. in Actes Soc. Hist. Nat. Paris 1: 114 
(1792) 

Toledo, E.J.F. Campbell s.n. (K!), Mitchell s.n. (K!). 
First collected from Belize in 1875 (Mitchell s.n., K!) but not 
recorded in any subsequent publication for the country either as 
an accepted name, a synonym or as a misapplied name. 
Trichomanes elegans has previously been recorded in Central 
America from Honduras to Panama (Pacheco, 1995ft: 75), from 
South America and the Caribbean region, and is to be expected 
from Belize. 

Trichomanes galeottii E. Fourn. in Bull. Soc. Bot. France 15: 147- 
148(1868) 85:76 

Cayo, H.H. Bartlett 11640 (MO 85 ' " 5 ); Stann Creek, W.A. Schipp 
482 (BM-000543305!, BRH 125 , NY); Toledo, G. Davidse&A.E. 
Brant 32238 (MO 115 ). 

Trichomanes godmanii Hook, in Baker in J. Linn. Soc., Bot. 9: 337, 
t. 8A(1866) 85:76 

Cayo, C.L. Lundell 6187 (MICH 28 , NY); Toledo, F. Boutin & 
Sc/z/oer5023(MO 85 - 115 ). 

Trichomanes hymenoides Hedw., Fil. gen. sp.: t. 3, f. 3 (1799) 85:77 
No specimen was listed for Belize in Flora Mesoamericana 
(Pacheco, 1995ft: 77), but reference is made to Stolze (1976: 81- 
82) who explicitly cited 'British Honduras' in the distribution for 
Flora of Guatemala, but did not mention any material. There 
may be a specimen at F to verify the occurrence in Belize. 

Trichomanes krausii Hook. & Grev., Icon.filic. 2: t. 149 (1830) 
Cayo, T.E. Hawkins 1179 (MO 115 ); Orange Walk, C. Whitefoord 
8095 (BM-000543310!); Toledo, C. Whitefoord 1984 (BM- 
000543309!). 

Recorded for Belize since Forests and flora of British Honduras 
(Standley & Record, 1936: 61). Stolze (1976: 83) gave the 
distribution as 'Mexico to Panama', which may be taken as an 
implicit reference to Belize though no material was listed for 
'British Honduras'. Pacheco (1995ft: 78) did not list the species 
for Belize in the published version of Flora Mesoamericana, 
though several specimens have been collected recently and 
appear on TROPICOS. 

Trichomanes membranaceum L., Sp. pi. 2: 1097 (1753) 85: 
Stann Creek, WA. Schipp S-108 (F 85 - 115 ). 

Trichomanes ovale (E. Fourn.) Wess. Boer in Acta Bot. Neerl. 11: 
296(1962) 

Toledo, C. Whitefoord 1244 (BM, MO 115 ). 
Not recorded for Belize in the published version of Flora 
Mesoamericana (Pacheco, 1995ft: 79) but material collected in 
1976 (C. Whitefoord 1244, MO) is added to the Internet version 
as an extended range record. 

Trichomanes pinnatum Hedw., Fil. gen. sp.: t. 4, f. 1 (1799) 85:8 
Cayo, H.H. Bartlett 11726 (NY); Stann Creek, W.A. Schipp 359 
(K!, MO 85 ' " 5 , NY); Toledo, E.J.F. Campbell 86 (K!). 

Trichomanes polypodioides L., Sp. pi 2: 1098 (1753) 85:8 

Stann Creek, WA. Schipp S-83 (F 5 - 115 , K!); Toledo, B.K. Hoist et 
al. 5 193 (MO 115 ). 

Trichomanes punctatum Poir. in Lam., Encycl. 8: 64 (1808) 85:8 

Trichomanes punctatum subsp. sphenoides (Kunze) Wess. Boer in 
Acta Bot. Neerl. 11: 301 (1962) 85:81 



Cayo, A. Hughes 129 (BM-000557729!); Toledo, B.H. Allen 

15442 (MO 3 - 85 -" 5 ). 
Trichomanes pyxidiferum L., Sp. pi. 2: 1098 (1753) 85:81 

Cayo, A. Hughes 126 (BM-000557730!); Toledo, B.K. Hoist 

43 10 (MO 85 - 91 -" 5 ). 
Trichomanes radicans Sw. in J. Bot. (Schroder) 1800(2): 97 (1801) 

Toledo, B.K. Hoist 5815 (MO 115 , UC), B.K. Hoist 5816 (MO 115 ). 

Not recorded for Belize in the published version of Flora 

Mesoamericana (Pacheco, 1 995ft: 8 1 ) but included in the Internet 

version. The first record for the country is material collected 

from Toledo District on vertical rocks in waterfall spray (B.K. 

Hoist 5815, UC) in 1997, and identified by A.R. Smith in 

December of the same year. 
Trichomanes rigidum Sw., Prodr.: 137 (1788) 85:82 

Cayo, B.H. Allen 15186 (MO 3 - 115 ); Toledo, B.K. Hoist ela\. 5197 

(MO 115 ). 
Trichomanes tuerckheimii H. Christ in Hedwigia 44: 361 (1905) 85:! 

Belize, C. Whitefoord 1298 (BM-000543313!, MO 115 ); Stann 

Creek, W.A. Schipp S-21 (F 5 - " 5 , US); Toledo, G. Davidse & M. 

Meadows 35841 (MO 115 ). 

ISOETACEAE 

Isoetes cubana Engelm. ex Baker in J. Bot. 18: 110 (1880) 21:42 
Toledo, M.E. Peck 420 (GH 21 , NY). 

LOMARIOPSIDACEAE 

Bolbitis bernoullii (Kuhn ex H. Christ) Ching in C. Chr., Index filic., 
Suppl. 3:47(1934) 20:248 
Toledo, WA. Schipp S-776 (GH 20 - 115 ). 

Bolbitis hastata (E. Fourn.) Hennipman in Amer. Fern J. 65: 1975 
(1975) 

Toledo, B.K. Hoist 5757 (MO 115 , UC). 

Not recorded for Belize in the published version of Flora 
Mesoamericana (Hennipman & Moran, 1995: 248), but included 
in the Internet version based on a specimen (B.K. Hoist 5757, 
UC) collected from Toledo District in 1997 and identified by 
A.R. Smith later that year. 

Bolbitis hemiotis (Maxon) Ching in C. Chr., Index filic., Suppl. 3: 48 

(1 934)20: 248 

Toledo, B.H. Allen 15441 (MO 3 - 20 - 115 ). 
Bolbitis pergamentacea (Maxon) Ching in C. Chr., Index filic., 

Suppl. 3:49(1934) 20:249 

Toledo, WA. Schipp S-764 (GH 20 - 115 ). 
Bolbitis portoricensis (Spreng.) Hennipman in Amer. Fern J. 65: 30 

(1 975 )20: 249 

Stann Creek, W.A. Schipp 526 (BRH 125 , NY); Toledo, G. Davidse 
& A.E. Brant 32381 (MO 20 - 115 ). 

Elaphoglossum christianeae Mickel in Novon 2: 371 (1992) 
Toledo, G. Davidse & H.B. Buchanan 36941 (MO 115 ). 
Not recorded for Belize in Flora Mesoamericana (Mickel, 1995a: 
263), this Central American endemic has been recorded formerly 
from Costa Rica and Panama but to that distribution can be added 
a specimen collected in Toledo District of Belize in 1997 and 
identified by A.F. Rojas-Alvarado in 1998. 

Elaphoglossum decursivum Mickel in Brittonia 32: 334 (1980) 
Toledo, G. Davidse & D.L. Holland 36729 (MO 115 ), D.L Hol- 
land & B. Kid 90 (MO 115 ), B.K. Hoist 5924 (MO 115 ). 
This taxon is not recorded for Central America in the published 
version of Flora Mesoamericana (Mickel, 1995a), but appended 
to the Internet version on the basis of recent identifications by 
A.R. Smith of material from Belize, Honduras and Costa Rica. 



88 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



All of the Belize material listed above was collected in 1997 from 
Toledo District. 

Elaphoglossum erinaceum (Fee) T. Moore, Index fil: 9 (1857) 
Toledo, G. Davidse & D.L. Holland 36764 (MO 115 ). 
Not included in the published version of Flora Mesoamericana 
(Mickel, 1995a: 265) but listed for Belize in the Internet version 
based on the collection listed above from Toledo District made in 
1997 and identified by A.R. Smith in 1998. 

Elaphoglossum eximiiforme Mickel in Novon 2: 374 (1992) 
Toledo, B.K. Hoist 5672 (MO 115 ). 

Not recorded for Belize in Flora Mesoamericana (Mickel, 1 995a: 
265), this Central American endemic was formerly known only 
from Costa Rica and Panama. The material included here was 
originally identified as E. latifolium (Sw.) Sm. by A.R. Smith but 
re-determined as E. eximiiforme by A.F. Rojas-Alvarado in 
October 1998. 

Elaphoglossum glaucum T. Moore, Index fii: 10 (1857) 
Toledo, B.K. Hoist 3869 (MO 91 115 ). 

Not recorded for Belize in Flora Mesoamericana (Mickel, 1 995a: 
267), though recorded from the neighbouring countries of Mexico, 
Guatemala and Honduras. The record here is tentative as it is 
based on material from Toledo District identifed by R. Moran in 
1992 and cited by Parker et al. (1993: 40), yet excluded from the 
flora. 

Elaphoglossum guatemalense (Klotzsch) T. Moore in Parker's Cat. 
(1858) 32:268 

Cayo, B.H. Allen 15175 (MO 3 ' 115 ); Toledo, G. Davidse & A.E. 
Brant 32239 (MO 3 - 115 ). 

Elaphoglossum herminieri (Bory ex Fee) T. Moore, Index fil.: xvi 

(1 8 57)32:268 

Toledo, F. Boutin & Schlosser 5088 (NY 32 ). 

Elaphoglossum herrerae A. Rojas in Brenesia 45-46: 13, f. 5 
(1996) 

Toledo, B.K. Hoist 5670 (MO 115 ). 

A new species provisionally included in the Internet version of 
Flora Mesoamericana based on a specimen from Toledo District 
listed above and identified by A.F. Rojas-Alvarado in October 
1998. This taxon was originally considered to be endemic to the 
Cordillera de Guanacaste in Costa Rica. 

Elaphoglossum latifolium (Sw.) J. Sm. in London J. Bot. 1: 197 



Toledo, P.H. Gentle 3756 (NY 32 ). 
Elaphoglossum latum (Mickel) Atehortiia ex Mickel in Fieldiana, 

Bot.n.s.,27: 123 (1991) 32:271 

Toledo, T.B. Croat 24313 (MO 32 - 115 ). 
Lomariopsis japurensis (Mart.) Sm., Hist, fil.: 140 (1875) 6 * 284 

Toledo, T.B. Croat 24393 (MO 66 - 115 ). 
Lomariopsis recurvata Fee, Mem. Foug. 2: 68 (1845) 66:284 

Cayo, T.B. Croat 23800 (MO 66 ); Stann Creek, R. Rivero et al. 

2524 (BRH 125 ); Toledo, B.K. Hoist 4417 (MO 91 ' 115 ). 
Lomariopsis vestita E. Fourn. in Bull. Soc. Bot. France 19: 250 

(1 872)66: 284 

Toledo, F. Boutin & Schlosser 5093 (MO 66 - 115 ). 
Peltapterispeltata (Sw.)C.V. Morton in Amer. FernJ. 45: 13 (1955) 
Cayo, B.H. Allen 15275 (BM-000543333, MO 3 - 115 ); Toledo, C. 
Whitefoord 1727 (BM-000543338!). 

Curiously omitted for Belize in Flora Mesoamericana (Mickel, 
1 995b: 285) although it was cited under the name Elaphoglossum 
peltatum (Sw.) Urb. by Parker et al. (1993: 40) and there are 
many specimens listed under the same synonym in TROPICOS. 
The earliest collection from Belize seen to date (C. W. Whitefoord 
1727, BM) was collected in May 1979. Mickel listed four 
separate forms of this species, but the northern material from 



Mexico, Guatemala and Honduras all falls within typical forma 
peltata. 

LOPHOSORIACEAE 

Lophosoria quadripinnata (J.F. Gmel.) C. Chr., Nat. Hist. Juan 
Fernandez 2: 16(1920) 

Lophosoria quadripinnata var. quadripinnata 
Toledo, T.E. Hawkins 1540 (MO 115 , UC). 
Not listed for Belize in the published version of Flora 
Mesoamericana (Riba, 1995a: 85), but included in the Internet 
version on the basis of a collection from Toledo District (T.E. 
Hawkins 1540, UC) made in 1996 and identified by A.R. Smith 
in January 1998. This is the first record of both the genus and the 
family for Belize. 

LYCOPODIACEAE 

Huperzia dichaeoides (Maxon) Holub in Folia Geobot. Phytotax. 

20:72(1985) 82:11 

Toledo, B.H. Allen 15384 (BM-000543283!, MO 3 - 82 - 115 ). 
Huperzia dichotoma (Jacq.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 

(1874) 82:11 

Toledo, P.H. Gentle 3034 (GH 82 - 115 ). 
Huperzia linifolia (L.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 

(1 874)82: 13 

Huperzia linifolia var. linifolia* 2 13 

Cayo, B.H. Allen 15280 (MO 3 - 115 ); Toledo, W.A. Schipp 811 
(US 82 - 115 ). 

Huperzia pithyoides (Schltdl. & Cham.) Holub in Folia Geobot. 
Phytotax. 20:76(1985) 
Toledo, B.K. Hoist 4381 (MO 91 ). 

Not listed for Belize in Flora Mesoamericana (011gaard, 1995a: 
15) though it was recorded from all surrounding countries and to 
be expected from Belize. Huperzia pithyoides was first cited for 
the country by Parker et al. (1993: 40) based on the specimen 
listed above collected from the Columbia River Forest Reserve in 
Toledo District. 

Huperzia reflexa (Lam.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 

(! 874)82: 16 

Huperzia reflexa var. reflexa* 2 - 16 

Toledo, M.C. Carlson 2617 (F 82 - 115 ). 

Huperzia taxifolia (Sw.) Trevis. in Atti Soc. Ital. Sci. Nat. 17: 248 
(1874) 82:17 

Cayo, C.L. Lundell 6258 (MICH 28 , NY 82 - 115 ); Toledo, G. Davidse 
36384 (MO 115 ). 

Lycopodiella caroliniana (L.) Pic. Serm., Webbia 23: 165 (1968) 83: 19 

Lycopodiella caroliniana var. meridionalis (Underw. & F.E. Lloyd) 
B. 011g. & P.O. Windisch in Bradea 5: 27 (1987) 83: 19 
Belize, C. Whitefoord 2405 (BM-000543290!); Cayo, G. Davidse 
&A.E. Brant 33079 (BM-000543288!, BM-000543289!, MO 115 ); 
Stann Creek, WA. Schipp51S (BM-000543286!, MO 115 , NY, S 83 - 115 ). 

Lycopodiella cernua (L.) Pic. Serm. in Webbia 23: 166 (1968) 83: 19 
Belize, C. Mute/o0frf2570(BM-000543294!); Cayo, J.N. Hedger 
127 (BM-000543297!); Stann Creek, W.A. Schipp 234 (BM- 
000543243!, NY); Toledo, P.H. Gentle 6781 (BM-000543244!, 
NY). 

MARATTIACEAE 

Danaea elliptica Sm. in Rees, Cycl. 11: Danaea no. 2 (1808) 9:49 
Cayo, C. Whitefoord 1215 (BM-000543246!); Stann Creek, T.B. 
Croat 24525 (MO 9 - 115 ); Toledo, B.K. Hoist 4305 (MO 91 - " 5 ). 



RECENT RECORDS OF PTERIDOPHYTES 



89 



Danaea nodosa (L.) Sm. in Mem. Acad. Roy. Sci. (Turin) 5(1790- 
1791): 420, t. 9, f. 11 (1793) 9:49 

Cayo, B.H. Allen 15 185 (MO 3 ' " 5 ); Stann Creek, W.A. Schipp 422 
(F- " 5 , US); Toledo, B.K. Hoist et al. 5503 (MO 115 ). 

Marattia excavata Underw. in Britton, N. Amer.fl. 16(1): 22 (1909) 
Toledo, B.K. Hoist et al. 5300 (BRH 115 , MO 115 , UC" 5 ). 
Not listed for Belize in the published version of Flora 
Mesoamericana, (Perez-Garcia, 1995a: 50) but included in the 
Internet version on the basis of a collection from Toledo District 
(B.K. Hoist et al. 5300, UC) made in 1996 and identified by A.R. 
Smith in April 1997. This is the first record of the genus in Belize. 
The specimen is cited as 'B (Hoist et al. 5300, US)' in the Internet 
description but it is not clear from the TROPICOS account if 
there is a duplicate at US or if this is an error for UC. 

METAXYACEAE 

Metaxya rostrata (Kunth) C. Presl, Tent, pterid.: 60, t. 1, f. 5 

(1836) 100:86 

Cayo, D. Burch 5881 (MO 115 , NY); Stann Creek, W.A. Schipp 89 
(BRH 125 , NY); Toledo, P.H. Gentle 2631 (GH 100 ). 

OPHIOGLOSSACEAE 

Cheiroglossa palmata (L.) C. Presl, Suppl. tent, pterid.: 57 (1845) 126:46 

Cayo, B.H. Allen 15170 (AAU 126 , MO 3 - 115 ); Toledo, B.K. Hoist 

5935 (MO 115 ). 
Ophioglossum nudicaule L.f., Suppl. pi: 433 (1782) 126:47 

Belize, C. Whitefoord 2605 (BM-000543245!); Stann Creek, 

P.H. Gentle 2997 (MO 115 - 126 ). 

POLYPODIACEAE 

Campyloneurum angustifolium (Sw.) Fee, Mem. Foug. 5: 257 

(1852) 25:335 

Cayo, D.R. Hunt 7054 (K!); Toledo, T.E. Hawkins 1348 (MO 115 ). 
Campyloneurum aphanophlebium (Kunze) T. Moore, Index fil. : 223 

(1861) 25:335 

Toledo, P.H. Gentle 7327 (US 25 ). 
Campyloneurum brevifolium (Lodd. ex Link) Link, Fil. spec.: 124 

(1841) 25:335 

Cayo, A. Hughes 96 (BM-000531759!, BM-000531835!); Stann 

Creek, W.A. Schipp 527 (BRH 125 , K); Toledo, P.H. Gentle 7835 

(F 5 , US). 
Campyloneurum costatum (Kunze) C. Presl, Tent, pterid.: 190 



Toledo, P.H. Gentle 6792 (US 25 ). 
Campyloneurum fasciale (Humb. & Bonpl. ex Willd.) C. Presl, Tent. 

pterid.: 190 (1836) 25:336 

Cayo, C. Whitefoord 2044 (BM, MO 25 ); Toledo, G. Davidse 

36003 (MO 115 ). 
Campyloneurum phyllitidis (L.) C. Presl, Tent, pterid.: 190 (1836) 25:33 ~ 

Belize, C.L. Lundell s.n. (K!); Cayo, A. Hughes 25b (BM- 

000531767!); Toledo, P.H. Gentle 1108 (F 25 , K). 
Campyloneurum repens (Aubl.) C. Presl, Tent, pterid.: 190 (1836) 25:331 

Cayo, C.L. Lundell 6262 (MICH 28 , US); Toledo, T.B. Croat 

24462 (MO 25 ). 
Campyloneurum xalapense Fee, Mem. Foug. 5: 258 (1852) 25: 

Toledo, P.H. Gentle 6515 (F 5 , US). 
Microgramma lycopodioides (L.) Copel., Gen. fil. (Ann. Cryptog. 

Phytopathol. 5): 185 (1947) 69:339 

Cayo, C.L Lundell 6287 (MICH 28 , NY); Stann Creek, R. Rivero 

et al. 2555 (BRH 125 ); Toledo, C. Whitefoord 1863 (BM, CR 69 ). 
Microgramma nitida (J. Sm.) A.R. Sm. in Proc. Calif. Acad. Sci. ser. 

4, 40(8): 230 (1975) 69:339 



Orange Walk, G. Davidse & A.E. Brant 32765 (MO 115 ); Toledo, 

D.L. Spellman & W.W. Newey 2105 (MO 69 ). 
Microgramma percussa (Cav.) de la Sola in Physis (A, B & C) 

44(106, Secc. C): 28 (1986) 69:339 

Cayo, A. Hughes 77 (BM-000531808!, BM-000557711! 22 ); To- 
ledo, G. Davidse & A.E. Brant 32027 (MO 69 - " 5 ). 
Microgramma reptans (Cav.) A.R. Sm. in Proc. Calif. Acad. Sci. ser. 

4,40(8):230(1975) 69:34 

Stann Creek, W.A. Schipp 210 (BRH 125 , NY); Toledo, C. 

Whitefoord 1590 (BM, CR 69 ). 
Neurodium lanceolatum (L.) Fee, Mem. Foug. 3: 28 (1852) 87:341 

Cayo, P.H. Gentle 2518 (MEXU 87 " 5 , NY); Toledo, G. Davidse 

& A.E. Brant 32346 (MO 115 ). 
Niphidium crassifolium (L.) Lellinger in Amer. Fern J. 62: 106 

(1 972)70: 341 

Cayo,A//wg/^21(BM-000531766! 22 ,BM-000531797!);Stann 

Creek, W.A. Schipp 88 (BRH 125 ); Toledo, P.H. Gentle 4964 

(MO 70 - 115 ). 
Niphidium oblanceolatum A. Rojas in Brenesia 45-46: 28, f. 1 

(1996) 

The protologue of this recently described species includes Belize 

in the distribution, and the taxon is provisionally accepted for the 

Internet version of Flora Mesoamericana. 
Pecluma atra (A.M. Evans) M.G. Price in Amer. Fern J. 73: 113 

(1 983)7 1:342 

Cayo, C.L. Lundell 6639 (GH, MICH 28 , US-1638286 15 - 71 ); To- 
ledo, M.E. Peck 820 (NY). 

Pecluma dispersa (A.M. Evans) M.G. Price in Amer. Fern J. 73: 114 
(1983) 71:343 

Cayo, A. Hughes 38 (BM-000531768! 22 , BM-000531855!); 
Toledo, T.B. Croat 24176 (MO 71 - 115 ). 

Pecluma divaricata (E. Fourn.) Mickel & Beitel in Mem. New York 
Bot. Card. 46: 269 (1988) 71:343 
Toledo, B.K. Hoist 4460 (MO 71 - 91 - 115 ). 

Pecluma pectinata (L.) M.G. Price in Amer. Fern J. 73: 115 (1983) 
Cayo, D.R. Hunt 605 (BM), G.R. Proctor 29841 (BRH 125 ). 
Not recorded for Belize in Flora Mesoamericana (Moran, 1 995ai: 
344) though specimens have been collected under the synonym 
Polypodiumpectinatum L. This record requires further investigation. 

Pecluma plumula (Humb. & Bonpl. ex Willd.) M.G. Price in Amer. 
Fern J. 73: 115 (1983) 71:345 

Belize, C. Whitefoord 1242 (BM); Cayo, A. Hughes 120 (BM- 
000557724! 22 ); Corozal, G. Davidse & A.E. Brant 32534 (MO 71 - 
I15 ); Orange Walk, C.L. Lundell 537 (NY); Toledo, T.E. Hawkins 
1717 (MO 115 ). 

Phlebodium decumanum (Willd.) J. Sm. in J. Bot. (Hooker) 4: 59 

(1 84 1)72: 345 

Toledo, A.H. Gentry 88 (F 2 ). 

Pleopeltis astrolepis (Liebm.) E. Fourn., Mexic. pi. 1: 87 (1872) 26:347 
Cayo, C.L. Lundell 6450 (MICH 28 , NY 26 ); Toledo, W.A. Schipp 
923 (BRH 125 , MO 115 ). 

Pleopeltis crassinervata (Fee) T. Moore, Index fil.: 345 (1862) 
Not recorded from Belize in Flora Mesoamericana (Lorea Hernan- 
dez, 1995: 348). This taxon was originally described from Mexico 
and listed by Weatherby (1922) from Veracruz, Chiapas and 
Guatemala. Lorea Hernandez expanded this distribution to include 
Honduras, Nicaragua and Costa Rica. Seymour (1975: 167) had 
listed Weatherby's combination for the taxon in his notes on the 
genus Polypodium in Nicaragua, but with the incorrect epithet 
'crassinervum'. It is clear from the exact reference to Weatherby 
that this taxon is intended, and Seymour listed Central American 
material including a specimen from 'British Honduras' at GH. 
This material needs locating and the identification verifying. 



90 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



x Pleuroderris michleriana (D.C. Eaton) Maxon in J. Wash. Acad. 
Sci. 24: 551, f. 1-2 (1934), pro sp. 

Stann Creek, R. Rivero et al. 2571 (BRH 125 ), 2608 (BRH 125 ). 
Hybridization between Tectaria incisa andDictyoxiphiumpanam- 
ense produces the very variable intrageneric hybrid x Pleuroderris 
michleriana (Wagner et al., 1978). It was not recorded for Belize 
by Moran (1995f: 201, 1995v: 207) though both parental species 
are present in the area. 

Polypodium dissimile L., Syst. nat. 10th ed., 2: 1325 (1759) 73;355 
Cayo, B.H. Allen 15173 (MO 3 - 115 ); Toledo, B.K. Hoist 4034 
(MO 73 ' 91 ' 115 ). 

Polypodium dulce Poir. in Lam., Encycl. 5: 523 (1804) 73:352 
Toledo, F. Boutin & Schlosser 5022 (MO 73 ' 115 ). 

Polypodium fallax Schltdl. & Cham, in Linnaea 5: 609 (1830) 73:361 
Cayo, C.L. Lundell 6216 (MICH 28 , NY); Toledo, C. Whitefoord 
1745 (BM, NY). 

Polypodiumfraternum Schltdl. & Cham, in Linnaea 5: 608 (1830) 73:358 
Cayo, A. Hughes 121 (BM-000532009!). 

Polypodium fraxinifolium Jacq., Collectanea 3: 187 (1791) 

Toledo, G. Davidse & D.L. Holland 36729A (MO 115 ), T.E. 
Hawkins 1427 (MO 115 ), 1429 (MO 115 ), B.K. Hoist et al. 5257 
(MO 115 ). 

Not listed for Belize in the published version of Flora 
Mesoamericana (Moran, \995ak: 355), but included in the 
Internet version on the basis of a collection from Toledo District 
(Davidse & Holland 36729 A, MO) made in 1997 and identified 
by A.R. Smith in January 1998. 

Polypodium hispidulum Bartlett in Proc. Amer. Acad. Arts 43: 48 
(1907) 

Cayo, T.E. Hawkins 1078 (MO 115 , UC); Toledo, T.E. Hawkins 
1330 (MO 115 ). 

Not listed for Belize in the published version of Flora 
Mesoamericana (Moran, \995ak: 358), but included in the 
Internet version on the basis of a collection from Cayo District 
(T.E. Hawkins 1078, UC) made in 1996 and identified by A.R. 
Smith in December 1997. 

Polypodium lindenianum Kunze, Farrnkrduter 2: 83 (1849) 73:362 
Cayo, A. Hughes 131 (BM-000557728! 22 ); Toledo, G. Davidse et 
al. 35704 (MO 115 ). 

Polypodium polypodioides (L.) Watt in Canad. Naturalist & Quart. 
J. Sci. ser. 2, 3: 158 (1867) 73:363 

Polypodium polypodioides var. aciculare Weath. in Contr. Gray 
Herb. 124: 33 (1939) 

Not recorded for Belize in Flora Mesoamericana, though present 
in Chiapas, Guatemala and Honduras and to be expected from 
Belize. Seymour (1975: 160) cited material of P. polypodioides 
var. burchellii (Baker) Weath. from countries of Central America 
in his treatment of the genus Polypodium for Nicaragua, and 
included reference to 'British Honduras' based on material at 
GH. Moran (\995ak: 364) considered that var. burchellii did not 
occur in Central America, but there is a note describing the 
confusion between this South American variety and var. aciculare. 
It is evident that Seymour's concept of var. burchellii falls under 
var. aciculare in Flora Mesoamericana, though the Belizean 
material at GH should be located to verify the identity. 

Polypodium polypodioides var. polypodioides 1 ^ 364 

Belize, C.L. Lundell 3884 (K!, NY); Cayo, J.D. Dwyer & R.L. 
Liesner 12092 (MO 73 ); Stann Creek, W.A. Schipp 349 (BRH 125 , 
K!, NY); Toledo, P.H. Gentle 906 (K!, NY). 

Polypodium triseriale Sw. in/ Bot. (Schroder) 1800(2): 26 (1801) 73:357 
Cayo, A. Hughes 117 (BM-000531901!, BM-000557725! 22 ); 
Toledo, P.H. Gentle 5014 (MO 73 - l15 ). 

Pseudocolysis bradeorum (Rosenst.) L.D. Gomez in Brenesia 10- 



11: 116(1977) 74:365 

RecordedforBelizeinF/oraMesoam^ncana (Moran, 1995a/: 365) 
on the basis of the entry in Flora of Guatemala (Stolze, 1981:381) 
where the distribution explicitly included 'British Honduras'. 
Stolze did not cite any material but made reference to Evans & 
Mickel (1969). Material was cited at GH from 'British Honduras' 
by Seymour ( 1 975 : 1 67), but without detail. This or other material 
should be sought to verify the distributional record for Belize. 

PSILOTACEAE 

Psilotum nudum (L.) P. Beauv. , Prodr. Aetheogam. : 1 06, 1 1 2 ( 1 805) 88; 3 
Belize, G. Davidse & D.L. Holland 37046 (MO 115 ); Corozal, G. 
Davidse & A.E. Brant 32533 (MO 115 ); Stann Creek, R. Rivero et 
al. 2613 (BRH 125 ); Toledo, W.A. Schipp S-261 (MO 88 - 115 ). 

PTERIDACEAE 

Acrostichum aureum L., Sp. pi: 1069 (1753) 41; 105 

Belize, M.-H. Sachet & D.R. Stoddart 1631 (BRH 125 ); Stann 

Creek, F.R. Fosberg & M.-H. Sachet 53849 (BRH 125 , MO 41 - " 5 ); 

Toledo, D.L Spellman & D.R. Stoddart 2299 (BRH 125 ). 
Acrostichum danaeifolium Langsd. & Fisch., PI. Voy. Russes monde 

1:5, t. 1 (1810) 41:105 

Belize, J.D. Dwyer 11449 (MO 115 ); Toledo, D.L. Spellman & 

D.R. Stoddart 2477 (MO 41 - 115 ). 
Adiantopsis radiata (L.) Fee, Mem. Foug. 5: 145 (1852) 86: 106 

Cayo, T.B. Croat 23775 (MO 86 - 115 ); Toledo, P.H. Gentle 6086 

(MO 115 ). 
Adiantum capillus-veneris L., Sp. pi.: 1096 (1753) 78: 108 

Toledo, B.K. Hoist 4014 (MO 78 91 - 115 ). 
Adiantum concinnum Humb. & Bonpl. ex Willd., Sp. pi. 5: 451 

(1810 )78:109 

Cayo, A. Hughes 146 (BM-000528104! 22 , BM-000531933!). 
Adiantum decoratum Maxon & Weath. in Amer. J. Bot. 19: 165 

(1 932)78: 113 

Toledo, T.B. Croat 24179 (MO 78 - 115 ). 
Adiantum fructuo sum Poepp. ex Spreng., Syst. veg. 16th ed., 4(1): 

113(1827) 78;114 

Toledo, T.B. Croat 24373 (BRH 125 , MO 78 - 115 ). 
Adiantum humile Kunze in Linnaea 9: 80 (1834) 78: " 4 

Toledo, G. Davidse 36082 (MO 115 ). 
Adiantum latifolium Lam., Encycl. 1(1): 43 (1783) 78: 114 

Cayo, C. Whitefoord 2096 (BM-000543184! 78 ' 115 ); Orange Walk, 

T. Arnason & J. Lambert 17176 (MO 115 ); Stann Creek, W.A. 

Schipp 77 (BM-000543 186!, BRH 125 , NY); Toledo, C. Whitefoord 

1812 (BM-000543 188!, MEXU). 
Adiantum macrophyllum Sw., Prodr.: 135 (1788) 78: 112 

Cayo, A. Hughes 14 (BM-000528105! 22 ); Stann Creek, W.A. 

Schipp 340 (BM-000543254!, BRH 125 , NY); Toledo, C. 

Whitefoord 1848 (BM-000543 190!, MO 78 - 115 ). 
Adiantum obliquum Willd., Sp. pi. 5: 429 (1810) 78: 112 

Belize, C. Whitefoord 2366 (BM); Cayo, A. Hughes 70 (BM- 

000528 106! 22 , BM-000531796!); Stann Creek, W.A. Schipp 275 

(BM, BRH 125 , MO 115 , NY); Toledo, C. Whitefoord 1521 (BM- 

000543253 ! 115 ). 
Adiantum petiolatum Desv. in Ges. Naturf. Freunde Berlin Mag. 

Neuesten Entdeck. Gesammten Naturk. 5: 326-327 (1811) 78: 112 

Belize, C. Whitefoord 2569 (BM); Cayo, A. Hughes 62 (BM- 

000528 100! 22 , BM-000531788!); Stann Creek, R. Rivero et al. 

2595 (BRH 125 ); Toledo, G.R. Proctor 36007 (BM-000543 194!). 
Adiantum princeps T. Moore in Card. Chron. n.s., 4: 197, f. 43^4 

(1875) 

Cayo, M.J. Balick et al. 3140 (MO 115 , NY). 



RECENT RECORDS OF PTERIDOPHYTES 



91 



Not recorded for Belize by Moran in Flora Mesoamericana 

(Moran, Zimmer & Jermy 1995: 109), though the specimen listed 

here was collected in 1991 and identifed by J.T. Mickel in 1993. 
Adiantum pulverulentum L., Sp. pi. 2: 1096 (1753) 78: " 5 

Cayo,A//Hg/2^56(BM-000528101! 22 ,BM-000531791!);Stann 

Creek, W.A. Schipp 241 (BM-000543 199!, BRH 125 , NY); Toledo, 

C. Whitefoord 1524 (NY, BM-000543196!). 
Adiantum tenerum Sw., Prodr.: 135 (1788) 78: " 

Cayo, DA. Sutton et al. 3 (BM-000543 1 80!); Orange Walk, M.J. 

Balick 3219 (BRH 125 , NY); Stann Creek, M.J. Balick 3091 

(BRH 125 , NY); Toledo, G. Davidse&A.E. Brant 32302 (MO 78 ' 115 ). 
Adiantum terminatum Kunze ex Miq. in Verslagen Meded. Vier Kl. 

Kon. Inst. Wetensch. Letterk. Schoone Kunsten 1842: 187 (1843) 78: " 6 

Toledo, G.R. Proctor 35922 (BRH 125 , F 8 - 115 ). 
Adiantum tetraphyllum Humb. & Bonpl. ex Willd., Sp. pi 5: 441 

(1810) 78:116 

Orange Walk, C.L. Lundell 404 (NY); Stann Creek, R. Rivero et 

al. 2523 (BRH 125 ); Toledo, B.K. Hoist 4360 (MO 91 - 115 ). 
Adiantum trapeziforme L., Sp. pi. 2: 1097 (1753) 78: " 6 

Cayo, P.H. Gentle 2342 (F 8 - " 5 , MO 115 , NY); Toledo, G. Davidse 

36198 (MO 115 ). 
Adiantum trichochlaenum Mickel & Beitel in Mem. New York Bot. 

Card. 46:29, f. 41L (1988) 

Toledo, B.K. Hoist 5873 (MO 115 ). 

Jermy did not record this taxon from Belize in the published 

version of Flora Mesoamericana (Moran et al., 1995: 116), but it 

was included in the Internet version based on a collection made 

in 1997 from Toledo District and identified by A.R. Smith in the 

same year. 
Adiantum tricholepis Fee, Mem. Foug. 8: 72 (1857) 78: uo 

Cayo, M. Brunt 2219 (BM-000543178!); Orange Walk, T. 

Arnason & J. Lambert 17177 (MO 78 - " 5 , NY); Toledo, T.E. 

Hawkins 1676 (MO 115 ). 
Adiantum villosum L., Syst. nat. 10th ed., 2: 1328 (1759) 78: 117 

Cayo, A. Hughes 135 (BM-000528108! 22 , BM-000531916!); 

Orange Walk, M.J. Balick 3220 (BRH 125 , NY); Toledo, P.H. 

Genr/e2409(F 8 - 115 , NY). 
Adiantum wilesianum Hook., Sp.fil. 2: 50 (1851) 78: 117 

Belize, P.H. Gentle 1569 (MO 115 , US); Cayo, A. Hughes 144b 

(BM-000528109!, BM-000531930!); Stann Creek, W.A. Schipp 

48 (BM-000543267!, BRH 125 , US); Toledo, G. Davidse et al. 

36450 (BM-B000531990, BRH, MO 115 ). 
Adiantum wilsonii Hook., Sp.fil. 2: 6 (1851) 78: 113 

Toledo, C. Whitefoord 1855 (BM-000543 176! 78 -" 5 , NY). 
Cheilanthes microphylla (Sw.) Sw., Syn.fil.: 111 (1806) 131: 127 
Cheilanthes microphylla var. microphylla 131 

Cayo, C.L. Lundell 6542 (MICH 28 - 121 , US). 
Cheilanthes notholaenoides (Desv.) Maxon ex Weath. in Contr. 

Gray Herb. 114: 34 (1936) 131: 128 

Cayo, A. Hughes 92 (BM-000557720! 22 , BRH, RNG). 
Hemionitis palmata L., Sp. pi.: 1077 (1753) 97: 132 

Stann Creek, W.A. Schipp 250 (BM-000543336!, BRH 125 , 

MICH 97 , MO 115 ); Toledo, C. Whitefoord 1904 (BM- 

000543266!). 

Pityrogramma calomelanos (L.) Link, Handbuch 3: 20 (1833) 42: 
Pityrogramma calomelanos var. calomelanos 42 

Belize, P.H. Gentle 1548 (K, NY); Cayo, T.B. Croat 24862 

(MO 42 - 115 ); Stann Creek, P.H. Gentle 8372 (BM-000543264!, 

NY); Toledo, B.K. Hoist et al. 35596 (MO 115 ). 
Pteris altissima Poir. in Lam., Encycl. 5: 722 (1804) 43: 141 

Cayo, A. Hughes 72 (BM-000531762! 22 , BM-000531800!, BM- 

000531801!); Stann Creek, /. Robertson23\ (BM-000543 191!); 

Toledo, T.B. Croat 23808 (MO 43 - 115 ). 



Pteris biaurita L., Sp. pi: 1076 (1753) 43: 141 

Cayo, A. Hughes 114 (BM-000531903!, BM-000557721! 22 ); 
Toledo, P.H. Gentle 2376 (US). 

Pteris grandifolia L., Sp. pi. 2: 1073 (1753) 43 l41 

Cayo, A. Hughes 37 (BM-000531758! 22 , BM-000531856!, BM- 
000531857!); Orange Walk, C. Whitefoord 8169 
(BM-000543166!); Toledo, P.H. Gentle 1523 (K). 

Pteris longifolia L., Sp. pi.: 1074 (1753) 43: 142 

Cayo, A. Hughes 81 (BM-000557714! 22 ); Orange Walk, C. 
Whitefoord 8168 (BM-000543 171!); Stann Creek, W.A. Schipp 
428 (BM-000543 173!, BRH 125 ); Toledo, J.D. Dwyer et al. 261 
(MO 43 - 115 ). 

Pteris propinqua J. Agardh, Recens. spec. Pter.: 65 (1839) 
Toledo, D.L. Holland 14A (MO 115 , UC). 
Not included for Belize in the published version of Flora 
Mesoamericana (Moran, 1995g: 143) but included in the Internet 
version based on a collection from Toledo District (D.L. Holland 
14A, UC) made in 1997 and determined by A.R. Smith. 

Pteris pungens Willd., Sp. pi. 5: 387 (1810) 43: 144 

Cayo, D.R. Hunt 607 (BM-000543 170!); Stann Creek, W.A. 
Schipp 289 (BM-000543 167!, BRH 125 ); Toledo, C. Whitefoord 
1849(BM-000543169!). 

Pteris quadriaurita Retz., Observ. bot. 6: 38 (1791) 43: 144 
Toledo, B.K. Hoist 3900 (MO 43 - 91 - 115 ). 

SALVINIACEAE 

Salvinia auriculata Aubl., Hist. pi. Guiane 2: 969 (1775) 75: 3% 
Orange Walk, G. Davidse & A.E. Brant 32892 (MO 75 - 115 ). 

Salvinia minima Baker in J. Bot. 24: 98 (1886) 75:396 

Belize, G. Davidse & A.E. Brant 33130 (MO 115 ); Toledo, C. 
Whitefoord 2145 (BM, MO 75 - 115 ). 

SCHIZAEACEAE 

Actinostachys germanii Fee, Mem. Foug. 11: 123 (1866) I02; 
Toledo, M.E. Peck 936 (GH 120 ). 

Anemia adiantifolia (L.) Sw., Syn.fil.: 157 (1806) 76:53 

Belize, P.H. Gentle 1316 (K!, MO 76 - 115 ); Cayo, A. Hughes 83 
(BM-000557706! 22 ); Toledo, G. Davidse & A.E. Brant 32322 
(MO 115 ). 

Anemia bartlettii Mickel in Iowa State J. Sci. 36: 420 (1962) 76:54 
Cayo, H.H. Bartlett 1 1898 (MICH-holotype 76 - " 5 , MICH-isotype, 
UC, US), D.R. Hunt 428 (BM, BRH, UCWI, US 120 ), J.R. Wiley 
402 (MO 76 - 115 ). 

One of very few Belizean endemic taxa, this locally distributed 
fern from the Mountain Pine Ridge area in Cayo District was first 
collected in 1931. It was identified as A. flexuousa by Maxon 
(1944a: 17) and distributed under that name, but not recognized 
as a new species until 1962. The specimen cited in the distribu- 
tional statement in Flora Mesoamericana (Moran & Mickel, 
1995: 54) as 'B (Wiley 4402, MO)' is incorrect as the collections 
made by J.R. Wiley in August 1970 were less than 500 numbers 
and another record on TROPICOS correctly lists this as 402. 

Anemia hirta (L.) Sw., Syn.fil.: 155 (1806) 76;54 
Cayo, D.A. Sutton et al. 4 (BM-000543239!). 

Anemia mexicana Klotzsch in Linnaea 18: 526 (1844) 76:55 

Anemia mexicana var. makrinii (Maxon) Mickel in Brittonia 33: 42 1 
(1981) 76:55 

Cayo, A. Hughes 99 (BM-000531834!, BM-000557705! 22 ); 
Toledo, G. Davidse & A.E. Brant 32127 (MO 76 - 115 ). 
Described originally from Mexico, this taxon has only been 
recorded from Belize for the Flora Mesoamericana area (Moran 
& Mickel, 1995: 55). 



92 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



Anemia oblongifolia (Cav.) Sw., Syn.fil.: 156 (1806) 76:55 

Cayo, J.N. Hedger 319 (BM-000543241!); Toledo, J.R. Wiley 

351 (MO 76 - 115 ). 
Anemia pastinacaria Moritz ex Prantl, Unters. Morph. Gefasskrypt. 

2: 110(1881) 76:55 

Cayo, T.E. Hawkins 1031 (MO 115 ); Stann Creek, W.A. Schipp 366 

(K!, MO 76 ' 115 , NY). 
Anemia speciosa C. Presl, Suppl. tent, pterid.: 89 (1846) 76:56 

Cayo, C.L. Lundell 6906 (MICH 28 , MO 76 " 5 , NY); Toledo, B.K. 

Hoist Qtal 35855 (MO 115 ). 
Lygodium heterodoxum Kunze, Farrnkrduter 2: 32 (1849) 37:56 

Cayo, A. Hughes 55 (BM-000531763! 22 ); Toledo, G. Davidse 

36891 (BM-000531988!, MO 115 ). 
Lygodium venustum Sw. in/. Bot. 1801(2): 303 (1803) 37;57 

Belize, P.P. Barlee s.n. (K!); Cayo, A. Hughes 16 (BM- 

000531764! 22 , BM-000531880!); Corozal, M.J. Balick et al. 

2190 (MO 115 ); Orange Walk, C. Whitefoord 8054 (BM- 

000543272!, F); Stann Creek, G.R. Proctor 36566 (MO 115 ); 

Toledo, W.A. Schipp 328 (MO 76 ' 115 ). 
Lygodium volubile Sw. in /. Bot. 1801(2): 304 (1803) 37:57 

Cayo, C.L. Lundell 6613 (MICH 28 , NY); Stann Creek, D.R. Hunt 

355 (BM-000543271 !); Toledo, B.K. Hoist et al. 35597 (MO 115 ). 
Schizaea elegans (Vahl) Sm. in Mem. Acad. Roy. Sci. (Turin) 

5(1790-1791): 419 (1793) 103:57 

Stann Creek, PH. Gentle 3487 (MEXU 103 - 115 , NY, US); Toledo, 

E.J.F. Campbell 10 (K!). 
Schizaea poeppigiana J.W. Sturm in Mart., Fl. bras. 1(2): 181 

(1859) 103:57 

Toledo, G. Davidse & A.E. Brant 32268 (MO 103 ' 115 ). 

SELAGINELLACEAE 

Selaginella apoda (L.) Spring in Mart., Fl. bras. 1(2): 119(1 840), as 
'Selaginella apus' 

Toledo, C. Whitefoord 3300 (BM-000543202!). 
Not recorded for Belize by Fraile in Flora Mesoamericana 
(Fraile et al., 1995: 29) but present in Chiapas and Guatemala. 
The specimen listed here was identified by Fraile in 1987 yet not 
included in the flora; further investigation is required. 

Selaginella cladorrhizans A. Braun in Ann. Sci. Nat., Bot. ser. 5, 3: 
282(1865) 17:3 

Cayo, H.H. Bartlett 11457 (BM 17 ' 115 ); Toledo, C. Whitefoord 
8359(BM-000543203!). 

Selaginella diffusa (C. Presl) Spring in Bull. Acad. Roy. Sci. Bruxelles 
10: 143(1843) 17:26 
Cayo, B.H. Allen 15208 (BM-000543206!, MO 3 ' 17 ' 115 ). 

Selaginella eurynota A. Braun in Ann. Sci. Nat., Bot. ser. 5, 3: 293 
(1865) 

Toledo, G. Davidse 35970 (MO 115 ), G.R. Proctor 36009 (BM- 
000543200!). 

This Central American endemic was not recorded by Fraile for 
Belize in Flora Mesoamericana (Fraile et al., 1995: 26) but 
material was collected under this name in 1976 (G.R. Proctor 
36009, BM!) and a subsequent collection in 1996 (Davidse 
35970, MO) was determined as this taxon by A.R. Smith in 1997. 

Selaginella flagellata Spring in Bull. Acad. Roy. Sci. Bruxelles 10: 
228 (1843) 

Toledo, B.K. Hoist et al. 5522 (MO 115 ). 

Fraile (Fraile et al., 1995: 32) did not record this taxon for Belize 
in Flora Mesoamericana, though it was listed from Chiapas and 
Guatemala. It is included here on the basis of a specimen col- 
lected from Toledo District in 1997 and determined by A.R. 
Smith in the same year. 



Selaginella guatemalensis Baker in J. Bot. 21: 243 (1883) I7:3 

Toledo, C. Whitefoord 1664 (BM-000543249! 17 ' 115 , NY). 
Selaginella harrisii Underw. & Hieron. in Urb., Symb. antill. 7: 162 

(1912) 17:33 

Fraile (Fraile et al., 1995: 33) did not cite a specimen from Belize 

but referred to extralimital distribution in the Pteridophyte flora 

ofOaxaca (Mickel & Beitel, 1988: 341). Specimens need to be 

located to confirm the record. 
Selaginella hoffmannii Hieron. in Hedwigia 41: 41 (1902) 17:33 

Cayo, C. Whitefoord 2059 (BM-000543204!). 
Selaginella huehuetenangensis Hieron. in Hedwigia 43: 32 (1904) 17;34 

Belize, C. Whitefoord 2492 (BM-000543152!); Cayo, H.H. 

Bartlett 13032 (BM-000543154!); Toledo, M.E. Peck 634 (BM- 

000543155!). 
Selaginella idiospora Alston in Bull. Brit. Mus. (Nat. Hist.), Bot. 1: 

246, t. 6(1955) 17:34 

Cayo, C. Whitefoord 1253 (BM-000543207! 17 - 1!5 ); Toledo, G. 

Davidse 36100 (MO 115 ). 
Selaginella microdendron Baker in /. Bot. 23: 116 (1885) 17:35 

Stann Creek, W.A. Schipp 99 (BM-000543205! 17 ' 115 ); Toledo, 

J.R. Wiley 356 (MO 115 ). 
Selaginella mollis A. Braun in Ann. Sci. Nat., Bot. ser. 5, 3: 276 

(1865) I7:36 

Belize, C. Whitefoord 2765 (BM-000543201!); Toledo, W.A. 

Schipp 925 (NY). 
Selaginella ovifolia Baker in J. Bot. 22: 90 (1884) 17:37 

Toledo, W.A. Schipp 924[a] (BM 17 - I15 ). 
Selaginella pallescens (C. Presl) Spring in Mart., Fl. bras. 1(2): 132 

(1840) 17:37 
Selaginella pallescens var. acutifolia Stolze in Amer. Fern 7. 71: 51 

(1981) 17:37 

Cayo, C. Whitefoord 1255 (BM-000543211! 17 ' 115 ). 
Selaginella pallescens var. pallescens 11 37 

Belize, T.B. Croat 23835 (NY); Cayo, C. Whitefoord 1941 (BM- 

000543212! 17 - 115 ); Toledo, B.K. Hoist 3881 (MO 91 - 115 ). 
Selaginella sertata Spring in Mem. Acad. Roy. Soc. Belgique 24: 104 

(1 849) 17: 28 

Belize, P.H. Gentle 1396 (BM 17 , NY); Cayo, C. Whitefoord 1091 
(BM-000543209!); Orange Walk, C.L. Lundell 394 (NY); To- 
ledo, B.K. Hoist 4341 (MO 91 - 115 ). 

Selaginella silvestris Aspl in Ark. Bot. 20A(7): 30, f. 3-5 (1926) 
Cayo, G.R. Proctor 30105 (BM-000543226!); Toledo, C. 
Whitefoord 1983 (BM-000543213!, NY). 
Not recorded for Belize by Somers and Moran in the published 
version of Flora Mesoamericana (Fraile et al., 1995: 28), though 
many specimens have been collected recently from Toledo Dis- 
trict and material from as early as 1969 (G.R. Proctor 30105, 
BM!) is known from the country. 

Selaginella stellata Spring in Flora 21: 194 (1838) 17:28 

Stann Creek, WA. Schipp 52 (BM-000543224!, MO 17 , NY); 
Toledo, C. Whitefoord 1776 (BM-000543223!). 

Selaginella umbrosa Lem. ex Hieron. in Engler & Prantl, Nat. 
Pflanzenfam. 1(4): 683, f. 404 (1901) 17:41 
Belize, P.H. Gentle 1552 (K, NY); Cayo, D.J. Lewis 43 (BM- 
00054322 1 !); Orange Walk, C.L. Lundell 395 (NY); Stann Creek, 
W.A. Schipp 51 (BM-000543216!, NY); Toledo, C. Whitefoord 
1851 (BM, MEXU 17 , NY). 

TECTARIACEAE 

Ctenitis equestris (Kunze) Ching in Sunyatsenia 5(4): 250 (1940) 54: 

197 

Ctenitis equestris var. equestris 54 197 



RECENT RECORDS OF PTERIDOPHYTES 



93 



Stann Creek, W.A. Schipp 276 (BM-000543331 !, BM- 

000543332!, BRH 125 , MO 54 ' 115 ). 
Ctenitis excelsa (Desv.) Proctor in Rhodora 63: 34 (1961) 54: 198 

Cayo, J.D. Dwyer 11200 (MO 54 ' 115 ); Toledo, G. Davidse 36202 

(MO 115 ). 
Ctenitis interjecta (C. Chr.) Ching in Sunyatsenia 5(4): 250 (1940) 54: 

198 

Toledo, T.B. Croat 24223 (MO 54 ' 115 ). 
Ctenitis melanosticta (Kunze) Copel., Gen. fil. (Ann. Cryptog. 

Phytopathol. 5): 124 (1947) 54: '" 

Cayo, T.B. Croat 23320 (MO 54 - l15 ); Toledo, B.K. Hoist et al. 

35883 (MO 115 ). 
Ctenitis nigrovenia (H. Christ) Copel., Gen. fil. (Ann. Cryptog. 

Phytopathol. 5): 124 (1947) 54:199 

Cayo, D.A. Sutton et al. 7 (BM-000543335!, MO 54 - l15 ). 
Ctenitis salvinii (Baker) Stolze inAmer. Fern J. 67: 43 (1977) 54: '" 

Toledo, B.K. Hoist 3890 (MO 91 - 115 ), WA. Schipp S-773 (GH 54 

115 ). 
Cyclopeltis semicordata (Sw.) Sm. in Bot. Mag. 72(Companion): 36 

(1846) 55:20 

Orange Walk, M.J. Balick 3222 (BRH 125 , NY); Toledo, J.D. 

Dwyer 9921 (MO 55 - 115 ). 
Dictyoxiphium panamense Hook., Gen. fil: 62 (1840) 56:201 

Stann Creek, W.A. Schipp 228 (BRH 125 , NY); Toledo, G. Davidse 

&A.E. Brant 32211 (MO 56 - 115 ). 
Lastreopsis effusa (Sw.) Tindale in Victoria Naturalist 73: 184 

(1 957)57:201 

Lastreopsis effusa subsp. divergens (Willd. ex Schkuhr) Tindale in 
Contr. New South Wales Natl. Herb. 3: 299, t. 21 (1965) 57: 201 
Cayo, A. Hughes 133 (BM-000557727! 22 ); Toledo, T.B. Croat 
24463 (MO 57 ' 115 ). 

Lastreopsis exculta (Mett.) Tindale in Victoria Naturalist 73: 185 
(1957) 

Lastreopsis exculta subsp. exculta 

Toledo, G. Davidse 35648 (MO 115 ), G. Davidse 35799 (MO 115 ), 
G. Davidse 36208 (MO 115 ). 

Recorded from Chiapas, Honduras and Guatemala in Flora 
Mesoamericana (Moran, 1995w: 202), but not from Belize. 
Included here on the basis of material collected from Toledo 
District and identified as L. exculta (Mett.) Tindale by A.R. 
Smith in 1997. Further material from Belize was identified as L. 
exculta subsp. guatemalensis (Baker) Tindale, though this name 
is included as a synonym of subsp. exculta by Moran. 

Megalastrum lunense (H. Christ) A.R. Sm. & R.C. Moran in Amer. 
Fern J. 77: 128(1987) 
Toledo, B.K. Hoist 5820 (MO 115 , UC). 

This Central American endemic was not recorded for Belize in 
the published version of Flora Mesoamericana (Smith & 
Moran, 19950: 203), but included in the Internet version on the 
basis of a collection from Toledo District (B.K. Hoist 5820, UC) 
made in February 1997 and identified by A.R. Smith later the 
same year. This is the first record of the genus from Belize. 

Tectaria heracleifolia (Willd.) Underw. in Bull. Torrey Bot. Club 33: 
200(1906) 58:207 

Tectaria heracleifolia var. heracleifolia 20 

Belize, C. Whitefoord 1163 (BM-000543323!); Cayo, C. 
Whitefoord 2034 (BM, MO 58 - " 5 ); Orange Walk, C.L. Lundell 338 
(NY); Toledo, G. Davidse & D.L. Holland 36495 (BM- 
000531986!, MO 115 ). 

Tectaria incisa Cav., Descr. pi: 249 (1802) 58:207 

Cayo, D.L. Spellman & W.W. Newey 1849 (MO 58 - l15 ); Stann 
Creek, W.A. Schipp 272 (BM-0005433 15!, BRH 125 , NY); Toledo, 
C. Whitefoord 1583 (BM-0005433 16!). 



Tectaria mexicana (Fee) C.V. Morton in Amer. Fern J. 56: 133 

(1 966) 58:207 

Belize, P.H. Gentle 1551 (K, NY); Cayo, A. Hughes 51 (BM- 
000531 775 ! 22 , BM-000531780!); Toledo, G. Davidse & A.E. 
Brant 32371 (MO 58 ' 115 ). 

Tectaria nicotianifolia (Baker) C. Chr., Index filic., Suppl. 3: 182 
(1934) 

Toledo, G. Davidse & D.L Holland 36603 (UC 115 ). 
Not recorded for Belize in Flora Mesoamericana (Moran, 1 995 v: 
208), but present in Guatemala, Honduras and from Nicaragua to 
Panama. Included in the Internet version based on the specimen 
from Toledo District listed above collected in 1997 and deter- 
mined by A.R. Smith. 

Tectaria pilosa (Fee) R.C. Moran in Novon 2: 138 (1992) 
Toledo, G.R. Proctor 36155 (BM-0005433 14!). 
Only recorded for Costa Rica and Panama in Flora 
Mesoamericana (Moran, 1995v: 208), T. pilosa is listed here 
based on a collection from high forest in the Columbia Forest 
Reserve of Toledo, originally identified as T. incisa var. pilosa by 
G.R. Proctor. It differs from T. heracleifolia in the oblique or 
decurrent base to the terminal segment of the lamina and rather 
sparsely pilose upper and lower faces. Further investigation of 
this material is required. 

Tectaria plantaginea (Jacq.) Maxon in Contr. U.S. Natl. Herb. 10: 
494(1908) 58:208 
Stann Creek, W.A. Schipp 465 (BRH 125 , MO 58 - " 5 , NY). 

Tectaria rivalis (Mett. ex Kuhn) C. Chr., Index filic., Suppl. 3: 184 

(1 934)58: 208 

Cayo, T.E. Hawkins 1273 (MO 115 ); Stann Creek, W.A. Schipp S- 
66 (BM-000543327!, F 58 - 115 , NY). 

Tectaria vivipara Jermy & T.G. Walker in Bull. Brit. Mus. (Nat. 
Hist.), Bot. 13: 274, f. 15 (1985) 58:209 

Belize, P.H. Gentle 1542 (K, MO 58 " 5 , NY); Toledo, C. Whitefoord 
1583 (BM, NY). 

THELYPTERIDACEAE 

Macrothelypteris torresiana (Gaud.) Ching in Acta Phytotax. Sin. 

8:310(1963) 

Cayo, A. Hughes 143 (BM-000531924!, BM-000557726! 22 ); 

Toledo, G. Davidse 35918 (MO 115 ). 

Not recorded for Belize in Flora Mesoamericana (Smith, 1995a: 

164). The species and genus were first listed for Belize by 

Hughes (1998) for the Chiquibul Forest Reserve and additional 

material is cited in TROPICOS based on identifications by A.R. 

Smith in 1997 and 1999. 
Thelypteris balbisii (Spreng.) Ching in Bull. Fan. Mem. Inst. Biol, 

Bot. 10:250(1941) 110:17 

Stann Creek, P.H. Gentle 2720 (NY); Toledo, A.H. Gentry 7900 

(UC 110 ). 
Thelypteris biolleyi (H. Christ) Proctor in Bull. Inst. Jamaica, Sci. 

Ser. 5: 58 (1953) 110 183 

Toledo, G. Davidse & A.E. Brant 32208 (MO 115 , UC 11 115 ). 
Thelypteris blanda (Fee) C.F. Reed in Phytologia 17: 264 (1968) 110: 

183 

Cayo, A. Hughes 113 (BM-000531751! 22 ); Toledo, M.E. Peck 

s.n. (US 110 - 115 ). 
Thelypteris decussata (L.) Proctor in Bull. Inst. Jamaica, Sci. Ser. 5: 

59(1953) 
Thelypteris decussata var. costaricensis A.R. Sm.in Univ. Calif. 

Publ. Bot. 76: 16(1980) 

Toledo, B.K. Hoist 5919 (MO 115 ). 

This Central American endemic was not recorded for Belize in 



94 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



the published version of Flora Mesoamericana (Smith, 1995ft: 
194), but was recorded from Honduras, Costa Rica and Panama. 
Belize was included in the Internet version based on a specimen 
collected from Toledo District listed here and identified by A.R. 
Smith in 1997. A second specimen (Hoist 5819, MO), identified 
by A.R. Smith as T. decussata (L.) Proctor, may also be referable 
to var. costaricensis. 
Thelypteris dentata (Forssk.) E.P. St. John in Amer. Fern J. 26: 44 

(1936)110:181 

Toledo, G. Davidse 36886 (MO 115 ), R. Rivero 2539 (UC). 
Not recorded for Belize in the published version of Flora 
Mesoamericana (Smith, 1995ft: 181) but included in the Internet 
version based on a collection (Rivero et al. 2539, UC) identified 
by A.R. Smith in March 1996. 
Thelypteris falcata (Liebm.) R.M. Tryon in Rhodora 69: 6 (1967)" 

192 

Stann Creek, W.A. Schipp 97 (BRH 125 , NY); Toledo, P.H. Gentle 

8693 (NY, UC"). 
Thelypteris ghiesbreghtii (Hook.) C.V. Morton in Contr. U.S. Natl. 

//<?rft. 38:45 (1967)" 0:186 

Toledo, W.A. Schipp S-935 (GH", NY). 
Thelypteris glandulosa (Desv.) Proctor in Rhodora 61: 306 ( 1 960)" 0: 

194 

Thelypteris glandulosa var. brachyodus (Kunze) A.R. Sm. in 

Phytologia 34: 233 (1976) 110: l94 

Toledo, WA. Schipp 362 (NY, UC 110 - 115 ). 
Thelypteris hispidula (Decne) C.F. Reed in Phytologia 17: 283 

(1968) ,,0:,81 

Belize, P.H. Gentle 1460 (K!); Cayo, A. Hughes 88 (BM- 

000531822!, BM-000531828!, BM-000531829!, 

BM-000557715! 22 ); Toledo, A.H. Gentry 7978 (MO" a " 5 , NY). 

Thelypteris hondurensis L.D. Gomez in Phytologia 50: 458 ( 1 982)" 

186 

Toledo, W.B. Crankshaw s.n. (CR lia " 5 , NY). 

Thelypteris kunthii (Desv.) C.V. Morton in Contr. U.S. Natl. Herb. 
38:53(1967) 110:I82 

Belize, T.B. Croat 23987 (UC" a 115 ); Cayo, A. Hughes 50a (BM- 
000531750! 22 , BM-000531773!); Toledo, B.K. Hoist 6020 
(MO 115 ). 

Thelypteris leprieurii (Hook.) R.M. Tryon in Rhodora 69: 6 (1967) 

Thelypteris leprieurii var. subcostalis A.R. Sm. in Univ. Calif. Publ. 
Bot. 76:26(1980) 

Toledo, G. Davidse & D.L. Holland36188 (MO 115 ), T.E. Hawkins 
1536 (MO 115 ). 

Not recorded for Belize in the published version of Flora 
Mesoamericana (Smith, 1995ft: 194), but included in the Internet 
version based on a subsequent collection from Toledo District 
(G. Davidse & D.L. Holland 36788, MO) identified by A.R. 
Smith in January 1998. 

Thelypteris linkiana (C. Presl) R.M. Tryon in Rhodora 69: 6 (1967) 
Toledo, D.L. Holland 29 (MO 115 , UC). 

Not recorded for Belize in the published verion of Flora 
Mesoamericana, (Smith, 1995ft: 174), but included in the Internet 
version based on a specimen collected subsequently from Toledo 
District (D.L. Holland 29, UC) and identified by A.R. Smith in 
December 1997. 

Thelypteris meniscioides (Liebm.) C.F. Reed in Phytologia 17: 292 
(1968) 

Toledo, C. Whitefoord 1518 (BM). 

Not recorded for Belize in Flora Mesoamericana (Smith, 1995ft: 
186-187) though both T. meniscioides var. meniscioides (with a 
pinnate lamina) and the Guatemalan endemic T. meniscioides var. 
ternata A.R. Sm. (with a ternate lamina) are known from Guatemala. 



Thelypteris nicaraguensis (E. Fourn.) C.V. Morton in Contr. U.S. 
Natl. Herb. 38:55(1967) 
Toledo, B.K. Hoist 4324 (MO 91 - 115 ). 

This Central American endemic was not recorded from Belize in 
Flora Mesoamericana (Smith, 1995ft: 187) although it was listed 
for Chiapas and Honduras. The material listed here, collected in 
1992, was identified by R.C. Moran under this name and cited by 
Parker et al. (1993: 41). The identity of this material requires 
confirmation. 

Thelypteris obliterata (Sw.) Proctor in Bull. Inst. Jamaica, Sci. Ser. 
5:62(1953) 110:187 

Stann Creek, W.A. Schipp 83 (BRH 125 , NY); Toledo, P.H. Gentle 
1458 (K, UC 110 ' 115 ). 

Thelypteris ovata R.P. St. John in Small, Ferns s.e. states: 230 

(193 0)110:182 

Thelypteris ovata var. lindheimeri (C. Chr.) A.R. Sm. in Amer. Fern 

J. 61:30 (1971) 110:182 

Cayo, P.H. Gentle 2323 (GH 110 115 , K!, NY). 
Thelypteris patens (Sw.) Small, Ferns s.e. states: 243 (1938) 110: 182 
Thelypteris patens var. patens 110 182 

Cayo, A. Hughes 36a (BM-000531757! 22 , BM-000531850!); 

Toledo, T.B. Croat 24272 (UC" a 115 ). 
Thelypteris patens var. smithiana Ponce in Darwiniana 28: 373 

(1987) 

Cayo, A. Hughes 85 (BM-000532010!). 

Central American material formerly identified as var. scabriuscula 

(C. Presl) A.R. Sm. was referred to var. smithiana by Smith 

( 1 995ft: 1 82) on the basis of a sudy by Ponce ( 1 987) who asserted 

that the type of var. scabriuscula was equivalent to T. patens var. 

patens. Material from Cayo District collected and identified as 

var. scabriuscula is included here in var. smithiana. 
Thelypteris paucipinnata (Donn. Sm.) C.F. Reed in Phytologia 17: 

302(1968) 110:188 

Cayo, A. Hughes 112 (BM-000531896!, BM-000531897!, BM- 

000557704! 22 ); Toledo, P.H. Gentle 6550 (US 110 ). 
Thelypteris poiteana (Bory) Proctor in Bull. Inst. Jamaica, Sci. Ser. 

5:63(1953) 110;188 

Cayo, P.H. Gentle 1457 (K, US lia 115 ); Orange Walk, C.L. Lundell 

406 (US); Toledo, C. Whitefoord 1640 (BM). 
Thelypteris praetermissa (Maxon) A.R. Sm. in Phytologia 34: 232 

(1976) .10:.88 

Belize, P.H. Gentle 9720 (US); Cayo, H.H. Bartlett 13104 
(MICH 29 ' lia " 5 , US- photograph lia 115 ); Stann Creek, T.B. Croat 
24548 (UC 110 - !15 ); Toledo, G. Davidse & M. Meadows 35842 
(MO 115 ). 

Based on a Belizean type (Bartlett 1 3 1 04, MICH), T. praetermissa 
forms putative hybrids with T. obliterata and several specimens 
from Belize are interpreted as of hybrid origin by Smith (1995ft: 
188). Amongst several paratypes listed by Maxon (1944a: 20) is 
another Belizean specimen from close to the type locality (H.H. 
Bartlett 11878, MICH), yet Smith interpreted a duplicate at US 
as a hybrid. 

Thelypteris reptans (J.F. Gmel.) C.V. Morton in Fieldiana, Bot. 28: 
12(1951) 110:189 

Cayo, T.B. Croat 23489 (UC 110 - 115 ); Toledo, B.K. Holstetal.5496 
(MO 115 ). 

Thelypteris resinifera (Desv.) Proctor in Bull. Inst. Jamaica, Sci. Ser. 
5:63(1953)" 0:177 

Cayo, D.R. Hunt 255 (US 1 . 10 - " 5 ); Stann Creek, P.H. Gentle 2759 
(US); Toledo, G. Davidse & A.E. Brant 31881 (MO 115 ). 

Thelypteris sancta (L.) Ching in Bull. Fan. Mem. Inst. BioL, Bot. 10: 
254(1941) 
Toledo, B.K. Hoist 5968 (MO 115 ), C. Whitefoord 1697 (BM). 



RECENT RECORDS OF PTERIDOPHYTES 



95 



Not recorded for Belize in the published version of Flora 
Mesoamericana (Smith, 19956: 178), though included in the 
Internet version based on a specimen collected by Hoist in 1997 
and determined by A.R. Smith in the same year. 
Thelypteris schippii (Weath.) A.R. Sm. in Phytologia 34: 233 

(1976) l.0:,89 

Toledo, W.A. Schipp S-782 (GH 108 - "- " 5 - l29 ). 
Endemic to Toledo District in Belize; the holotype at GH is 
incorrectly cited by Weatherby (1935: 52) and Smith (19816: 
505) as 'Schipp 8-782' which is a common error for Schipp 
specimens prefixed by 'S-'. The correct citation is 'S-782' as 
amended by Smith (19956: 189). 
Thelypteris skinneri (Hook.) C.F. Reed in Phytologia 17: 314 

(1968) 110:189 

Toledo, W.A. Schipp S-797 (GH 110 - 115 ). 

Thelypteris struthiopteroides (C. Chr.) C.F. Reed in Phytologia 17: 
316(1968) 

Stann Creek, W.A. Schipp 926 (BRH 125 ). 
Not recorded for Belize in Flora Mesoamericana (Smith, 1 9956: 
178), though known from Oaxaca to Guatemala and tentatively 
included here on the basis of a record in Vargas & Shawe (1997: 
48). Further investigation of the Schipp material at BRH or other 
duplicates is required to confirm the record. 

Thelypteris tetragona (Sw.) Small, Ferns s.e. states: 256 (1938)" 0: 189 
Cayo, P.M. Gentle 9021 (US 108 ' "- 115 ); Stann Creek, R. Rivero et 
al. 2530 (BRH 125 ); Toledo, G. Davidse 35647 (MO 115 ). 

Thelypteris toganetra A.R. Sm. in Amer. FernJ. 63: 118(1973) 110:19 
Toledo, B.D. Vanderveen 587 (UC 110 - 115 ). 

VITTARIACEAE 

Ananthacorus angustifolius (Sw.) Underw. & Maxon in Contr. Gray 

Herb. 10: 487 (1908)" : 148 

Cayo, A. Hughes 124 (BM!, BM-000557722! 22 ); Toledo, T.B. 

Croat 24380 (MO 46 - 115 ). 

As Vittaria costata Kunze in Flora Mesoamericana (Moran, 

1 995.;': 148), but treated here as a species of Ananthacorus 

following the recent revision by Crane (1997). 
Anetium citrifolium (L.) Splitg. in Tijdschr. Natuurl. Gesch. Physiol. 

7: 395(1 840) 4 * 145 

Toledo, B.K. Hoist 5940 (MO 115 ). 
Hecistopteris pumila (Spreng.) J. Sm. in London J. Bot. 1: 193 

( 1842 )45:148 

Stann Creek, WA. Schipp S-50 (US). 

Polytaenium cajenense (Desv.) Benedict in Bull. Torrey Bot. Club 
38: 169(1911) 45:146 

As Antrophyum cajenense (Desv.) Spreng. in Flora Mesoameric- 
ana (Moran, 1995/: 146) and cited for Belize on the basis of an 
extralimital distribution record in Flora of Chiapas (Smith, 
198 la: 33) though no supporting specimens are listed. The taxon 
is included here as a species of Polytaenium on the basis of the 
recent revision by Crane ( 1 997 : 5 1 6), though confirmation of the 
distribution from specimens is required. 

Polytaenium feei (W. Schaffn. ex Fee) Maxon in Sci. Surv. Porto 
Rico & Virgin Islands 6: 405 (1926) 

Cayo, C.L. Lundell 6223 (MICH 28 , US); Stann Creek, WA. 
Schipp S-70 (US); Toledo, P.M. Gentle 5020 (MO 45 - " 5 , US). 
Treated as Antrophyum lanceolatum (L.) Kaulf. in Flora Meso- 
americana (Moran, 1995/: 147), but included as a species of Poly- 
taenium here on the basis of the recent revision by Crane (1997). 

Polytaenium lineatum (Sw.) J. Sm. in J. Bot. (Hooker) 4: 68 (1841) 
Toledo, G. Davidse 36258 (MO 115 ), B.K. Hoist et al. 5402 
(MO 115 ). 



This taxon was treated as a species of Antrophyum by Moran 
( 1995/: 147) and not recorded from Belize, although it was listed 
for Chiapas and Guatemala. Stolze (1981: 442) gave the distri- 
bution as 'southern Mexico to Panama' in Flora of Guatemala 
but did not list supporting specimens, making this implicit refer- 
ence difficult to interpret. It is included here on the basis of 
material collected in Toledo District in 1996 and identified in 
1997 by A.R. Smith as Polytaenium lanceolatum (Sw.) Desv. 
The earlier name P. lineatum (Sw.) Sm. is used here following the 
revision of Vittariaceae by Crane (1997: 516). 

Radiovittaria stipitata (Kunze) E.H. Crane in Syst. Bot. 22: 515 
(1997) 

Toledo, B.K. Hoist 4Q1Q (MO 46 gi ),B.K. Holstetal 5501 (MO 115 ), 
B.K. Hoist 5943 (MO 115 ). 

Listed for Belize in Flora Mesoamericana (Moran, 1995/: 150) 
as Vittaria stipitata Kunze, the genus Radiovittaria is adopted 
here following the revision of the Vittariaceae by Crane (1997: 
515). 

Scoliosorus ensiforme (Hook.) T. Moore, Index fil. : xxix ( 1 857) 45: 147 
Toledo, F. Boutin & Schlosser 5116 (MO 45 - l15 ). 
Treated as a species of Antrophyum in Flora Mesoamericana 
(Moran, 1995/: 147), but included in Scoliosorus here following 
the recent revision by Crane (1997). 

Vittaria graminifolia Kaulf., Enum. filic.: 192 (1824) 46: 149 

Cayo, T.B. Croat 23767 (MO 46 115 ); Toledo, B.K. Hoist 3876 
(MO 91 - 115 ). 

Vittaria lineata (L.) Sm. in Mem. Acad. Roy. Sci. (Turin) 5(1790- 
1791):421,t. 9,f. 5(1793) 46:149 

Cayo, C.L. Lundell 6493 (MICH 28 ); Stann Creek, WA. Schipp 
152 (BM-000543175!, US); Toledo, J.D. Dwyer & R. Coomes 
12992 (MO 46 - 115 ). 

WOODSIACEAE 

Athyriumfllix-femina (L.) Roth, Tent. fl. Germ. 3: 65 (1799) 6 * 228 
Toledo, sine leg. 2 (US). 

While there is a doubt concerning the exact provenance of the 
material listed here, A. filix-femina is recorded from Chiapas, 
Guatemala and El Salvador. Moran (1995ac: 228) did not cite a 
specimen but referred to Flora of Guatemala where Stolze ( 198 1 : 
96) made explicit reference to 'British Honduras' under the 
synonym A. dombei Desv., but again without supporting material. 

Diplazium cristatum (Desr.) Alston in J. Bot. 74: 173 (1936) 1:23 
Toledo, P.H. Gentle 7357 (BM-000543334! 1 - ll5 ). 

Diplazium franconis Liebm.in Kongel. Danske Vidensk. Selsk. Skr., 
Naturvidensk. Math. Afd. ser. 5, 1(1): 256 (1849) 
Toledo, G. Davidse & D.L. Holland 36602 (MO 115 ), G. Davidse 
36869 (MO 115 , UC). 

Not recorded for Belize in the published version of Flora 
Mesoamericana (Adams, 1995a: 235) though listed for Chiapas, 
Guatemala and Honduras. It was included in the Internet version 
on the basis of material collected in 1997 (Davidse & Holland 
36869, UC) and identified by A.R. Smith. 

Diplazium grandifolium (Sw.) Sw. in 7. Bot. (Schroder) 1800(2): 62 
(1801) 1:236 
Toledo, WA. Schipp S-783 (BR 1 - 115 , NY). 

Diplazium neglectum (H. Karst.) C. Chr., Index filic.: 236 (1905)' 2V 
Stann Creek, WA. Schipp 535 (BM 1 , BRH 125 ). 

Diplazium plantaginifolium (L.) Urb., Symb. antill. 4: 31 (1903)' * 
Cayo, D.R. Hunt 602 (BM); Stann Creek, P.H. Gentle 8205 (BM 1 
115 ); Toledo, B.K. Hoist et al. 5506 (MO 115 ). 

Diplazium riedelianum (Bong, ex Kuhn) Kuhn ex C. Chr., Index 
filic.: 230 (1905) 1:241 



96 

Toledo, B.K. Hoist 4317 (MO 1 9I 115 ). 
Diplazium striatum (L.) C. Presl, Tent, pterid.: 114 (1836) 1:243 

Toledo, B.K. Hoist 4473 (MO 1 - 91 - 115 ). 
Diplazium urticifoliumH. Christ, Prim.fl. Costaric. 3(1): 29(1901) 1: 

245 

Adams (1995a: 245) did not list a specimen from Belize, refer- 
ring instead to Flora of Guatemala where Stolze (1981: 197) 
explicitly cited 'British Honduras' but without material. There 
may be supporting specimens at F. 
Diplazium werckleanum H. Christ in Bull. Herb. Boiss. ser. 2, 4: 969 

(1904 )l:246 

Toledo, F. Boutin & Schlosser 5052 (MO 1 115 ). 
Hemidictyum marginatum (L.) C. Presl, Tent, pterid. : 1 1 1 , t. 3, f. 24 
(1836) 

Standley & Record (1936: 63) included this taxon without a 
specimen reference. Stolze (1981: 271) gave the general distri- 
bution as 'southern Mexico to Panama' which may be taken as an 
implicit record for Belize, though without supporting specimens 
this is difficult to interpret. It was not listed for Belize in Flora 
Mesoamericana (Moran 1995ac: 246) though recorded from all 
surrounding countries. The only species of the genus widespread 
in the neotropics, H. marginatum is a robust plant unlikely to be 
misidentified as it is easily distinguished by the linear sori and 
venation which is anastomosed in the distal third. It is to be 
expected in Belize and there may be material seen by Standley at F. 



DISCUSSION 

To date, at least 319 species (using currently accepted taxonomy) of 
pteridophytes have been recorded for Belize. Put in context, Belize 
has almost half of the total of 652 species listed by Stolze (1983: 1) 
for the much larger and topographically diverse neighbouring coun- 
try of Guatemala. Compared to the United States, Belize has half as 
many species again, yet the United States has approximately 400 
times the land area. Despite its small size and lack of the high 
elevations found throughout much of Central America, Belize argu- 
ably has one of the richest pteridophyte floras in the world on an area 
basis. 

Data presented in this paper include conventional sources, par- 
ticularly herbaria and published literature, but also many digital 
sources (Web, CD-ROM, databases) for names, specimens, litera- 
ture and related information. Development of the World Wide Web 
version of Flora Mesoamericana provides an unparalleled opportun- 
ity to assess the dynamic changes to a tropical flora. Increasingly 
information on species distribution becomes available on the Internet 
long before it appears in published literature and it may be available 
several years earlier. In order to resolve the complex and time- 
consuming task of searching the Internet, a web-enabled information 
system has been built by the senior author. This embodies design 
aspects of a conventional botanical database, a management infor- 
mation system, a metadatabase and web browser, to locate new 
records semi-autonomously and track changes in known records 
from Internet sources on a regular basis. Using all of the available 
published citations, Internet sources, determinations and herbarium 
records, an analysis of the growth of knowledge over time of the 
pteridophyte flora of Belize is shown in Fig. 2. 

The graph covers records built up over nearly two centuries. 
During the nineteenth century there were very few relevant publica- 
tions, so all data are presented as a single datum point on the graph 
to provide a starting point. Data for the twentieth century are 
summarized for each decade with an error bar to indicate annual 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 

variation within the decade. An expected shape for the graph would 
be a curve flattening out and approaching an asymptote as all species 
for the area were discovered. In contrast, a trend line shows that the 
rate of discovery of species is still continuing to rise steeply. The 
number of pteridophyte taxa recorded for Belize has risen by 
approximately 20% since the publication of the most recent volume 
of Flora Mesoamericana, demonstrating that the flora of this coun- 
try has not been studied adequately. This percentage increase is 
similar to that for the flora taken as a whole, indicating that the 
deficiency is with collecting in general, rather than just pteridophytes. 

A few of the new records listed here represent widespread, 
distinctive taxa and have been known for a long time from Belize, 
but appear in relatively obscure or overlooked literature. But most 
result from recent collections and it is apparent that the wetter areas 
of Belize, particularly towards the higher reaches of the Maya 
Divide in Toledo and Cayo (Fig. 1), are yielding most of the new 
records for the country. However, much of the Maya Divide is 
unexplored, particularly from the Cayo side. Some vegetation classes 
such as the Liquidambar forest of the higher elevations of the Maya 
Mountains and features such as the karst sinkholes in the Chiquibul 
Forest Reserve have had virtually no collecting. There has been a 
strong historical bias towards collections from southern Belize, 
particularly as the two most famous collectors of the Belizean flora, 
William A. Schipp and Percy H. Gentle, were based mainly in the 
south. Approximately 80% of the pteridophyte specimens examined 
come from the southern districts of Toledo, Stann Creek and Cayo 
and, of these, over half come from Toledo District. Very few 
pteridophytes have been collected from Corozal District. It is to be 
expected that a greater number of pteridophytes will be present on 
the more impervious shales and granite of the south than the 
limestone plateau of the north. The north-south gradient in rainfall 
undoubtedly exacerbates this effect, with almost three times as 
much rain falling in the far south as in the north of the country. 
However, it is clear that collecting in the more agriculturally devel- 
oped north of the country has received far less effort and many more 
species are to be expected from this area, as well as from the 
floristically richer unexplored areas of the south. 

There is also a temporal element to past collecting effort. An 
analysis by altitude and month reveals that very few collections have 
been made from upland areas during the wet season (especially June 
and July), presumably due to the difficult access. Collecting fre- 
quency is greatest in the dry season (mainly February to April), 
when many pteridophytes are sterile. Many species of pteridophytes 
widespread in Central America are currently missing from the list 
for Belize. While the country does not have the high elevations of 
most of its neighbours, it is evident that this list of native ferns and 
fern allies is far from complete and the total will increase substan- 
tially with focussed collecting. 

Other taxa tentatively recorded for Belize in the literature and 
collections, but with insufficient information at present include: 
Adiantum andicola Liebm, A. urophyllum Hook., Anemia phyllitidis 
(L.) Sw., Asplenium munchii A.R. Sm., A. radicans L., Diplazium 
lonchophyllum Kunze, Elaphoglossum setigerum (Sodiro) Diels, E. 
vestitum (Schltdl. & Cham.) T. Moore, Lomariopsis fendleri D.C. 
Eaton, L. sorbifolia (L.) Fee, Loxogramme mexicana (Fee) C. Chr., 
Polypodium murorum Hook., Tectaria trifoliata (L.) Cav., Thelypteris 
interrupta (Willd.) K. Iwats., T. parasitica (L.) Tardieu, T. scalaris 
(H. Christ) Alston, and T. serrata (Cav.) Alston. 

This paper is the first in a series arising from research on Belizean 
floristics, and updates will appear on The Natural History Museum 
web site (http://www.nhm.ac.uk/) as specimens, publications and 
citations are added. For the first time, the Belizean collections held 
at The Natural History Museum are being located in the herbaria and 



RECENT RECORDS OF PTERIDOPHYTES 

100 

% taxa first recorded in each decade 
90 -I K>- cumulative % taxa recorded by each decade 

trendline (polynomial: y = 2.6442x' 5763 ) 
80 



70 



60 



50 



40 



30 



20 



10 



97 




1800- 
1890 



1900 



1910 



1920 



1930 



1940 



1950 



1960 



1970 



1980 



Checklists 



1990 



Internet 



decade 



World Wide Web 
databases 



Spellmanetal. (1975) 
Dwyer & Spellman (1981) 
Vargas & Shawe ( 1997) 



Regional floristics 



Botany of the Maya area (1940-1961) 
Flora of Guatemala (1946-1985) 
Flora Neotropica (1968-) 
Flora Mesoamericana (1994-) 



National floristics 

Standley & Record (1936) 



Preliminary checklists 

Campbell (1899) 



Fig. 2 Growth of knowledge of the pteridophyte flora of Belize. 



98 



D.A. SUTTON, A. HUGHES AND B. BULMER-THOMAS 



documented. The project currently seeks to collate basic informa- 
tion for vascular plant taxa from Belize, supported by relevant 
specimens and literature, and some 8000 names, 7000 references 
and 32 000 specimens are in the database at the time of submission 
of the paper. It is estimated that minimal information on at least 
45 000 specimens from Belize is comparatively readily available in 
the public domain, and that 75-100 000 specimens have been 
collected from the country. This material forms the basis for the 
cumulative knowledge on the flora of Belize, and provides a basis 
for further research. 



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39. 1995c. Sticherus C. Presl. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 59-62. 

40. 1 995d. Cyathea Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 93- 

103. 

41. 1995e. Acrostichum L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

105-106. 

42. 1 995/ Pityrogramma Link. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 137-140. 

43. 1995g. Pteris L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 140- 

144. 

44. 1995/1. Anetium Splitg. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

145-146. 

45. 1995/. Antrophyum Kaulf. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 146-147. 

46. 1995/. Vittaria Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 148- 

150. 

47. 1 995k. Dennstaedtia Bernh. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 151-153. 

48. 1995/. Hypolepis Bernh. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

153-157. 

49. 1995m. Lindsaea Dryand. ex Sm. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 157-160. 

50. 1995n. Lonchitis L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 160. 

51. 1995o. Odontosoria Fee. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 160-161. 

52. 1995/7. Pteridium Gled. ex Scop. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 161-162. 

53. 1995<?. Saccoloma Kaulf. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 162-163. 

54. 1995r. Ctenitis (C. Chr.) C. Chr. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 196-200. 

55. 1995s. Cyclopeltis ). Sm. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 200-201. 

56. 1 995t. Dictyoxiphium Hook. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 201. 

57. 1995. Lastreopsis Ching. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 201-202. 

58. 1995v. Tectaria Cav. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

204-209. 

59. 1 995 w. Arachnoides Blume. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 210-211. 

60. 1 995x Didymochlaena Desv. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 212. 

61. 1995y. Olfersia Raddi. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

214. 

62. 1995z. Polybotrya Humb. et Bonpl. ex Willd. In G. Davidse et al. (Eds), 

Flora Mesoamericana 1: 216-218. 

63. 1995aa. Stigmatopteris C. Chr. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 225-226. 

64. 1995a6. Athyrium Roth. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

227-228. 

65. 1995ac. Hemidictyum C. Presl. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 246. 



RECENT RECORDS OF PTERIDOPHYTES 



99 



66. I995ad. Lomariopsis Fee. In G. Davidse et al. (Eds), Flora Mesoamericana 100. 

1: 283-284. 

67. \995ae. Blechnum L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 101. 

325-332. 

68. 1995a/ SalpichlaenaL Sm. In G. Davidse etal. (Eds), Flora Mesoamericana 102. 

1: 332. 

69. 1995ag. Microgramma C. Presl. In G. Davidse et al. (Eds), Flora 103. 

Mesoamericana 1: 339-340. 

70. \995ah. Niphidium J. Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 104. 

1:341. 

7 1 . 1 995ai. Pecluma M.G. Price. In G. Davidse et al. (Eds), Flora Mesoamericana 1 05 . 

1: 341-345. 

72. \995aj. Phlebodium (R. Br.) J. Sm. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 345-346. 106. 

73. \995ak. Polypodium L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 107. 

349-365. 

74. 1995a/. Pseudocolysis L.D. Gomez. In G. Davidse et al. (Eds), Flora 108. 

Mesoamericana 1: 365. 

75. 1995am. Salvinia S6g. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 109. 

396-397. 

76. & Mickel, J.T. 1995. Anemia Sw. In G. Davidse et al. (Eds), Flora 110. 

Mesoamericana 1: 53-56. 

77. & Smith, A.R. 1 995. Lellingeria A.R. Sm. et R.C. Moran. In G. Davidse et 111. 

al. (Eds), Flora Mesoamericana 1: 376-380. 

78. Zimmer, B. & Jermy, A.C. 1995. Adiantum L. In G. Davidse et al. (Eds), 112. 

Flora Mesoamericana 1: 106-117. 

79. Morris, D. 1883. The colony of British Honduras, its resources and prospects 113. 
with particular reference to its indigenous plants and economic productions. 
London. 114. 

80. Nauman, C.E. 1995. Nephrolepis Schott. In G. Davidse et al. (Eds), Flora 
Mesoamericana 1: 286-289. 115. 

81. Ollgaard, B. 1983. Lycopodiaceae. In Stolze, R.G., Ferns and fern allies of 
Guatemala. Part III. Marsiliaceae, Salviniaceae and the Fern Allies. Fieldiana, 116. 
Botany new sen, 12: 20-44. 

82. 1995a. Huperzia Bernh. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

6-18. 117. 

83. 1995/7. Lycopodiella Holub. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 18-20. 118. 

84. Pacheco, L. 1995a. Hymenophyllum Sm. In G. Davidse et al. (Eds), Flora 
Mesoamericana 1: 63-71. 

85. 19956. Trichomanes L. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

71-83. 120. 

86. 1995c. Adiantopsis Fee. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

106. 121. 

87. \995d. Neurodium Fee. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 

341. 122. 

88. Palacios-Rios, M. 1995o. Psilotum Sw. In G. Davidse et al. (Eds), Flora 
Mesoamericana 1: 3-4. 

89. \995b. Sphaeropteris Bernh. InG. Davidse etal. (Eds), Flora Mesoamericana 123. 

1: 103-104. 

90. 1995c. Oleandra Cav. In G. Davidse et al. (Eds), Flora Mesoamericana 1: 289- 

290. 124. 

91. Parker, T.A. HI, Hoist, B.K., Emmons, L.H. & Meyer, J.R. 1993. A biological 
assessment of the Colombia River Forest Reserve, Toledo District, Belize Conser- 1 25. 
vation International, RAP Working Papers 3. Washington. 

92. Perez-Garcia, B. 1995a. 2. Marattia Sw. In G. Davidse et al. (Eds), Flora 
Mesoamericana 1: 50-51. 126. 

93. I995b. 1. Cibotium Kaulf. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 86-87. 127. 

94. Ponce, M.M. 1987. Revision de las Thelypteridaceae argentinas. Darwiniana 28: 
317-390. 128. 

95. Price, M.G. 1983. Pecluma, a new tropical American fern genus. American Fern 
Journal!!: 109-116. 

96. Proctor, G.R. 1985. Ferns of Jamaica: a guide to the pteridophytes. London. 129. 

97 . Ranker, T.A. 1 995. Hemionitis L.InG. Davidse et al. (Eds), Flora Mesoamericana 

1: 131-133. 130. 

98. Record, S.J. 1925. Preliminary check list of British Honduras woods. Tropical 
Woods 1: 14-16. 

99. Riba, R. 1995o. Lophosoria C. Presl. In G. Davidse et al. (Eds), Flora 131. 
Mesoamericana 1: 85. 



\995b. Metaxya C. Presl. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 85-86. 

1995c. Alsophila R. Br. In G. Davidse et al. (Eds), Flora Mesoamericana 

1: 88-90. 

& Pacheco, L. 1995a. Actinostachys Wall, ex Hook. In G. Davidse et al. 

(Eds), Flora Mesoamericana 1: 52-53. 

1 9956. Schizaea Sm. In G. Davidse et al. (Eds), Flora Mesoamericana 

1:57. 

Rojas-Alvarado, A.F. 1996. Twelve new species of Elaphoglossum 

(Elaphoglossaceae) from Costa Rica and Panama. Brenesia 45-46: 7-26. 

Schipp, W.A. 1934. Flora of British Honduras. Price list of seeds & herbarium 

material from William A. Schipp. Stann Creek. British Honduras. Anno 1933- 

34. Stann Creek. 

Seymour, F.C. 1975. Polypodium in Nicaragua. Phytologia 31: 129-192. 

Smith, A.R. 198 la. Part 2. Pteridophytes. In Breedlove, D.E. (Ed.), Flora of 

Chiapas 2: 1-370. San Francisco. 

19816. Thelypteris Schmidel. In Stolze, R.G., Ferns and fern allies of 

Guatemala. Part II. Polypodiaceae. Fieldiana, Botany new ser., 6: 473-514. 

1995a. Macrothelypteris (H. Ito) Ching. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 164. 

19956. Thelypteris Schmidel. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 164-195. 

1995c. Micropolypodium Hayata. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 383-385. 

& Bishop, L.E. 1995. Enterosora Baker. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 372-375. 

& Moran, R.C. 1995a. Megalastrum Holttum. In G. Davidse et al. (Eds), 

Flora Mesoamericana 1: 202-204. 

19956. Terpsichore A.R. Sm. In G. Davidse et al. (Eds), Flora 

Mesoamericana 1: 385-392. 

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Bull. nat. Hist. Mus. Land. (Bot.) 30(2): 101-130 



Issued 30 November 2000 



Collections of flowering plants by Francis 
Buchanan-Hamilton from Nepal, 1802-1803 



J. ROBERT PRESS 

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD 

KRISHNA K. SHRESTHA 

Central Department of Botany, Tribhuvan University, Kirtipur, Kathmandu 



X/ 330 I 



CONTENTS 



Introduction 

Correct citation of Hamilton (formerly Buchanan) 

Buchanan-Hamilton's collections 



The catalogue 

Localities of collections by Buchanan-Hamilton in Nepal (1802-3) 
Unpublished material relating to Buchanan-Hamilton's collections 

References 

Index ... 



101 
102 
102 
103 
125 
125 
126 
126 



SYNOPSIS. The herbarium of Francis Hamilton (formerly Buchanan) is one of the most important historical collections relating 
to the flora of Nepal. Containing the earliest plant collections from that country, and made by one of the most accomplished 
botanists in the Indian Subcontinent at that time, it has considerable intrinsic interest and value. Moreover, along with Wallich 
material, Hamilton's collections formed the basis for Don's Prodromus florae nepalensis ( 1 825) in which many new species were 
described: thus they include numerous types. Only two extensive sets exist, the top set at the Linnean Society of London and the 
second set, the basis of Don's work, at The Natural History Museum. Small numbers of Hamilton's Nepalese sheets also occur 
elsewhere, e.g. at Kew, Edinburgh and Liverpool. This paper enumerates Hamilton's flowering plant collections from Central 
Nepal, as well as providing some background to, and an explanation of, the name changes undergone by the man himself. 



INTRODUCTION 



The man variously referred to as Buchanan, Hamilton or Buchanan- 
Hamilton was born in 1762 at Branziet (on the Bardowie estate, near 
Glasgow) in Scotland and christened simply Francis Buchanan. The 
causes of the confusion surrounding his name, and the correct 
citation of it are explained later in this paper; but for the sake of 
simplicity he is referred to throughout the text as Buchanan, the 
name he bore for the greater part of his life. His life and career are 
covered in detail by Prain (1905) and in admirable precis by 
Mabberley (1977). Only a very brief outline is given here to provide 
some necessary background and to explain in more detail events 
involving his Nepalese collections. 

Buchanan was a physician who already had made two, possibly 
three, voyages to the East Indies before joining the Honourable East 
India Company as an Assistant Surgeon on the Bengal Establish- 
ment based at Calcutta. This marked the beginning of an 18-year 
period in the Indian Subcontinent and surrounding regions during 
which he visited much of India as well as Burma, the Andaman 
Islands, areas such as East Bengal and Chittagong now in Bangla- 
desh and Nepal (Table 1). Despite his position as a medical officer, 
Buchanan was highly regarded as a collector and natural historian. 
He was an astute observer of all aspects of the region and had already 
undertaken several official economic surveys before embarking on 



his first attempt to visit the Kingdom of Nepal in 1 800. This embassy 
was aborted but a year later he left Calcutta on a second attempt to 
visit Nepal, travelling via Saran and Tirhoot and collecting en route. 
By late January of 1802 he had reached the Indian village of Dacca 
on the Nepalese border, but after several weeks had only progressed 
some two miles into Nepal. The expedition had to return to Indian 
territory for a brief period, probably for diplomatic reasons, but by 
April was once again in Nepal and travelling towards Kathmandu. 
On first entering Nepal, Buchanan had remarked (in a letter to his 
friend and mentor William Roxburgh) that the 'pestilential season is 
fast approaching'. Once in Kathmandu, he found himself with many 
professional (i.e. medical) duties, contributing no doubt to his 
opinion of 'the country being most unhealthy' and one which he 
wished to leave as soon as possible. 

Notwithstanding his initial opinion, Buchanan spent 14 months in 
and around Kathmandu recording information on all forms of 
natural resources, manufacturing, climate and the people them- 
selves, as well as sending seeds and living plants to the Botanic 
Garden in Calcutta. In addition to all of these, Buchanan made a 
large collection of dried plant specimens accompanied by many 
drawings (by Indian artists) and plant descriptions. Much of the 
information was obtained by his companion and employee Babu 
Ramajai Bhattacharji, a Brahmin from Calcutta who accompanied 
Buchanan on this and other expeditions. In March 1803 the expedi- 
tion left Kathmandu to return to to Calcutta. 



The Natural History Museum, 2000 



102 



J.R. PRESS AND K.K. SHRESTHA 



In 1 805 Buchanan returned to England on leave. While in London 
he gave to his friend J.E. Smith his large collection of drawings and 
manuscripts relating to Indian plants, which included a large number 
collected on the survey of Mysore (S. India) in 1800-1801 along 
with his entire collection of specimens from Nepal. According to 
Smith the whole collection represented some 1500 species but it is 
more likely to have been 1500 specimens. 

Buchanan's time in the Kathmandu valley was his only visit in 
Nepal but he was able to augment his knowledge and collections on 
two further occsions. By 1807 he had returned to Calcutta as 
Surveyor for the Honourable East India Company charged with 
detailing the climate, topography, agriculture, natural resources, 
trade, culture, politics and religion of the Bengal Presidency and 
those adjacent countries and minor states with which the Company 
had no regular contact. With regard to areas outside the Company's 
possessions, information had to be obtained indirectly, as Buchanan 
himself was specifically barred from entering them. This did not 
prevent him, when based at Nathpur near the Nepalese border during 
the rainy season of 1810, from dispatching local collectors across 
the border into Eastern Nepal to bring back specimens. 

In 1814, he was again near the Nepalese border, this time at 
Gorakhpur, when further data on Nepal was gathered. Presumably 
such information was obtained by questioning natives of the country 
or other subjects of the East India Company who had travelled or 
stayed in Nepal, as required by Buchanan's instructions from the 
Company's Court of Directors. Again, local collectors were employed 
to bring back collections. 

Shortly after this Buchanan's health began to decline and after a 
very brief tenure as Superintendant of the Calcutta Botanic Garden 
he prepared to return to England and eventual retirement in his 
native Scotland. Here he continued to arrange his papers and prepare 
material for publication but was hampered by lack of access to his 
collections which, as the property of the East India Company, 
remained in London. Indeed, he clearly felt that as well as failing to 
appreciate his own efforts on their behalf, the Company was cheat- 
ing the scientific world by withholding his collections from wider 
study, and in a letter dated 14 February 1817 to his successor in 
Calcutta, Nathanial Wallich, warned against finding himself [Wallich] 
in a similar situation 'The Court of Directors [of the East India 
Company] has indeed received my collection with such contempt 
and arrogance that I would neither ask nor receive any favour. . . My 
collection would have been received with the utmost thankfulness 
by the most learned bodies here and might have gratified several of 
the most distinguished. Do not therefore throw any of your pearls 
before swine but collect largely and keep your collections for the 
learned of your own country [Wallich was Danish], who I have no 
doubt will be thankful'. 

Despite any difficulties caused by lack of access to his collec- 
tions, Buchanan prepared for publication an account of Nepal 
(Hamilton, 1819) and in 1819 he was able to obtain from the East 
India Company a duplicate set of his 1807-1814 botanical collec- 
tions which he immediately began to catalogue. However, the Nepal 
material was not among them, having been given to J.E. (now Sir 
James) Smith in 1805, in whose collection they remained little 
regarded. In another letter to Wallich dated 16 October 1821, 
Buchanan wrote 'I ... rejoice at your good luck in having access to 
the treasures of Nepal. A great part of what I have done there has 
been in a sort lost as having been given to Sir J.E. Smith who is rather 
indolent and not likely to publish any considerable part of what he 
has. A Mr. Don, however, who lives with Mr Lambert, to whom I 
gave duplicates of the collection presented to Sir J.E. Smith, is 
engaged in publishing an account of them together with those which 
you have sent, and I believe has both abilities and industry to 



produce a very valuable work. Whether or not Sir J.E. Smith will 
allow him the use of my drawings and written descriptions I have not 
learned. Your offer of joining me in a work on Nepal is very 
flattering, but I have no intention of taking upon myself such a 
labour; indeed I have not a single note respecting any of the plants I 
brought with me from Nepal-Smith has the whole.' While Buchanan 
never published any work based on his Nepal collections, Don did 
complete and publish his account. 

On the death of his eldest brother in 1818 Buchanan succeeded to 
the family properties, part of which comprised his mother's estate of 
Bardowie. In order to benefit from this portion of his inheritance and 
comply with the legalities attached to it, he was required to adopt his 
mother's family name of Hamilton. It was as Francis Hamilton that 
he died in 1829 at the age of 67. 



CORRECT CITATION OF HAMILTON 
(FORMERLY BUCHANAN) 

The problem of how to refer to a man who began his life under one 
name and ended it under another has attracted different solutions. In 
a document submitted to support his (successful) claim to be the 
chief of the Buchanans of Buchanan, he styled himself Francis 
Hamilton Buchanan, an appellation followed by many authors 
referring to his zoological works. Other documents name him as 
Buchanan Hamilton, the name used by many botanical authors. 
Prain ( 1 905) stated that the correct citation was simply Hamilton and 
condemmed zoologists for their 'erroneous and unnecessary prac- 
tice' of using both names and botanists for following suit but in 
reverse! Of the more modern reference works, Stafleu & Cowan 
(1979) use the form Francis Hamilton (ne Buchanan) with the 
abbreviation F. Ham. while Brummitt & Powell (1992) prefer the 
name Buchanan-Hamilton with the abbreviation Buch.-Ham. Thus, 
as an author in literature he is Hamilton, but as an authority in plant 
names he is Buchanan-Hamilton. There appears no definitive usage, 
but for botanical works at least it seems best to refer to Buchanan- 
Hamilton as the name least likely to cause further confusion. 



BUCHANAN-HAMILTON'S COLLECTIONS 

As already explained, Buchanan's Central Nepalese material col- 
lected during 1802-1803 and including the associated drawings and 
manuscripts, were given by him to J.E. Smith. Smith described 33 
species based on these collections in Rees' Cyclopaedia (1802- 
1820) and 12 others in his own Exotic Botany (1804-1805) but 
otherwise made little or no use of them. The entire collection of 433 
specimens collected in Nepal (Kara et al., 1978) remained in his 
herbarium, eventually passing to the Linnean Society in 1829. 
Duplicates distributed from Smith's herbarium may include small 
numbers of Buchanan sheets. For example, 14 Buchanan specimens 
of flowering plants and three specimens of bryophytes are held at 
Liverpool Museum (LIV) (Sedgwick, pers. com.). 

Buchanan gave a duplicate set, as complete as he could achieve, 
to his contemporary A.B. Lambert, a British botanist living in 
London. In turn, Lambert made the material available to his assistant 
David Don who began studying it in 1 820. This material, together 
with specimens from Nepal collected and sent to Lambert by 
Wallich, together amounted to some 2000 specimens (Miller, 1970) 
and formed the basis for Don's Prodromus florae nepalensis (1825) 
which contained descriptions of many new species. For reasons 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 
Table 1 Some significant dates in the life of Buchanan-Hamilton. 



103 



15 February 1762 Born Branziet, Stirlingshire. Fourth son of Thomas Buchanan of Spittal and Elizabeth Hamilton of Bardowie. 

1 7797-1783 Studied medicine at Edinburgh University. J.E. Smith who later founded the Linnean Society was a contemporary and friend. 

1 785 First voyage to Bombay via Cape and Comoros. 

1 788?- 1 789 Second voyage to Philippines and Moluccas. 

1791-?? Possible third voyage, to East Indies (not confirmed). 

26 September 1 794 Appointed Assistant Surgeon for Honourable East India Company on Bengal Establishment. 

1795 Visit to Rangoon, the Andamans, Pegu and Ava. Plants collected and drawn during this period were described and published by Sir 

Joseph Banks. 

1796-1798 Posted to SE Bengal. Excursions to the Eastern Sundribuns. 

1798 Undertook three-month economic survey of Chittagong. 

1798-1800 Several trips to the Western Sundribuns: descriptions and drawings but no collections of plants. Mainly worked on Gangetic fishes. 

1 800 Nepal. Start date of expedition unknown but by February Buchanan was waiting to cross the Indian/Nepalese border. He did not enter 

Nepal on this occasion. 
1 800 Survey of the Raj of Mysore and parts of Malabar. 

1 802 Second attempt to reach Nepal. First entered the country on 2 March 1 802. 

1803 Expedition leaves Kathmandu on 18 March 1803, reaching Segouli on 28 March 1803. 

1 805 Returned to England. Entire collection of Nepal plants given to J.E. Smith. 

1 806 Returned to India and in 1 807 begins the most detailed survey of India ever made. 

1810 Near the Pernea/Nepal border during the rainy season, at Nathpur. 

1811 Buchanan's health begins to decline, especially during the period 1813-14. 
1814 Near the Nepalese border, this time at Gorakhpur. 

1814 Returned to Calcutta. 

1814 Appointed Superintendant of the Calcutta Botanic Garden as successor to Roxburgh and Colebrooke. 

1815 Left for England aboard the Marchioness of Ely on 23 March 1815. The care of the Calcutta Botanic Garden passed to Nathanial 
Wallich. 

1815 Returns in August to England and shortly afterwards to his native Scotland. 

1818 Succeeds to the family properties on death of his elder brother on 10 January 1818; by April has adopted his mother's name of 
Hamilton. 

1819 Prepared for publication an account of Nepal. In this year he also visited London and was able to obtain from the East India 
Company his botanical collections. 

1829 Died on 15 June 1829 at Leny, in Scotland, aged 67. 



which remain unknown, Don was apparently unable to consult 
either the full set in Smith's herbarium or Buchanan's manuscript 
descriptions, also in Smith's possession. The type specimens thus 
probably belong to Lambert's set, with those in Smith's herbarium at 
the Linnean Society presumably representing isotypes. 

On Lambert's death his herbarium was divided into lots and sold 
at auction. Lot 286, described in the sale catalogue as containing a 
large collection of about 500 species of plants from Nepal, Mysore 
and Malabar made by Hamilton was purchased by Robert Brown for 
the British Museum and incorporated into the herbarium there. 
According to annotations by James Britten in The Natural History 
Museum's copy of Don's Prodromus florae nepalensis at least 332 
sheets of flowering plants (together with 31 sheets of pteridophytes) 
in Lot 286 were collected in Central Nepal in 1 802-1 803. These are 
only a part of the material available to Don in Lambert's herbarium 
and not all of the Buchanan-Hamilton specimens cited in Don's 
Prodromus have found their way into the collections at The Natural 
History Museum. Only 285 of the original 332 BM specimens have 
been traced. In such a large herbarium it is quite possible that a 
number have been refiled (possibly under new names) and remain to 
be found. The task of recognizing Buchanan-Hamilton specimens is 
made harder by the fact that a number were remounted and the 
original labels (if present) were not retained in some cases. One 
hundred and ninety-two Buchanan-Hamilton Nepalese sheets are 
kept in the Smith herbarium at the Linnean Society. 

While working on his Prodromus, Don apparently sent duplicates 
from Lambert's set of Buchanan's 1802-03 specimens to various 
colleagues. They include Proctor, von Martius and de Jussieu; these 
sheets are now held in Oxford (OXF), Meise (BR) and Paris (P-JU) 
respectively. 

Although Buchanan stated on several occasions that the whole of 
his early Nepal collections were presented to Smith, the duplicate set 



provided to Lambert indicates that Smith did not receive every 
specimen. It is possible that the gift to Lambert also did not exhaust 
the material and that at least some further specimens remained in the 
East India Company herbarium. If so, these would have been 
disposed of in the same way as Buchanan's later collections. 

Buchanan's small 1810 and 1813-14 collections from Eastern 
Nepal formed part of the East India Company's herbarium and were 
distributed along with this material by Wallich. Thus, Buchanan 
specimens from these periods may be held in several institutes. The 
greatest number appears to be in the Wallich Herbarium at Kew (K- 
W) with a smaller number in Edinburgh (E). Specimens from this 
period are not dealt with in this paper. 



THE CATALOGUE 

The Catalogue which follows covers only those flowering plant 
specimens collected by Buchanan-Hamilton in Central Nepal dur- 
ing the period 1 802-1 803 and housed in the herbaria at The Natural 
History Museum (BM), the Smith herbarium at the Linnean Society 
(LINN-SM) and Liverpool (LIV). 

Choosing the most useful arrangement for the entries posed 
problems. We feel that the most likely starting point for users will be 
either literature citing a Buchanan-Hamilton specimen or informa- 
tion taken from the specimen itself. Therefore the entries are arranged 
alphabetically by family, genus and species in descending order by 
the name under which the specimen was first cited in published 
literature. Two families, Amaryllidaceae and Liliaceae, are used in 
their broad sense with genera assigned accordingly. This is done to 
keep the arrangement in line with that used in An enumeration of the 
flowering plants of Nepal (Kara, Steam & Williams, 1978; Kara & 
Williams, 1979; Kara, Chater & Williams, 1982), up to now the 



104 



J.R. PRESS AND K.K. SHRESTHA 



standard reference for Nepalese plants. The names of the segregated 
families to which these genera are now commonly assigned are 
indicated as appropriate. As Buchanan's collections are so closely 
associated with D. Don's Prodromus florae nepalensls (1825), this 
work is also cited when relevant. Where a specimen has not been 
cited in literature, the Buchanan-Hamilton name on the sheet is used. 
Accepted names are shown in bold and synonyms in italics. Where 
the name is a synonym the currently accepted name is provided. The 
latter are mostly taken from Annotated checklist of the flowering 
plants of Nepal (Press et al., 2000), an updated version of the earlier 
An enumeration of the flowering plants of Nepal (Hara et al. 1978, 
1979, 1982). Various names not found in either of these works have 
been provided by Dr Henry Noltie (pers. com.). The authors have 
been unable to examine the material in the Smith herbarium as this 
is currently undergoing conservation treatment at the National 
Museums and Galleries on Merseyside, Liverpool. A large propor- 
tion of these specimens lack an accepted name, an omission which 
we hope will be rectified by appropriate experts when the material 
is, once again, accessible. Buchanan-Hamilton manuscript names 
taken up by other authors are not repeated in the entries. Otherwise, 
each entry includes any additional Buchanan-Hamilton manuscript 
names (using Buchanan-Hamilton's spellings), together with the 
locality and collection date as provided by the label on the specimen. 
Where these are not known, the relevant data from the literature 
citation is given in quotation marks. The herbarium where the 
specimen is held is also shown; where material has not been located 
by us in any of the herbaria examined, this is indicated by the 
comment 'Specimen not found'. These include specimens which 
may have been seen at The Natural History Museum (BM) by 
authors of accounts for An enumeration of the flowering plants of 
Nepal (Hara et al. 1978, 1979, 1982) but which we have been unable 
to locate. A number of specimens from The Natural History Mu- 
seum are known or presumed to be among material on loan at the 
time of this study. Information from specimens on loan to Edinburgh 
has kindly been provided by Dr Noltie (pers. com.): such specimens 
are annotated as BM but without an exclamation mark as they have 
not been seen by the authors. The remainder are indicated by the 
comment 'Material on loan: not seen'. An explanatory note may be 
appended to an entry. 

Many of the specimens represent type material, particularly for 
Don names. The situation here is complicated by Don's propensity 
for coining superfluous names for taxa already validly published 
elsewhere and often cited as synonyms of his new name. While it 
seems that Don studied the specimens now held in BM, those in 
other herbaria may also be original elements. We are very aware of 
the risks of inadvertently lectotypifying specimens by careless use 
of terms. We have, therefore, chosen to refer to original material 
throughout as 'syntypes', even when is it clear that only a single 
specimen is being referred to. It is not our intention to publish any 
novel typifications here and no statements in this publication 
should be interpreted as effecting typification for any name. We 
feel it is more appropriate for authors engaged in more detailed 
taxonomic studies of particular groups to make any necessary 
lectotypifications. 



ACANTHACEAE 

Barleria cristata L., D. Don, Prodr.fl. nepal.: 119 (1825). 
Narainhetty, 26 August 1803 (BM!). 

The date on the specimen is incorrect, Buchanan having left 
Nepal in March 1803. In all probability it should read 26 August 
1802. 



Lepidagathis incurva [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

119(1825). 

Ruellia recurva Buch.-Ham., in sched. 
Narainhetty, 27 January 1803 (BM!-syntype). 

Ruellia capitata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 120 

(1825). 
'ad Narainhetty Nepalensium. Floret Octobri' (BM?-syntype). 

Material on loan: not seen. 
= Strobilanthes pentastemonoides (Nees) T. Anderson 

ACERACEAE 

Acer laurifolium D. Don, Prodr.fl. nepal.: 249 (1825), nom. superfl. 
Acer buzimpala Buch.-Ham., in sched. 
Narainhetty, 24 February 1802 (BM!). 
= Acer oblongum Wall, ex DC. 

ALISMATACEAE 

Sagittaria lappula D. Don, Prodr.fl. nepal: 22 (1825). 

Alisma lappula Buch.-Ham., in sched. 

'in Nepalia' [No original label]. (BM!-syntype). 

= Sagittaria guyanensis subsp. lappula (D. Don) Bogin 

AMARYLLIDACEAE s.l. 

Narcissus tazetta L., Buch.-Ham., in sched. 
Narainhetty, 22 September 1802 (Sheet 573.11 LINN-SM, micro- 
fiche!). 

(ALLIACEAE) 

Allium sulvia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 53 (1825). 
Suembu, 11 July 1802 (Sheet 583.38 LINN-SM, microfiche!- 

syntype). 
= Allium tuberosum Rottler ex Spreng. 

ANACARDIACEAE 

Dobinea vulgaris [Buch.-Ham. ex] D. Don, Prodr. fl. nepal. : 249 

(1825). 
Narainhetty, 15 September 1802 (BM!-syntype?). 

Although the locality matches that given in the protologue, the 
collection date is September while Don gives the flowering time as 
August, suggesting that he saw a different specimen. 

APOCYNACEAE 

Alstonia lucida D. Don, Prodr.fl. nepal: 131 (1825). 

Echites triangularis Buch.-Ham., in sched. 

Sembu, 18 May 1802 (BM!-syntype; Sheet 444.18 LINN-SM- 

syntype, microfiche!). 
= Trachelospermum lucidum (D. Don) K. Schum. 

Echites fragrans Buch.-Ham., in sched. 

Suembu, 5 May 1802 (Sheet 444.20 LINN-SM, microfiche!). 

= Chonemorpha fragrans (Moon) Alston 

Echites tomentosa Buch.-Ham., in sched. 

Sembu, 7 July 1802 (Sheet 444.19 LINN-SM, microfiche!). 

Nerium niveum Buch.-Ham., in sched. 
Norcotera, 26 March 1802 (LINN microfiche!). 
= Vallaris solanacea (Roth) O. Kuntze 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 

AQUIFOLIACEAE 



105 



Ilex dipyrena Wall., D. Don, Prodr.fl. nepal.: 188 (1825). 
'in Nepalia' [No original label]. (BM!). 

Ilex odorata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 189 (1825). 
Chitlong, 12 April 1802 (BM!-syntype). 

Ilex rotunda sensu D. Don, Prodr.fl. nepal. : 1 89 ( 1 825), non Thunb 

(1784). 

Ilex saysia Buch.-Ham., in sched. 
Suemby, 9 May 1802 (BM!). 
= Ilex excelsa (Wall.) Hook.f. 

ARALIACEAE 

Hedera aculeata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 187 

(1825). 
'in Nepalia ad Narainhetty. Floret Octobri' (BM?-syntype). Material 

on loan: not seen. 
= Brassaiopsis aculeata (D. Don) Seem. 

Hedera elata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 187 (1825). 
Narainhetty, 13 November 1802 (BM!-syntype). 
= Schefflera elata (D. Don) Harms 

Hedera hainla [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 187 

(1825). 
'ad Narainhetty Nepaliae. Floret Februario' (BM?-syntype). Material 

on loan: not seen. 
= Brassaiopsis hainla (D. Don) Seem. 

Hedera helix sensu D. Don, Prodr. fl. nepal.: 187 (1825), non L. 

(1751). 

Narainhetty, 16 September 1802 (BM!). 
= Hedera nepalensis K. Koch 

Hedera parasitica [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 188 

(1825). 

Aralia parasitica Buch.-Ham., in sched. 
'in Nepalia ad Narainhetty. Floret Novembri' (BM?-syntype). 

Material on loan: not seen. 
= Pentapanax parasiticus (D. Don) Seem. 

Hedera tomentosa [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 187 

(1825), non Schefflera tomentosa (Seem.) Harms (1894). 
'in Nepalia' [No original label] (BM!-syntype). 
= Schefflera impressa (C.B. Clarke) Harms 

ASCLEPIADACEAE 

Ceropegia candelabrum L., Buch.-Ham., in sched. 

Narainhetty, 13 August 1802 (Sheet 405.2 LINN-SM, microfiche!). 

Cyananchus foetidum Buch.-Ham., in sched. 

Bassaria, 12 March 1802 (Sheet 454.17 LINN-SM, microfiche!). 

BALSAMINACEAE 

Impatiens odorata D. Don, Prodr. fl. nepal.: 203 (1825), nom. 

superfl. 

Balsamina odorata Buch.-Ham., in sched. 
'in Nepaliae ruderatis ad Narainhetty. Floret Septembri' (BM?). 

Material on loan: not seen. 
= Impatiens leptoceras DC. 



Impatiens racemosa D. Don, Prodr.fl. nepal.: 203 (1825), non DC. 

(1824), nom. superfl. 

Balsamina racemosa Buch.-Ham., in sched. 
'in ruderatis apricis Nepaliae ad Narainhetty. Floret Augusto' (BM?). 

Material on loan: not seen. 
= Impatiens insignis DC. 

BEGONIACEAE 

Begonia dioica [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 223 

(1825). 
'ad Narainhetty Nepaliae. Floret Augusto'. (Syntype: specimen not 

found). 

Begonia hatacoa [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 223 

(1825). 
Sembu, 8 July 1802 (BM!-syntype). 

Begonia picta Sm., Exot. hot. 2: 81, t. 101 (1805). 
Sembu, 21 July 1802 (BM!-syntype). 
The specimen is cited as holotype by Z. Badcock in sched. (1998). 

Begonia rubella [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 223 

(1825). 
Narainhetty, 10 September 1802 (BM!-syntype). 

BERBERIDACEAE 

Berberis aristata DC., Syst. nat. 2: 8 (1821). 

Berberis chitria Buch.-Ham., in sched. 

'in Napaulia'. (Syntype: specimen not found). 

The specimen appears also to be the basis for Berberis chitria 
Lindl. (see below). 

Berberis chitria [Buch.-Ham. ex] Lindl. in Bot. Reg. 9, t. 729 

(1823). 

Berberis paniculata Buch.-Ham., in sched. 
'Nepal'. (Syntype?: specimen not found). 
Chitlong, Upper Napaul, 10 April 1802 (Sheet 622.7 LINN-SM, 

microfiche!). 

Lindley coined this name in preference to B. aristata DC. and 
states that it is based on 'samples in the Lambertian Herbarium, 
collected by Dr. Hamilton in Nepal'. The names B. aristata and B. 
chitria are therefore based on the same type material. However, both 
taxa are currently accepted species. 

The sheet is annotated by C.K. Schneider 'Differt a B. aristata 
DC., inflores. compositis et a B. Chitria ramis grinis angularis 
glabris. Sp. nov. videter.' 

Mahonia napaulensis DC., Syst. nat. 2: 21 (1821). 

Berberis miccia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 205 
(1825), nom. superfl. 

Berberis pinnata Buch.-Ham., in sched.; Berberis miccia Buch.- 
Ham., in sched. 

Narainhetty, 15 November 1802 (BM!). 

Berberis miccia Buch.-Ham., in sched; Leontia fruticosa Buch.- 
Ham., in sched. 

Chitlong, 10 May 1802 (Sheet 622.10.1 LINN-SM, microfiche!). 

Narainhetty, 23 December 1802 (Sheet 622.9 LINN-SM, micro- 
fiche !-syntype). 
DeCandolle cites both Buchanan and Roxburgh sheets in Herb. 

Lambert with the locality given as Harain-Netty and flowering in 

December. No material dated December has been located at BM. 



106 
BETULACEAE 

AInus nepalensis D. Don, Prodr.fl. nepal: 58 (1825). 
Betula boshia Buch.-Ham., in sched. 
Narainhetty, 25 October 1802 (BM!-syntype). 

Betula alnoides [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 58 

(1825). 
'in sylvis ad Narainhetty Nepaliae superioris. Floret Octobri'. 

(Syntype: specimen not found). 

BORAGINACEAE 

Cynoglossum prostratum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

100(1825). 

'ad Baguanpur. Floret Martio'. (Syntype: specimen not found). 
= Bothriospermum zeylanicum (J. Jacq.) Druce 

Heliotropium obovatum [Roxb. ex] D. Don, Prodr.fl. nepal.: 101 

(1825). 
'versus ripas fluminis infra Morshidabad. Floret Aprili'. (Syntype: 

specimen not found). 
= Heliotropium ovalifolium Forssk. 

Although included by Don this specimen is not from Nepal but 
was collected in India. 

BURMANNIACEAE 

Burmannia disticha L., Buch.-Ham., in sched. 
Narainhetty, 15 August 1802 (Sheet 565.2.2 LINN-SM, micro- 
fiche!). 

BUTOMACEAE 

Butomus latifolius D. Don, Prodr.fl. nepal.: 22 (1825). 
'in Nepalia'. (Syntype: specimen not found). 
= Butomopsis latifolia (D. Don) Kunth 

CAMPANULACEAE 

Lobelia begonifolia Wall., D. Don, Prodr.fl. nepal: 158 (1825). 

Lobelia obliqua Buch.-Ham., in sched. 

Sembu, 5 June 1802 (BM!). 

= Pratia nummularia (Lam.) A. Braun & Asch. 

Lobelia pyramidalis Wall., D. Don, Prodr.fl. nepal: 157 (1825). 
Lobelia stimulans Buch.-Ham., in sched. 

'in Nepaliae montosis ad Narainhetty. Floret Februario'. (Specimen 
not found). 

Lobelia secundeil Buch.-Ham., in sched. 
Narainhetty, 17 February 1803 (BM!) 
= Lobelia rosea Wall. 

Lobelia trialata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 157 

(1825), nom. superfl. 
'in Nepalia' [No original label]. (BM!). 
= Lobelia heyneana Roem. & Schult. 

CAPRIFOLIACEAE 

Caprifolium macranthum D. Don, Prodr.fl. nepal.: 140 (1825). 

Xylosteum scandens Buch.-Ham., in sched. 

Sembu, 2 June 1802 (BM!). 

= Lonicera macrantha (D. Don) Spreng. 



J.R. PRESS AND K.K. SHRESTHA 

Viburnum cylindricum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 

142(1825). 
Narainhetty, 10 August 1802 (BMNsyntype). 

Viburnum mullaha [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 141 

(1825). 

Viburnum lantanal Buch.-Ham., in sched. 
Sembu, 11 July 1802 (BM!-syntype). 

Viburnum punctatum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 

142(1825). 
Suembu, 28 April 1802 (BM!-syntype). 

Xylosteum ligustrinum (Wall.) D. Don, Prodr.fl. nepal : 140 ( 1 825), 

nom. superfl. 

Xylosteon naisoca Buch.-Ham., in sched. 
Chitlong, 11 April 1802 (BM!). 
= Lonicera ligustrina Wall. 

CARYOPHYLLACEAE 

Arenaria serpyllifolia L., D. Don, Prodr.fl. nepal: 215 (1825). 
Narainhetty, 1 January 1802 (BM!). 

Cerastium grandiflorum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 

216(1825). 

Narainhetty, 11 March 1803 (BM!-syntype). 
= Cerastium fontanum subsp. grandiflorum (D. Don) H. Hara 

Stellaria monosperma [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 

215(1825). 

Stellaria saccurcunda Buch.-Ham., in sched. 
Narainhetty, 16 October 1802 (BM!-syntype). 

Cited as lectotype by Hara in Enum.fl. pi Nepal 2: 58 (1979). 

Stellaria saxatilis [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 215 

(1825). 

Bimpedi, 8 April 1802 (BM!-syntype). 
= Stellaria vestita Kurz 

CELASTRACEAE 

Euonymus lacerus [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 191 

(1825). 

Euonymus madana Buch.-Ham., in sched. 
Suembu, 5 May 1802 (BM!-syntype). 
= Euonymus grandiflorus Wall. 

COMBRETACEAE 

Combretum? appendiculatum Buch.-Ham., in sched. 
Hettaura, 24 March 1803 (Sheet 658.14 LINN-SM, microfiche!); 
Hettaurei (Sheet 658.15 LINN-SM, microfiche!). 

Combretum nanum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.:2l9 

(1825). 

Bassaria, 4 March 1802 (BM!-syntype); Bassaria, 9 March 1802 
(Sheet 586.16.2 LINN-SM, microfiche!-syntype); Terriany for- 
est, 30 March 1802 (Sheet 586.16.1 LINN-SM, 
microfiche ! -syntype) . 

Combretum spicatum Buch.-Ham., in sched. 

Suembu, 17 May 1802 (Sheet 658.17 LINN-SM, microfiche!). 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



107 



COMMELINACEAE 

Aneilema hispida D. Don, Prodr.fl. nepal: 45 (1825). 
Commelina hispida Buch.-Ham., in sched. 
Narainhetty, 22 September 1802 (BM!-syntype). 
= Floscopa scandens Lour. 

Commelina obliqua [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 45 

(1825), nonVahl( 1806). 
'in Nepalia.' (Syntype: specimen not found). 
= Commelina paludosa Blume 

COMPOSITAE 

Antennaria contorta D. Don in Bot. Reg. 7: t. 605 (1821). 
Gnaphalium contortum Buch.-Ham., in sched. 
Narainhetty, 14 October 1802 (BM!). 
= Anaphalis contorta (D. Don) Hook.f. 

Don states that 'The species was originally observed by Dr. 
Hamilton (then Buchanan) near the town of Narainhetty in Nepal, 
and samples collected in the native spot are preserved in Mr. 
Lambert's Herbarium'. However, the species was described from 
material grown at Spofforth in England. The original source was 
probably Buchanan-Hamilton, according to Don via the Calcutta 
Botanic Garden. 

Antennaria timmua D. Don, Prodr.fl. nepal: 174 (1825). 
Gnaphalium timmua Buch.-Ham., in sched. 
'in Nepalia'. No original label (BM!-syntype). 
= Anaphalis margaritacea (L.) Benth. 

Antennaria triplinervis Sims in Bot. Mag. 51: t. 2468 (1824). 
Gnaphalium quintuplinerve Buch.-Ham., in sched. 
Narainhetty, 30 August 1802 (BM!). 
= Anaphalis triplinervis (Sims) C.B. Clarke 

Artemisia parviflora [Roxb. ex] D. Don, Prodr. fl. nepal: 181 

(1825). 
Artemisia parviflora Buch.-Ham., in sched.; Artemisia chinense 

L.?, Buch.-Ham., in sched. 
Narainhetty, 15 September 1802 (BM!-syntype?). 
= Artemisia japonica Thunb. 

Roxburgh, Hort. bengal: 61 (1814) also cites a Buchanan speci- 
men from Nepal, but without date or locality. 

Aster trinervius [Roxb. ex] D. Don, Prodr.fl. nepal: 111 (1825). 
Aster asper [Buch.-Ham. ex] DC., Prodr. 5: 277 (1836), nom. 

superfl. 
Narainhetty, 23 October 1802 (BM!-syntype). 

Roxburgh, Hort. bengal.: 61 (1814) also cites a Buchanan speci- 
men from Nepal, 1802 but without further details. 

Cacalia cusimbua D. Don, Prodr.fl. nepal: 179 (1825). 
Cacalia cusimbium Buch.-Ham., in sched. 
'in Nepalia.' (Syntype: specimen not found). 
= Gynura bicolor (Willd.) DC. 

Chaptalia maxima D. Don, Prodr.fl. nepal: 166 (1825). 
Perdicium semiflosculare? Buch.-Ham., in sched. 
Narainhetty, 17 November 1802 (BM-syntype). 
= Gerbera maxima (D. Don) Beauverd 

Cnicus verutus D. Don, Prodr.fl. nepal: 167 (1825). 
Carduus trilobus Buch.-Ham., in sched. 



Narainhetty, 11 March 1803 (BM-syntype). 
= Cirsium verutum (D. Don) Spreng. 

Conyza cappa [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 176 

(1825). 

Conyza? cappa Buch.-Ham., in sched. 
Suembu, 7 July 1802 (BM-syntype). 
= Duhaldea cappa (D. Don) Anderberg 

Erigeron alatum [Roxb. ex] D. Don, Prodr.fl. nepal: 171 (1825). 
Conyza alata Buch.-Ham., in sched. 
Narainhetty, 18 October 1802 (BM!-syntype?). 
= Laggera alata (D. Don) Sch. Bip. ex Oliv. 

Roxburgh, Hort. bengal: 61 (1814), cites a Buchanan specimen 
from Nepal but without date or locality. 

Erigeron falcatum D. Don, Prodr.fl. nepal: 172 (1825). 

Conyza falcata Buch.-Ham., in sched. 

'ad Bassaria Nepalensium. Floret Februario'. (Syntype: specimen 

not found). 
= Blumeopsis falcata (D. Don) Merrill 

This specimen is also the syntype of Conyza falcata [Buch.-Ham. 
ex] Spreng., Sys. veg. 2, 513 (1825). 

Erigeron leucanthum D. Don, Prodr.fl. nepal: 171 (1825). 

Conyza leucanthemea Buch.-Ham., in sched. 

Bassaria, 1 March 1802 (BM-syntype). 

= Conyza leucantha (D. Don) Ludlow & P.H. Raven 

Erigeron pinnatifidum [Roxb. ex] D. Don, Prodr. Fl Nepal: 172 

(1825). 

Conyza triflda Buch.-Ham., in sched. 
'in Nepalia'. (Syntype: specimen not found). 
= Conyza stricta var. pinnatifida (D. Don) Kitam. 

Roxburgh, Hort. bengal: 61 (1814) cites Buchanan from Nepal 
but without further data. 

Erigeron hieracifolium D. Don, Prodr.fl. nepal: 172 (1825). 
'in Nepalia'. (Syntype: specimen not found). 
= Blumea hieracifolia (D. Don) DC. 

Eupatorium acuminatum D. Don, Prodr.fl. nepal: 171 (1825). 
Eupatorium bhuibu? Buch.-Ham., in sched. 
Narainhetty, 26 January 1803 (BM!-syntype). 
= Vernonia extensa DC. 

Eupatorium pyramidale D. Don, Prodr.fl. nepal: 170 (1825). 
Eupatorium bhuibu Buch.-Ham., in sched. 
Narainhetty, 19 November 1802 (BM!-syntype). 
= Vernonia aspera Buch.-Ham. 

Gnaphalium busua [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 173 

(1825). 

Gnaphalium busua Buch.-Ham., in sched. 
Sembu, 22 June 1802 (BM-syntype). 
= Anaphalis busua (D. Don) DC. 

Liatris latifolia D. Don, Prodr.fl. nepal: 169 (1825). 
Perdicium? triflorum Buch.-Ham., in sched. 
Narainhetty, 11 February 1803 (BM-syntype). 
= Ainsliaea latifolia (D. Don) Sch. Bip. 

Senecio buimalia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 178 
(1825). 



108 

Cacalia volubilis Buch.-Ham., in sched. 

'ad Narainhetty Nepalensium. Floret Novembri et Decembri'. 

(Syntype: specimen not found). 
= Cissampelopsis buimala (D. Don) C. Jeffrey & Y.L. Chen 

Senecio cappa [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 179 

(1825). 

Senecio cappa Buch.-Ham., in sched. 
Narainhetty, 15 November 1802 (BM-syntype). 
= Synods cappa (D. Don) C. Jeffrey & Y.L. Chen 

Senecio denudata D. Don, Prodr. fl. nepal.: 179 (1825). 
Cineraria denudata Buch.-Ham., in sched. 
Chitlong, 16 April 1802 (BM-syntype). 
= Senecio nudicaulis [Buch.-Ham. ex] D. Don 

Senecio jacobaea L., D. Don, Prodr. fl. nepal.: 179 (1825). 
Senecio musuca Buch.-Ham., in sched. 
Narainhetty, 18 November 1802 (BM!). 
= Senecio chrysanthemoides DC. 

Senecio triligulatus [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 178 

(1825) 'triligulata'. 

Senecio 3-ligulata Buch.-Ham., in sched. 
Narainhetty, 3 December 1802 (BM-syntype). 
= Synods triligulata (D. Don) C. Jeffrey & Y.L. Chen 

CONVOLVULACEAE 

Parana racemosa Roxb., D. Don, Prodr. fl. nepal.: 98 (1825). 
Parana dichotoma Buch.-Ham., in sched.; Dinetus [no species 

name given] Buch.-Ham., in sched. 
Narainhetty, 29 September 1802 (BM!). 
= Dinetus racemosus (Wall.) Sweet 

Roxburgh, Fl. Ind. 2: 41 (1824) based his description on plants 
introduced by Buchanan to the Calcutta Botanic Garden. 

CORDIACEAE 

Ehreda laevis Roxb., D. Don, Prodr. fl. nepal.: 102 (1825). 
Gorasan, 11 February 1802 (BM!). 

CORNACEAE 

Cornus oblonga Wall, in Roxb., D. Don, Prodr. fl. nepal: 140 

(1825). 

Cornus paniculata Buch.-Ham., in sched. 
Narainhetty, 3 September 1802 (BM!). 
= Swida oblonga (Wall.) Sojak 

CRUCIFERAE 

Cardamine debilis D. Don, Prodr. fl. nepal.: 201 (1825). 
Cardamine resedifolia L.?, Buch.-Ham., in sched. 
'Narainhetty. Octobri' (Syntype: specimen not found). 
= Cardamine flexuosa With. 

Cardamine nasturtioides D. Don, Prodr. fl. nepal.: 201 (1825). 
'in Nepalia'. (Syntype: specimen not found). 

Kara, in Enum.fl. pi. Nepal 2: 41 (1979), suggested this taxon 
might represent a form of Cardamine scutata subsp. flexuosa (With.) 
H. Kara i.e. C. flexuosa With. 

CUPRESSACEAE 

Juniperus chinensis? Buch.-Ham., in sched. 



J.R. PRESS AND K.K. SHRESTHA 

Narainhetty, 17 August 1802 (BM!). 
= Juniperus indica Bertol. 

Juniperus recurva [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 55 

(1825). 
'ad Narainhetty Nepaliae. Floret Februario'. (Syntype: specimen 

not found). 

CYPERACEAE 

Eleocharis congesta D. Don, Prodr. fl. nepal.: 41 (1825). 

Scirpus congestus Buch.-Ham., in sched. 

'in Nepalia'. [No original label]. (BM!-syntype). 

Fimbristylis diphylla (Retz.) Vahl, Enum. pi. 2: 289 (1806). 

Bassaria, 9 March 1802 (BM!). 

= Fimbristylis dichotoma (L.) Vahl 

Scirpus elongatus [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 40 

(1825). 

Narainhetty, 9 November 1802 (BM!-syntype). 
= Eriophorum comosum (Wall.) Wall, ex C.B. Clarke 

DATISCACEAE 

Datisca nepalensis D. Don, Prodr. fl. nepal.: 203 (1825). 
'in Nepalia'. (Syntype: specimen not found). 
= Datisca cannabina L. 

DIOSCOREACEAE 

Dioscorea obtusangula Buch.-Ham., in sched. 

Suembu, 17 July 1802 (Sheet 1544.15 LINN-SM, microfiche!). 

= Dioscorea bulbifera L. 

Dioscorea pisiformis Buch.-Ham., in sched. 
Narainhetty, 12 October 1802 (Sheet 1544.14 LINN-SM, micro- 
fiche!). 

Rajania? cordata L., Buch.-Ham., in sched. 
Suembu, 15 May 1802 (Sheet 1543.2 & 1543.3 LINN-SM, micro- 
fiche!). 

DROSERACEAE 

Drosera lunata [Buch.-Ham. ex] DC., Prodr. 1: 319 (1824). 

Sembu, 19 July 1802 (BM!-syntype). 

= Drosera peltata var. lunata (DC.) C.B. Clarke 

ELAEAGNACEAE 

Elaeagnus arborea Roxb., D. Don, Prodr. fl. nepal.: 67 (1825). 
Elaeagnus armata Buch.-Ham., in sched. 

Narainhetty, 22 November 1802 (Sheet 228.6 LINN-SM, micro- 
fiche!); 4 March 1803 (Sheet 228.7 LINN-SM, microfiche!). 
= Elaeagnus infundibularis Momiy. 

Elaeagnus umbellata Thunb., D. Don, Prodr. fl. nepal.: 68 (1825). 
Elaeagnus umbellatus Thunb., Buch.-Ham., in sched. 
Chitlong, 13 April 1802 (Sheet 228.8 LINN-SM, microfiche!). 
= Elaeagnus parvifolia Wall, ex Royle 

ERICACEAE 

Andromeda ovalifolia Wall., D. Don, Prodr. fl. nepal.: 148 (1825). 
Andromeda capricida Buch.-Ham., in sched. 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



109 



Suembu, 30 April 1802 (Sheet 789.35 LINN-SM, microfiche!). 
= Lyonia ovalifolia (Wall.) Drude 

Gaultheria fragrans D. Don, Prodr. fl. nepal.: 151 (1825), nom. 

superfl. 

Arbutus laurifolia Buch.-Ham., in sched. 
Narainhetty, 10 March 1803 (BM!); Chitlong, 10 April 1802 (Sheet 

792.9 LINN-SM, microfiche!). 
= Gaultheria fragrantissima Wall. 

Brossaea procumbens Buch.-Ham., in sched. 

Narainhetty, 3 February 1803 (Sheet 790.3 LINN-SM, microfiche!). 

= Gaultheria nummularioides D. Don 

Rhododendron arboreum Sm., Exot. hot. 1: 9, t. 6 (1805). 
Rhododendrum album [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 

154(1825), nom. superfl. 
Nepal (Sheet 788.6 LINN-SM, microfiche!); Narainhetty, 3 March 

1803 (Sheet 788.8 LINN-SM, microfiche!). 
Don's citation of R. purpureum Buch.-Ham., in sched. suggests 
that there is one (or more) other Buchanan specimen(s) annotated 
with this name. None have been located at BM. 

EUPHORBIACEAE 

Euphorbia angustifolia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 

62(1825). 

Norcotera, 26 March 1802 (BM!-syntype). 
= Euphorbia dracunculoides Lam. 

Euphorbia fusiformis [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 

62(1825). 
'in Nepalia" [No original label]. (BM!-syntype). 

Euphorbia prolifera [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 62 

(1825). 
'in Nepalia' [No original label]. (BM!-syntype). 

Euphorbia tenuis [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 61 

(1825). 

Bassaria, 10 March 1803 (BM!-syntype). 
= Euphorbia parviflora L. 

Myrica octandra [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 56 

(1825). 

Ettaura, 2 April 1802 (BM!-syntype). 
= Aporusa octandra (D. Don) A.R. Vickery ex M. Short & A.R. 

Vickery 

Phyllanthus parvifolius [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 

63 (1825). 'parvifolia'. 
Sembu, 5 June 1802 (BM!-syntype). 

FAGACEAE 

Quercus armata Roxb., D. Don, Prodr. fl. nepal.: 56 (1825). 

Quercus catungea Buch.-Ham., in sched. 

Narainhetty, 19 November 1 802; Narainhetty, 26 July 1 802; 29 July 

1802; Suembu, 1 May 1802 (BM!-syntypes). 
= Castanopsis tribuloides (Sm.) A. DC. 

Quercus cassura [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 57 

" (1825). 

'in Nepalia' [No original label]. (BM!-syntype). 

= Quercus semecarpifolia Sm. 



Quercus imbricata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 57 

(1825). 

Narainhetty, 7 November 1802 (BM!-syntype). 
= Cyclobalanopsis lamellosa (Sm.) Oerst. 

Quercus lanuginosa D. Don, Prodr. fl. nepal.: 57 (1825), non Thuill. 

(1799). 

Quercus banga Buch.-Ham., in sched. 
Narainhetty, 17 December 1802 (BM!-syntype). 
= Quercus lanata Sm. 

Quercus phullata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 57 

~ (1825). 

Narainhetty, 5 December 1802 (BM!-syntype); Suembu, 5 May 

1802 (BM!-syntype). 
= Cyclobalanopsis glauca (Thunb.) Oerst. 

Quercus spicata Sm. in Rees, Cycl 29: n. 12 (1814), non Humb. & 

Bonpl. (1809). 

Quercus arcaula Buch.-Ham., in sched. 
Suembu, 5 May 1802 (BM!-syntype). 
= Lithocarpus grandifolius var. brevipetiolatus (A. DC.) S.N. 

Biswas 

FLACOURTIACEAE 

Flacourtia infrafoliacea Buch.-Ham., in sched. 

Malucona communis Buch.-Ham., in sched. 

Sembu, 21 April 1802 (Sheet 1555.5.1 LINN-SM, microfiche!); 
Narainhetty, 14 October 1802 (Sheet 1555.5.2 LINN-SM, micro- 
fiche!). 

Flacourtia sepiaria Roxb., Buch.-Ham., in sched. 

Norcotera, 26 March 1802 (Sheet 1555.3 LINN-SM, microfiche!). 

Flacourtia [no species name given] 

Malucona communis? Buch.-Ham., in sched. 

Tancote, 16 April 1802 (Sheet 1555.4 LINN-SM, microfiche). 

GENTIANACEAE 

Exacum tetragonum Roxb., Buch.-Ham., in sched. 

Narainhetty, 21 August 1802 (Sheet 203.7 LINN-SM, microfiche!). 

Gentiana capitata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 126 

(1825). 

Narainhetty, 3 February 1803 (BM!-syntype); Narainhetty, 12 Janu- 
ary 1803 (Sheet 476.56.1 LINN-SM, microfiche!); Narainhetty, 
6 March 1803 (Sheet 476.56.2 LINN-SM, microfiche!). 

Gentiana decemfida [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 
127(1825). 

Narainhetty, 5 February 1803 (Sheet 476.55 LINN-SM, micro- 
fiche!). 
Although the locality matches that given in the protologue, Don 

states 'Floret Martio'. No specimen with this date has been located. 

Menyanthes indica L., Buch.-Ham., in sched. 

Sembu, 27 June 1802 (Sheet 276.7 LINN-SM, microfiche!). 

= Nymphoides indica (L.) Kuntze 

Swertia angustifolia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 

127(1825). 
Narainhetty, 13 September 1802 (BM!-syntype; Sheet 475.9 LINN- 

SM-syntype, microfiche!) 



110 



J.R. PRESS AND K.K. SHRESTHA 



Swertia ciliata (G. Don) B.L. Burtt in Notes Roy. Bot. Card. 

Edinburgh 26: 272(1965). 
Narainhetty, 7 October 1 802 (Sheet 475. 1 1 LINN-SM, microfiche!). 

Swertia quadriculata Buch.-Ham., in sched. 
Narainhetty, 12 October 1802 (Sheet 475.10 LINN-SM, micro- 
fiche!). 

GERANIACEAE 

Geranium nepalense Sweet, D. Don, Prodr.fl. nepal.: 208 (1825). 
Geranium quinquenerve Buch.-Ham., in sched. 
'in Nepalia, ad ripas umbrosas fluvii Kuli Khana dicti. Floret 
Aprili'. (Specimen not found). 

GUTTIFERAE 

Brathys nepalensis Blume, Mus. hot. 2: 19 (1856). 
Catmandu, 13 May 1802 (BM!-syntype; L,-lectotype). 
= Hypericum japonicum Thunb. 

Lectotype designated by N.K.B. Robson in Bull. Brit. Mus. (Nat. 
Hist.), Bot. 20: 129(1990). 

Hypericum bracteatum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

217 (1825), nom. superfl. 
Tancote, 16 April 1802 (BMMectotype). 
H. [Hypericum] lungusum Buch.-Ham., in sched. 
Narainhetty, 10 October 1802 (BM!-syntype). 
= Hypericum cordifolium Choisy 

BM specimen designated as lectotype by N.K.B. Robson in Bull. 
Brit. Mus. (Nat. Hist.), Bot. 12: 213 (1985). 

Hypericum japonicum Thunb., D. Don, Prodr. fl. nepal.: 219 

(1825). 

Hypericum dichotomum Buch.-Ham., in sched. 
'in Nepalia in scaturiginosis prope urbem Katmandu. Floret Maio'. 

[No original label]. (BM!). 

Hypericum nervosum D. Don, Prodr.fl. nepal.: 219 (1825), nom. 

superfl. 

Narainhetty, 23 September 1802 (BM!). 
= Hypericum elodeoides Choisy 

Hypericum uralum [Buch.-Ham. ex] D. Don in Bot. Mag. 50: t. 

2375 (1823). 
'in Nepalia ad Narainhetty'. [No original label]. (BMMectotype). 

Lectotype designated by N.K.B. Robson in Bull. Br. Mus. Nat. 
Hist. (Bot.) 12 : 268 (1985). 

HYDRANGEACEAE 

Hydrangea aspera [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 211 

(1825). 
Narainhetty, 26 September 1802 (BM!-syntype). 

LABIATAE 

Ajuga decumbens sensu D. Don, Prodr.fl. nepal.: 108 (1825), non 

Thunb. (1784). 

Bassaria, 21 February 1802 (BM!). 
= Ajuga macrosperma var. breviflora Hook.f. 

Ajuga integrifolia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 108 
(1825). 

Ajuga alpina ? Buch.-Ham., in sched.; Ajuga pyramidalis L. ? Buch.- 
Ham., in sched. 



Narainhetty, 18 September 1802 (BM!-syntype). 
= Ajuga bracteosa Wall, ex Benth. 

Colebrookea oppositifolia Sm., Exot. hot. 2: 111, t. 115 (1805). 
Selago? oppositifolia Buch.-Ham., in sched. 
Muking, 21 January 1802 (BM!-syntype). 

Smith cites Buchanan material collected 'by roadsides in Nepal, 
1 802' but without precise locality or date. 

Perilla elata D. Don, Prodr.fl. nepal.: 115 (1825). 
'in Nepalia'. (Syntype: specimen not found) 
= Elsholtzia blanda (Benth.) Benth. 

Perilla fruticosa D. Don, Prodr.fl. nepal.: 115 (1825). 

Mentha fruticosa Buch.-Ham., in sched. 

'in Nepalia' (BM!-syntype). 

= Elsholtzia fruticosa (D. Don) Rehder 

The original label gives a description of the plant but no date or 
locality. 

Perilla leptostachya D. Don, Prodr.fl. nepal.: 115 (1825). 

Nepeta imubus Buch.-Ham., in sched. 

Narainhetty, 20 October 1802 (BM!-syntype). 

= Elsholtzia stachyodes (Link) Raizada & Saxena 

Perilla ocimoides L., D. Don, Prodr.fl. nepal.: 114 (1825). 
Mentha perilloides L., Buch.-Ham., in sched. 
Narainhetty, 1 October 1802 (BM!). 
= Perilla frutescens (L.) Britton 

Perilla polystachya D. Don, Prodr.fl. nepal: 114 (1825). 
Mentha? cristata Buch.-Ham., in sched. 
'in Nepalia'. (Syntype: specimen not found) 
= Elsholtzia fruticosa (D. Don) Rehder 

Plectranthus coetsa [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 117 

(1825). 

Germanea coetsa Buch.-Ham., in sched. 
Narainhetty, 21 October 1802 (BM!-syntype). 
= Isodon coetsa (D. Don) Kudo 

Plectranthus ternifolius D. Don, Prodr. fl. nepal.: 117 (1825), 

'ternifolia'. 
Lavendula ternifolia Buch.-Ham., in sched.; Plectranthus buchia 

Buch.-Ham., in sched. 

Narainhetty, 18 September 1802 (BM!-syntype). 
= Isodon ternifolius (D. Don) Kudo 

Plectranthus virgatus D. Don, Prodr.fl. nepal.: 1 16 (1825) 'virgata', 

non Ocimum virgatum Thunb. ex Murray (1784). 
Ocymum virgatum Thunb., Buch.-Ham., in sched. 
Ettaura, 2 April 1803 (BM!-syntype). 
= Orthosiphon rubicundus (D. Don) Benth. 

Scutellaria indica sensu D. Don, Prodr.fl. nepal. : 109 ( 1 825), non L. 

(1751). 

Narainhetty, 3 August 1802 (BM!). 
= Scutellaria discolor Colebr. 

Scutellaria repens [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 110 

(1825). 
Narainhetty, 4 October 1802 (BM!-syntype). 

Scutellaria scandens [Buch.-Ham. ex] D. Don, Prodr. fl. nepal. : 
110(1825). 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



111 



Scutellaria albida? Buch.-Ham., in sched. 

Tomba Cana, Nepalia, 22 March 1803 (BM!-syntype). 

Teucrium laxum D. Don, Prodr.fl. nepal: 109 (1825). 

Ajuga laxa Buch.-Ham., in sched. 

'in Nepalia'. [No original label]. (BM!-syntype). 

Teucrium quadrifarium [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

108 (1825). 
Narainhetty, 27 August 1802 (BM!-syntype). 

Thytnus piperitus D. Don, Prodr.fl. nepal.: 112 (1825). 
Marrubium piperitum Buch.-Ham., in sched. 
Narainhetty, 8 March 1803 (BM!-syntype). 
= Clinopodium piperitum (D. Don) Murata 

Don also cites Roxburgh, Hort. bengal.: 44 (1814), who lists a 
Buchanan-Hamilton specimen collected from Nepal in 1802. 

Thymus repens D. Don, Prodr.fl. nepal.: 113 (1825). 
Clinopodium repens Buch.-Ham., in sched. 
Sembu, 25 May 1802 (BM!-syntype). 
= Clinopodium umbrosum (M. Bieb.) K. Koch 

LAURACEAE 

Cinnamomum cathia D. Don, Prodr.fl. nepal.: 66 (1825). 

Laurus cathia Buch.-Ham., in sched. 

Suembu, 1 June 1802 (BM!; Sheet 707.42 LINN-SM, microfiche!- 

syntypes). 
= Phoebe cathia (D. Don) Kosterm. 

Cinnamomum tomentosum D. Don, Prodr.fl. nepal: 66 (1825). 
Laurus tomentosa Buch.-Ham., in sched. 

Suembu, 19 April 1802 (BM!; Sheet 707.41 LINN-SM, micro- 
fiche !-syntypes). 
= Phoebe cathia (D. Don) Kosterm. 

Laurus cuneata Buch.-Ham., in sched. 

Chitlong, 10 April 1802 (Sheet 707.44 LINN-SM, microfiche!). 

Laurus cuspidata D. Don, Prodr.fl. nepal.: 64 (1825). 

Tomex bolo Buch.-Ham., in sched. 

Narainhetty, 11 March 1803 (Sheet 847.4 LINN-SM, microfiche !- 
syntypes); Narainhetty, 3 January 1803 (Sheet 847.5 LINN-SM, 
microfiche!); Kargoo, Nepal, 21 March 1803 (Sheet 847.6 LINN- 
SM, microfiche!). 

= Lindera melastomacea (Nees) Villar 

Laurus gushia Buch.-Ham., in sched. 

Narainhetty, 17 November 1802 (Sheet 707.24 LINN-SM, micro- 
fiche!). 

Laurus jacaricata Buch.-Ham., in sched. 

Catmandu, 12 May 1802 (Sheet 707.43 LINN-SM, microfiche!). 

Laurus^ lateralis Buch.-Ham., in sched. 

Catmandu, 13 May 1802 (Sheet 707.23 LINN-SM, microfiche!). 

Laurus nacusua D. Don, Prodr.fl. nepal.: 64 (1825). 

Laurus nancushia Buch.-Ham., in sched. 

Narainhetty, 10 March 1803 (BM!; Sheet 707.22 LINN-SM, micro- 

fiche!-syntypes). 
= Lindera nacusua (D. Don) Merr. 



Laurus umbellata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 64 
(1825), nonThunb. ( 1784). 

Chisapany, 9 April 1802 (BM!; Sheet 707.21 LINN-SM, micro- 
fiche! -syntypes). 

= Lindera nacusua (D. Don) Merr. 

Tetranthera cuipala [Buch.-Ham. ex] D. Don, Prodr. fl. nepal. : 65 

(1825). 

Laurus cuipala Buch.-Ham., in sched. 
Narainhetty, 27 February 1803 (BM!; Sheet 707.25 LINN-SM, 

microfiche! -syntypes); Narainhetty, 9 January 1 803 (Sheet 707.26 

LINN-SM, microfiche!). 
= Neolitsea cuipala (D. Don) Kosterm. 

Sheet 707.26 LINN-SM is not a syntype as Don gives the flower- 
ing time as February. 

Tetranthera doshia D. Don, Prodr.fl. nepal.: 65 (1825). 
Tomex doshia Buch.-Ham., in sched. 

Narainhetty, 1 November 1802 (BM!; Sheet 847.7 LINN-SM, mi- 
crofiche ! -syntypes). 
= Litsea doshia (D. Don) Kosterm. 

Tomex llampatia Buch.-Ham., in sched. 

Narainhetty, 17 February 1803 (Sheet 847.3 LINN-SM, micro- 
fiche!). 

LEGUMINOSAE 

Aspalanthus cuneata D. Don, Prodr.fl. nepal.: 246 (1825). 
Anthyllis cuneata Buch.-Ham., in sched.; Hedysarumjunceum L.f., 

Buch.-Ham., in sched. 

Narainhetty, 2 October 1802 (BM!-syntype). 
= Lespedeza juncea var. sericea (Thunb.) F.B. Forbes & Hemsl. 

Astragalus stipulatus [D. Don ex] Sims in Bot. Mag. 50: t. 2380 

(1823), D. Don, Prodr.fl. nepal.: 246 (1825). 
Astragalus lanceolatus Buch.-Ham., in sched.; Coronilla stipulata 

Buch.-Ham., in sched. 
Gorasan, 18 March 1802 (BM!-syntype). 

Cassia dimidiata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 247 

(1825). 

Narainhetty, 8 July 1802 (BM!-syntype). 
= Cassia mimosoides subsp. lechenaultiana (DC.) H. Ohashi 

Don also cites Roxburgh, Hort. bengal.: 32 (1814), who lists a 
Buchanan-Hamilton specimen collected from Nepal in 1801. 

Crotalaria alata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 241 

(1825). 
Sembu, 17 July 1802 (BM!-syntype). 

Crotalaria anthylloides Lam., D. Don, Prodr.fl. nepal.: 241 (1825). 
Crotalaria salicifolia Buch.-Ham., in sched. 
'in Nepalia ad Narainhetty. Floret Octobri'. (Specimen not found). 
= Crotalaria sessiliflora L. 

Crotalaria linifolia sensu D. Don, Prodr.fl. nepal.: 241 (1825), non 

L.f. (1782). 

Crotalaria pilosa Buch.-Ham., in sched. 
Narainhetty, 23 September 1802 (BM!). 

Crotalaria poly galifolia Buch.-Ham., in sched.; Crotalaria prostrata 

Buch.-Ham., in sched. 
Bassaria, 4 March 1802 (BM!). 
= Crotalaria albida [Heyne ex] Roth 



112 



J.R. PRESS AND K.K. SHRESTHA 



Crotalaria prostrata sensu D. Don, Prodr.fl. nepal. : 241 ( 1 825), non 

Rottb. (1809). 

Crotalaria ciliata Buch.-Ham., in sched. 
'in Nepalia'. (Specimen not found). 
= Crotalaria humifusa Benth. 

Crotalaria psoralioides D. Don, Prodr.fl. nepal.: 242 (1825), non 

Lam. (1786), nom. superfl. 
Crotalaria ?cytissoides Buch.-Ham., in sched. 
Narainhetty, 10 August 1802 (BM!). 
= Crotalaria cytisoides Roxb. ex DC. 

Don also cites Roxburgh, Hon. bengai.: 54 (1814), who lists a 
Buchanan-Hamilton specimen collected from Nepal in 1801. 

Crotalaria tuberosa [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 241 

(1825). 

Borbonia tuberosa Buch.-Ham., in sched. 
Narainhetty, 28 September 1802 (BM!-syntype). 
= Eriosema himalaicum H. Ohashi 

Dalbergia sericea G. Don, Gen. hist. 2: 375 (1832). 
Churiaghant hills, 31 March 1802 (BM!-syntype). 

Hedysarum dioicum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 244 

(1825). 

Narainhetty, 25 September 1802 (BM!-syntype). 
= Desmodium confertum DC. 

Hedysarum retusum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 243 

(1825). 

Narainhetty, 2 September 1802 (BM!-syntype). 
= Desmodium concinnum var. retusum (D. Don) H. Ohashi 

Hedysarum sambuense D. Don, Prodr.fl. nepal.: 243 (1825). 
Hedysarum suembum Buch.-Ham., in sched. 
Suembu, 16 July 1802 (BM!-syntype). 
= Desmodium mult ilium m DC. 

Hedysarum tenellum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 243 

(1825). 

Narainhetty, 6 October 1802 (BM!-syntype). 
= Desmodium microphyllum (Thunb.) DC. 

Indigofera atropurpurea [Buch.-Ham. ex] Hornem., Hort. hot. 

Hafh.Suppl.: 152(1819). 
Indigofera atropurpurea [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

244(1825), nom. superfl. 
Narainhetty, 7 February 1803 (BM!-syntype). 

Don also cites Roxburgh, Hort. bengai.: 57 (1814), who lists a 
Buchanan specimen collected from Nepal but without date or other 
locality information. 

Indigofera dosua [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 244 

(1825). 
Suembu, 7 July 1802 (BM!-syntype). 

Indigofera trifoliata L., Buch.-Ham., in sched. 
Sembu, 23 July 1802(BM!). 

Manna nepalensis D. Don, Prodr.fl. nepal.: 247 (1825). 
Genista juasi Buch.-Ham., in sched.; Hedysarum alhagi L., Buch.- 
Ham., in sched. 

Sitaucund, 21 April 1803 (BM!-syntype). 
= Alhagi nepalensis (D. Don) Shap. 



Sitacund is in Chittagong (now Bangladesh). Either the date on 
the specimen is incorrect, (Buchanan having left Nepal in March 
1 803) and probably should read 2 1 April 1 802, or this plant was not 
described from Nepal. 

Parochetus communis [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

240(1825). 
Glycine proscimum Buch.-Ham.?, in sched.; Parochetus communis 

Buch.-Ham., in sched. 
Narainhetty, 24 October 1802 (BM!-syntype). 

Parochetus major D. Don, Prodr.fl. nepal.: 241 (1825). 

'in Nepalia'. (Syntype: specimen not found). 

= Parochetus communis [Buch.-Ham. ex] D. Don 

LENTIBULARIACEAE 

Utricularia confervaefolia [Jackson ex] D. Don, Prodr.fl. nepal.: 84 

(1825). 

Utricularia vulgaris Buch.-Ham., in sched. 
Narainhetty, 3 September 1802 (BM!-syntype); Narainhetty (Sheet 

52.13 LINN-SM-syntype, microfiche!). 
= Utricularia aurea Lour. 

The name is apparently based on a Jackson name on a specimen 
in Herb Lambert. 

LILIACEAE s.l. 
(ASPARAGACEAE) 

Asparagus albus Buch.-Ham., in sched. 

Gorasan, 20 March 1802 (Sheet 600.10 LINN-SM, microfiche!). 

Asparagus curillus [Buch.-Ham. ex] Roxb., Fl. ind. ed. 1832 2: 152 

(1832). 

Sembu, 22 June 1 802 (BM? specimen not found; Sheet 600.9 LINN- 
SM, microfiche!). 

Hara, in Enum.fl. pi. Nepal 1: 71 (1978) cites 'Buch.Ham. (Herb. 
Roxb., type of A. curillus)' and notes that 'Hamilton's A. curillus 
(BM & E) seems to be a mixture of this plant and A. racemosus [i.e. 
A. racemosus Willd.], but Roxburgh's plate of A. curillus at Kew 
shows short flattish cladophylla' . Hara may well have meant that the 
Roxburgh icones should be taken as the type. In any case, no 
Buchanan-Hamilton specimen has been located at BM. 

Roxburgh, Fl. ind. ed. 1832 2: 152 (1832) cites material grown in 
the Calcutta Botanic Garden from seed supplied by Buchanan. 

Asparagus filicinus [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 49 

(1825). 
Sembu, 2 June 1802 (BM!; Sheet 600.6 LINN-SM, microfiche !;- 

syntypes). 

(UVULARIACEAE) 

Disporum pitsutum D. Don, Prodr.fl. nepal.: 50 (1825). 

Uvularia pitsutu Buch.-Ham., in sched. 

Chitlong, 10 April 1802 (BM!-syntype); Chitlong, 11 April 1802 

(Sheet 586.3 LINN-SM, microfichei-syntypes). 
= Disporum cantoniense var. parviflorum (Wall.) H. Hara 

(LILIACEAE) 

Lilium japonicum Thunb., D. Don, Prodr.fl. nepal.: 52 (1825). 
Lilium batisua Buch.-Ham., in sched. 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 

LOGANIACEAE 



113 



Narainhetty, 2 August 1802 (BM!); 26 July 1802 (Sheet 584.12 

LINN-SM, microfiche!). 
= Lilium \\allichiiiiiiiiii Schult. & Schult.f. 

(TRILLIACEAE) 

Paris polyphylla Sm., in Rees, Cycl. 26: Paris n. 2 (1813). 
Paris diasua Buch.-Ham., in sched. 

Narainhetty, 13 March 1803 (Sheet 703.4.1 LINN-SM, microfiche!- 
syntype); Narainhetty, 17 March 1803 (BM!-syntype). 

(ANTHERICACEAE) 

Phalangium anceps Buch.-Ham., in sched. 

Suembu, 4 July 1802 (Sheet 596.9 LINN-SM, microfiche!). 

= Chlorophytum nepalense (Lindl.) Baker 

(SMILACACEAE) 

Smilax maculata [Roxb. ex] D. Don, Prodr.fl. nepal: 49 (1825). 
Smilax capitata Buch.-Ham., in sched. 

Narainhetty, 18 September 1802 (BM!-syntype); Narainhetty, 17 
September 1802(Sheet 1542. 14 LINN-SM, microfiche !-syntype). 
= Smilax aspera L. 

Don also cites Roxburgh, Hort. bengal: 72 (1814), who lists a 
Buchanan-Hamilton specimen collected from Nepal but without 
further data. 

Smilax ovalifolia [Roxb. ex] D. Don, Prodr.fl. nepal.: 49 (1825). 

Smilax columnifera Buch.-Ham., in sched. 

'ad Narainhetty Nepalensium. Floret Septembri'. Narainhetty, 7 

February 1803 (Sheet 1542.15 LINN-SM, microfiche !-syntype). 

Both specimens are from the type locality but Don gives the 
flowering time as February. 

(CONVALLARIACEAE) 

Sultea elatior Buch.-Ham., in sched. 

Narainhetty, 2 October 1 802 (Sheet 601 . 13 LINN-SM, microfiche!). 

= Ophiopogon clarkei Hook.f. 

Sultea humilior Buch.-Ham., in sched. 

Narainhetty, 27 July 1 802 (Sheet 601 . 14. 1 LINN-SM, microfiche!); 
Narainhetty, 21 August 1802 (Sheet 601.14.2 LINN-SM, micro- 
fiche!) 

= Ophiopogon wallichianus (Kunth) Hook.f. 

Ttopistra aurantica ([Wall, ex] Baker) Hook.f., Fl Brit. India. 6: 

325(1892). 

Pilcusta sylvatica Buch.-Ham., in sched. 
Narainhetty, 13 March 1803 (BM!); 16 March 1803 (Sheets 614.4 & 

614.5 LINN-SM, microfiche!). 



Buddleja paniculata Wall., D. Don, Prodr.fl. nepal: 92 (1825). 
Budleja shina Buch.-Ham., in sched. 
Narainhetty, 7 February 1803 (BM!) 
Budleja fastigiata Buch.-Ham, in sched. 
Tombah Cana, 22 March 1803 (BM!). 

Budleja tomentosa Buch.-Ham., in sched; Budleja nimda Buch.- 
Ham., in sched. 
Narainhetty, 30 December 1802 (BM!). 

Buddleja subserrata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 92 

(1825), nom. superfl. 
Bassaria, 27 February 1802 (BM!); 
Budleia shina Buch.-Ham., in sched.; Budleia simba Buch.-Ham., 

in sched. 

Narainhetty, 3 November 1802 (BM!). 
= Buddleja asiatica Lour. 

Don's name is a superfluous epithet for B. neemda Roxb., Fl ind. 
1: 411 (1820), based in part on material introduced to Calcutta 
Botanic Garden by Buchanan-Hamilton. 

LORANTHACEAE 

Loranthus odoratus Wall., D. Don, Prodr.fl. nepal: 143 (1825). 
Loranthus hexapetala Buch.-Ham., in sched. 
Narainhetty, 13 November 1802 (BM!). 

Viscum dichotomum D. Don, Prodr.fl. nepal: 142 (1825). 
Viburnum dichotomum Buch.-Ham., in sched. 
Narainhetty, 17 December 1802 (BM!-syntype). 
= Viscum articulatum Burm.f. 

Viscum stellatum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 142 

(1825). 

Suembu, 3 June 1802 (BM!-syntype). 
= Viscum album L. 

LYTHRACEAE 

Ammannia rubra [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 220 

(1825). 

'in Nepalia' [No original label]. (BM!-syntype). 
= Rotala rubra (D. Don) H. Hara 

Ammannia rotundifolia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 

220(1825). 

'in Nepalia' (Syntype: specimen not found). 
= Rotala rotundifolia (D. Don) Koehne 

Don also cites Roxburgh, Hort. bengal: 11 (1814) which lists a 
Buchanan-Hamilton specimen from Chittagong, 1796. The only 
specimen located at BM lacks an original label but is annotated 
'Mysoor', without further information. 



LINACEAE 

Linum cicanobum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 217 

(1825). 

Narainhetty, 5 December 1802 (BM!-syntype). 
= Reinwardtia cicanoba (D. Don) H. Hara 

Linum repens [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 217 (1825). 
Linum semitrigynum Buch.-Ham., in sched. 
Narainhetty, 5 November 1802 (BM!-syntype). 
= Reinwardtia indica Dumort. 



MAGNOLIACEAE 

Michelia doltsopa [Buch.-Ham. ex] DC., Syst. not. 1: 448 (1817). 
Narainhetty, 9 February 1803 (BM!-syntype). 

A specimen at BM collected by Buchanan-Hamilton from C. 
Nepal was identified by Dandy as a hybrid between Michelia 
doltsopa and Michelia champaca L. (cf. Dandy in /. Bot. 65: 278 
(1927). It has not been located. 

Michelia kisopa [Buch.-Ham. ex] DC., Syst. nat. 1: 448 (1817). 
Narainhetty, 22 October 1802 (BM!-syntype). 



114 



J.R. PRESS AND K.K. SHRESTHA 



MELASTOMATACEAE 

Arthrostemma paniculatum D. Don in Mem. Wernerian. nat. Hist. 

Soc. 4: 299 (1822). D. Don, Prodr.fl. nepal: 222 (1825). 
Rhexia hari Buch.-Ham., in sched.; Rhexia paniculata Buch.-Ham., 

in sched. 

Narainhetty, 6 October 1802 (BM!-syntype). 
= Oxyspora paniculata (D. Don) DC. 

Melastoma normale D. Don, Prodr. fl. nepal.: 220 (1825), 

'normalis'. 
'in Nepalia'. (Syntype: specimen not found). 

Hara, Enum. fl. pi. Nepal 2: 170 (1979) cites Wallich s.n., ann. 
1 8 1 8 as 'possibly type of M. normale'' . This suggestion is erroneous 
as Don cites only the Buchanan-Hamilton collection. 

Osbeckia chulesis D. Don, Prodr.fl. nepal: 221 (1825). 
Melastoma chulese Buch.-Ham., in sched. 
Narainhetty, 27 July 1802 (BM!-syntype). 
= Osbeckia nepalensis Hook. 

Cited as type of O. chulesis by Hara, Enum. Fl. PI. Nepal 2: 170 
(1979). 

Osbeckia rostrata D. Don, Prodr.fl. nepal.: 221 (1825). 
Melastoma? rostratum Buch.-Ham., in sched. 
Narainhetty, 11 August 1802 (BM!-syntype). 

Osbeckia speciosa D. Don, Prodr. fl. nepal: 222 (1825), nom. 

superfl. 

Melastoma humile Buch.-Ham., in sched. 
Narainhetty, 28 July 1802 (BM! synype?). 
= Osbeckia nepalensis Hook. 

One of several elements in the protologue of 0. nepalensis Hook., 
a name cited by Don. 

Osbeckia stellata [Buch.-Ham. ex] D. Don in Bot. Reg. 8: t. 674 

(1822). 

Melastoma stellatum Buch.-Ham., in sched. 
Narainhetty, 10 August 1802 (BMl-lectotype). 

Lectotype cited by Hara, Enum.fl. pi Nepal 2: 171 (1979). 

Osbeckia ternifolia D. Don, Prodr.fl. nepal: 221 (1825). 

Osbeckia quaterna Buch.-Ham., in sched. 

'in Nepalia' [No original label]. (BM!-syntype). 

The specimen was annotated as holotype of Osbeckia stellata var. 
rostrata (D. Don) by Carlo Hansen, 1974. 
= Osbeckia rostrata D. Don 

MENISPERMACEAE 

Cissampelos angulata Buch.-Ham., in sched. 

Cissampelos bahapo Buch.-Ham., in sched. 

Sembu, 28 May 1802 (Sheet 1567.5 LINN-SM, microfiche!). 

Cissampelos hirsuta [Buch.-Ham. ex] DC., Syst. nat. 1: 535 (1817). 
Suembu,21 May 1 802 (BM!; Sheet 1567.6 LINN-SM, microfiche !- 

syntypes). 
= Cissampelos pareira var. hirsuta (DC.) Forman 

MORACEAE 

Ficus cabur [Buch.-Ham. ex] Sm. in Rees, Cycl. 14: Ficus n. 47 
(1810). 

Narainhetty, 10 February 1803 (Sheet 1610.41 LINN-SM, micro- 
fiche !-syntype). 

= Ficus sarmentosa Sm. 



Ficus citrifolia? Roxb., Buch.-Ham., in sched. 
Narainhetty, 21 February 1803 (Sheet 1610.39 LINN-SM, micro- 
fiche!). 

Ficus infrafoliacea [Buch.-Ham. ex] Sm., in Rees, Cycl. 14: Ficus n. 

31 (1810). 
Lohiar, by roadsides in Nepal, 29 March 1803 (Sheet 1610.44 

LINN-SM, microfichei-syntype). 
= Ficus virens Ait. 

Ficus neriifolia Sm. in Rees, Cycl. 14: Ficus n. 21 (1810). 
'Narainhetty, Upper Nepal, 10 February 1803'. (Specimen not 

found). 

This specimen would be expected to be in LINN-Smith but no 
sheet has been located. The details given in the protologue exactly 
match those on the sheet of F. cabur. 

Ficus pilashi Sm., in Rees, Cycl. 14: Ficus n. 3 (1810). 
Narainhetty, 28 December 1802 (Sheet 1610.43 LINN-SM, micro- 
fiche !-syntype). 
= Ficus virens var. sublanceolata (Miq.) Corner 

Ficus sarmentosa [Buch.-Ham. ex] Sm. in Rees, Cycl 14: Ficus n. 
45(1810). 

Sembu, woods in Upper Nepal, 28 May 1802 (Sheet 16 10.40 LINN- 
SM, microfiche !-syntype). 

Ficus semicordata [Buch.-Ham. ex] Sm. in Rees, Cycl. 14: Ficus n. 

71(1810). 

Ficus cunea Buch.-Ham., in sched. 
Hettaura, woods in Upper Nepal, 4 April 1802 (Sheet 1610.26 

LINN-SM, microfichei-syntype). 

Ficus subincisa [Buch.-Ham. ex] Sm. in Rees, Cycl. 14: Ficus n. 91 

(1810). 
Narainhetty, on rocks, 28 January 1803 (Sheet 1610.42 LINN-SM, 

microfiche !-syntype). 

MYRICACEAE 

Myrica esculenta [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 56 

(1825). 
Narainhetty, 3 September 1802 (BM!-syntype). 

MYRSINACEAE 

Baebotrys gocala Buch.-Ham., in sched. 

Narainhetty, 17 February 1803 (Sheet 346.8 LINN-SM, micro- 
fiche!). 

Embelia esculenta D. Don, Prodr. fl. nepal: 147 (1825), nom. 

superfl. 

Samara? esculenta Buch.-Ham., in sched. 
Narainhetty, 5 January 1803 (BM!) 
= Embelia floribunda Wall. 

Embelia nagushia D. Don, Prodr. fl. nepal: 147 (1825), nom. 

superfl. 

Samara? nagushia Buch.-Ham., in sched. 
Narainhetty, 20 October 1802 (BM!). 
= Embelia vestita Roxb. 

Maesa chisia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 148 
(1825). 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



115 



Baebotrys chishia Buch.-Ham., in sched. 

Narainhetty, 10 January 1803 (BM!); Narainhetty, 9 February 1803 
(Sheet 346.4 LINN-SM, microfiche!). 

The type locality is Narainhetty but Don gives the flowering time 
as April. No specimens with this date have been located. 

Maesa tomentosa D. Don, Prodr.fl. nepal: 148 (1825). 

Baebotrys tomentosa Buch.-Ham., in sched. 

Churiaghant Hills, 31 March 1802 (BM!; Sheet 346.10 LINN-SM, 

microfiche !-syntypes). 
= Maesa macrophylla (Wall.) A. DC. 

Maesa viridiflora Buch.-Ham., in sched. 

Pherphing, 20 March 1803 (Sheet 392.6 LINN-SM, microfiche!). 

Manglilla bilrhi Buch.-Ham., in sched. 
Narainhetty, 17 October 1802 (LINN microfiche!). 
= Myrsine capitellata Wall. 

Myrsine excelsa D. Don, Prodr.fl. nepal.: 147 (1825). 
Manglillal [no species given] Buch.-Ham., in sched. 
Narainhetty, 17 October 1802 (BM!-syntype). 
= Myrsine capitellata Wall. 

Myrsine potama D. Don, Prodr.fl. nepal.: 146 (1825). 

Samara? potama Buch.-Ham., in sched. 

'an Narainhetty Nepaliae. Floret Februario'. (Syntype: specimen 

not found). 
= Myrsine africana L. 

Myrsine sessilis D. Don, Prodr.fl. nepal: 146 (1825). 
Samara sessilis Buch.-Ham., in sched. 

'in Nepaliae sylvis ad Narainhetty. Floret Octobri'. (Syntype: speci- 
men not found). 
= Myrsine semiserrata Wall. 

Myrsine subspinosa D. Don, Prodr.fl. nepal.: 147 (1825). 
Samara? subspinosa Buch.-Ham., in sched. 
Narainhetty, 1 February 1803 (BM!-syntype) . 
= Myrsine semiserrata Wall. 

OCHNACEAE 

Ochnapumila [Buch.-Ham. ex] DC., Prodr. 1: 736 (1824). 
Terriany forest, Nepalia, 30 March 1802 (BM!-syntype). 
= Ochna obtusata var. pumila (DC.) Kanis 

OLACACEAE 

Schoepfia fragrans Wall., D. Don, Prodr.fl. nepal: 145 (1825). 
Symphoricarpos ?odoratus Buch.-Ham., in sched. 
Narainhetty, 28 September 1802 (BM!). 

OLEACEAE 

Jasminum dichotomum D. Don, Prodr. fl. nepal.: 105 (1825), non 

Vahl (1790). 

Mogorium dichotomum Buch.-Ham., in sched. 
Sembu, 26 May 1802 (BM!-syntype). 
= Jasminum nepalense Spreng. 

Jasminum dispermum Wall., D. Don, Prodr.fl. nepal: 105 (1825). 
Jasminum latifolium Buch.-Ham., in sched. 
Tancote, 15 April 1802 (BM!). 



Jasminum heterophyllum Roxb., D. Don, Prodr. fl. nepal: 106 

(1825). 

Jasminum arboreum Buch.-Ham., in sched. 
Suembu, 30 April 1802 (BM!-syntype?). 
= Jasminum subhumile W.W. Sm. 

Roxburgh in Fl ind. 1: 411 (1820) based this name on material 
introduced to Calcutta Botanic Garden by Buchanan. The BM 
specimen may represent an original voucher of this material and is 
possibly a syntype. 

Jasminum pubescens (Retz.) Willd., D. Don, Prodr.fl. nepal: 105 

(1825). 

'in Nepalia ad Narainhetty. Floret Octobri' (Specimen not found). 
A specimen at BM from Narainhetty, 28 October 1802, is anno- 
tated by Buch.-Ham. 'Mogorium hirsutum Plant. Luccass.' This 
may be the specimen referred to by Don, despite his omission of the 
epithet. 
= Jasminum multiflorum (Burm.f.) Andrews 

Ligustrum bracteolatum D. Don, Prodr.fl. nepal: 107 (1825). 
Ligustrum? japonicum? Buch.-Ham., in sched. 
Suembu, 17 June 1802 (BM!-syntype?). 
= Ligustrum nepalense Wall. 

Buchanan-Hamilton, Sambu [sic] 17 & 18 June 1802 are given as 
syntypes by P.S. Green in Kew Bull 50: 380 (1995). The specimen 
dated 1 8 June has not been located. 

Ligustrum bracteolatum is based on a manuscript name in Herb. 
Lambert but no sheet with this annotation has been found. 

Ligustrum spicatum [Buch.-Ham. ex] D Don, Prodr.fl. nepal: 107 

(1825), nom. superfl. 

'in Nepaliae montosis' (BM?). Material on loan: not seen. 
= Ligustrum nepalense Wall. 

Notelaea posua D. Don, Prodr.fl. nepal: 107 (1825). 

Olea posua Buch.-Ham., in sched. 

Narainhetty, 18 October 1802 (BM!-syntype). 

= Osmanthus fragrans var. longifolius (DC.) H. Kara 

ONAGRACEAE 

Epilobium brevifolium D. Don, Prodr.fl. nepal: 222 (1825). 

Epilobium montanum L.? Buch.-Ham., in sched. 

'in Nepalia superiore' (BM?-syntype). Material on loan: not seen. 

ORCHIDACEAE 

Aerides calceolaris [Buch.-Ham. ex] Sm. in Rees, Cycl. 39: Aerides 

n. 11 (1819). 

Gastrochilus calceolaris (Sm.) D. Don, Prodr.fl. nepal: 32 (1825). 
Epidendrum calceolare Buch.-Ham., in sched. 
Narainhetty, 15 February 1803 (Sheet 1404.29 LINN-SM, micro- 

fiche!-syntype). 
= Gastrochilus calceolaris (Sm.) D. Don 

Aerides dasypogon Sm. in Rees, Cycl. 39: Aerides n. 10(1819). 
Napaul (Sheet LINN-SM, microfiche !-syntype). 
= Gastrochilus dasypogon (Sm.) O. Kuntze 

Cited as holotype by Ji in Guihaia 16: 142 (1996). 

Aerides rigida [Buch.-Ham. ex] Sm. in Rees, Cycl 39: Aerides n. 12 

(1819). 

Napaul (Sheet LINN-SM, microfiche !-syntype). 
= Acampe rigida (Sm.) P.P. Hunt 



116 



J.R. PRESS AND K.K. SHRESTHA 



Aerides spicatum D. Don, Prodr.fl. nepal.: 31 (1825). 
Epidendrum hippium Buch.-Ham., in sched. 
Suembu, 22 June 1802 (BM!; Sheet 1404.28 LINN-SM, micro- 
fiche !-syntypes). 
= Rhynchostylis retusa (L.) Blume 

Bletia bicallosa D. Don, Prodr.fl. nepal.: 30 (1825). 

Limodorum bicallosum Buch.-Ham., in sched. 

'in Nepalia' [No original label]. (BM!-syntype); Terriany forest, 30 

March 1802 (Sheet 1402.8 LINN-SM, microfichel-syntype). 
= Eulophia bicallosa (D. Don) P.P. Hunt & Summerh. 

Bletia dabia D. Don, Prodr.fl. nepal.: 30 (1825). 

Limodorum dabia Buch.-Ham., in sched. 

'in Nepalia' [No original label]. (BM!-syntype); Gorasan, Lower 

Nepal, 30 January 1802 (Sheet 1402.6 LINN-SM, microfiche!- 

syntype). 
= Eulophia dabia (D. Don) Hochr. 

Bletia graminifolia D. Don, Prodr.fl. nepal: 29 (1825). 

Limodorum graminifolium Buch.-Ham., in sched.; 

Suembu, 15 July 1802 (BM! Sheet 1397.38 LINN-SM, microfiche!- 

syntypes);Narainhetty, 8 August 1802 (Sheet 1397.39 LINN-SM, 

microfiche!). 
= Arundina graminifolia (D. Don) Hochr. 

Bletia masuca D. Don, Prodr.fl. nepal: 30 (1825). 

Zoduba masuca Buch.-Ham., in sched. 

Narainhetty, 21 February 1803 (BM!; Sheet 1403.11.1 LINN-SM, 

microfiche !-syntypes). 
= Calanthe masuca (D. Don) Lindl. 

Epidendrum angustifolium Swartz, Prodr.: 123 (1783). 
Narainhetty, 10 October 1802 (Sheet 1397.35.1 LINN-SM, micro- 
fiche!). 

Epidendrum bifarium Swartz, Schrad. J. Bot. 2: 212 (1799). 

Pholiota imbricata Buch.-Ham., in sched. 

Suembu, 16 June 1802 (Sheet 1397.25.1 LINN-SM, microfiche!). 

Epidendrum cuybua Buch.-Ham., in sched. 

Suembu, 22 June 1802 (Sheet 1404.17.1 LINN-SM, microfiche!). 

Epidendrum damun-sultea Buch.-Ham., in sched. 

Suembu, 15 May 1802 (Sheet 1404.21 LINN-SM, microfiche!). 

Epidendrum geniculatum Buch.-Ham. in Hook.f, Fl Brit. India 6: 

45 (1890), nom. nud. in syn. 
Epidendrum gemellum Buch.-Ham., in sched. 
Hettaura, 2 April 1802 (Sheet 1404.18 LINN-SM, microfiche!). 
= Aerides multiflora Roxb. 

Epidendrum humile Sm., Exot. hot. 2: 75, t. 98 (1806). 
'Upper Nepal' (Sheet LINN-SM, microfiche-syntype). 
= Pleione humilis (Sm.) D. Don 

Epidendrum praecox Sm., Exot. hot. 2: 73, t. 97 (1806). 
Narainhetty, 30 October 1802 (Sheet 1404.24.1 LINN-SM, micro- 

fichei-syntype). 
= Pleione praecox (Sm.) D. Don 

According to the protologue, Smith's epithet is based on a 
Buchanan-Hamilton specimen from 'Upper Nepal' . No other data is 
given. 



Epidendrum teres Thunb., Buch.-Ham., in sched. 
Narainhetty, 10 January 1803 (Sheet 1404.22 LINN-SM, micro- 
fiche!). 

Habenaria uniflora D. Don, Prodr.fl. nepal: 25 (1825). 
Orchis uniflora Buch.-Ham., in sched. 

'in Nepalia' [No original label]. (BM!-syntype). Narainhetty, 15 
August 1802 (Sheet 1381.2.1 LINN-SM, microfiche !-syntype). 
= Pecteilis triflora (D. Don) T. Tang & F.T. Wang 

Limodorum asperifolium Buch.-Ham., in sched. 
Narainhetty, 11 November 1802 (Sheet 1397.37 LINN-SM, micro- 
fiche!). 

Limodorum lechmana Buch.-Ham., in sched. 

Gorasan, 31 January 1802 (Sheet 1402.7 LINN-SM, microfiche!). 

Malaxis cordifolia Sm. in Rees, Cycl. 22: Malaxis n. 12(1812), non 
Liparis cordifolia Hook.f. (1889). 

Ophrys monophylla L., Buch.-Ham., in sched. 

Narainhetty, 2 October 1802 (Sheet 1396.15 LINN-SM, micro- 
fichel-syntype). 

= Liparis petiolata (D. Don) P.P. Hunt & Summerh. 

Malaxis ensiformis Sm. in Rees, Cycl 22: Malaxis n. 14 (1812). 

Pinalia ensiformis Buch.-Ham., in sched. 

Narainhetty, 30 November 1802 ( Sheet 1396.11.1 LINN-SM, mi- 
crofiche ! -syntype?) . 

= Oberonia ensiformis (Sm.) Lindl. 

In the protologue, Smith cites a Buchanan-Hamilton specimen 

dated 13 th November 1802. The only specimen at LINN matching 

the other protologue details is that cited above but which is dated 30 

November 1802. 

Malaxis lancifolia Sm. in Rees, Cycl 22: Malaxis n. 7 (1812). 

Ophrys egaleata Buch.-Ham., in sched. 

Suembu, 17 July 1802 (Sheet 1396.5 LINN-SM, microfiche !- 

syntype). 
= Liparis nervosa (Thunb.) Lindl. 

Malaxis latifolia Sm. in Rees, Cycl. 22: Malaxis n. 3 (1812). 
Pinalia trifida Buch.-Ham., in sched. 

Narainhetty, 12 th August 1802 (Sheet 1396.3.1 LINN-SM, micro- 
ficheJ-syntype). 

Neottiaflexuosa Sm. in Rees, Cycl 24: Neottia n. 9 (1813). 
Ophrys spiralis Buch.-Ham., in sched. 

Suembu, 1 May 1802 (Sheet 1389.3 LINN-SM microficheJ-syntype). 
= Spiranthes sinensis (Pers.) Ames 

Neottia parviflora Sm. in Rees, Cycl 24: Neottia n. 10 (1813). 
Ophrys spiralis flore purpureo, Buch.-Ham., in sched. 
Suembu, 23 July 1802 (Sheet 1389.4.1 LINN-SM, microficheJ- 
syntype). 
= Spiranthes sinensis (Pers.) Ames 

Octomeria spicata D. Don, Prodr.fl. nepal: 31 (1825). 

Pinalia alba Buch.-Ham., in sched. 

Narainhetty, 15 August 1802 (BM!; Sheet 1396.16.1 LINN-SM, 

microfiche ! -sy ntypes) . 
= Eria spicata (D. Don) Hand.-Mazz. 

Ophrys alata L.f.?, Buch.-Ham., in sched. 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 

PINACEAE 



117 



Narainhetty, 1 September 1802 (Sheet 1386.21 LINN-SM, micro- 
fiche!). 

Ophrys cernua L.?, Buch.-Ham., in sched. 

Gorasan, 31 January 1802 (Sheet 1386.20 LINN-SM, microfiche!). 

Ophrys sulcata Roxb., Buch.-Ham., in sched. 

Gorasan, 31 January 1802 (Sheet 1386.22 LINN-SM, microfiche!). 

Orchis bicornuta Buch.-Ham., in sched. 

Narainhetty, 17 September 1802 (Sheet 1383.5.1 LINN-SM, micro- 
fiche!). 

Orchis flexuosa L.f., Buch.-Ham., in sched. 

Suembu, 17 July 1802 (Sheet 1381.98 LINN-SM, microfiche!). 

Orchis gigantea Sm., Exot. hot. 2: 79, t. 100 (1806). 
'Upper Nepal' (Sheet LINN-SM, microfiche-syntype). 
= Pecteilis susannae (L.) Raf. 

Orchis obcordata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 23 

(1825), nonWillem. ( 1796). 
'in Napaliae alpibus' (BM!-syntype); Suembu, 23 July 1802 (Sheet 

1381.86.1 LINN-SM, microfiche-syntype). 
= Brachycorythis obcordata (Lindl.) Summerh. 

Orchis pectinata [Buch.-Ham. ex] Sm., Exot. hot. 2: 77, t. 99 (1805). 
Secumbu, 18 July 1802 (Sheet 1381.22.1 LINN-SM, microfiche !- 

syntype). 
= Habenaria pectinata D. Don 

Satyrium latifolium Buch.-Ham., in sched. 

Narainhetty, 5 August 1802 (Sheet 1381.97 LINN-SM, microfiche). 

Stelis biflora Sm. in Rees, Cycl. 34: Stelis n. 13 (1816). 
'On the mossy banks of Upper Napaul'. 

H. Kara in Enum. fl. pi. Nepal 1: 56 (1978) noted that The 
application of this name is uncertain. There is no specimen in the 
Smith herbarium at the Linnean Society of London'. 

Stelis hirta Sm. in Rees, Cycl. 34: Stelis n. 11 (1816). 
Sunipia hirta Buch.-Ham., in sched. 

Narainhetty, 30 January 1803 (Sheet 1405.4 LINN-SM, micro- 
fiche! -syntype). 
= Bulbophyllum hirtum (Sm.) Lindl. ex Wall. 

Stelis mucronata D. Don, Prodr. fl. nepal.'. 32 (1825). 
Oberonia iridifolia Lindl., in Wall. (1829) 
'in Nepalia' [No original label]. (BM!-syntype). 
= Oberonia ensiformis (Sm.) Lindl. 

Stelis odoratissima Sm. in Rees, Cycl. 34: Stelis n. 12 (1816). 

'Native of mossy rocks in Upper Nepaul'. 

= Bulbophyllum odoratissimum (Sm.) Lindl. ex Wall. 

Stelis racemosa Sm. in Rees, Cycl. 34: Stelis n. 10 (1816). 

'on trees in Upper Napaul'. 

= Sunipia racemosa (Sm.) T. Tang & FT. Wang 

PALMAE 

Elate bulbifera Buch.-Ham., in sched. 

Baguanpur, 16 March 1802 (Sheet 1612.11 LINN-SM, microfiche!). 

= Phoenix cf. acaulis Roxb. 



Pinus excelsa Wall, ex D. Don in Lamb, Descr. Pinus 2: 5, t. 3 

( 1 824), non Lam. (1779). 
Pinus cembra L?, Buch.-Ham., in sched. 
Narainhetty, 9 November 1802 (BM!-syntype). 
= Pinus wallichiana A.B. Jacks. 

PIPERACEAE 

Piper guigual [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 20 (1825). 
Narainhetty, 6 February 1803 (BM!-syntype). 
= Piper mullesua [Buch.-Ham. ex] D. Don 

Piper mullesua [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 20 

(1825). 
Narainhetty, 17 January 1803 (BM!-syntype). 

Piper suipigua [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 20 

(1825). 
Narainhetty, 21 August 1802 (BM!-syntype). 

PLANTAGINACEAE 

Plantago erosa Wall., D. Don, Prodr. fl. nepal.: 11 (1825). 
Plantago filiformis Buch.-Ham., in sched. 
Suembu, 19 April 1802 (BM!). 

POLYGALACEAE 

Polygala arillata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 199 

(1825). 
'in Nepalia ad Narainhetty. Floret Augusto' . (Syntype: specimen not 

found). 

Polygala buchanani D. Don, Prodr. fl. nepal.: 199 (1825), nom. 

superfl. 

Polygala monspeliaca Buch.-Ham., in sched. 
Narainhetty, 1 September 1802 (BM!). 
= Polygala persicariifolia DC. 

Polygala crotalarioides [Buch.-Ham. ex] DC., Prodr. 1 : 327 ( 1 824). 
'in Napalia' [No original label]. (BM!-syntype). 

Polygala discolor [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 199 

(1825), nom. superfl. 

'in Nepalia superiore' [No original label]. (BM!). 
= Polygala longifolia Poir. 

Polygala triphylla [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 200 

(1825), non Burm.f. (1768). 
Narainhetty, 1 September 1802 (BM!-syntype). 
= Polygala furcata Royle 

Salomonia petiolata in D. Don, Prodr. fl. nepal.: 200 (1825), nom. 

superfl. 

'in Nepalia' [No original label]. (BM!). 
= Salomonia cantoniensis Lour. 

POLYGONACEAE 

Coccoloba totnea [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 74 

(1825). 

'in Nepalia'. (Syntype: specimen not found). 
= Aconogonum molle (D. Don) H. Kara 



118 



J.R. PRESS AND K.K. SHRESTHA 



Polygonum capitatum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 73 

(1825). 

'in Nepalia'. (Syntype: specimen not found). 
= Persicaria capitata (D. Don) H. Gross 

Polygonum dibotrys D. Don, Prodr.fl. nepal.: 73 (1825). 

Polygonum fagopyrum Buch.-Ham., in sched. 

'in Nepalia ad Narainhetty. Floret Septembri' (BM?-syntype). 

Material on loan: not seen. 
= Fagopyrum dibotrys (D. Don) H. Hara 

Polygonum posumbu [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 71 

(1825). 

'in Nepalia'. (Syntype: specimen not found). 
= Persicaria posumbu (D. Don) H. Gross 

Polygonum runcinatum [Buch.-Ham. ex] D. Don, Prodr. fl. nepal. : 

73 (1825). 

'in Nepalia' (Syntype: specimen not found). 
= Persicaria runcinata (D. Don) H. Gross 

Polygonum viscosum [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 71 
(1825). 

'ad Suembu Nepaliae superioribus. Floret Aprili'. (Syntype: speci- 
men not found). 

= Persicaria viscosa (D. Don) Nakai 

PRIMULACEAE 

Anagallis alternifolia Buch.-Ham., in sched. 

Sembu, 25 May 1802 (Sheet 284.11 LINN-SM, microfiche). 

Anagallis caerulea L., Buch.-Ham., in sched. 

Nepal, ann. 1802 (Sheet 284.7 LINN-SM, microfiche!). 

Anagallis mauritiana Buch.-Ham., in sched. 
Sembu, 1 1 June 1802 (Sheet 284.8.1 LINN-SM, microfiche); Nepal, 
9 July 1802 (Sheet 284.8.2 LINN-SM, microfiche!). 

Anagallis multiangularis Buch.-Ham. in Hook.f., Fl. Brit. India. 3: 

503 (1882), nom. nud. in syn. 

Sembu, 26 May 1802 (Sheet 284.9 LINN-SM, microfiche!). 
= Lysimachia pyramidalis Wall. 

Anagallis teres Buch.-Ham., in sched. 

Sembu, 18 May 1802 (Sheet 284.10 LINN-SM, microfiche!). 

Androsace rotundifolia sensu Sm., Exot. Bot. 2: 107, t. 113 (1806), 

nonHardw. (1795). 

Androsace villosa? Buch.-Ham., in sched. 
Narainhetty, 8 January 1802 (BM!-syntype); Nepal (Sheet 270. 15 1 

LINN-SM, microfiche-syntype). 
= Androsace umbellata (Lour.) Merr. 

Lysimachia secunda [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 84 

(1825), nom. superfl. 
Sembu, 5 June 1802 (BM!); Sembu, 2 June 1802 (Sheet 282.14 

LINN-SM, microfiche!). 
= Lysimachia lobelioides Wall. 

Lysimachia tetragona D. Don, Prodr.fl. nepal.: 83 (1825), nom. 

superfl. 
Lysimachia quinquangularis Buch.-Ham., in sched. 



Narainhetty, 8 August 1802 (BM!). 
= Lysimachia alternifolia Wall. 

Lysimachia triangularis Buch.-Ham., in sched. 

Sembu, 10 June 1802 (Sheet 282.15 LINN-SM, microfiche). 

Primula denticulata Sm., Exot. hot. 2: 109, t. 114 (1806). 

Primula elata Buch.-Ham., in sched. 

Chitlong, 10 April 1802 (Sheet 271.2.2 LINN-SM, microfiche!- 
syntype); Chitlong, 1 1 April 1802 (BM!-syntype); Narainhetty, 5 
February 1803 (Sheet 271.2.1 LINN-SM, microfiche !-syntype). 

Primula tridentata D. Don, Prodr. fl. nepal.: 77 (1825), nom. 

superfl. 

Primula cushia Buch.-Ham., in sched. 
Narainhetty, 10 February 1803 (BM! Sheet 271.13.1 LINN-SM, 

microfiche!); Narainhetty, 3 March 1803 (LINN microfiche!); 

source of the Bagmutty, Nepal, 28 December 1 802 (Sheet 27 1 . 1 2. 1 

LINN-SM, microfiche!). 
= Primula petiolaris Wall. 

Wallich in Roxburgh, Fl. ind. 2: 22 (1824), cites no specimen but 
states 'I have had frequent supplies of specimens of this very 
distinctive Primrose from the vicinity of Katumanda and from 
Gosain-Than'. It is possible that Buchanan's Narainhetty material 
could constitute part of these supplies. 

RANUNCULACEAE 

Anemone rivularis [Buch.-Ham. ex] DC., Syst. not. 1: 211 (1817). 
Chitlong, 12 April 1802 (BM!; Sheet 972.40 LINN-SM, micro- 

fichei-syntypes). 

Anemone vitifolia [Buch.-Ham. ex] DC., Syst. nat. 1: 211 (1817). 
Sembu, 18 July 1 802 (BM!; Sheet 972.39.2 LINN-SM, microfiche !- 

syntypes); Narainhetty, 1 September 1802 (Sheets 972.38 & 

972.39.1 LINN-SM, microfiche!). 

DeCandolle correctly cites the flowering time as July but gives 
the locality as Lamba. This is almost certainly a result of misreading 
Buchanan-Hamilton's handwritten Suembu. 

Clematis acuminata DC., Syst. nat. 1: 148 (1817). 
Clematis trinervis Buch.-Ham., in sched. 

Narainhetty, 31 December 1802 (BM!; Sheet 974.21 LINN-SM, 
microficheJ-syntypes). 

Clematis buchananiana [Buch.-Ham. ex] DC., Syst. nat. 1: 140 
(1817). 

Clematis bucamara Buch.-Ham., in sched. 

Narainhetty, 30 October 1802 (BM!-syntype); Narainhetty, 26 Oc- 
tober 1802 (Sheets 974.23 & 974.24 LINN-SM, 
microfiche!-syntype). 
DeCandolle gives the locality as 'Harain-Hetty'. 

Clematis grewiiflora DC., Syst. nat. 1: 140 (1817), 'grewiaeflora'. 

Clematis vitalba? Buch.-Ham., in sched. 

Narainhetty, 27 November 1802 (BM!; Sheet 974.22 LINN-SM, 

microfiche! -syntypes); Narainhetty, 14 November 1802 (Sheet 

974.26 LINN-SM, microfiche !-syntypes). 

Clematis loasaefolia DC., Syst. nat. 1: 140 (1817). 

Clematis bucamara var. loasaefolia Buch.-Ham., in sched. 

Narainhetty, 30 October 1802 (BM!-syntype); Narainhetty, 26 Oc- 
tober 1802 (Sheet 974.27 LINN-SM, microfiche !-syntype). 

= Clematis grewiiflora DC. 

DeCandolle gives the locality as 'Harain-Hetty'. 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



119 



Clematis montana [Buch.-Ham. ex] DC., Syst. nat. 1: 164 (1817). 
Clematis anemoniflora D. Don, Prodr.fl. nepal.: 192 (1825), nom. 

superfl. 
Chitlong, 13 April 1802 (BM!-syntype?); Chitlong, 11 April 1802 

(Sheet 974.25 LINN-SM, microfiche,-syntype?). 

Clematis napaulensis DC., Syst. nat. 1: 164 (1817). 

Clematis diphylla Buch.-Ham., in sched. 

'Napaulia' [No original label]. (BM!-syntype?); Narainhetty, 2 
February 1803 (Sheet 974. 19 LINN-SM, microfichei-syntype?); 
Narainhetty, 3 February 1803 (Sheet 974.20 LINN-SM, micro- 
fiche !-syntype?). 

Clematis montana D. Don, Prodr. fl. nepal: 192 (1825), nom. 
superfl. 

Clematis montana var. ? Buch.-Ham., in sched. 

'in montanis ? Nepaliae' [No original label]. (BM!-syntype?). 

Delphinium carela [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 195 

(1825), nom. superfl. 
Narainhetty, 10 August 1802 (Sheets 965.7 & 965.8 LINN-SM, 

microfiche!). 
= Delphinium scabriflorum D. Don 

Delphinium pauciflorum D. Don, Prodr. fl. nepal.: 196 (1825). 

Delphinium consolida? Buch.-Ham., in sched. 

Catmandu, 9 May 1802 (Sheet 965.5 LINN-SM, microfichel- 
syntype); Narainhetty, 28 February 1 803 (Sheet 965.6 LINN-SM, 
microfiche ! -sy ntype) . 

Ranunculus indicus Roxb., Buch.-Ham., in sched. 
Napaul, ann. 1802 (Sheet 987.23 LINN-SM, microfiche). 
= Ranunculus sceleratus L. 

Ranunculus sceleratus L., D. Don, Prodr.fl. nepal: 195 (1825). 
'in Nepaliae inundatis'. (Specimen not found). 

Ranunculus ternatus Thunb., Buch.-Ham., in sched. 

Chitlong, 11 April 1802 (Sheet 987.24 LINN-SM, microfiche!). 

Ranunculus umbellatus Roxb., Buch.-Ham., in sched. 
Ranunculus buchiana Buch.-Ham., in sched. 
(BM!). No date or locality on original label. 
= Ranunculus sceleratus L. 

Thalictrum foliolosum DC., Syst. nat. 1: 175 (1817). 

Thalictrum dalingo Buch.-Ham., in sched. 

Suembu, 1 May 1802 (BM!; Sheet 984.16 LINN-SM, microfiche!- 

syntypes); Suembu, 15 May 1802 (Sheet 984.17 LINN-SM, 

microfiche ! -syntype) . 

Thalictrum rotundifolium DC., Syst. nat. 1: 185 (1817). 

Thalictrum batula Buch.-Ham., in sched. 

(BM!-syntype) No date or locality on original label; Narainhetty, 10 

August 1802 (Sheet 984.18 LINN-SM, microfiche! -syntype); 

Sembu, 21 July 1802 (Sheet 984.19 LINN-SM, microfiche!- 

sy ntype). 

RHAMNACEAE 

Hovenia dulcis sensu Roxb., Fl. ind. 2: 414 (1824), non Thunb. 

(1798). 

Narainhetty, 13 November 1802 (BM!). 
= Hovenia acerba Lindl. 



Roxburgh cites material introduced to the Calcutta Botanic Gar- 
den by Buchanan and collected on November lOand 14 1802, but 
not on November 13th. 

Rhamnus trigynus D. Don, Prodr. fl. nepal: 190 (1825), nom. 

superfl. 

Rhamnus terminalis Buch.-Ham., in sched. 
Narainhetty, 3 September 1802 & 10 September 1802 (BM!). 
= Sageretia filiformis (Roth ex Schult.) G. Don 

Rhamnus virgatus Roxb., Fl ind. 2: 351 (1824). 
Rhamnus catharticus L.?, Buch.-Ham., in sched. 
Sembu, 18 April 1802 (BM! synype?). 

Roxburgh cites material grown from seeds introduced to the 
Calcutta Botanic Garden by Buchanan. 

Zizyphus incurva Roxb., Fl ind. 2: 364 (1824). 
Zizyphus paniculata Buch.-Ham., in sched. 
Sembu, 18 May 1802 (BM!-syntype?). 

Roxburgh cites material introduced to the Calcutta Botanic Gar- 
den by Buchanan. 

ROSACEAE 

Agrimonia nepalensis D. Don, Prodr.fl. nepal: 229 (1825). 

Agrimonia repens L.?, Buch.-Ham., in sched. 

'in Nepalia". (Syntype: specimen not found). 

= Agrimonia pilosa var. nepalensis (D. Don) Nakai 

Comarumflavum [Buch.-Ham. ex] Roxb., Fl. ind. ed. 1832 2: 521 

(1832). 

Bassaria, 13 March 1802 (Sheet 908.3 LINN-SM, microfiche!). 
= Potent ilia supina L. 

Cotoneaster affinis Lindl. in Trans. Linn. Soc. London 13: 101 

(1821). 

Mespilus affinis (Lindl.) D. Don, Prodr.fl. nepal: 238 (1825). 
Crataegus cotoneaster Buch.-Ham., in sched.; Mespilus integerrima 

Buch.-Ham., in sched. 
Chitlong, 14 April 1802; Chitlong, 15 April 1802 (BM!-syntypes). 

Lindley mistakenly cites the locality as Chittong, a misreading of 
Buchanan-Hamilton's handwritten Chitlong. 

Cydonia sumboshia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 237 

(1825). 

Narainhetty, 11 March 1803 (BM! -syntype). 
= Cydonia oblonga Mill. 

Eriobotrya elliptica Lindl. in Trans. Linn. Soc. London 13: 102 

(1821). 
Mespilus cuila [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 238 

(1825), nom. superfl. 

Narainhetty, 1 February 1803 (BM!-syntype). 
= Eriobotrya elliptica Lindl. 

Fragaria tuberosa Buch.-Ham., in sched. 

Hettaura, Nepal, 1 April 1802 (Sheet 904.1 LINN-SM, micro- 
fiche!). 

Mespilus tinctoria D. Don, Prodr. fl. nepal: 238 (1825), nom. 

superfl. 

Crataegus? shicola Buch.-Ham., in sched. 
'ad Narainhetty Nepaliae. Floret Novembri'. (Specimen not found). 
= Eriobotrya dubia (Lindl.) Decne. 



120 



J.R. PRESS AND K.K. SHRESTHA 



Neillia thrysiflora D. Don, Prodr.fl. nepal.: 228 (1825). 

'in Nepalia'. (Syntype: specimen not found). 

Potentilla exarata Sm., in sched.; Potentilla rupestris? Buch.-Ham., 
in sched. 

Suembu, 20 July 1802 (Sheet 903.8.1 LINN-SM, microfiche); 
Narainhetty, 8 August 1802 (Sheet 903.8.2 LINN-SM, micro- 
fiche). 

Potentilla opaca L., Buch.-Ham., in sched. 

Narainhetty, 6 March 1803 (Sheet 903.15 LINN-SM, microfiche). 

Potentilla splendens Wall, ex D. Don, Prodr.fl. nepal.: 230 (1825). 

Potentilla naspata Buch.-Ham., in sched. 

Mn Nepalia alpibus' [No original label]. (BM!-syntype). 

[Potentilla} rupestris? Buch.-Ham., in sched. 

Suembu (Sheet 903.59 LINN-SM, microfiche!). 

= Potentilla fulgens Wall, ex Hook. 

Potentilla verna L., Buch.-Ham., in sched. 

Sembu, 15 July 1802 (Sheet 903.16 LINN-SM, microfiche!). 

Primus cerasoides D. Don, Prodr.fl. nepal.: 239 (1825). 
Cerasus phoshia Buch.-Ham., in sched. 
Narainhetty, 26 October 1802 (BM!-syntype). 

Prunus undulata [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 239 

(1825). 
Narainhetty, 14 November 1802 (BMMectotype). 

Lectotype designated by H. Hara in J. Jap. Dot. 48: 97 (1973). 

Pyrus crenata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 237 
(1825), nonLindl. (1835). 

'ad Suembu Nepaliae superioribus. Floret Julio'. (Syntype: speci- 
men not found). 

Pyrus nussia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 237 (1825). 
Nilcunt, 26 April 1802 (BM!-syntype). 
= Stranvaesia nussia (D. Don) Decne. 

Pyrus pashia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 236 

(1825). 
Narainhetty, 6 November 1 802 (BM!-syntype); Narainhetty, 3 March 

1803(BM!-syntype). 

Rosa brunonii Lindl., Ros. monogr: 120, t. 14 (1820). 
Rosa glutinosa Buch.-Ham., in sched. 
Sambu, Nepalia, 18 April 1802 (BM!-syntype). 

Rosa involucrata Roxb. ex Lindl., Ros. monogr.: 8 (1820). 
Rosa palustris Buch.-Ham., in sched. 
'in Nepalia' (BM!-syntype). 
= Rosa clinophylla Thory 

The original label gives the taxon name but no other data. 

Rubus acuminatus Sm. in Rees, Cycl. 30: Rubus n. 43 (1819). 
Sembu, 4 July 1802 (Sheet 902.92 LINN-SM, microfiche !-syntype). 

Rubus betulinus D. Don, Prodr.fl. nepal.: 233 (1825). 
Rubus triflorus Buch.-Ham., in sched. 
Sembu, 8 July 1802 (BM!-syntype). 
= Rubus acuminatus Sm. 

Rubus biflorus [Buch.-Ham. ex] Sm. in Rees, Cycl. 30: Rubus n. 9 
(1819). 



Chitlong, Upper Napaul, 13 April 1802 (Sheet 902.66 LINN-SM, 
microfiche !-syntype). 

H. Hara, in Enum.fl. pi. Nepal 2: 144 (1979), noted that The type 
specimen differs from the common form of the species illustrated in 
B. Mag. t. 4678 in having smaller leaflets, smaller flowers, and 
densely pubescent calyces'. 

Rubus ellipticus Sm. in Rees, Cycl. 30: Rubus n. 16(1819). 

Rubus ishia Buch.-Ham., in sched. 

Hettaura, stony banks of rivulets in Nepal, 9 April 1802 (Sheet 

902.71 LINN-SM, microfiche !-syntype). 

A second specimen, dated January 1 802, is cited in the protologue 
but has not been located. 

Rubus flavus [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 234 (1825). 
Rubus parviflorus L., Buch.-Ham., in sched.; Rubus ischia Buch.- 
Ham., in sched. 

Ethaura, 1 April 1802 (BM!-syntype). 
= Rubus ellipticus Sm. 

Rubus hamiltonianus Sen in DC., Prodr. 2: 566 (1825), p.p. 
'ad Suembu Napaliae super' [No original label]. (BM!-syntype). 
= Rubus rugosus Sm. 

Rubus paniculatus Sm., in Rees, Cycl. 30: Rubus n. 41 (1819). 
Narainhetty, 18 November 1802 (Sheet 902.87 LINN-SM, micro- 
fiche ! -syntype). 

Rubus parvifolius sensu Sm. in Rees, Cycl. 30: n. 21 (1819), non L. 

(1751). 
Chittlang, 10 April 1802 (LEG no 7129 LIV-syntype); Nepaul 

(Sheet 902.72 LINN-SM, microficheJ-syntype?). 
= Rubus foliolosus D. Don 

Referring to specimens in the protologue Smith cites Chittlang, 
10 April 1802 but also says 'we have . . . ones' implying there were 
several sheets. 

Rubus pedunculosus D. Don, Prodr.fl. nepal: 234 (1825). 
Rubus biflorus? Buch.-Ham., in sched. 
Chitlong, 10 April 1802 (BM!-syntype). 
= Rubus foliolosus D. Don 

Rubus rugosus Sm. in Rees, Cycl. 30: Rubus n. 34 (1819). 

Rubus rugosus [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 234 
(1825), nom. superfl. 

Rubus cumbata Buch.-Ham., in sched. 

Narainhetty, 11 August 1802 (Sheet 902.88 LINN-SM, micro- 
fiche!); Sembu, 18 July 1802 (LEG no 7131 LIV-syntype); 
Suembu, 18 July 1802 (BM!-syntype). 

Rubus tiliaceus Sm. in Rees, Cycl. 30: Rubus n. 35 (1819). 
Sembu, 2 June 1 802 (Sheet 902.86 LINN-SM, microfiche ! -syntype). 
= Rubus paniculatus forma tiliaceus (Sm.) H. Hara 

RUBIACEAE 

Cuncea trifida [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 135 

(1825). 

Narainhetty, 11 August 1802 (BM!-syntype). 
= Knoxia sumatrensis (Retz.) DC. 

Galium ciliatum [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 133 
(1825), non Ruiz & Pav. (1798). 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



121 



Narainhetty, 6 August 1802 (BM!-syntype). 
= Galium hirtiflorum Req. ex DC. 

Galium latifolium [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 133 

(1825). 

Sembu, 15 July 1802 (BM!-syntype). 
= Galium elegans Wall, ex Roxb. 

Galium parviflorum D. Don, Prodr. fl. nepal.: 133 (1825). 
Narainhetty, 7 August 1802 (BM!-syntype). 
= Galium asperifolium Wall. 

Gardenia rigida [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 138 

(1825). 

Suembu, 30 April 1802 (BM!-syntype). 
= Himalrandia tetrasperma (Roxb.) T. Yamazaki 

Hamiltonia scabra D. Don, Prodr. fl. nepal: 137 (1825). 
Hamiltonia dulina Buch.-Ham., in sched.; Nonatellia? filamentosa 

Buch.-Ham., in sched. 

Narainhetty, 11 October 1802 (BM!-syntype). 
= Spermadictyon suaveolens Roxb. 

Hedyotis fusca [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 134 

(1825), nom. superfl. 

Gorasan, 16 February 1802 (BM!-syntype). 
= Kohautia gracilis (Wall.) DC. 

Hedyotis lineata Roxb., Fl. ind. 1: 369 (1820). 
Spermacoce lineata Buch.-Ham., in sched. 
Narainhetty, 13 October 1802 (BM!). 

Mussaenda hispida D. Don, Prodr. fl. nepal.: 139 (1825). 
Mm saenda f rondo sa L?, Buch.-Ham., in sched. 
Narainhetty, 19 August 1802 (BM!-syntype). 
= Mussaenda macrophylla Wall. 

Mussaenda luculia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal. : 1 39 

(1825), nom. superfl. 
Narainhetty, 7 October 1802 (BM!). 
= Luculia gratissima (Wall.) Sweet 

Ophiorrhiza fasciculata D. Don, Prodr. fl. nepal.: 136 (1825). 
Virecta fasciculata Buch.-Ham., in sched. 
Suembu, 22 June 1802 (BM!-syntype). 

Ophiorrhiza prostrata D. Don, Prodr. fl. nepal: 136 (1825). 
Virecta? prostrata Buch.-Ham., in sched.; Virecta? suffruticosa 

Buch.-Ham., in sched. 

Narainhetty, 21 August 1802 (BM!-syntype). 
= Ophiorrhiza rugosa Wall. 

Don cites the flowering time as June. No specimen with this date 
has been located. 

Randia triflora [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 138 

(1825). 

Hethaura, April 1802 (Syntype: specimen not found). 
= Fagerlindia fasciculata (Roxb.) Tirveng. 

Rondeletia coriacea Wall, in Roxb., D. Don, Prodr. fl. nepal: 138 

(1825). 
Rondeletia asiatica L.?, Buch.-Ham., in sched.; Rondeletia budda 

Buch.-Ham., in sched. 



Narainhetty, 23 February 1803 (BM!). 
= Wendlandia coriacea (Wall.) DC. 

Spermacoce pusilla Wall, in Roxb., D. Don, Prodr. fl. nepal: 134 

(1825). 
'in Nepalia' [No original label]. (BM!). 

RUTACEAE 

Bergera integerrima [Buch.-Ham. ex] Colebr. in Trans. Linn. Soc. 

London 15: 367 (1827). 
Narainhetty, 4 March 1803 (BM!-syntype). 
= Micromelum integerrimum (Colebr.) Wight & Arn. ex M. 

Roem. 

SALICACEAE 

Salix disperma [Roxb. ex] D. Don, Prodr. fl. nepal.: 58 (1825). 
'in Nepalia'. (Syntype: specimen not found). 

A. Kimura, in Enum. fl. pi Nepal 3: 218 (1982) cites Wallich 
3700A as the lectotype of 5. disperma. 

Salix japonica sensu D. Don, Prodr. fl. nepal: 59 (1825), non 

Thunb. (1784). 

Narainhetty, 16 February 1803 (BM!). 
= Salix babylonica L. 

SANTALACEAE 

Osyris chama Buch.-Ham., in sched. 

Suembu, 15 May 1802 (Sheet 1518.4 LINN-SM, microfiche!). 

= Osyris wightiana Wall, ex Wight 

Viscum latifolium [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 142 

(1825), non Lam. (1789). 
Bimpedi Nepaliae, 8 April 1803 (BM!-syntype). 
= Dufrenoya platyphylla (Spreng.) Stauffer 

SAPOTACEAE 

Bassia butyracea Roxb., Buch.-Ham., in sched. 
Narainhetty, 9 November 1802 (Sheet 849.3 LINN-SM, micro- 
fiche!). 
= Diploknema butyracea (Roxb.) HJ. Lam 

Sideroxylon arboreum Buch.-Ham., in sched. 

Narainhetty, 24 July 1803 (Sheet 382.7 LINN-SM, microfiche!). 

SAURAUIACEAE 

Ternstroemia racemosa D. Don, Prodr. fl. nepal: 225 (1825), nom. 
superfl. 

Dillenia racemosa Buch.-Ham., in sched.; Tonshia polypetala Buch.- 
Ham., in sched. 

'ad Narainhetty Nepalensium. Floret Augusto' (BM!). 

The original label gives the Buchanan-Hamilton manuscript 
names but no other data. 

= Saurauia napaulensis DC. 

SAXIFRAGACEAE 

IThalictrum digynum, Buch.-Ham., in sched. 
Narainhetty, 30 August 1802 (Sheets 948.31 & 948.32 LINN-SM, 
microfiche!). 



122 

Astilbe rivularis [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 211 

(1825). 
'ad Narainhetty Nepalensium. Floret Septembri' (Specimen not 

found); Narainhetty, 14 October 1802 (Sheet 984.30 LINN-SM, 

microfiche!). 

The specimen cited by Don ('Floret Septembri') has not been 
found. 

Saxifraga ligulata Wall., D. Don, Prodr. fl. nepal.: 209 (1825). 
Saxifraga pacumbis Buch.-Ham., in sched. 
Narainhetty, 22 January 1803 (BM!). 
= Bergenia ciliata forma ligulata Yeo 

The specimen cited by Don ('Floret Martio') has not been found. 

SCROPHULARIACEAE 

Buchnera cruciata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 91 

(1825). 
Narainhetty, 27 November 1802 (BM!-syntype). 

Buchnera hispida [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 91 

(1825). 
Narainhetty, 21 October 1802 (BM!-syntype). 

Centranthera nepalensis D. Don, Prodr. fl. nepal.: 88 (1825). 
'in Nepalia' (Syntype: specimen not found). 

Cybbanthera connata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal: 87 

(1825). 
'in Nepalia ad Narainhetty. Floret Septembri'. (Syntype: specimen 

not found). 
= Limnophila connata (D. Don) Hand.-Mazz. 

Gmelina speciosissima D. Don, Prodr. fl. nepal.: 104 (1825). 

Gmelina? tacabushia Buch.-Ham., in sched. 

'in Nepalia' (BM!-syntype). 

= Wightia speciosissima (D. Don) Merr. 

The original label gives no date or locality data. 

Gratiola cordifolia Vahl, D. Don, Prodr. fl. nepal.: 85 (1825). 

'in Nepalia'. (Specimen not found). 

= Lindernia anagallis (Burm.f.) Pennell 

Rhinanthus bifldus [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 94 
(1825). 

'ad Narainhetty Nepalensium. Floret Septembri'. (Syntype: speci- 
men not found). 

= Pedicularis hi fid a (D. Don) Pennell 

Sopubia triflda [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 88 

(1825). 

Manulea sopubia Buch.-Ham., in sched. 
Sembu, 12 June 1802 (BM!-syntype). 

Stemodia grandiflora [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 89 

(1825). 
'in Nepalia ad Narainhetty. Floret Octobri'. (Syntype: specimen not 

found). 
= Lindenbergia grandiflora (D. Don) Benth. 

Stemodia muraria Roxb. ex D. Don, Prodr. fl. nepal.: 89 (1825). 
'in Nepalia'. (Syntype: specimen not found). 
= Lindenbergia indica (L.) Vatke 



J.R. PRESS AND K.K. SHRESTHA 

Veronica punctata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal. : 93 

(1825). 
'ad Narainhetty Nepaliae, in scaturiginosis. Floret Novembri'. 

(Syntype: specimen not found). 
= Veronica undulata Wall. 

SIMAROUBACEAE 

Simaba quassioides [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 248 

(1825). 

Nima quassioides Buch.-Ham., in sched. 
Tancote, 16 April 1802 (BM!-syntype). 
= Picrasma quassioides (D. Don) Benn. 

SOLANACEAE 

Physalis divaricata D. Don, Prodr. fl. nepal.: 97 (1825). 
Physalis angulata (var.) Buch.-Ham., in sched. 
'ad pagum Bassaria Nepalensibus dictum. Floret Martio'. (Syntype: 
specimen not found). 

Solanum biflorum sensu D. Don, Prodr. fl. nepal.: 96 (1825), non 

Lour. (1790). 

Solanum multifldum Buch.-Ham., in sched. 
Narainhetty, 19 August 1802 (BM!). 
= Lycianthes macrodon (Wall, ex Nees) Bitter 

SYMPLOCACEAE 

Symplocos crataegoides [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 

145 (1825). 

Palura odorata Buch.-Ham., in sched. 
Tancote, 16 April 1802 (BM!-syntype). 
= Symplocos paniculata (Thunb.) Miq. 

Symplocos loha [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 144 
(1825). 

Symplocos subspinosa Buch.-Ham., in sched. 

Narainhetty, 26 October 1802 (BM!-syntype); Narainhetty, 13 Oc- 
tober 1802 (BM!-syntype); Narainhetty, 17 October 1802 
(BM!-syntype). 

= Symplocos cochinchinensis (Lour.) S. Moore 

Symplocos sumuntia [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 

145 (1825). 
Narainhetty, 9 March 1802 (BM!-syntype). 

Symplocos theifolia D. Don, Prodr. fl. nepal. : 145 (1 825) 'theaefolia' . 
Narainhetty, 3 November 1802 (BM!-syntype). 
= Symplocos lucida (Thunb.) Siebold & Zucc. 

THEACEAE 

Camellia keina [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 224 

(1825), nom. superfl. 
Narainhetty, 9 August 1802 (BM!). 
= Camellia kissi Wall. 

Diospyros serrata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 143 

(1825). 

Ternstroemia bifaria Buch.-Ham., in sched. 
Narainhetty, 30 August 1802 (BM!-syntype). 
= Eurya acuminata DC. 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



123 



Gordonia chilaunea [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 225 

(1825). 

Suembu, 17 May 1802 (BM!-syntype). 
= Schima wallichii (DC.) Korth. 

Temstroemia lushia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 225 

(1825), nom. superfl. 
Suembu, 5 June 1802 (BM!-syntype). 
= Cleyera japonica var. wallichiana (DC.) Sealy 

THYMELAEACEAE 

Dais bamutis Buch.-Ham., in sched. 

Narainhetty, 27 January 1803 (Sheet 686.28 LINN-SM, micro- 
fiche). 
= Edgeworthia gardneri (Wall.) Meisn. 

Daphne bholua [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 68 
(1825). 

Daphne botlua Buch.-Ham., in sched. 

Narainhetty, 25 January 1803 (BM!; Sheet 686.31 LINN-SM, mi- 
crofiche ! -sy ntypes) . 

Daphne cannabina Lour, ex Wall., Asiat. Res. 13: 385 (1820). 

Daphne odora sensu D. Don, Prodr. fl. nepal.: 68 (1825), non 
Thunb. (1784). 

Daphne papyri/era [Buch.-Ham. ex] Meissn. in DC., Prodr. 14: 537 
(1857). 

Narainhetty, 12 January 1803 (Sheet 686.30.1 LINN-SM, micro- 
fiche). 

Chitlong, 12 April 1802 (Sheet 686.30.2 LINN-SM, microfiche). 

= Daphne papyracea Wall, ex Steud. 

Daphne sericea D. Don, Prodr. fl. nepal.: 69 (1825), non Vahl 
(1790). 

Daphne oppositifolia Buch.-Ham., in sched. 

Narainhetty, 13th August 1802 (BM! syntype; Sheet 686.35 LINN- 
SM, microfiche !-syntype). 

= Wikstroemia canescens Meisn. 

TILIACEAE 

Grewia asiatica ?, Buch.-Ham., in sched. 

Gorasan, 22 March 1802 (Sheet 1423.17 LINN-SM, microfiche!). 

= Grewia helicterifolia Wall, ex G. Don 

Grewia oppositifolia [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 221 

(1825). 
Sumbu, 18 May 1802 (Sheet 1423.16 LINN-SM, microfiche!- 

syntype). 
= Grewia optiva J.R. Drumm. ex Burrett 

Grewia pumila [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 227 

(1825). 

Grewia hirsuta W?, Buch.-Ham., in sched. 
Bassaria (Sheet 1423.18 LINN-SM, microfiche !-syntype). 
= Grewia sapida Roxb. ex DC. 

Triumfetta annua L., D. Don, Prodr.fl. nepal.: 227 (1825). 
Narainhetty, 17 August 1802 (Sheet 863.7 LINN-SM, microfiche!). 

Triumfetta oblonga Hornem. ex Schrank. in Syll. PI. Nov. 1: 213 
(1824); [Wall ex] D. Don, Prodr.fl. nepal.: 221 (1825), nom. 
superfl. 



Narainhetty, 29 September 1802 (Sheet 863.6 LINN-SM, micro- 
fiche!). 
= Triumfetta pilosa Roth 

UMBELLIFERAE 

Bupleurum tenue [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 182 

(1825), non Salisb. ( 1796). 
Narainhetty, 8 September 1802 (BM! -syntype). 
= Bupleurum hamiltonii N.P. Balakr. 

This specimen is also a type of B. hamiltonii N.P. Balakr. in 
J. Bombay Nat. Hist. Soc. 63: 328 (1967). 

Hydrocotyle hispida [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 182 

(1825), nom. superfl. 

'in Nepalia' [No original label]. (BM-syntype). 
= Hydrocotyle nepalensis Hook. 

Hydrocotyle tenella D. Don, Prodr.fl. nepal.: 183 (1825). 
'in Nepalia'. (Syntype: specimen not found). 
= Hydrocotyle sibthorpioides Lam. 

Pimpinella anethifolia D. Don, Prodr.fl. nepal.: 184 (1825). 
'ad Narainhetty Nepalensium'. (Syntype: specimen not found). 
= Trachyspermum anethifolium (D. Don) H. Wolff 

Sanicula elata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 183 

(1825). 
Narainhetty, 18 November 1802 (BM!-syntype). 

Sanicula hermaphrodita [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 

183(1825). 

Suembu, 1 July 1802 (BM!-syntype). 
= Sanicula elata [Buch.-Ham. ex] D. Don 

URTICACEAE 

Boehmeriafrutescens sensu D. Don, Prodr.fl. nepal. : 59 ( 1 825), non 

Thunb. (1784). 

Urticafrutescens Thunb., Buch.-Ham., in sched. 
Sembu, 23 July 1802 (BM!). 
= Maoutia puya (Hook.) Wedd. 

Boehmeria macrophylla D. Don, Prodr.fl. nepal.: 60 (1825), non 

Hornem (1815), nom. superfl. 
Urtica angustifolia Buch.-Ham., in sched. 
Narainhetty, 5 September 1802 (BMi-lectotype). 

Lectotype designated by D.G. Long in Notes Roy. Bot. Card. 

Edinburgh 40: 130 (1982). 
= Boehmeria penduliflora [Wedd. ex] D.G. Long 

Boehmeria nana D. Don, Prodr.fl. nepal.: 60 (1825). 
Urtica nana Buch.-Ham., in sched. 
Narainhetty, 21 October 1802 (BM!-syntype). 
= Pouzolzia zeylanica (L.) Benn. & R. Br. 

Boehmeria platyphylla D. Don, Prodr.fl. nepal.: 60 (1825). 
Urtica platyphylla Buch.-Ham., in sched. 
Narainhetty, 3 September 1802 (BM!-syntype). 
= Boehmeria macrophylla Hornem. 

Boehmeria rotundifolia D. Don, Prodr.fl. nepal.: 60 (1825). 
Urtica rotundifolia Buch.-Ham., in sched. 
Sembu, 23 July 1802 (BM!-syntype). 
= Boehmeria macrophylla Hornem. 



124 



J.R. PRESS AND K.K. SHRESTHA 



Boehmeria salicifolia D. Don, Prodr.fl. nepal.: 60 (1825). 
Urtica arbuscula Buch.-Ham., in sched. 
Chisa Pany Hill (BM!-syntype). 

Determined as Debrageasia saeneb (Forssk.) Hepper & Wood by 
C. Wilmott-Dear in 1987 and indicated on the sheet by her as the 
syntype of B. salicifolia D. Don. 
= Debregeasia salicifolia (D. Don) Rendle 

Boehmeria ternifolia D. Don, Prodr.fl. nepal.: 59 (1825). 

Urtica ternifolia Buch.-Ham., in sched. 

'in Nepalia' [No original label]. (BM.'-syntype). 

Procris monandra [Buch.-Ham. ex] D. Don, Prodr.fl. nepal: 61 

(1825). 

'in Nepalia' [No original label]. (BM!-syntype). 
= Elatostema monandrum (D. Don) H. Hara 

Procris punctata [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 61 

(1825). 

Procris docum Buch.-Ham., in sched. 
Sembu, 23 July 1802 (BM!-syntype). 
= Elatostema sessile var. polycephalum Wedd. 

Procris rupestris [Buch.-Ham. ex] D. Don, Prodr. fl. nepal.: 60 

(1825). 

'in Nepalia. Floret Julio' [No original label]. (BM!-syntype). 
= Elatostema rupestre (D. Don) Wedd. 

Urtica scripta [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 59 (1825). 
Narainhetty, 8 August 1802 (BM!-syntype); 
Narainhetty, 22 September 1802 (BM!). 
= Pilea scripta (D. Don) Wedd. 

VERBENACEAE 

Clerodendrum foetidum D. Don, Prodr.fl. nepal.: 103 (1825). 
Volkameria foetida Buch.-Ham., in sched. 
Narainhetty, 20 March 1803 (BM!-syntype). 
= Caryopteris foetida (D. Don) Thell. 

Clerodendrum odoratum D. Don, Prodr.fl. nepal.: 102 (1825). 

Volkameria odorata Buch.-Ham., in sched. 

'ad Gorasan Nepalensium. Floret Februario'. (Syntype: specimen 

not found). 
= Caryopteris bicolor (Hardwicke) Mabberley 

Don also cites Roxburgh, Hon. bengai: 46 (1814), who lists a 
Buchanan-Hamilton specimen collected in Nepal in 1802. 

Clerodendrum serratum (L.) Moon, D. Don, Prodr.fl. nepal.: 103 

(1825). 

Volkameria serrata L., Buch.-Ham., in sched? 
'in Nepalia. Floret Septembri'. (Specimen not found). 

Clerodendrum verticillatum [Roxb. ex] D. Don, Prodr. fl. nepal. : 

102(1825). 

'in Nepalia'. (Syntype?: specimen not found). 
= Clerodendrum itulic um (L.) Kuntze 

Verbena sororia [Roxb. ex] D. Don, Prodr.fl. nepal.: 104 (1825), 

nom. superfl. 

Verbena officinalis Buch.-Ham., in sched. 
Bassaria, 7 March 1802 (BM!). 
= Verbena officinalis L. 



Don also cites Roxburgh, Hort. bengai.: 4 (1814), who lists a 
Buchanan-Hamilton specimen collected in Nepal in 1802. 

VIOLACEAE 

Viola caespitosa D. Don, Prodr.fl. nepal.: 205 (1825), nom. superfl. 
Viola patrinii var. napaulensis DC., Prodr. 1: 293 (1824). 
Viola primulifolia Buch.-Ham., in sched., non L. 
Chitlong, 12 April 1802 (BM! syntype of Viola patrinii var. 

napaulensis DC. and V. caespitosa). 
= Viola betonicifolia Sm. subsp. betonicifolia 

Viola hamiltoniana D. Don, Prodr.fl. nepal.: 206 (1825). 
'in Nepalia' [No original label]. (BM!-syntype) 
Cited as lectotype by H. Hara in Enum.fl. pi Nepal 2: 47 (1979). 

Viola palmaris [Buch.-Ham. ex] DC., Prodr. 1: 298 (1824). 
Bimpedi, 8 April 1802 (BM!-syntype). 
= Viola pilosa Blume 

VITACEAE 

Vitispurani [Buch.-Ham. ex] D. Don, Prodr.fl. nepal.: 188 (1825). 
Tancot, 15 April 1802 (BM!-syntype). 
= Vitis parvifolia Roxb. 

ZINGIBERACEAE 

Globba racemosa Sm., Exot. hot. 2: 115, t. 117 (1808). 
Hedychoum deosara Buch.-Ham., in sched. 
Suembu, 28 June 1802 (Sheet 67.5.1 LINN-SM, microfiche!- 
syntype). 

Hedychium coccineum [Buch.-Ham. ex] Sm. in Rees, Cycl. 17: 

Hedychiumn. 5(1811). 
Suembu, 8 June 1802 (Sheet 8.22 LINN-SM, microfiche !-syntype). 

Hedychium gandasulium Buch.-Ham. in Hook, in Hooker's J. Bot. 

Kew Card. Misc. 5: 325 (1853), nom. nud. in syn. 
Narainhetty, 2 September 1 802 (Sheet 8.37 LINN-SM, microfiche!). 
= Hedychium coronarium J. Koenig 

Hedychium ellipticum Sm. in Rees, Cycl. 17: Hedychium n. 2 

(1811). 
Narainhetty, 26 July 1802 (Sheet 8.31 LINN-SM, microfiche !- 

syntype). 

Hedychium spicatum Sm., in Rees, Cycl. 17: Hedychium n. 3 

(1811). 
Narainhetty, 10 August 1802 (Sheet 8.27 LINN-SM, microfiche!- 

syntype). 

Hedychium thyrsiforme Sm. in Rees, Cycl 17: Hedychium n. 4 

(1811). 
Narainhetty, 21 August 1802 (Sheet 8.34 LINN-SM, microfiche!- 

syntype). 

Roscoea purpurea Sm., Exot. hot. 2: 97, t. 108 (1806). 
Narainhetty, 8 August 1802 (Sheet 9.2.1 LINN-SM, microfiche!- 

syntype). 

In the protologue Smith mentions a Buchanan-Hamilton drawing 
and specimens, suggesting he saw several sheets; only one has been 
located. 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



125 



Localities of collections by Buchanan-Hamilton in 
Nepal (1802-3) 

(Present-day names, if different, are given in parentheses.) 

Bagmutty (Bagmati River) 

Baguanpur (Makwanpur) 

Bassaria 

Bimpedi (Bhimphedi) 

Bheempedi (Bhimphedi) 

Catmandu (Kathmandu) 
Chisa Pany Hill (Chisapani) 
Chisapani 

Chisapany (Chisapani) 
Chitlong (Chitlang) 
Chittlang (Chitlang) 
Chittong (Chitlang) 
Churiaghant hills (Churia hills) 
Culi Khana (Kulekhani) 

Ethaura (Hetaunda) 
Ettaura (Hetaunda) 

Gorasaan (Gorasan) 
Gorasan 

Harain-Hetty (Narayanhiti) 
Harain-Netty (Narayanhiti) 
Hethaura (Hetaunda) 
Hettaura (Hetaunda) 
Hettaurei (Hetaunda) 

Kargoo 

Katmandu (Kathmandu) 

Kuli Khana (Kulekhani) 

Lamba (Swayambhu) 
Lohiar 

Muking 

Narainhetty (Narayanhiti) 
Nilcunt (Budanilkantha) 
Norcotera 

Pherphing (Pharping) 

Sambu (Swayambhu) 
Secumbu (Swayambhu) 
Sembu (Swayambhu) 
Suembu (Swayambhu) 
Suemby (Swayambhu) 

Tancot (Thankot) 
Tancote (Thankot) 
Terriany forest (Terai forest) 
Tomba Cana (Tombu Cana) 
Tombha Cana (Tombu Cana) 
Tombu Cana 



Unpublished material relating to Buchanan- 
Hamilton's collections 

There is various unpublished material relating to Buchanan-Hamil- 
ton's Nepalese collections, including letters, manuscripts and 
drawings. These form part of the large volume of material given by 
Buchanan-Hamilton to J.E. Smith in 1 805 and are held in the library 
of the Linnean Society in London. They are listed here, with 
information taken from the Linnean Society library catalogue. 

1. Letters of . . . F. Buchanan (afterwards Hamilton) addressed to 
... W. Roxburgh. 1795-1812. Typescript: 165 leaves. There is an 
unnumbered leaf between ff. 76 & 77. 

2. Flora Nepalensis. 1802-1803. 168 leaves. Incomplete Flora of 
Nepal, commencing with cryptogams and terminating with 
Lysimachiae, made in 1802-1803 and based on collections taken 
chiefly near Kathmandu. Presented to J.E. Smith in 1805 together 
with Nepal Buchanan-Hamilton's herbarium. Accompanied by a 
typescript note on the Flora by A.H.G. Alston, 1945. 

3. Plantarum Nepalensium Icones Pictae. No date. Large folio of 
182 sheets of water-colour drawings by native artists, mostly with 
Buchanan-Hamilton determinations in ink and some in pencil (by 
J.E. Smith?). (Title taken from original wrapper), cf. Buchanan- 
Hamilton's manuscript Flora Nepalensis which cites some of these 
drawings. 

4. Water colour drawings of Indian plants. No date. Large folio of 
95 sheets of water-colour drawings of Indian plants, numbered vii to 
ci, by native artists, with Buchanan-Hamilton determinations in ink 
and some pencilled determinations (by J.E. Smith?), cf. Buchanan- 
Hamilton's Animalium et Plantarum descriptiones which cites some 
of these drawings. 

5. Catalogue of dried plants, c. 1 822. Folio. Catalogue of Buchanan- 
Hamilton's collections presented to the Museum of the Honourable. 
East lindia Company, and arranged according to the system of 
Linnaeus (Ff. 193), & Herbarii Heyneani enumeration (Ff. 30). Fair 
copies of original mss dates 1822. 

6. Manuscript notes on Indian plants. No date. c. 400 sheets in a 
quarto box. Field notes relating to the years 1802-1803 and lists of 
seeds sent to Dr Roxburgh & Co. Each leaf has the place and date on 
it with 3 different index numbers at the head, pro J.E. Smith. 

7. Index slips. 1802-1803. Several hundred slips in a quarto box, 
each with plant name and 3 different index numbers at the head. 
They probably relate to the field notes on Indian plants. 



ACKNOWLEDGEMENTS. We wish to acknowledge the help of various 
colleagues during the course of this work, especially Dr C.E. Jarvis for his 
advice on typification and nomenclatural matters, Dr J. Edmondson and Miss 
C. Sedgwick for information on specimens in the Smith Herbarium which is 
currently undergoing conservation treatment at the National Museums and 
Galleries on Merseyside, Liverpool, and the librarians at The Natural History 
Museum for help with bibliographic problems. We are especially grateful to 
Dr Henry Noltie for his pertinent comments on and numerous corrections to 
the manuscript, and for supplying considerable additional information. 

This study arose from a collaborative project between The Natural History 
Museum and Tribhuvan University, Kathmandu dealing with Nepalese plant 
information and funded under the DETR Darwin Initiative (Round 5). 



126 



J.R. PRESS AND K.K. SHRESTHA 



REFERENCES 



B rum mitt. R.K. & Powell, C.E. 1992. Authors of plant names. London. 

Don, D. 1825. Prodromus florae nepalensis. London. 

Hara, H., Stearn, W.T. & Williams, L.HJ. 1978. An enumeration of the flowering 

plants of Nepal 1. London. 
Hara, H. & Williams, L.HJ. 1979. An enumeration of the flowering plants of Nepal 

2. London. 
Hara, H., Chater, A.O. & Williams, L.HJ. 1982. An enumeration of the flowering 

plants of Nepal 3. London. 
Hamilton, F. 1 8 1 9. An account of the Kingdon of Nepal, and of the Territories annexed 

to this domain by the House ofGorkha. Edinburgh. 



Mabberley, DJ. 1977. Francis Hamilton's commentaries with particular reference to 

Meliaceae. Taxon 26: 523-540. 
Miller, H.S. 1970. The herbarium of Aylmer Bourke Lambert. Notes on its acquisition, 

dispersal and present whereabouts. Taxon 19: 489-553. 
Prain, D. 1905. A sketch of the life of Francis Hamilton (once Buchanan) sometime 

Superintendent of the Honourable Company's Botanic Garden, Calcutta. Annals of 

the Royal Botanic Garden, Calcutta 10(2) I: i-ixxv. 
Press, J.R., Shrestha, K.K. & Sutton, D.A. 2000. Annotated checklist of the flowering 

plants of Nepal. London. 

Rees, A. 1802-1820. The Cyclopaedia. 1-39. London. 
Smith, J.E. 1804-1805. Exotic botany. 1, 2. London. 
Stafleu, F.A. & Cowan, R.S. 1979. Taxonomic literature 2nd ed., 3: 35. Utrecht. 



INDEX 



Acampe rigida (Sm.) P.P. Hunt 115 

Acanthaceae 104 

Acer buzimpala Buch.-Ham. 104 

Acer laurifolium D. Don 104 

Acer oblongum Wall, ex DC. 104 

Aceraceae 104 

Aconogonum molle (D. Don) H. Hara 117 

Aerides calceolaris [Buch.-Ham. ex] Sm. 1 15 

Aerides dasypogon Sm. 1 15 

Aerides multiflora Roxb. 116 

Aerides rigida [Buch.-Ham. ex] Sm. 115 

Aerides spicatum D. Don 116 

Agrimonia nepalensis D. Don 119 

Agrimonia pilosa var. nepalensis (D. Don) Nakai 1 19 

Agrimonia repens L. 119 

Ainsliaea latifolia (D. Don) Sch. Bip. 107 

Ajuga alpina? Buch.-Ham. 110 

Ajuga bracteosa Wall, ex Benth. 110 

Ajuga decumbens sensu D. Don, non Thunb. 110 

Ajuga integrifolia [Buch.-Ham. ex] D. Don 110 

Ajuga laxa Buch.-Ham. Ill 

Ajuga macrosperma var. breviflora Hook.f. 110 

Ajuga pyramidalis L? 110 

Alhagi nepalensis (D. Don) Shap. 1 1 2 

Alisma lappula Buch.-Ham. 104 

Alismataceae 104 

Alliaceae 104 

Allium sulvia [Buch.-Ham. ex] D. Don 104 

Allium tuberosum Rottler ex Spreng. 104 

Alnus nepalensis D. Don 106 

Alstonia lucida D. Don 104 

Amaryllidaceae s.l. 104 

Ammannia rotundifolia [Buch.-Ham. ex] D. Don 113 

Ammannia rubra [Buch.-Ham. ex] D. Don 113 

Anacardiaceae 104 

Anagallis alternifolia Buch.-Ham. 1 1 8 

Anagallis caerulea L. 118 

Anagallis mauritiana Buch.-Ham. 1 1 8 

Anagallis multiangularis Buch.-Ham. 1 1 8 

Anagallis teres Buch.-Ham. 1 1 8 

Anaphalis busua (D. Don) DC. 107 

Anaphalis contorta (D. Don) Hook.f. 107 

Anaphalis margaritacea (L.) Benth. 107 

Anaphalis triplinervis (Sims) C.B. Clarke 107 

Andromeda capricida Buch.-Ham. 108 

Andromeda ovalifolia Wall. 108 

Androsace rotundifolia sensu Sm., non Hardw. 118 

Androsace umbellata (Lour.) Merr. 118 

Androsace villosa? Buch.-Ham. 118 

Aneilema hispida D. Don 107 

Anemone rivularis [Buch.-Ham. ex] DC. 118 

Anemone vitifolia [Buch.-Ham. ex] DC. 118 

Antennaria contorta D. Don 107 

Antennaria timmua D. Don 107 

Antennaria triplinervis Sims 107 

Anthericaceae 113 

Anthvllis cuneata Buch.-Ham. 1 1 1 



Apocynaceae 1 04 

Aporusa octandra (D. Don) A.R. Vickery ex M. Short 

&A.R. Vickery 109 
Aquifoliaceae 105 
Aralia parasitica Buch.-Ham. 105 
Araliaceae 105 

Arbutus laurifolia Buch.-Ham. 109 
Arenaria serpyllifolia L. 106 
Artemisia chinense L.? 107 
Artemisia japonica Thunb. 107 
Artemisia parviflora [Roxb. ex] D. Don 107 
Artemisia parviflora Buch.-Ham. 107 
Arthrostemma paniculatum D. Don 114 
Arundina graminifolia (D. Don) Hochr. 1 16 
Asclepiadaceae 105 
Aspalanthus cuneata D. Don 1 1 1 
Asparagaceae 1 1 2 
Asparagus albus Buch.-Ham. 112 
Asparagus curillus [Buch.-Ham. ex] Roxb. 112 
Asparagus filicinus [Buch.-Ham. ex] D. Don 112 
Asparagus racemosus Willd. 112 
Aster asper [Buch.-Ham. ex] DC. 107 
Aster trinervius [Roxb. ex] D. Don 107 
Astilbe rivularis [Buch.-Ham. ex] D. Don 122 
Astragalus lanceolatus Buch.-Ham. Ill 
Astragalus stipulatus [D. Don ex] Sims 1 1 1 

Baebotrys chishia Buch.-Ham. 1 1 5 

Baebotrys gocala Buch.-Ham. 114 

Baebotrys tomentosa Buch.-Ham. 115 

Balsamina odorata Buch.-Ham. 105 

Balsamina racemosa Buch.-Ham. 105 

Balsaminaceae 105 

Barleria cristata L. 104 

Bassia butyracea Roxb. 121 

Begonia dioica [Buch.-Ham. ex] D. Don 105 

Begonia hatacoa [Buch.-Ham. ex] D. Don 105 

Begonia picta Sm. 105 

Begonia rubella [Buch.-Ham. ex] D. Don 105 

Begoniaceae 105 

Berberidaceae 105 

Berberis aristata DC. 105 

Berberis chitria [Buch.-Ham. ex] Lindl. 105 

Berberis chitria Buch.-Ham. 105 

Berberis miccia [Buch.-Ham. ex] D. Don 105 

Berberis miccia Buch.-Ham. 105 

Berberis paniculata Buch.-Ham., in sched. 105 

Berberis pinnata Buch.-Ham. 105 

Bergenia ciliata forma ligulata Yeo 122 

Bergera integerrima [Buch.-Ham. ex] Colebr. 121 

Betula alnoides [Buch.-Ham. ex] D. Don 106 

Betula boshia Buch.-Ham. 106 

Betulaceae 106 

Bletia bicallosa D. Don 116 

Bletiadabia D. Don 116 

Bletia graminifolia D. Don 1 16 

Bletia masuca D. Don 116 

Blumea hieracifolia (D. Don) DC. 107 



Blumeopsis falcata (D. Don) Merrill 107 

Boehmeria frutescens sensu D. Don, non Thunb. 123 

Boehmeria macrophylla D. Don, non Hornem. 1 23 

Boehmeria macrophylla Hornem. 123 

Boehmeria nana D. Don 123 

Boehmeria penduliflora [Wedd. ex] D.G. Long 123 

Boehmeria platyphylla D. Don 123 

Boehmeria rotundifolia D. Don 123 

Boehmeria salicifolia D. Don 124 

Boehmeria ternifolia D. Don 124 

Boraginaceae 106 

Borbonia tuberosa Buch.-Ham. 1 1 2 

Bothriospermum zeylanicum (J. Jacq.) Druce 106 

Brachycorythis obcordata (Lindl.) Summerh. 117 

Brassaiopsis aculeata (D. Don) Seem. 105 

Brassaiopsis hainla (D. Don) Seem. 105 

Brathys nepalensis Blume 110 

Brossaea procumbens Buch.-Ham. 109 

Buchnera cruciata [Buch.-Ham. ex] D. Don 122 

Buchnera hispida [Buch.-Ham. ex] D. Don 122 

Buddleja asiatica Lour. 113 

Buddleja paniculata Wall. 113 

Buddleja subserrata [Buch.-Ham. ex] D. Don 113 

Budleia shina Buch.-Ham. 113 

Budleia simba Buch.-Ham. 113 

Budleja fastigiata Buch.-Ham. 113 

Buddleja neemda Roxb. 1 1 3 

Budleja nimda Buch.-Ham. 1 1 3 

Budleja shina Buch.-Ham. 113 

Budleja tomentosa Buch.-Ham. 113 

Bulbophyllum hirtum (Sm.) Lindl. ex Wall. 117 

Bulbophyllum odoratissimum (Sm.) Lindl. ex 

Wall. 117 

Bupleurum hamiltonii N.P. Balakr. 123 
Bupleurum tenue [Buch.-Ham. ex] D. Don, non 

Salisb. 123 

Burmannia disticha L. 106 
Burmanniaceae 106 
Butomaceae 106 

Butomopsis latifolia (D. Don) Kunth 106 
Butomus latifolius D. Don 106 

Cacalia cusimbium Buch.-Ham. 107 

Cacalia cusimbua D. Don 107 

Cacalia volubilis Buch.-Ham. 108 

Calanthe masuca (D. Don) Lindl. 116 

Camellia keina [Buch.-Ham. ex] D. Don 122 

Camellia kissi Wall. 122 

Campanulaceae 106 

Caprifoliaceae 106 

Caprifolium macranthum D. Don 106 

Cardamine debilis D. Don 108 

Cardamihe flexuosa With. 108 

Cardamine nasturtioides D. Don 108 

Cardamine resedifolia L. 108 

Cardamine scutata subsp. flexuosa (With.) H. 

Hara 108 
Carduus trilobus Buch.-Ham. 107 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



127 



Caryophyllaceae 106 

Caryopteris bicolor (Hardwicke) Mabberly 124 
Caryopteris foetida (D. Don) Thell. 124 
Cassia dimidiala [Buch.-Ham. ex] D. Don 1 1 1 
Cassia mimosoides subsp. lechenaultiana (DC.) H. 

Ohashi 111 

Castanopsis tribuloides (Sm.) A. DC. 109 
Celastraceae 106 

Centranthera nepalensis D. Don 122 
Cerastium fontanum subsp. grandiflorum (D. Don) H. 

Hara 106 

Cerastium grandiflorum [Buch.-Ham. ex] D. Don 106 
Cerasus phoshia Buch.-Ham. 120 
Ceropegia candelabrum L. 105 
Chaptalia maxima D. Don 107 
Chlorophytum nepalense (Lindl.) Baker 113 
Chonemorpha fragrans (Moon) Alston 104 
Cineraria denudata Buch.-Ham. 108 
Cinnamomum cathia D. Don 111 
Cinnamomum tomentosum D. Don 1 1 1 
Cirsium verutum (D. Don) Spreng. 107 
Cissampelopsis buimala (D. Don) C. Jeffrey & Y.L. 

Chen 108 

Cissampelos angulata Buch.-Ham. 114 
Cissampelos bahapo Buch.-Ham. 114 
Cissampelos hirsuta [Buch.-Ham. ex] DC. 114 
Cissampelos pareira var. hirsuta (DC.) Forman 1 14 
Clematis acuminata DC. 118 
Clematis anemoniflora D. Don 119 
Clematis bucamara Buch.-Ham. 118 
Clematis bucamara var. loasaefolia Buch.-Ham. 118 
Clematis buchananiana [Buch.-Ham. ex] DC. 118 
Clematis diphylla Buch.-Ham. 119 
Clematis grewiiflora DC. 118 
Clematis loasaefolia DC. 118 
Clematis montana [Buch.-Ham. ex] DC. 119 
Clematis montana D. Don 119 
Clematis montana var. ? Buch.-Ham. 119 
Clematis napaulensis DC. 119 
Clematis trinervis Buch.-Ham. 118 
Clematis vitalba? Buch.-Ham. 118 
Clerodendrum foetidum D. Don 124 
Clerodendrum indicum (L.) Kuntze 124 
Clerodendrum odoratum D. Don 124 
Clerodendrum serratum (L.) Moon 124 
Clerodendrum verticillatum [Roxb. ex] D. Don 124 
Cleyera japonica var. wallichiana (DC.) Sealy 123 
Clinopodium piperitum (D. Don) Murata 1 1 1 
Clinopodium repens Buch.-Ham. Ill 
Clinopodium umbrosum (M. Bieb.) K. Koch 1 1 1 
Cnicus verutus D. Don 107 
Coccoloba totnea [Buch.-Ham. ex] D. Don 117 
Colebrookea oppositifolia Sm. 110 
Comarum flavum [Buch.-Ham. ex] Roxb. 119 
Combretaceae 106 

Combretum nanum [Buch.-Ham. ex] D. Don 106 
Combretum spicatum Buch.-Ham. 106 
Combretum? appendiculatum Buch.-Ham. 106 
Commelina hispida Buch.-Ham. 107 
Commelina obliqua [Buch.-Ham. ex] D. Don 107 
Commelina paludosa Blume 107 
Commelinaceae 107 
Compositae 107 
Convallariaceae 113 
Convolvulaceae 108 
Conyza alata Buch.-Ham. 107 
Conyza cappa [Buch.-Ham. ex] D. Don 107 
Conyza falcata [Buch.-Ham. ex] Spreng. 107 
Conyza falcata Buch.-Ham. 107 
Conyza leucantha (D. Don) Ludlow & P.H. Raven 107 
Conyza leucanthemea Buch.-Ham. 107 
Conyza stricta var. pinnatifida (D. Don) Kitam. 107 
Conyza triflda Buch.-Ham. 107 
Conyza? cappa Buch.-Ham. 107 
Cordiaceae 108 
Cornaceae 108 
Comus oblonga Wall. 108 
Cornus paniculata Buch.-Ham. 108 



Coronilla stipulata Buch.-Ham. 1 1 1 

Cotoneaster affinis Lindl. 119 

Crataegus cotoneaster Buch.-Ham. 1 19 

Crataegus? shicola Buch.-Ham. 119 

Crotalaria ?cytissoides Buch.-Ham. 1 1 2 

Crotalaria alata [Buch.-Ham. ex] D. Don 1 1 1 

Crotalaria albida [Heyne ex] Roth 1 1 1 

Crotalaria anthylloides Lam. 1 1 1 

Crotalaria ciliata Buch.-Ham. 112 

Crotalaria cytisoides Roxb. ex DC. 112 

Crotalaria humifusa Benth. 112 

Crotalaria linifolia sensu D. Don, non L.f . 1 1 1 

Crotalaria pilosa Buch.-Ham. 1 1 1 

Crotalaria polygalifolia Buch.-Ham. 1 1 1 

Crotalaria prostrata Buch.-Ham. 1 1 1 

Crotalaria prostrata sensu D. Don 112 

Crotalaria psoralioides D. Don, non Lam. 1 12 

Crotalaria salicifolia Buch.-Ham. 1 1 1 

Crotalaria sessiliflora L. Ill 

Crotalaria tuberosa [Buch.-Ham. ex] D. Don 112 

Cruciferae 108 

Cuncea triflda [Buch.-Ham. ex] D. Don 120 

Cupressaceae 108 

Cyananchus foetidum Buch.-Ham. 105 

Cybbanthera connata [Buch.-Ham. ex] D. Don 122 

Cyclobalanopsis glauca (Thunb.) Oerst. 109 

Cyclobalanopsis lamellosa (Sm.) Oerst. 109 

Cydonia oblonga Mill. 119 

Cydonia sumboshia [Buch.-Ham. ex] D. Don 119 

Cynoglossum prostratum [Buch.-Ham. ex] D. Don 106 

Cyperaceae 108 

Dais bamutis Buch.-Ham. 123 

Dalbergia sericea G. Don 1 1 2 

Daphne bholua [Buch.-Ham. ex] D. Don 123 

Daphne botlua Buch.-Ham. 123 

Daphne cannabina Lour, ex Wall. 123 

Daphne odora sensu D. Don, non Thunb. 123 

Daphne oppositifolia Buch.-Ham. 123 

Daphne papyracea Wall, ex Steud. 123 

Daphne papyri/era [Buch.-Ham. ex] Meissn. 123 

Daphne sericea D. Don, non Vahl 123 

Datisca cannabina L. 108 

Datisca nepalensis D. Don 108 

Datiscaceae 108 

Debrageasia saeneb (Forssk.) Hepper & Wood 124 

Debregeasia salicifolia (D. Don) Rendle 124 

Delphinium carela [Buch.-Ham. ex] D. Don 119 

Delphinium consolida? Buch.-Ham. 119 

Delphinium pauciflorum D. Don 119 

Delphinium scabriflorum D. Don 119 

Desmodium concinnum var. retusum (D. Don) H. 

Ohashi 112 

Desmodium confertum DC. 112 
Desmodium microphyllum (Thunb.) DC. 112 
Desmodium multiflorum DC. 112 
Dillenia racemosa Buch.-Ham. 121 
Dinetus racemosus (Wall.) Sweet 108 
Dioscorea bulbifera L. 108 
Dioscorea obtusangula Buch.-Ham. 108 
Dioscorea pisiformis Buch.-Ham. 108 
Dioscoreaceae 108 

Diospyros serrata [Buch.-Ham. ex] D. Don 122 
Diploknema butyracea (Roxb.) H.J. Lam 121 
Disporum cantoniense var. parviflorum (Wall.) H. 

Hara 112 

Disporum pitsutum D. Don 1 12 
Dobinea vulgaris [Buch.-Ham. ex] D. Don 104 
Drosera lunata [Buch.-Ham. ex] DC. 108 
Drosera peltata var. lunata (DC.) C.B. Clarke 108 
Droseraceae 108 

Dufrenoya platyphylla (Spreng.) Stauffer 121 
Duhaldea cappa (D. Don) Anderberg 107 

Echites fragrans Buch.-Ham. 104 
Echites tomentosa Buch.-Ham. 104 
Echites triangularis Buch.-Ham. 104 
Edgeworthia gardneri (Wall.) Meisn. 123 



Ehretia laevis Roxb. 108 

Elaeagnaceae 108 

Elaeagnus arborea Roxb. 108 

Elaeagnus armata Buch.-Ham. 108 

Elaeagnus infundibularis Momiy. 108 

Elaeagnus parvifolia Wall, ex Royle 108 

Elaeagnus umbellata Thunb. 108 

Elaeagnus umbellatus Thunb. 108 

Elate bulbifera Buch.-Ham. 117 

Elatostema monandrum (D. Don) H. Hara 124 

Elatostema rupestre (D. Don) Wedd. 124 

Elatostema sessile var. polycephalum Wedd. 124 

Eleocharis congesta D. Don 108 

Elsholtzia blanda (Benth.) Benth. 1 10 

Elsholtzia fruticosa (D. Don) Rehder 110 

Elsholtzia stachyodes (Link) Raizada & Saxena 110 

Embelia esculenta D. Don 1 14 

Embelia floribunda Wall. 1 14 

Embelia nagushia D. Don 1 14 

Embelia vestita Roxb. 114 

Epidendrum angustifolium Swartz 116 

Epidendrum bifarium Swartz 116 

Epidendrum calceolare Buch.-Ham. 1 1 5 

Epidendrum cuybua Buch.-Ham. 116 

Epidendrum damun-sultea Buch.-Ham. 116 

Epidendrum gemellum Buch.-Ham. 116 

Epidendrum geniculatum Buch.-Ham. 116 

Epidendrum hippium Buch.-Ham. 116 

Epidendrum humile Sm. 116 

Epidendrum praecox Sm. 116 

Epidendrum te res Thunb. 116 

Epilobium brevifolium D. Don 115 

Epilobium montanum L. 115 

Eria spicata (D. Don) Hand.-Mazz. 116 

Ericaceae 108 

Erigeron alatum [Roxb. ex] D. Don 107 

Erigeron falcatum D. Don 107 

Erigeron hieracifolium D. Don 107 

Erigeron leucanthum D. Don 107 

Erigeron pinnatifidum [Roxb. ex] D. Don 107 

Eriobotrya dubia (Lindl.) Decne. 1 19 

Eriobotrya elliptica Lindl. 119 

Eriophorum comosum (Wall.) Wall, ex C.B. 

Clarke 108 

Eriosema himalaicum H. Ohashi 112 
Eulophia bicallosa (D. Don) P.P. Hunt & 

Summerh. 116 

Eulophia dabia (D. Don) Hochr. 116 
Euonymus grandiflorus Wall. 106 
Euonymus lacerus [Buch.-Ham. ex] D. Don 106 
Euonymus madana Buch.-Ham. 106 
Eupatorium acuminatum D. Don 107 
Eupatorium bhuibu Buch.-Ham. 107 
Eupatorium bhuibu? Buch.-Ham. 107 
Eupatorium pyramidale D. Don 107 
Euphorbia angustifolia [Buch.-Ham. ex] D. Don 109 
Euphorbia dracunculoides Lam. 109 
Euphorbia fusiformis [Buch.-Ham. ex] D. Don 109 
Euphorbia parviflora L. 109 
Euphorbia prolifera [Buch.-Ham. ex] D. Don 109 
Euphorbia tenuis [Buch.-Ham. ex] D. Don 109 
Euphorbiaceae 109 
Eurya acuminata DC. 122 
Exacum tetragonum Roxb. 109 

Fagaceae 109 

Fagerlindia fasciculata (Roxb.) Tirveng. 121 

Fagopyrum dibotrys (D. Don) H. Hara 1 1 8 

Ficus cabur [Buch.-Ham. ex] Sm. 114 

Ficus citrifolia? Roxb. 114 

Ficus cunea Buch.-Ham. 1 14 

Ficus infrafoliacea [Buch.-Ham. ex] Sm. 1 14 

Ficus neriifolia Sm. 1 14 

Ficus pilashi Sm. 1 14 

Ficus sarmentosa [Buch.-Ham. ex] Sm. 1 14 

Ficus sarmentosa Sm. 114 

Ficus semicordata [Buch.-Ham. ex] Sm. 114 

Ficus subincisa [Buch.-Ham. ex] Sm. 1 14 



128 



J.R. PRESS AND K.K. SHRESTHA 



Ficus virens Ait. 1 14 

Ficus virens var. sublanceolata (Miq.) Corner 1 14 

Fimbristylis dichotoma (L.) Vahl 108 

Fimbristylis diphylla (Retz.) Vahl 108 

Flacourtia 109 

Flacourtia infrafoliacea Buch.-Ham. 109 

Flacourtia sepiaria Roxb. 109 

Flacourtiaceae 109 

Floscopa scandens Lour. 107 

Fragaria tuberosa Buch.-Ham. 119 

Galium asperifolium Wall. 121 

Galium ciliatum [Buch.-Ham. ex] D. Don, non Ruiz & 

Pav. 120 

Galium elegans Wall, ex Roxb. 121 
Galium hirtiflorum Req. ex DC. 121 
Galium latifolium [Buch.-Ham. ex] D. Don 121 
Galium pan'iflorum D. Don 121 
Gardenia rigida [Buch.-Ham. ex] D. Don 121 
Gastrochilus calceolaris (Sm.) D. Don 1 15 
Gastrochilus dasypogon (Sm.) O. Kuntze 1 15 
Gaultheria fragrans D. Don 109 
Gaultheria fragrantissima Wall. 109 
Gaultheria nummularioides D. Don 109 
Genista juasi Buch.-Ham. 112 
Gentiana capitata [Buch.-Ham. ex] D. Don 109 
Gentiana decemfida [Buch.-Ham. ex] D. Don 109 
Gentianaceae 109 
Geraniaceae 110 
Geranium nepalense Sweet 1 10 
Geranium quinquenerve Buch.-Ham. 110 
Gerbera maxima (D. Don) Beauverd 107 
Germanea coetsa Buch.-Ham. 110 
Globba racemosa Sm. 124 
Glycine proscimum Buch.-Ham. 112 
Gmelina speciosissima D. Don 122 
Gmelina? tacabushia Buch.-Ham. 122 
Gnaphalium busua [Buch.-Ham. ex] D. Don 107 
Gnaphalium busua Buch.-Ham. 107 
Gnaphalium contortum Buch.-Ham. 107 
Gnaphalium qulntuplinerve Buch.-Ham. 107 
Gnaphalium timmua Buch.-Ham. 107 
Gordonia chilaunea [Buch.-Ham. ex] D. Don 123 
Gratiola cordifolia Vahl 122 
Grewia asiatica ?, Buch.-Ham. 123 
Grewia helicterifolia Wall, ex G. Don 123 
Grewia hirsuta W?, Buch.-Ham. 123 
Grewia oppositifolia [Buch.-Ham. ex] D. Don 123 
Grewia optiva J.R. Drumm. ex Burrett 1 23 
Grewia pumila [Buch.-Ham. ex] D. Don 123 
Grewia sapida Roxb. ex DC. 123 
Guttiferae 110 
Gynura bicolor (Willd.) DC. 107 

Habenaria pectinata D. Don 117 

Habenaria uniflora D. Don 116 

Hamiltonia dulina Buch.-Ham. 121 

Hamiltonia scabra D. Don 121 

Hedera aculeata [Buch.-Ham. ex] D. Don 105 

Hedera elata [Buch.-Ham. ex] D. Don 105 

Hedera hainla [Buch.-Ham. ex] D. Don 105 

Hedera helix sensu D. Don, non L. 105 

Hedera nepalensis K. Koch 105 

Hedera parasitica [Buch.-Ham. ex] D. Don 105 

Hedera tomentosa [Buch.-Ham. ex] D. Don 105 

Hedychium coccineum [Buch.-Ham. ex] Sm. 124 

Hedychium coronarium J. Koenig 124 

Hedychium ellipticum Sm. 124 

Hedychium gandasulium Buch.-Ham. 124 

Hedychium spicatum Sm. 1 24 

Hedychium thyrsiforme Sm. 1 24 

Hedychoum deosara Buch.-Ham. 124 

Hedyotis fusca [Buch.-Ham. ex] D. Don 121 

Hedyotis lineata Roxb. 121 

Hedysarum alhagi L. 112 

Hedysarum dioicum [Buch.-Ham. ex] D. Don 1 12 

Hedysarum junceum L.f. Ill 

Hedysarum retusum [Buch.-Ham. ex] D. Don 112 



Hedysarum sambuense D. Don 1 1 2 

Hedysarum suembum Buch.-Ham. 112 

Hedysarum tenellum [Buch.-Ham. ex] D. Don 112 

Heliotropium obovatum [Roxb. ex] D. Don 106 

Heliotropium ovalifolium Forssk. 106 

Himalrandia tetrasperma (Roxb.) T. Yamazaki 121 

Hovenia acerba Lindl. 119 

Hovenia dulcis sensu Roxb., non Thunb. 119 

Hydrangea aspera [Buch.-Ham. ex] D. Don 1 10 

Hydrangeaceae 110 

Hydrocotyle hispida [Buch.-Ham. ex] D. Don 123 

Hydrocotyle nepalensis Hook. 123 

Hydrocotyle sibthorpioides Lam. 123 

Hydrocotyle tenella D. Don 123 

Hypericum bracteatum [Buch.-Ham. ex] D. Don 110 

Hypericum cordifolium Choisy 1 10 

Hypericum dichotomum Buch.-Ham. 110 

Hypericum elodeoides Choisy 110 

Hypericum japonicum Thunb. 110 

Hypericum lungusum Buch.-Ham. 110 

Hypericum nervosum D. Don 1 10 

Hypericum uralum [Buch.-Ham. ex] D. Don 1 10 

Ilex dipyrena Wall. 105 

Ilex excelsa (Wall.) Hook.f. 105 

Ilex odorata [Buch.-Ham. ex] D. Don 105 

Ilex rotunda sensu D. Don, non Thunb. 105 

Ilex saysia Buch.-Ham. 105 

Impatiens insignis DC. 105 

Impatiens leptoceras DC. 105 

Impatiens odorata D. Don 105 

Impatiens racemosa D. Don 105 

Indigofera atropurpurea [Buch.-Ham. ex] D. Don 112 

Indigofera atropurpurea [Buch.-Ham. ex] Hornem. 112 

Indigofera dosua [Buch.-Ham. ex] D. Don 1 1 2 

Indigofera trifoliata L. 112 

Isodon coetsa (D. Don) Kudo 110 

Isodon ternifolius (D. Don) Kudo 110 

Jasminum arboreum Buch.-Ham. 115 
Jasminum dichotomum D. Don, non Vahl 115 
Jasminum dispermum Wall. 115 
Jasminum heterophyllum Roxb. 115 
Jasminum latifolium Buch.-Ham. 115 
Jasminum multiflorum (Burm.f.) Andrews 115 
Jasminum nepalense Spreng. 115 
Jasminum pubescens (Retz.) Willd. 115 
Jasminum subhumile W.W. Sm. 115 
Juniperus chinensis? Buch.-Ham. 108 
Juniperus indica Bertol. 108 
Juniperus recurva [Buch.-Ham. ex] D. Don 108 

Knoxia sumatrensis (Retz.) DC. 120 
Kohautia gracilis (Wall.) DC. 121 

Labiatae 110 

Laggera alata (D. Don) Sch. Bip. ex Oliv. 107 

Lauraceae 111 

Laurus cathia Buch.-Ham. 1 1 1 

Laurus cuipala Buch.-Ham. 1 1 1 

Laurus cuneata Buch.-Ham. Ill 

Laurus cuspidata D. Don 1 1 1 

Laurus gushia Buch.-Ham. 1 1 1 

Laurus jacaricata Buch.-Ham. Ill 

Laurus nacusua D. Don 1 1 1 

Laurus nancushia Buch.-Ham. Ill 

Laurus tomentosa Buch.-Ham. Ill 

Laurus umbellata [Buch.-Ham. ex] D. Don, non 

Thunb. Ill 

Laurusl lateralis Buch.-Ham. Ill 
Lavendula ternifolia Buch.-Ham. 110 
Leguminosae 111 
Lentibulariaceae 112 
Leontia fruticosa Buch.-Ham. 105 
Lepidagathis incurva [Buch.-Ham. ex] D. Don 104 
Lespedeza juncea var. sericea (Thunb.) F.B. Forbes & 

Hemsl. Ill 
Liatris latifolia D. Don 107 



Ligustrum bracteolatum D. Don 1 1 5 

Ligustrum nepalense Wall. 115 

Ligustrum spicatum [Buch.-Ham. ex] D Don 1 15 

Ligustrum? japonicum? Buch.-Ham. 115 

Liliaceae 112 

Liliaceae s.l. 112 

Lilium batisua Buch.-Ham. 112 

Lilium japonicum Thunb. 1 1 2 

Lilium wallichianum Schult. & Schult.f. 113 

Limnophila connata (D. Don) Hand.-Mazz. 122 

Limodorum asperifolium Buch.-Ham. 116 

Limodorum bicallosum Buch.-Ham. 116 

Limodorum dabia Buch.-Ham. 116 

Limodorum graminifolium Buch.-Ham. 116 

Limodorum lechmana Buch.-Ham. 116 

Linaceae 113 

Lindenbergia grandiflora (D. Don) Benth. 122 

Lindenbergia indica (L.) Vatke 122 

Lindera melastomacea (Nees) Villar 1 1 1 

Lindera nacusua (D. Don) Merr. 1 1 1 

Lindernia anagallis (Burm.f.) Pennell 122 

Linum cicanobum [Buch.-Ham. ex] D. Don 113 

Linum repens [Buch.-Ham. ex] D. Don 113 

Linum semitrigynum Buch.-Ham. 113 

Liparis cordifolia Hook.f. 116 

Liparis nervosa (Thunb.) Lindl. 1 16 

Liparis petiolata (D. Don) P.P. Hunt & Summerh. 116 

Lithocarpus grandifolius var. brevipetiolatus (A. DC.) 

S.N. Biswas 109 

Litsea doshia (D. Don) Kosterm. 1 1 1 
Lobelia begonifolia Wall. 106 
Lobelia heyneana Roem. & Schult. 106 
Lobelia obliqua Buch.-Ham. 106 
Lobelia pyramidalis Wall. 106 
Lobelia rosea Wall. 106 
Lobelia secundeil Buch.-Ham. 106 
Lobelia stimulans Buch.-Ham. 106 
Lobelia trialata [Buch.-Ham. ex] D. Don 106 
Loganiaceae 113 
Lonicera ligustrina Wall. 107 
Lonicera macrantha (D. Don) Spreng. 107 
Loranthaceae 113 

Loranthus hexapetala Buch.-Ham. 113 
Loranthus odoratus Wall. 113 
Luculia gratissima (Wall.) Sweet 121 
Lycianthes macrodon (Wall, ex Nees) Bitter 122 
Lyonia ovalifolia (Wall.) Drude 109 
Lysimachia alternifolia Wall. 118 
Lysimachia lobelioides Wall. 1 18 
Lysimachia pyramidalis Wall. 118 
Lysimachia quinquangularis Buch.-Ham. 1 1 8 
Lysimachia secunda [Buch.-Ham. ex] D. Don 1 1 8 
Lysimachia tetragona D. Don 1 1 8 
Lysimachia triangularis Buch.-Ham. 118 
Lythraceae 113 

Maesa chisia [Buch.-Ham. ex] D. Don 114 
Maesa macrophylla (Wall.) A. DC. 115 
Maesa tomentosa D. Don 1 15 
Maesa viridiflora Buch.-Ham. 115 
Magnoliaceae 113 
Mahonia napaulensis DC. 105 
Malaxis cordifolia Sm. 116 
Malaxis ensiformis Sm. 116 
Malaxis lancifolia Sm. 116 
Malaxis latifolia Sm. 116 
Malucona communis Buch.-Ham. 109 
Malucona communis? Buch.-Ham. 109 
Manglilla? Buch.-Ham. 115 
Manglilla bilrhi Buch.-Ham. 115 
Manna nepalensis D. Don 1 1 2 
Manulea sopubia Buch.-Ham. 122 
Maoutia puya (Hook.) Wedd. 123 
Marrubium piperitum Buch.-Ham. Ill 
Melastoma chulese Buch.-Ham. 114 
Melastoma humile Buch.-Ham. 114 
Melastoma normale D. Don 114 
Melastoma stellatum Buch.-Ham. 114 



COLLECTIONS OF FLOWERING PLANTS BY FRANCIS BUCHANAN-HAMILTON 



129 



Melastomu? rostratum Buch.-Ham. 114 

Melastomataceae 114 

Menispermaceae 114 

Mentha fruticosa Buch.-Ham. 110 

Menlha perilloides L. 110 

Mentha? cristata Buch.-Ham. 110 

Menyanthes indica L. 109 

Mespilus affinis (Lindl.) D. Don 1 19 

Mespilus cuila [Buch.-Ham. ex] D. Don 1 19 

Mespilus integerrima Buch.-Ham. 119 

Mespilus tinctoria D. Don 119 

Michelia champaca L. 113 

Michelia doltsopa [Buch.-Ham. ex] DC. 113 

Michelia kisopa [Buch.-Ham. ex] DC. 113 

Micromelum integerrimum (Colebr.) Wight & Arn. ex 

M. Roem. 121 

Mogorium dichotomum Buch.-Ham. 115 
Mogorium hirsutum 115 
Moraceae 114 
Mussaenda frondosa L? 121 
Mussaenda hispida D. Don 121 
Mussaenda luculia [Buch.-Ham. ex] D. Don 121 
Mussaenda macrophylla Wall. 121 
Myrica esculenta [Buch.-Ham. ex] D. Don 114 
Myrica octandra [Buch.-Ham. ex] D. Don 109 
Myricaceae 114 
Myrsinaceae 114 
Myrsine africana L. 115 
Myrsine capitellata Wall. 115 
Myrsine excelsa D. Don 115 
Myrsine potama D. Don 1 15 
Myrsine semiserrata Wall. 115 
Myrsine sessilis D. Don 1 1 5 
Myrsine subspinosa D. Don 115 

Narcissus tazetta L. 104 
Neillia thrysiflora D. Don 120 
Neolitsea cuipala (D. Don) Kosterm. 1 1 1 
Neottia flexuosa Sm. 116 
Neottia parviflora Sm. 116 
Nepeta imubus Buch.-Ham. 110 
Nerium niveum Buch.-Ham. 104 
Nima quassioides Buch.-Ham. 122 
Nonatellia ? fdamentosa Buch.-Ham. 121 
Notelaea posua D. Don 115 
Nymphoides indica (L.) Kuntze 109 

Oberonia ensiformis (Sm.) Lindl. 116, 117 

Oberonia iridifolia Lindl. 117 

Ochna obtusata van pumila (DC.) Kanis 115 

Ochna pumila [Buch.-Ham. ex] DC. 1 15 

Ochnaceae 115 

Ocimum virgatum Thunb. ex Murray 110 

Octomeria spicata D. Don 116 

Ocymum virgatum Thunb. 110 

Olacaceae 115 

Olea posua Buch.-Ham. 115 

Oleaceae 115 

Onagraceae 115 

Ophiopogon clarkei Hook.f. 113 

Ophiopogon wallichianus (Kunth) Hook.f. 113 

Ophiorrhiza fasciculata D. Don 121 

Ophiorrhiza prostrata D. Don 121 

Ophiorrhiza rugosa Wall. 121 

Ophrys alata L.f. 1 16 

Ophrys cernua L. 117 

Ophrys egaleata Buch.-Ham. 116 

Ophrys monophylla L. 116 

Ophrys spiralis Buch.-Ham. 116 

Ophrys spiralis flore purpureo, Buch.-Ham. 116 

Ophrys sulcata Roxb. 117 

Orchidaceae 115 

Orchis bicornuta Buch.-Ham. 117 

Orchis flexuosa L.f. 117 

Orchis gigantea Sm. 117 

Orchis obcordata [Buch.-Ham. ex] D. Don, non 

Willem. 117 
Orchis pectinata [Buch.-Ham. ex] Sm. 117 



Orchis uniflora Buch.-Ham. 116 

Orthosiphon rubicundus (D. Don) Benth. 110 

Osbeckia chulesis D. Don 1 1 4 

Osbeckia nepalensis Hook. 1 14 

Osbeckia quaterna Buch.-Ham. 114 

Osbeckia rostrata D. Don 1 14 

Osbeckia speciosa D. Don 1 14 

Osbeckia stellata [Buch.-Ham. ex] D. Don 1 14 

Osbeckia stellata var. rostrata (D. Don) 1 14 

Osbeckia ternifolia D. Don 1 14 

Osmanthus fragrans var. longifolius (DC.) H. 

Hara 115 

Osyris chama Buch.-Ham. 121 
Osyris wightiana Wall, ex Wight 121 
Oxyspora paniculata (D. Don) DC. 1 14 

Palmae 117 

Palura odorata Buch.-Ham. 122 

Paris diasua Buch.-Ham. 1 1 3 

Paris polyphylla Sm. 113 

Parochetus communis [Buch.-Ham. ex] D. Don 112 

Parochetus communis Buch.-Ham. 112 

Parochetus major D. Don 1 1 2 

Pecteilis susannae (L.) Raf. 1 17 

Pecteilis triflora (D. Don) T. Tang & FT. Wang 116 

Pedicularis bifida (D. Don) Penned 122 

Pentapanax parasiticus (D. Don) Seem. 105 

Perdicium semiflosculare? Buch.-Ham. 107 

Perdicium? triflorum Buch.-Ham. 107 

Perillaelata D. Don 110 

Perilla frutescens (L.) Britton 1 10 

Perilla fruticosa D. Don 110 

Perilla leptostachya D. Don 110 

Perilla ocimoides L. 110 

Perilla polystachya D. Don 110 

Persicaria capitata (D. Don) H. Gross 1 1 8 

Persicaria posumbu (D. Don) H. Gross 118 

Persicaria runcinata (D. Don) H. Gross 1 1 8 

Persicaria viscosa (D. Don) Nakai 1 1 8 

Phalangium anceps Buch.-Ham. 113 

Phoebe cathia (D. Don) Kosterm. 1 1 1 

Phoenix cf. acaulis Roxb. 117 

Pholiota imbricata Buch.-Ham. 116 

Phyllanthus parvifolius [Buch.-Ham. ex] D. Don 109 

Phy salts angulata (var.) Buch.-Ham. 122 

Physalis divaricata D. Don 122 

Picrasma quassioides (D. Don) Benn. 122 

Pilcusta sylvatica Buch.-Ham. 113 

Pilea scripta (D. Don) Wedd. 124 

Pimpinella anethifolia D. Don 123 

Pinaceae 117 

Pinalia alba Buch.-Ham. 116 

Pinalia ensiformis Buch.-Ham. 116 

Pinalia trifida Buch.-Ham. 116 

Pinus cembra L. 117 

Pinus excelsa Wall, ex D. Don, non Lam. 117 

Pinus wallichiana A.B. Jacks. 117 

Piper guigual [Buch.-Ham. ex] D. Don 117 

Piper mullesua [Buch.-Ham. ex] D. Don 117 

Piper suipigua [Buch.-Ham. ex] D. Don 117 

Piperaceae 117 

Plantaginaceae 117 

Plantago erosa Wall. 117 

Plantago filiformis Buch.-Ham. 117 

Plectranthus buchia Buch.-Ham. 110 

Plectranthus coetsa [Buch.-Ham. ex] D. Don 1 10 

Plectranthus ternifolius D. Don 110 

Plectranthus virgatus D. Don 110 

Pleione humilis (Sm.) D. Don 116 

Pleione praecox (Sm.) D. Don 1 16 

Polygala arillata [Buch.-Ham. ex] D. Don 117 

Poly gala buchanani D. Don 117 

Polygala crotalarioides [Buch.-Ham. ex] DC. 117 

Polygala discolor [Buch.-Ham. ex] D. Don 117 

Polygala furcata Royle 117 

Polygala longifolia Poir. 117 

Polygala monspeliaca Buch.-Ham. 117 

Polygala persicariifolia DC. 117 



Polygala triphylla [Buch.-Ham. ex] D. Don, non 

Burm.f. 117 
Polygalaceae 117 
Polygonaceae 117 

Polygonum capitatum [Buch.-Ham. ex] D. Don 1 18 
Polygonum dibotrys D. Don 1 1 8 
Polygonum fagopyrum Buch.-Ham. 118 
Polygonum posumbu [Buch.-Ham. ex] D. Don 118 
Polygonum runcinatum [Buch.-Ham. ex] D. Don 118 
Polygonum viscosum [Buch.-Ham. ex] D. Don 118 
Parana dichotoma Buch.-Ham. 108 
Parana racemosa Roxb. 108 
Potentilla exarata Sm. 120 
Potentilla fulgens Wall, ex Hook. 120 
Potentilla naspata Buch.-Ham. 120 
Potentilla opaca L. 120 
Potentilla rupestris? Buch.-Ham. 120 
Potentilla splendens Wall, ex D. Don 120 
Potentilla supina L. 119 
Potentilla verna L. 120 
Pouzolzia zeylanica (L.) Benn. & R. Br. 123 
Pratia nummularia (Lam.) A. Braun & Asch. 106 
Primula cushia Buch.-Ham. 118 
Primula denticulata Sm. 118 
Primula elata Buch.-Ham. 1 1 8 
Primula petiolaris Wall. 118 
Primula tridentata D. Don 1 1 8 
Primulaceae 1 1 8 
Procris docum Buch.-Ham. 124 
Procris monandra [Buch.-Ham. ex] D. Don 124 
Procris punctata [Buch.-Ham. ex] D. Don 124 
Procris rupestris [Buch.-Ham. ex] D. Don 1 24 
Prunus cerasoides D. Don 120 
Prunus undulata [Buch.-Ham. ex] D. Don 120 
Pyrus crenata [Buch.-Ham. ex] D. Don 120 
Pyrus nussia [Buch.-Ham. ex] D. Don 120 
Pyrus pashia [Buch.-Ham. ex] D. Don 120 

Quercus arcaula Buch.-Ham. 109 

Quercus armata Roxb. 109 

Quercus banga Buch.-Ham. 109 

Quercus cassura [Buch.-Ham. ex] D. Don 109 

Quercus catungea Buch.-Ham. 109 

Quercus imbricata [Buch.-Ham. ex] D. Don 109 

Quercus lanata Sm. 109 

Quercus lanuginosa D. Don 109 

Quercus phullata [Buch.-Ham. ex] D. Don 109 

Quercus semecarpifolia Sm. 109 

Quercus spicata Sm. in Rees 109 

Rajania? cordata L. 108 

Randia triflora [Buch.-Ham. ex] D. Don 121 

Ranunculaceae 118 

Ranunculus buchiana Buch.-Ham. 119 

Ranunculus indicus Roxb. 119 

Ranunculus sceleratus L. 119 

Ranunculus ternatus Thunb. 119 

Ranunculus umbellatus Roxb. 119 

Reinwardtia cicanoba (D. Don) H. Hara 113 

Reinwardtia indica Dumort. 1 1 3 

Rhamnaceae 119 

Rhamnus catharticus L. 119 

Rhamnus terminalis Buch.-Ham. 119 

Rhamnus trigynus D. Don 119 

Rhamnus virgatus Roxb. 1 19 

Rhexia hari Buch.-Ham. 1 14 

Rhinanthus bifidus [Buch.-Ham. ex] D. Don 122 

Rhododendron arboreum Sm. 109 

Rhododendrum album [Buch.-Ham. ex] D. Don 109 

Rhododendron purpureum Buch.-Ham. 109 

Rhynchostylis retusa (L.) Blume 116 

Rondeletia asiatica L. 121 

Rondeletia budda Buch.-Ham. 121 

Rondeletia coriacea Wall. 121 

Rosa brunonii Lindl. 120 

Rosa clinophylla Thory 120 

Rosa glutinosa Buch.-Ham. 120 

Rosa involucrata Roxb. ex Lindl. 120 



130 



J.R. PRESS AND K.K. SHRESTHA 



Rosa palustris Buch.-Ham. 120 

Rosaceae 119 

Roscoea purpurea Sm. 124 

Rotala rotundifolia (D. Don) Koehne 113 

Rotala rubra (D. Don) H. Hara 113 

Rubiaceae 1 20 

Rubus acuminatus Sm. 120 

Rubus betulinus D. Don 120 

Rubus biflorus [Buch.-Ham. ex] Sm. 120 

Rubus cumbata Buch.-Ham. 120 

Rubus ellipticus Sm. 120 

Rubus flavus [Buch.-Ham. ex] D. Don 120 

Rubus foliolosus D. Don 120 

Rubus hamiltonianus Ser. 120 

Rubus ishia Buch.-Ham. 120 

Rubus paniculatus forma tiliaceus (Sm.) H. Hara 120 

Rubus paniculatus Sm. 120 

Rubus pan'iflorus L. 1 20 

Rubus parvifolius sensu Sm., non L. 120 

Rubus pedunculosus D. Don 120 

Rubus rugosus [Buch.-Ham. ex] D. Don 120 

Rubus rugosus Sm. 120 

Rubus tiliaceus Sm. 1 20 

Rubus triflorus Buch.-Ham. 120 

Ruellia capitata [Buch.-Ham. ex] D. Don 104 

Ruetlia recurva Buch.-Ham. 104 

Rutaceae 121 

Sageretia filiformis (Roth ex Schult.) G. Don 119 
Sagittaria guyanensis subsp. lappula (D. Don) 

Bogin 104 

Sagittaria lappula D. Don 104 
Salicaceae 121 
Salix babylonica L. 121 
Salix disperma [Roxb. ex] D. Don 121 
Salix japonica sensu D. Don 121 
Salomonia cantoniensis Lour. 117 
Salomonia petiolata in D. Don 117 
Samara sessilis Buch.-Ham. 115 
Samara? esculenta Buch.-Ham. 114 
Samara? nagushia Buch.-Ham. 114 
Samara ? potama Buch.-Ham. 1 1 5 
Samara? subspinosa Buch.-Ham. 115 
Sanicula elata [Buch.-Ham. ex] D. Don 123 
Sanicula hermaphrodita [Buch.-Ham. ex] D. Don 123 
Santalaceae 121 
Sapotaceae 121 

Satyrium latifolium Buch.-Ham. 117 
Saurauia napaulensis DC. 121 
Saurauiaceae 121 
Saxifraga ligulata Wall. 122 
Saxifraga pacumbis Buch.-Ham. 122 
Saxifragaceae 1 2 1 
Schefflera elata (D. Don) Harms 105 
Schefflera impressa (C.B. Clarke) Harms 105 
Schima wallichii (DC.) Korth. 123 
Schoepfia fragrans Wall. 115 
Scirpus congestus Buch.-Ham. 108 
Scirpus elongatus [Buch.-Ham. ex] D. Don 108 
Scrophulariaceae 122 
Scutellaria albida? Buch.-Ham. 110 
Scutellaria discolor Colebr. 110 
Scutellaria indica sensu D. Don, non L. 110 
Scutellaria repens [Buch.-Ham. ex] D. Don 110 
Scutellaria scandens [Buch.-Ham. ex] D. Don 1 10 
Selago? oppositifolia Buch.-Ham. 110 
Senecio 3-ligulata Buch.-Ham. 108 
Senecio buimalia [Buch.-Ham. ex] D. Don 107 
Senecio cappa [Buch.-Ham. ex] D. Don 108 
Senecio cappa Buch.-Ham. 108 
Senecio chrysanthemoides DC. 108 
Senecio denudata D. Don 108 
Senecio jacobaea L. 108 
Senecio musuca Buch.-Ham. 108 



Senecio nudicaulis [Buch.-Ham. ex] D. Don 108 

Senecio triligulatus [Buch.-Ham. ex] D. Don 108 

Sideroxylon arboreum Buch.-Ham. 121 

Simaba quassioides [Buch.-Ham. ex] D. Don 122 

Simaroubaceae 122 

Smilacaceae 1 1 3 

Smilax aspera L. 113 

Smilax capitata Buch.-Ham. 1 1 3 

Smilax columnifera Buch.-Ham. 113 

Smilax maculata [Roxb. ex] D. Don 113 

Smilax ovalifolia [Roxb. ex] D. Don 113 

Solanaceae 1 22 

Solanum biflorum sensu D. Don 1 22 

Solanum multifidum Buch.-Ham. 122 

Sopubia trifida [Buch.-Ham. ex] D. Don 122 

Spermacoce lineata Buch.-Ham. 121 

Spermacoce pusilla Wall. 121 

Spermadictyon suaveolens Roxb. 121 

Spiranthes sinensis (Pers.) Ames 116 

Stelis biflora Sm. 117 

Stelis hirta Sm. 117 

Stelis mucronata D. Don 117 

Stelis odoratissima Sm. 117 

Stelis racemosa Sm. 117 

Stellaria monosperma [Buch.-Ham. ex] D. Don 106 

Stellaria saccurcunda Buch.-Ham. 106 

Stellaria saxatilis [Buch.-Ham. ex] D. Don 106 

Stellaria vestita Kurz 106 

Stemodia grandiflora [Buch.-Ham. ex] D. Don 122 

Stemodia muraria Roxb. ex D. Don 122 

Stranvaesia nussia (D. Don) Decne. 120 

Strobilanthes pentastemonoides (Nees) T. 

Anderson 104 

Sultea elatior Buch.-Ham. 113 
Sultea humilior Buch.-Ham. 113 
Sunipia hirta Buch.-Ham. 117 
Sunipia racemosa (Sm.) T. Tang & F.T. Wang 117 
Swertia angustifolia [Buch.-Ham. ex] D. Don 109 
Swertia ciliata (G. Don) B.L. Bum 1 10 
Swertia quadriculata Buch.-Ham. 110 
Swida oblonga (Wall.) Sojak 108 
Symphoricarpos ?odoratus Buch.-Ham. 115 
Symplocaceae 122 

Symplocos cochinchinensis (Lour.) S. Moore 122 
Symplocos crataegoides [Buch.-Ham. ex] D. Don 122 
Symplocos loha [Buch.-Ham. ex] D. Don 122 
Symplocos lucida (Thunb.) Siebold & Zucc. 122 
Symplocos paniculata (Thunb.) Miq. 122 
Symplocos subspinosa Buch.-Ham. 122 
Symplocos sumuntia [Buch.-Ham. ex] D. Don 122 
Symplocos theifolia D. Don 122 
Synotis cappa (D. Don) C. Jeffrey & Y.L. Chen 108 
Synotis triligulata (D. Don) C. Jeffrey & Y.L. Chen 108 

Ternstroemia bifaria Buch.-Ham. 122 

Temstroemia lushia [Buch.-Ham. ex] D. Don 123 

Ternstroemia racemosa D. Don 121 

Tetranthera cuipala [Buch.-Ham. ex] D. Don 1 1 1 

Tetranthera doshia D. Don 1 1 1 

Teucrium laxum D. Don 1 1 1 

Teucrium quadrifarium [Buch.-Ham. ex] D. Don 111 

Thalictrum batula Buch.-Ham. 119 

Thalictrum dalingo Buch.-Ham. 1 19 

'^Thalictrum digynum, Buch.-Ham. 121 

Thalictrum foliolosum DC. 119 

Thalictrum rotundifolium DC. 119 

Theaceae 122 

Thymelaeaceae 123 

Thymus piperitus D. Don 1 1 1 

Thymus repens D. Don 1 1 1 

Tiliaceae 123 

Tomex llampatia Buch.-Ham. Ill 

Tomex bolo Buch.-Ham. 1 1 1 

Tomex doshia Buch.-Ham. 1 1 1 



Tonshia polypetala Buch.-Ham. 121 

Trachelospermum lucidum (D. Don) K. Schum. 104 

Trachyspermum anethifolium (D. Don) H. Wolff 123 

Trilliaceae 113 

Triumfetta annua L. 1 23 

Triumfetta oblonga Hornem. ex Schrank. 123 

Triumfetta pilosa Roth 123 

Tupistra aurantica ([Wall, ex] Baker) Hook.f. 113 

Umbelliferae 123 

Urtica angustifolia Buch.-Ham. 123 

Urtica arbuscula Buch.-Ham. 124 

Urtica frutescens Thunb. 1 23 

Urtica nana Buch.-Ham. 123 

Urtica platyphylla Buch.-Ham. 123 

Urtica rotundifolia Buch.-Ham. 123 

Urtica scripta [Buch.-Ham. ex] D. Don 124 

Urtica ternifolia Buch.-Ham. 124 

Urticaceae 123 

Utricularia aurea Lour. 112 

Utricularia confervaefolia [Jackson ex] D. Don 112 

Utricularia vulgaris Buch.-Ham. 112 

Uvularia pitsutu Buch.-Ham. 1 1 2 

Uvulariaceae 112 

Vallaris solanacea (Roth) O. Kuntze 104 

Verbena offlcinalis Buch.-Ham. 1 24 

Verbena officinalis L. 124 

Verbena sororia [Roxb. ex] D. Don 124 

Verbenaceae 124 

Vernonia aspera Buch.-Ham. 107 

Vernonia extensa DC. 107 

Veronica punctata [Buch.-Ham. ex] D. Don 122 

Veronica undulata Wall. 122 

Viburnum cylindricum [Buch.-Ham. ex] D. Don 106 

Viburnum dichotomum Buch.-Ham. 113 

Viburnum lantanal Buch.-Ham. 106 

Viburnum mullaha [Buch.-Ham. ex] D. Don 106 

Viburnum punctatum [Buch.-Ham. ex] D. Don 106 

Viola betonicifolia Sm. subsp. betonicifolia 124 

Viola caespitosa D. Don 124 

Viola hamiltoniana D. Don 124 

Viola palmaris [Buch.-Ham. ex] DC. 124 

Viola patrinii var. napaulensis DC. 124 

Viola pilosa Blume 1 24 

Viola primulifolia Buch.-Ham. 124 

Violaceae 124 

Virecta fasciculata Buch.-Ham. 121 

Virecta? prostrata Buch.-Ham. 121 

Virecta? suffruticosa Buch.-Ham. 121 

Viscum album L. 113 

Viscum articulatum Burm.f. 113 

Viscum dichotomum D. Don 1 1 3 

Viscum latifolium [Buch.-Ham. ex] D. Don 121 

Viscum stellatum [Buch.-Ham. ex] D. Don 113 

Vitaceae 124 

Vitis parvifolia Roxb. 124 

Vitis purani [Buch.-Ham. ex] D. Don 124 

Volkameria foetida Buch.-Ham. 124 

Volkameria odorata Buch.-Ham. 124 

Volkameria serrata L. 124 

Wendlandia coriacea (Wall.) DC. 121 
Wightia speciosissima (D. Don) Merr. 122 
Wikstroemia canescens Meisn. 1 23 

Xylosteon naisoca Buch.-Ham. 106 
Xylosteum ligustrinum (Wall.) D. Don 106 
Xylosteum scandens Buch.-Ham. 106 

Zingiberaceae 124 
Zizyphus incurva Roxb. 119 
Zizyphus paniculata Buch.-Ham. 119 
Zoduba masuca Buch.-Ham. 116 



Bulletin of The Natural History Museum 
Botany Series 

Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, 
this may also be obtained from the same address. 



Volume 4 

No. 1 Cuticular studies as an aid to plant taxonomy. C.A. Stace. 1965. 
Pp. 1-78, 5 plates, 10 figs. Facsimile edition. 7.20 

No. 2 The genus Elaphoglossum in the Indian peninsula and Ceylon. 
W.A. Sledge. 1967. Pp. 79-96. Facsimile edition. 3.25 

No. 3 Fungi of recent Nepal expeditions. F.L. Balfour-Browne. 1968. 
Pp. 97-141 , 4 figs. Facsimile edition. 3.75 

No. 4 A synopsis of Jamaican Myrsinaceae. W.T. Stearn. 1969. Pp. 

143-178, 8 plates, 25 figs. 4.55 

No. 5 The Jamaican species of Columnea and Alloplectus 

(Gesneriaceae). W.T.Stearn. 1969. Pp. 179-236, 8 plates, 29 
figs- 6.40 

No. 6 New or little known Himalayan species of Swertia and 

Veratrilla (Gentianaceae). H. Smith. 1970. Pp. 237-258, 16 
plates, 7 figs. 8.25 

No. 7 A survey of the tropical genera Oplonia and Psilanthele 

(Acanthaceae). W.T. Stearn. 1971. Pp. 259-323, 10 plates, 18 
figs. 12.40 

No. 8 Angiosperms of the islands of the Gulf of Guinea (Fernando 
Po, Principe, S. Tome, and Annobon). A.W. Exell. 1973. Pp. 
325-411. 12.50 

Volume 5 

No. 1 The dryopteroid ferns of Ceylon. W.A. Sledge. 1973. Pp. 1-43, 

4 figs. 6.90 

No. 2 New Himalayan and Tibetan species of Corydalis 

(Papaveraceae). F. Ludlow & W.T. Stearn. 1975. Pp. 45-69, 15 
plates, 14 figs. 8.40 

No. 3 The marine algae of Trinidad, West Indies. W.D. Richardson. 

1975. Pp. 71-143, 12 plates, 2 figs. 13.45 

No. 4 A revision of the Macaronesian genus Argyranthemum Webb ex 
Schultz Bip. (Compositae-Anthemideae).C.J. Humphries. 1976. 
Pp. 145-240, 2 plates, 26 figs. 14.20 

No. 5 Frank Ludlow (1885-1972) and the Ludlow-Sherriff expedi- 
tions to Bhutan and south-eastern Tibet of 1933-1950. W.T. 
Stearn. 1976. Pp. 243-268, 1 fig. Reliquiae botanicae 
himalaicae. F. Ludlow. 1976. Pp. 269-289, 7 plates, 8 figs. 
Facsimile edition. 11.10 

No. 6 Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric 
classification. N.K.B. Robson. 1977. Pp. 291-355, 9 figs. 

14.20 

No. 7 Sphagnales of tropical Asia. A.Eddy. 1 977. Pp. 357^45 , 4 

plates, 17 maps, 25 figs. 17.80 

Volume 6 

No. 1 The handwriting of Joseph Banks, his scientific staff and 

amanuenses. J.B. Marshall. 1978. Pp. 1-85, 62 figs. 18.30 

No. 2 Seaweeds of the western coast of tropical Africa and adjacent 
islands: a critical assessment. II. Phaeophyta. J.H. Price, D.M. 
John & G.W. Lawson. 1978. Pp. 87-182, 1 fig. 24.40 

No. 3 The lichenicolous Hyphomycetes. D.L. Hawksworth. 1979. Pp. 
183-300, 47 figs. 24.40 



No. 4 



The species of Chisocheton (Meliaceae). D.J. Mabberley. 1979. 
Pp. 301-386, 3 plates, 10 figs. 24.40 



Volume 7 

No. 1 The distribution of Padina pavonica (L.) Lamour. (Phaeophyta: 
Dictyotales) on British and adjacent European shores. J.H. 
Price, I. Tittley & W.D. Richardson. 1979. Pp. 1-67, 3 plates, 2 
figs. 17.40 

No. 2 Seaweeds of the western coast of tropical Africa and adjacent 
islands: a critical assessment. 111. Rhodophyta 
(Bangiophyceae). D.M. John, J.H. Price, C.A. Maggs, G.W. 
Lawson. 1979. Pp. 69-82, 1 fig. 5.40 

No. 3 A revision of the genus Anacyclus L. (Compositae: Anthemi- 

deae). C.J. Humphries. 1979. Pp. 83-142, 27 figs. 15.60 

Volume 8 

No. 1 The Thelypteridaceae of Ceylon. W.A. Sledge. Pp. 1-54, 5 figs. 
1981. 15.15 

No. 2 Studies in the genus Hypericum L. (Guttiferae) 2. Characters of 
the genus. N.K.B. Robson. 1981. Pp. 55-226, 73 figs. 33.55 

No. 3 A revision of the lichen family Thelotremataceae in Sri Lanka. 
M.E. Hale, Jr. 1981. Pp. 227-332, 20 figs. 24.80 

No. 4 Vascular plant collections from the Tristan da Cunha group of 
islands. E.W. Groves. Pp. 333^20, 33 figs. 21.40 

Volume 9 

No. 1 The lichenicolous Coelomycetes. D.L. Hawksworth. 1981. Pp. 

1-98, 36 figs. 22.70 

No. 2 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the 
British Isles. P.S. Dixon & J.H. Price. 1981. Pp. 99-141,5 figs. 

12.50 

No. 3 Parmelia subgenus Amphigymnia (lichens) in East Africa. H. 

Krog & T.F.V. Swinscow. 1981. Pp. 143-231, 31 figs. 21.05 

No. 4 The genus Selaginella in tropical South America. A.H.G. 

Alston, A.C. Jermy & J.M. Rankin. 1981. Pp. 233-330, 18 figs. 

23.05 

Volume 10 

No. 1 Taxonomic studies in the Labiatae tribe Pogostemoneae. J.R. 
Press. 1982. Pp. 1-89, 33 figs. 21.85 

No. 2 The typification of Hudson's algae: a taxonomic and 

nomenclatural reappraisal. L.M. Irvine & P.S. Dixon. 1982. Pp. 
91-105. 5.40 

No. 3 Seaweeds of the Faroes. Various authors. 1982. Pp. 107-225, 

13 figs. 27.15 

No. 4 The lichen genus Steinera. A.M. Henssen & P.W. James. 1982. 
Pp. 227-256, 24 figs. 9.70 

Volume 11 

No. 1 The algae of Lightfoot's Flora scotica. P.S. Dixon. 1983. Pp. 1- 
15, 2 figs. 5.55 

No. 2 A taxonomic study of the lichen genus Micarea in Europe. B.J. 
Coppins. 1 983 . Pp. 1 7-2 1 4, 57 figs, 28 maps. 37.75 



No. 3 The hepatics of Sierra Leone and Ghana. E.W. Jones & A.J. 

Harrington. 1983. Pp. 215-289, 8 figs. 18.15 

No. 4 Studies in the Corallinaceae with special reference to Fosliella 
and Pneophyllum in the British Isles. Y.M. Chamberlain. 1983. 
Pp. 29 1^63, 89 figs. 33.45 

Volume 12 

No. 1 A revision of the Morinaceae (Magnoliophyta-Dipsacales). 

M.J. Cannon & J.F.M. Cannon. 1984. Pp. 1-35, 9 figs. 11.40 

No. 2 An introduction to fem genera of the Indian subcontinent. C.R. 
Fraser- Jenkins. 1984. Pp. 37-76, 1 fig. 12.50 

No. 3 A revision of African Sphagnales. A. Eddy. 1985. Pp. 77-162, 
47 figs. 23.05 

No. 4 Studies in the genus Hypericum L. (Guttiferae) 3. Sections 1. 
Campylosporus to 6a. Umbmculoides. N.K.B. Robson. 1985. 
Pp. 1 63-325, 24 figs, 34 maps. 39.20 

Volume 13 

No. 1 The lichen genus Usnea subgenus Neuropogon. F.J. Walker. 

1985. Pp. 1-130, 39 figs. 31.85 

No. 2 Cytotaxonomic studies of the ferns of Trinidad. A.C. Jermy & 
T.G. Walker. 1985. Pp. 13 1-276, 69 figs. 31.50 

No. 3 Some genera of the Biddulphiaceae (diatoms) with interlocking 
linking spines. R. Ross & PA. Sims. 1985. Pp. 277-381, 33 
plates. 28.75 

Volume 14 

No. 1 Cytological observations on Indian subcontinent and Chinese 
Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae). 
M. Gibby. 1985. Pp. 1-42, 78 figs. 12.50 

No. 2 A redisposition of the species referred to the ascomycete genus 
Microthelia. D.L. Hawksworth. 1985. Pp. 43-181, 73 figs. 

34.25 

No. 3 A classification of the genus Dryopteris (Pteridophyta: 

Dryopteridaceae). C.R. Fraser-Jenkins. 1986. Pp. 183-218, 4 
figs. 11.75 

No. 4 Evolutionary cladistics of marattialean ferns. C.R. Hill & J.M. 
Camus. 1986. Pp. 219-300, 27 figs. 24.65 

Volume 15 

No. 1 Seaweeds of the western coast of tropical Africa and adjacent 

islands: a critical assessment. IV. Rhodophyta (Florideae) 1 . 
Genera A-F. J.H. Price, D.M. John & G.W. Lawson. 1986. Pp. 
1-122, 1 fig. 33.60 

No. 2 Cytology of the fern flora of Madeira. I. Manton, J.D. Lovis, G. 
Vida & M. Gibby. 1986. Pp. 123-161, 12 plates. 14.70 

No. 3 A revision of the lichen genus Xanthoparmelia in Australasia. 
J.A. Elix, J. Johnston & P.M. Armstrong. 1986. Pp. 163-362, 
42 figs, 117 maps. 42.90 

Volume 16 

No. 1 Studies in the genus Hypericum L. (Guttiferae) 7. Section 29. 

Brathys (part 1). N.K.B. Robson. 1987. Pp. 1-106, 14 figs, 25 
maps. 28.75 

No. 2 The lichen genus Ramalina in Australia. G.N. Stevens. 1987. 

Pp. 1 07-233, 1 5 plates, 3 1 figs. 32.20 

No. 3 An annotated list of vascular plants collected in the valleys 
south of Mt Everest. G. Miehe. 1987. Pp. 225-268, 4 figs. 

16.50 

No. 4 Further genera of the Biddulphiaceae (diatoms) with interlock- 
ing linking spines. R. Ross & A.Sims. 1987. Pp. 269-31 1, 13 
plates. 15.60 



Volume 17 

No. 1 Studies in Pseudocyphellaria (lichens) 1 . The New Zealand 

species. D.J. Galloway. 1988. Pp. 1-267, 124 figs. 55.65 

Volume 18 

No. 1 An illustrated catalogue of the type specimens in the Greville 

diatom herbarium. D.M. Williams. 1988. Pp. 1-148, 74 plates. 

38.00 

No. 2 Erik Acharius and his influence on English lichenology. D.J. 

Galloway. 1988. Pp. 149-194, 18 figs. 17.05 

No. 3 Seaweeds of the western coast of tropical Africa and adjacent 
islands: a critical assessment. IV. Rhodophyta (Florideae) 2. 
Genera G. J.H. Price, D.M. John & G.W. Lawson. 1988. Pp. 
195-273, 1 fig. 25.10 

No. 4 Some Cretaceous and Palaeogene Trinacria (diatom) species. 

PA. Sims & R. Ross. 1988. Pp. 275-322, 13 plates. 17.55 

No. 5 A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in 
the Indian subcontinent. C.R. Fraser-Jenkins. 1989. Pp. 323- 
477, 79 figs. 34.10 

No. 6 Corydalis (Papaveraceae: Fumarioideae) in Nepal. M. Liden. 

1989. Pp. 479-538, 26 figs. 19.35 

Volume 19 

A new species of Maytenus (Celastraceae) in Ethiopia. Sebsebe 

Demissew. 1989. Pp. 1-3, 1 fig. 

Central American Araliaceae - a precursory study for the Flora 

Mesoamericana. M.J. Cannon & J.F.M.Cannon. 1989. Pp. 5- 

6 1,36 figs. 

A revision of the Solarium nitidum group (section Holophylla 

pro parte): Solanaceae. S. Knapp. 1989. Pp. 63-102, 21 figs. 

Six new species of Solarium sect. Geminata from South 

America. S. Knapp. 1989. Pp. 103-1 12, 8 figs. 

The application of names of some Indian species of Ocimum 

and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan. 

1989. Pp. 11 3- 11 6, 4 figs. 

Revision of Piper (Piperaceae) in the New World 1 . Review of 

characters and taxonomy of Piper section Macrostachys. M.C. 

Tebbs. 1989. Pp. 1 17-158, 41 figs. Facsimile edition. 52.45 

Volume 20 

No. 1 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. 

Brathys (part 2) and 30. Trigynobrathys. N.K.B. Robson. 1990. 
Pp. 1-1 5 1 , 22 figs, 46 maps. Facsimile edition. 49.25 

No. 2 The marine algal flora of Namibia: its distributions and 

affinities. G.W. Lawson, R.H. Simons and W.E. Isaac. 1990. 

Pp. 153-168, 1 fig, 7 plates. 

The infrageneric classification of Gentiana (Gentianaceae). T- 

N. Ho and S.-W. Liu. 1990. Pp. 169-192, 13 figs. 

Revision of Piper (Piperaceae) in the New World. 2. The 

taxonomy of Piper section Churumayu. M.C. Tebbs. 1990. Pp. 

193-236, 49 figs. 34.10 

Volume 21 

No. 1 Historical and taxonomic studies in the genus Titanoderma 

(Rhodophyta, Corallinales) in the British Isles. Y.M. Chamber- 
lain. 1991. Pp. 1-80, 247 figs. 42.35 

No. 2 Early collections of the Holy Thorn (Crataegus monogyna cv. 
Biflora). A.R. Vickery. 1991. Pp. 81-83, 1 fig. 
A taxonomic study of the species referred to the ascomycete genus 
Leptorhaphis. B. Aguirre-Hudson. 1991. Pp. 85-192, 76 figs. 
The typification and identification of Calymperes 
crassilimbatum Renauld & Cardot (Musci: Calymperaceae). 
L.T. Ellis. 1991. Pp. 193-194, 1 fig. 42.35 



Volume 22 

No. 1 An account of southern Australian species of Lithophyllum 

(Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. 
Campbell. 1992. Pp. 1-107, 63 figs. 41.25 

No. 2 Palynological evidence for the generic delimitation of Sechium 

(Cucurbitaceae) and its allies. J.L. Alvarado, R. Lira-Saade & J. 

Caballero. 1992. Pp. 109-121. 

Seaweeds of the western coast of tropical Africa and adjacent 

islands: a critical assessment. IV. Rhodophyta (Florideae) 3. 

Genera H-K. J.H. Price, D.M. John & G.W. Lawson. 1992. Pp. 

123-146. 

Two new species of Solarium section Geminata (Solanaceae) 

from Cerro del Torr in western Colombia. S. Knapp. 1992. Pp. 

147152. 

Fissidens ceylonensis Dozy & Molkenb. (Musci: 

Fissidentaceae) and some allied taxa from southern India. L.T. 

Ellis. 1992. Pp. 153-156, 2 figs. 

New species of Piper (Piperaceae) from Central America. M. 

Tebbs. 1992. Pp. 157-158. 

Studies on the Cretan flora 1. Floristic notes. N.J. Turland. 

1992. Pp. 159-164. 

Studies on the Cretan flora 2. The Dianthus juniperinus 

complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169. 

41.25 
Volume 23 
No. 1 Revision of Piper (Piperaceae) in the New World 3. The 

taxonomy of Piper sections Lepianthes and Radula. M.C. 

Tebbs. 1993. Pp. 1-50, 18 figs. 

Mounting techniques for the preservation and analysis of 

diatoms. S.J. Russell. 1993. Pp. 51-54. 1 fig. 43.25 

No. 2 New taxa of Gentiana (Gentianaceae) from Western China and 
the Himalayan region. T.-N. Ho and S.-W. Liu. 1993. Pp. 55- 
60, 2 figs. 

New combinations, names and taxonomic notes on Gentianella 
(Gentianaceae) from South America and New Zealand. T.-N. 
Ho and S.-W. Liu. 1993. Pp. 61-66. 
Studies in Hypericum: validation of new names. N.K.B. 
Robson. 1993. Pp. 67-70. 

Generic monograph of the Asteraceae-Anthemideae. K. 
Bremer and C.J. Humphries. 1993. Pp. 71-177, 12 figs. 43.25 

Volume 24 

No. 1 Pre-Linnaean references for the Macaronesian flora found in 

Leonard Plukenet's works and collections. J. Francisco-Ortega, 
A. Santos-Guerra and C.E. Jarvis. 1994. Pp. 1-34, 16 figs. 
Studies on the lichen genus Sticta (Schreber) Ach.: II. 
Typification of taxa from Swartz's Prodromus of 1788. D.J. 
Galloway. 1993. Pp. 35^8, 9 figs. 

Seaweeds of the western coast of tropical Africa and adjacent 
islands: a critical assessment. IV. Rhodophyta (Florideae) 4. 
Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F. 
Prud'homme van Reine and W.J. Woelkerling. 1994. Pp. 49-90, 
Ifig. 

Studies on the Cretan flora 3. Additions to the flora of 
Karpathos. N.J. Turland and L. Chilton. 1994. Pp. 91-100, 
1 fig. 43.25 

No. 2 Observations on the benthic marine algal flora of South 

Georgia: a floristic and ecological analysis. D.M. John, P.J.A. 

Pugh and I. Tittley. 1994. Pp. 101-1 14, 8 figs. 

Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical 

species (excluding Australia). D.J. Galloway. 1994. Pp. 1 15- 

160, 36 figs. 

Morphology and ecology of seedlings, fruits and seeds of 

Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. 

1994. Pp. 161-1 72, 2 figs. 

A study of Bixa (Bixaceae), with particular reference to the leaf 
undersurface indumentum as a diagnostic character. R.E. 
Dempsey and N.C. Garwood. 1994. Pp. 173-180, 2 figs. 

43.40 

Volume 25 

No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. 

1995. Pp. 1-94, 76 figs, 20 plates. 



William Roxburgh's St Helena plants. Q.C.B. Cronk. 1995. 

Pp. 95-98. 

43.40 
No. 2 Seaweeds of the western coast of tropical Africa and adjacent 

islands: a critical assessment. IV. Rhodophyta (Florideae) 5. 

Genera P. G.W. Lawson, W.J. Woelkerling, J.H. Price, W.F. 

Prud'homme Van Reine and D.M. John. 1995. Pp. 99-122, 

Ifig. 

A new species of Odontorrhynchos (Orchidaceae, 

Spiranthinae) from Boliva. D.L. Szlachetko. 1995. Pp. 123- 

125, 1 fig. 

Linnaeus's interpretation of Prospero Alpino's De plantis 

exoticis, with special emphasis on the flora of Crete. N.J. 

Turland. 1995. Pp. 127-159, 27 figs. 

Book review. M.G. Gilbert. 1995. P. 161 . 43.40 

Volume 26 
No. 1 A morphological study of Chaetoceros species 

(Bacillariophyta) from the plankton of the Pacific ocean of 

Mexico. D.U. Hernandez-Becerril. 1996. Pp. 1-73, 52 plates. 

43.40 

No. 2 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. 
Myriandra to 28. Elodes. N.K.B. Robson. 1996. Pp. 75-217, 
43 maps, 29 figs. 43.40 

Volume 27 

No. 1 Notes on the diatom species Tetracyclus castellum (Ehrenb.) 

Grunow with a description of Tetracyclus pseudocastellum nov. 

sp. D.M. Williams. 1997. Pp. 1-5, 8 figs. 

A new species of Calymperes (Musci: Calymperaceae) from 

Peninsular Malaysia. L.T. Ellis. 1997. Pp. 7-9, 1 fig. 

A phylogenetic conspectus of the tribe Hyoscyameae 

(Solanaceae). A.L. Hoare and S. Knapp. 1997. Pp. 1 1-29, 

7 figs. 

A revision of Solarium section Pteroidea: Solanaceae. S. Knapp 

and T. Helgason. 1997. Pp. 3 1-73, 23 figs. 43.40 

No. 2 Systematics of Pogostemon (Labiatae) G.R. Bhatti and M. 

Ingrouille. 1997. Pp. 77-147, 40 figs. 43.40 

Volume 28 

No. 1 Morphology and ecology of seedlings, fruits and seeds of 

Panama: Vochysiaceae. N.C. Garwood. 1998. Pp. 1-16, 3 figs. 
A revision of the genus Mandragora (Solanaceae). S. Ungricht, 
S. Knapp and J.R. Press. 1998. Pp. 17-40, 9 figs. 
The pteridophytes of Sao Tome and Principe (Gulf of Guinea). 
E. Figueiredo. 1998. Pp. 41-66, 2 figs. 43.40 

No. 2 A revision of Brillantaisia (Acanthaceae). K. Sidwell. 1998. 
Pp. 67-1 13, 5 maps, 16 figs. 

Seaweeds of the western coast of tropical Africa and adjacent 
islands: a critical assessment. IV. Rhodophyta (Florideae) 6. 
Genera [Q] R-Z, and an update of current names for non- 
geniculate Corallinales. W.J. Woelkerling, G.W. Lawson, J.H. 
Price, D.M. John and W.F. Prud'homme van Reine. 1998. 
Pp. 115-150, Ifig. 43.40 

Volume 29 

No. 1 The moss family Calymperaceae (Musci) in the Philippines. 
L.T. Ellis. 1999. Pp. 1-46, 25 figs. 

Revision of Hibiscus section Furcaria (Malvaceae) in Africa 
and Asia. F.D. Wilson. 1999. Pp. 47-79, 6 figs. 43.40 

No. 2 Catalogue of the holdings in The Natural History Museum 
(London) of the Australian botanical drawings of Ferdinand 
Bauer ( 1 760-1 826) and cognate materials relating to the 
Investigator voyage of 1801-1805. D.J. Mabberley and D.T. 
Moore. 1 999. Pp. 8 1 -226, 268 figs. 43.40 

Volume 30 

No. 1 A new species of Heisteria (Olacaceae) from Mesoamerica . Q. 
Jimenez and S. Knapp. 2000. Pp. 1-6. 
Three new species of Pilea (Urticaceae) from Costa Rica and 
Panama. A.K. Monro. 2000. Pp. 7-12. 
A revision of Solanum thelopodium species group (section 
Anthoresis sensu Seithe, pro parte): Solanaceae. S. Knapp. 
2000. Pp. 13-30. 






33 The genus Polystichum (Dryopteridaceae) in Africa 

J.P. Roux 
81 Recent records of pteridophytes for Belize, Central America 

D.A. Sutton, A. Hughes and B. Bulmer-Thomas 
101 Collections of flowering plants by Francis Buchanan-Hamilton from Nepal, 1802-1803 

J.R. Press and K.K. Shrestha 




jral History Mus 

BOTANY SERIES 

Vol. 30, No. 2, November 2000