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VOLUME 28 NUMBER 1 25 JUNE 1998
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© The Natural History Museum, 1998
Botany Series
ISSN 0968-0446 Vol. 28, No. 1, pp. 1-66
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Bull. nut. Hist. Mus. Land. (Bot.) 28(1): 1-16
Issued 25 June 1998
Morphology and ecology of seedlings, fruits
and seeds of Panama: Vochysiaceae
THE NATURAL
HISTORY
NANCY C. GARWOOD ^
Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD
CONTENTS
18 JUN 1998
pnr^cu*rgn>
GENERAL LIBRARY
Introduction
Methods
Vochysiaceae
Vochysia ferruginea Mart
Vochysia guatemalensis Donn. Sm
Vochysia sp
Key to known seedlings of Vochysia in Panama and Central America .
Discussion ...
Fruit
Seed
Embryo
Seedling ................................................................................................................................................................................. 10
Phylogeny .............................................................................................................................................................................. 12
Ecology .................................................................................................................................................................................. 14
References ... ... 15
SYNOPSIS Seedlings, saplings, fruits and seeds of Vochysia ferruginea (Vochysiaceae) are described, and ecological data on
these stages summarized, as part of the ongoing Seedling Flora Project centred on Barro Colorado Island, Panama. Seedlings of
two other species reported from Panama are also described. The morphology of these stages is compared to that of other taxa in
the family. The potential use of seed and seedling characters in phylogenetic studies within the family is discussed. Special
attention is drawn to unusual two-parted cotyledonary stipules, ontogenetic changes in leaf hair morphology, and complex ptyxis
and vernation of the embryonic cotyledons.
INTRODUCTION
The aim of the Seedling Flora Project, initiated in 1985, is to
produce an illustrated flora of seedlings, saplings, fruits and seeds
for the neotropical forest centred on Barro Colorado Island, Panama
(see Garwood & Humphries, 1993;Garwood, 1994, 1995, in prep.).
This is the account of Vochysia ferruginea Mart. (Vochysiaceae). It
is the most abundant Vochysia in Panama, common in the younger
forests of Barro Colorado Island and Central Panama; four other
species are known, but only from a few collections (Robyns, 1967a,
b\ Croat, 1978; D'Arcy, 1987). Ranging from Central America
through the Amazon basin, V. ferruginea is also the most widespread
species in the genus, in contrast to most species which have rela-
tively limited distributions (Stafleu, 1948). Because of its fast
growth rate and valuable timber, V. ferruginea has been identified as
a potentially important forestry species for use in plantations or
managed secondary forests in the neotropics (Finegan, 1992; Condit
et al., 1993; Butterfield & Fisher, 1994).
In this account, I describe the seedlings, saplings, fruits and seeds
of V. ferruginea, and the seedlings of two other species known from
Panama, then compare these stages to those of other species in the
family to identify characters of potential phylogenetic importance.
Fruit and seed characters have been important in delimiting the
© The Natural History Museum, 1998
tribes and genera of Vochysiaceae; subgeneric classifications and
keys to species, however, have been based almost solely on floral
and vegetative characters (Warming, 1875; Stafleu, 1948-1954;
Keay & Stafleu, 1953). There have been few comparative studies of
the gross morphology or anatomy of fruits and seeds (Corner, 1976;
Boesewinkel & Venturelli, 1987). Seedlings of few species have
been described in detail (delAmo, 1979; Ricardi et al., 1987; Flores,
19930, b).
METHODS
Standard methods for germinating seeds, producing voucher speci-
mens for each seed collection, growing and harvesting seedlings,
collecting unvouchered seedlings (wildlings) from the forest, and
manipulating the descriptive database were developed for the Seed-
ling Flora Project (Garwood, 1994, 1995) and were used for this
account. The detailed descriptions follow the standard format devel-
oped for the Project. Descriptive terms have been standardized for
use across all families in the Project, but generally follow common
use. See de Vogel (1980) for definitions of seedling-specific terms
and Corner (1976) and Boesewinkel & Bouman (1984) for seed-
specific terms. To facilitate identifying isolated seeds, the base of the
N.C. GARWOOD
seed is arbitrarily defined as the part nearest the hilum, rather than
using a reference point within the fruit. Leaf nodes of seedlings are
numbered sequentially from the first leaf-bearing node above the
cotyledons. The following less conventional terms are used: for
length of pubescence or size of other surface features - minute (< 0. 1
mm), short (0. 1 -0.5 mm), long (0.5- 1 .0 mm), very long (> 1 .0 mm);
for relative thickness of the seed coat or pericarp layers compared to
total width of the structure - very thick (> 25% width), thick (10-
25%), thin (1-10%), and very thin (< 1%).
VOCHYSIACEAE
HABIT AND DISTRIBUTION. Trees or shrubs, rarely subshrubs.
Neotropical except for one very small African genus; 6-8 genera and
c. 200 species, 3 genera and 8 species from Panama, 1 genus and
species known from Barro Colorado Island.
TAXONOMIC REFERENCES. Stafleu ( 1 948), Keay & Stafleu ( 1 953),
Robyns (1967a, b), Cronquist (1981), D'Arcy (1987), Kawasaki (in
press).
Vochysia Aubl.
HABIT AND DISTRIBUTION. Trees, shrubs or rarely subshrubs.
Neotropical; c. 100 species, 5 species in Panama, 1 species known
from Barro Colorado Island.
Vochysia ferruginea Mart, in Mart. & Zucc., Nov. Gen. sp. pi. 1:
151,1.92(1824).
Fig. 1.
HABIT AND DISTRIBUTION. Trees, 6-35 m tall, of moist to wet
lowland and premontane forests, usually in secondary forest. Nica-
ragua to Peru and Brazil.
COLLECTIONS. Panama. Barro Colorado Island: Garwood 1 630A
(F, BM, PMA, adult vouchers), Garwood 1630B-F (seedlings),
Garwood 2005 (seedlings), Foster s.n. (seedlings with fruit voucher);
50 seedlings to 37 cm tall examined.
Fruits
Infructescences terminal and upper-axillary thyrses, of 1- to 5-flow-
ered helicoid cymose branches, usually with < 1 fruit maturing per
branch. Rachis densely short-pubescent, with hairs 2-branched, ±
sessile to short-stalked, appressed to ascending, straight to sinuous;
lateral branches (including pedicels) 8-15 mm long, slightly to
moderately pubescent. Receptacles undeveloped. Stipes absent. Fruits
dry capsules, from superior ovaries. Capsules ± woody, loculicidally
dehiscent; completely septate; 17-26 x 8-12 x 8-12 mm; straight;
oblong to obovate in outline, 3-lobed to 3-cleft in t.s., not com-
pressed; margins entire; base rounded to abruptly truncate; apex
rounded to slightly retuse, with stylar scar to 1 mm wide; surface
dark brown to black-brown, dull, slightly to prominently warty,
often irregularly clear- to white-scaly, glabrous. Seeds 1-3, 1 per
locule, intermediary, separated by mesocarp and endocarp, com-
pressed laterally; long axis of seed radially parallel to long axis of
fruit. Placentae axile. Funicle absent. Valves 3, dehiscing from apex,
slightly spreading apart then each folding back, remaining firmly
attached at base. Fruit wall thick, 0.4-1.5 mm wide, thickest cen-
trally, 3-layered. Exocarp very thin, = 0.05 mm wide, ± hard-papery,
dark brown to black-brown. Mesocarp thick, 0.2-1.3 mm wide,
thickest centrally between locules, hard-spongy to ± woody, with 2
large longitudinal resin-filled canals near axis between each pair of
locules and several smaller chambers along locules (resin drying
red-brown and glassy), light to medium brown. Endocarp thin, ~ 0. 1
mm wide, crustaceous, cream to light brown; inner surface smooth,
minutely transversely fibrous, glabrous, glossy, medium
brown-orange. When fruit unripe, exocarp grey-green.
Seeds
Seeds 20-25 x 5-6 x 1-2 mm; hilar-anatropous; 1 -winged; trans-
versely half-obovate in outline, compressed and elliptic in t.s. in
seminiferous area, planar in wing; margins constricted distally in
seminiferous area on hilar side, constricted below wing on opposite
side; base widely truncate; apex widely rounded; outer surface ±
smooth to slightly rough, minutely rough under hairs at higher
magnification, densely long- to very long-pubescent, with hairs
sinuous, appressed, light to medium brown-orange; glossy, medium
brown to brown-orange, dark brown under hairs; not exfoliating.
Wing lateral, fibrous, ~ 15-17 x 5-6 mm, ± half-elliptic in outline,
planar, asymmetrically rounded at apex, entire along margins, with
curved ridge slightly raised, ± flattened, extending from base to 3/4
length of wing and forming thickened straight margin; formed from
very long, interwoven, testal hairs. Seminiferous area lateral, ~ 10-
1 1 x 4-5 mm, ± elliptic in outline, compressed, darker than wings,
with short fringe of interwoven hairs on side opposite wing. Aril
absent. Sarcotesta absent. Hilum basal, along straight side, exposed,
flush, light brown, lighter than testa, linear, 7-10 mm long. Micro-
pyle subbasal to marginal (visible internally at tip of radicle, possibly
continuing through thickened ridge of wing and terminating along
straight side past hilum), concealed externally. Lens absent. Storage
reserves absent. Embryo large, = 10-11 mm long, filling seminifer-
ous area, straight, ± oblong or ellipsoid, compressed; axis
well-developed, with long 4-winged hypocotyl separated from very
short radicle by annular ridge at collet, half-exposed; cotyledons
well-developed, thin, with compressed obliquely supervolute ptyxis
and convolute (sensu imbricate) vernation, with wider side of each
cotyledon angled down and partially enclosing hypocotyl, with
shape obscured by folding and base hidden within folded blade;
plumule rudimentary. Vascular bundle half-lateral on hilar side,
unbranched, thin, with tip exposed at end of hilum and fragments of
the funicular bundle often exposed in hilum; passing through short
raphe, terminating marginally at tip of seminiferous area. Seed coat
very thin, ± papery, undifferentiated at maturity. Testa very thin, ±
thicker under hilum and around micropyle, ± papery, 1 -layered,
medium to dark brown, densely long-pubescent (see above); inner
surface irregular and minutely rough, glabrous, glossy, dark brown.
Tegmen absent.
Fig. 1 Vochysia ferruginea Mart. Fruit: A, undehisced fruit; B, dehisced capsule; C, t.s. Seed: D, seed, lateral view; E, embryo in seminiferous region,
with seed coat partially removed; F, embryonic cotyledon and radicle, with one side of cotyledon unfolded and second cotyledon removed to show
primary veins, radicle, and angle of folding; G, t.s. through seminiferous region at level marked by arrow in E-F, view toward radicle and wing. Seedling:
H, seedling; J, cotyledon; K, cotyledonary stipules, hypocotyl, and petiole ridges; L-M, cotyledonary stipules after epicotyl expands; N, first leaf; O,
stipules at first node. Sapling: P, young sapling; Q, branched sapling; R, leaf at node 15; S, stipules at about node 20. Scale bars: black or black and
white, 1 mm units; white only, 1 cm units. Symbols: ac, annulate collet; ab, abaxial surface of cotyledon; ad, adaxial surface of cotyledon; ^-c,,
cotyledons 1 and 2; en, endocarp; ex, exocarp; m, midvein of cotyledon; ms, mesocarp; r, radicle; re, resin canals; sd, seed; so, outline of seed (dotted
line); ts, testa; f,-f4, folds of cotyledon.
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
N.C. GARWOOD
Seedlings and saplings
Seedlings phanero-epigeal with photosynthetic cotyledons.
DEVELOPMENT. Radicle emerges from hilar side of seed near
wing. Hypocotyl elongates into loop, then straightens, often carry-
ing seed upward. Cotyledons emerge from seed and fully expand
before first leaves are initiated. Shoot rest period is short during the
cotyledon stage, about 1 month in sun and 3 months in shade. Seed
reserves are exhausted by the end of this period, as albino seedlings
die at the cotyledon stage without initiating any leaves. Growth is
continuous, with a pair of leaves initiated about every month in sun
and every 2-3 months in shade, although shade-produced leaves are
smaller.
ROOTS. Primary- root 30-50 x 0.6-0.8 mm, circular in t.s., slightly
sinuous, cream to white, becoming medium orange-brown; narrow
annular ridge encircles axis at collet, becoming indistinct with age;
surface texture and colour between hypocotyl and root change
conspicuously and abruptly at collet. Secondary roots to 0.2 mm
wide, sinuous, very sparsely scattered along primary root. Root
hairs appear abruptly and densely on primary root just below collet.
Sapling root system: taproot thick, long-tapering; secondary roots ±
thin and common along primary root; tertiary and higher order roots
very fine and numerous.
STEMS. Hypocotyl 15-67 x 0.25-0.50 mm, ± square in t.s., be-
coming circular in t.s. and hollow with age; 4-ridged to 4-winged,
each wing to 0.5 mm tall, continuing at apex into lower margin of
cotyledon petiole and terminating at base abruptly above collet;
glabrous; medium yellow-green to green, becoming dark brown or
red-brown. Epicotyl 3-12 x 0.3-0.5 mm, ± square in t.s., becoming
circular with age, 4-winged, each 0.3-1.0 mm tall and terminating
apically in stipules, glabrous or sparsely short-pubescent, medium
green to yellow-green. Hairs short, T-shaped (ratio of arm lengths =
1 : 1 ), ± sessile, appressed, straight, and light brown to orange-brown.
Cataphylls absent. Internodes 1-10, 2-15 mm long; later internodes
to 34 mm; similar to epicotyl except later internodes becoming
densely short- to long-pubescent, with hairs long, unequally
2-branched (arm length ratio to 8:1), short-stalked, ascending, and
curved or sinuous or coiled. Older stems dark brown, with wings
often red-brown, hollow, often sparsely pubescent with age; wood
cream-coloured. Lenticels appearing on stems > 2 mm wide, sparse
on stems 2—4 mm wide; mostly 0.5 mm long, ± round to elliptic,
light brown, lighter on dark stems. Buds: Cotyledonary buds hidden
in axils and inconspicuous above cotyledon scars, rarely protruding
to == 0.5 mm in older seedlings, moderately short-pubescent. Lateral
buds usually slightly protruding out of axils, to 1 mm long at later
nodes, moderately pubescent at early nodes to densely pubescent at
later nodes, with accessory bud above sylleptic branches at later
nodes. Terminal buds naked, partially covered by stipules, moder-
ately short-pubescent at early nodes to densely long-pubescent at
later nodes, with hairs light brown to orange-brown.
COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, stipu-
late, emergent from seed, spreading apart, persistent at least until
node 3 matures, falling sometime before node 9 matures (between 9
and 20 months). Blades 10-15 x 13-23 mm; straight, flat, depressed
obovate, with right half (viewed from the lower abaxial side) usually
slightly larger, 2-lobed to 2-cleft; apex of lobes obtuse to broadly
obtuse; base emarginate or cordate, often asymmetrical with right
lobe slightly larger, or half-rounded/half-cordate; margins entire.
Blades glabrous; green above, lighter below. Venation palmate,
3-nerved; acro-brochidodromous, basal and perfect. Primary veins
3; moderate-sized; impressed to slightly raised above, moderately
raised below; central primary vein forked, with straight stem '/2-3/4
distance to apex and branches slightly recurved; lateral primary
veins basal, = 3/4 length of blade, straight to curved, diverging at
narrowly acute angle, looped, joining branches of central vein to
form large ± triangular to obovate intercostal area, the pair of
intercostal areas usually spanning about 60-70% of the width of the
cotyledons. Secondary veins moderate-sized basally along lateral
primaries becoming fine apically; 16-24, 2-4 along midvein, 6-10
along lateral primaries, 6-10 along branches of midvein; mostly
marginal, alternate along midvein; 'A>-3/4 length of blade (mostly 'A>-
'/4 along branches of midvein), longest basally; mostly curved or
arched along lateral primaries, hooked along branches of midvein,
curved to zigzag along midvein, rarely scythe-shaped; diverging at
narrowly acute angles (< 45°) basally on lateral primaries to perpen-
dicular angles along branches of midvein; looped marginally,
sometimes percurrent in intercostal area, mostly obtusely joined;
slightly raised above, slightly raised below. Inter-secondary veins
absent. Tertiary veins reticulate within intercostal area, looped in
marginal area. Petioles 0.5-1 .5 mm long, transversely oblong in t.s.,
straight to slightly curved, 2-ridged above, with ridges continuing
into lateral primary veins, 4-winged along margin, with lower wings
continuing into hypocotyl wings and upper wings terminating in
fleshy knobs near centre of hypocotyl, glabrous. Stipules usually
fused and interpetiolar, sometimes separate and paired; composed of
two structures, both variable in shape and development. The lower
part usually a ± conical projection between the petioles at the base of
the upper marginal ridge of each petiole, sometimes with two or
three separate projections or these absent, mostly 0.125-0.25 mm
long, usually perpendicular to the stem, sometimes gland-like
apically. The upper part usually a flattened interpetiolar scale-like
outgrowth just above the petioles, initially folded over the terminal
bud, becoming erect and visible as the epicotyl expands, 0.25-0.5
mm tall and 0.5-0.75 mm wide, ± transversely oblong to depressed
ovate, usually 2- to 5-lobed or -cleft, but sometimes divided into two
separate parts or unlobed, the apex of lobes gland-like when young;
persistent, papery, green becoming light brown.
LEAVES. Opposite, decussate on trunk axis, distichous on branches;
simple; short-petiolate, stipulate. Leaf blades 17-32 x 3-9 mm at
first node; 20-67 x 5-20 mm at nodes 2-10; to 142 x 49 mm at later
nodes; 65-125 x 25-37 mm on branches; straight; narrowly elliptic
at nodes 1-3, narrowly elliptic or narrowly obovate at later nodes,
elliptic or narrowly elliptic or narrowly obovate or linear-obovate on
branches; apex narrowly obtuse or acuminate at nodes 1-10, acumi-
nate to long-acuminate at later nodes and on branches, mucronate at
most nodes; base mostly attenuate at early nodes, sometimes nar-
rowly cuneate at later nodes, mostly acute on branches; margins
entire, sometimes undulate. Blades green above, lighter below; flat
between secondary veins above when fresh; papery; densely minutely
pusticulate below (from internal crystals), but weakly so at nodes 1-
2; densely minutely pusticulate above, with pustules usually white
on larger leaves; slightly to moderately short- to long-pubescent on
blade and veins below at early nodes becoming densely long- to very
long-pubescent only on the midvein at later nodes; glabrous on blade
above, slightly to moderately ± short-pubescent on midvein at early
nodes to densely long-pubescent at later nodes; slightly to moder-
ately short- to long-pubescent along margin, often also densely
minute-pubescent when young at later nodes; pubescence light
brown to orange-brown. Hairs on midvein above simple, ± erect,
straight to slightly curved, mostly 0.5-0.75 mm long on first node, to
1-2 mm long on later nodes; those on blade and veins below
T-shaped (ratio of arm lengths ~ 1:1), ± sessile, appressed and
straight at early nodes, mostly 0.25-0.75 mm long, becoming pro-
gressively more unequally 2-branched (ratio of arm lengths mostly
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
< 4:1 on blades but to 8:1 on midvein), short-stalked, ascending,
curved or sinuous or coiled at later nodes; those on margins T-shaped
or 2-branched (as above) and, at later nodes, also minute, simple, ±
erect and sinuous. Young leaves conduplicate, spreading open before
half-grown, densely pubescent, light green, rigid, erect to horizon-
tal. Venation pinnate; brochidodromous. Midvein moderate-sized;
straight; flush to slightly raised above, impressed at later nodes and
when fresh, prominently raised below. Secondary veins ± fine; 14—
17 on leaves at first node, 15-30 at nodes 2-10, to 30 at later nodes
and on branches; alternate; mostly '/6-'/3 length of blade, longest
centrally; mostly hooked or arched, sometimes also slightly zigzag;
diverging at moderately acute angles (45"-65°); looped, sometimes
diffusely looped basally on larger leaves, obtusely joined; slightly
raised above, slightly raised below. Inter-secondary veins infre-
quent. Tertiary veins reticulate within intercostal area, looped in
marginal area. Petioles 0.5^.0 mm long, < 2 mm long at nodes 1-
1 0; ± circular in t.s., straight to slightly curved; shallowly channelled,
with erect narrow margins above; sparsely to moderately pubescent.
Stipules paired; 0.5-1 .0 mm long at nodes 1-6, to 3 mm long at later
nodes and on branches; simple, triangular to narrowly triangular;
persistent, ± papery; densely pubescent, pale against dark older
stems; the inner margin and base continuing to and perhaps above
the petiole; the outer margin and base usually terminating abruptly,
but sometimes forming a low interpetiolar ridge with the stipule of
the opposite leaf as the stem thickens at early but not later nodes.
ARCHITECTURE. Trunk axis monopodial to at least 37 cm tall
(based on one individual). Initial axis orthotropic. Branches
plagiotropic but ± ascending, with leaves distichous through twist-
ing of internodes; sylleptic; alternate and perpendicular at two
consecutive nodes (semi-decussate); first produced 3 1-34 cm above
the collet (at about nodes 17-18). First branch internode 60-75 mm
long, much longer than superadjacent trunk internode (10-30 mm).
First branch leaf half- to full-size of later branch leaves. Single or
paired sylleptic secondary branches were produced on the primary
branches 24-29 cm from the trunk. Leaves held horizontal. Several
factors indicate that growth of the trunk axis above the branches
stopped or slowed considerably while the branches were develop-
ing, so that overall growth of the trunk was rhythmic (see Fig. 1Q):
the trunk axis was very short (about 4 cm long) compared to the
length of the branches (28-38 cm long); it was non-woody and very
thin compared to the woody internodes on the trunk below the
branches and on the lower parts of the branches; and it had matured
only one pair of leaves compared to 7-8 pairs on the branches . At the
time of harvest, the trunk axis had re-initiated growth, although the
primary and secondary branches were still elongating. Rhythmic
growth, if substantiated, would suggest Massart's model.
Ecology
PHENOLOGY. In Panama, Vochysiaferruginea usually flowers from
late March to early July, and sometimes to a lesser extent in
September-October, and its fruits mature August-October (Croat,
1978). Ripe fruits were dehiscing and releasing mature seeds in
October (Garwood 1630A). Elsewhere in Central America, it prima-
rily flowers in April-June and October-December and fruits
July-September and December-January (Croat, 1978;Flores, 19936;
Moreira & Arnaez, 1994).
DISPERSAL. When ripe, valves of the dark brown capsules open
and fold back, exposing the glossy bright orange-brown endocarp
and releasing the seeds. Seeds are wind-dispersed.
VIABILITY. Seeds remain viable for 2-3 months when stored at
ambient temperature (24-26°C) and moisture levels if well-aerated
(Flores, 1993ft). Flores (1993ft) considers the seeds orthodox, al-
though the initial moisture content of the seeds (23%) is rather high.
GERMINATION. Seeds germinate rapidly (13 days) at moderate
percentages (39-50%) in both sun and shade in the growing house
(this study). Other studies also report rapid germination (8-14 days)
in the forest or growing house, but higher germination percentages
(80-95%) (Flores, 1993ft; Moreira & Arnaez, 1994).
ESTABLISHMENT AND GROWTH. The earliest stages of natural seed-
ling establishment and growth have not been studied, although
Flores (1993ft) noted that germinating seeds and young seedlings
suffered high predation from ants and other herbivores. Boucher et
al. (1994) reported exceptional seedling densities and sapling growth
after Hurricane Joan in Nicaragua in October 1989, which killed all
adults in the population. Four months after the hurricane, there were
94 500 seedlings and saplings (0.1-1.5 m tall) per hectare, but few
young trees (3.2-10 cm DBH). The extremely open canopy, created
by the hurricane through defoliating and felling trees, favoured
survivorship and growth of these juveniles, such that young trees
were abundant four years later and were growing on average at about
1 .4 cm DBH/year. Under the less open canopy of Barro Colorado
Island, not recently if ever disturbed by hurricanes, similar-sized
individuals (1-9.9 cm DBH) grew on average only 0.45 cm DBH/
year, but larger individuals (> 10 cm DBH) grew more rapidly, 1.3-
1.7 cm DBH/year (Condit et al., 1993).
The population dynamics and appropriate management strategies
for this species may differ greatly in hurricane-prone areas such as
Nicaragua compared to hurricane-free areas such as Panama. Where
hurricanes are common in the Caribbean, they occur mostly August-
October during the period Vochysia ferruginea is maturing and
dispersing its seeds. A hurricane occurring late in the season might
blow the wind-dispersed seeds over large areas of disturbed forest,
expanding the population. An earlier hurricane might kill adult trees
before seeds could be matured and dispersed, preventing regenera-
tion from seed and eliminating the local population, unless most
regeneration comes from established shade-tolerant juveniles. It is
not yet known whether seedling recruitment after disturbance is
primarily from recently germinated seedlings from the current
year's seed crop or older shade-tolerant juveniles surviving from
previous years.
Vochysia guatemalensis Donn. Sm. in Bot. Gaz. 12: 131, pi. 22
(1887).
Vochysia hondurensis Sprague in Bull. Misc. Inform., Kew 1922:
183(1922).
Fig. 2A-D.
HABIT AND DISTRIBUTION. Trees, 30-55 m tall. Veracruz, Mexico,
to Panama.
COLLECTIONS. Belize. Cayo District, Chiquibul: Palmar, Garwood
403 1 (BM, seedlings with seed voucher); New Maria, Monro 867 to
871, Monro, Howe & King M4162X, M7231X, M7340X (seed-
lings) and M6002X (sapling); 28 seedlings to 17 cm tall examined,
sapling height about 2 m.
TAXONOMIC NOTE. The seeds associated with Garwood 4031
unmistakenly identify the seedlings to genus. If one accepts the
synonymy of Vochysia guatemalensis and V. hondurensis (Standley
& Steyermark, 1949; Gaos, 1978; Flores, 1993a), then only one
species has been reported from Belize (Stafleu, 1948; Standley &
Steyermark, 1949). That the mature leaves and stems are nearly
glabrous supports the identification of these collections as V.
guatemalensis.
N.C. GARWOOD
Fig. 2 Vochysia guatemalensis Donn. Sm. Seedling: A, cotyledon; B, first leaf; C-D, cotyledonary stipules. Vochysia sp. Seedling: E-F, cotyledons; G,
cotyledonary stipules; E, Hampshire & Whitefoord 269; F-G, Garwood 135. Scale bars: black, 1 mm; white, 1 cm.
Seedlings and saplings
Seedlings phanero-epigeal with photosynthetic cotyledons.
ROOTS. Primary root 30-70 x 0.8-1 .2 mm, circular in t.s., slightly
sinuous, cream-coloured; narrow annular ridge encircles axis at
collet, becoming indistinct with age. Secondary roots sinuous,
scattered along primary root.
STEMS. Hypocotyl 50-85 x 1 mm, square to rectangular in t.s.,
becoming circular with age; 4-ridged or 4-winged, each wing < 0.25
mm tall, continuing at apex into lower margin of cotyledon petiole;
glabrous; green. Epicotyl 5-22 x 1 mm, ± square in t.s. when young,
becoming circular with age; 4-winged, each wing< 0.25 mm tall and
terminating apically in a stipule; glabrous; green. Cataphylls absent.
Internodes 1-16, 3-5 mm long; 4-winged, each wing about 0.25 mm
tall and terminating apically in a stipule; glabrous; green. Older
stems light brown to grey-brown, hollow. Buds: Cotyledonary buds
hidden in axils, sparsely pubescent. Lateral buds protruding out of
axils, < 0.5 mm long at nodes 1-15, sparsely short-pubescent at first
node, becoming densely pubescent at later nodes. Terminal buds
naked, sparsely pubescent at first node, becoming moderately pu-
bescent at later nodes. Hairs on all buds clear to white, mostly < 0.25
mm long.
COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, stipu-
late, emergent from seed, spreading apart, persistent until at least
leaves at nodes 2-3 mature. Blades 15-26 x 25-37 mm; straight,
flat, ± depressed obovate, with right half (seen from lower or abaxial
side) usually larger at both base and apex; apex truncate to emarginate,
often obliquely so, rarely ± rounded; base cordate, rounded, to ±
truncate, often obliquely so; margins entire. Venation palmate, 3-
nerved; acro-brochidodromous, basal and perfect. Primary veins 3,
± fine, slightly raised above, moderately raised below; central
primary vein forked, with straight stem '/2 to 3A distance to apex and
branches slightly recurved; lateral primary veins basal, about 3A
length of blade, mostly curved, diverging at narrowly acute angle,
looped, joining branches of central vein and forming ± small obovate
or triangular intercostal area or rarely branching before joining the
central vein and forming a ± indistinct intercostal area, the pair of
intercostal areas, when distinct, usually spanning about 30-50% of
the width of the cotyledon. Secondary veins moderate-sized basally
along lateral primaries becoming fine apically; 12-16, 0-1 along
midvein, 8-10 along lateral primaries, 4-5 along branches of midvein;
mostly marginal; to 2/3 length of blade, longest basally; mostly
arched or hooked; diverging at narrowly acute angles basally on
lateral primaries to perpendicular angles along branches of midvein;
looped marginally, acutely to obtusely joined; flush above, moder-
ately raised below. Petioles 2-3 mm long, transversely oblong in t.s.,
2-ridged above, with ridges continuing into lateral primary veins; 4-
ridged along margins, the lower ridges continuing down into the
ridges or wings on the hypocotyl and the upper ridges terminating at
the stipules; glabrous. Stipules separate and paired or sometimes
fused and interpetiolar, composed of two structures, both variable in
shape and development. The lower part a pair of ± conical projec-
tions between the petioles near the base of the upper marginal ridge
of each petiole, one or both sometimes absent, mostly 0.25-0.75 mm
long, usually perpendicular to the stem, sometimes gland-like
apically. The upper part usually a pair of flattened scale-like
outgrowths between and just above the petioles, initially folded over
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
the terminal bud but becoming erect and visible as the epicotyl
expands, each ± ovate to rhombic, 0.5-0.75 mm tall and wide, the
apex often 3-lobed and gland-like, the pair sometimes fused to-
gether, then 0.5-0.75 x 0.75-1.5 mm tall and wide and the apex
multilobed or irregular.
LEAVES. Opposite, decussate on seedling trunk axis, sometimes 3-
whorled on sapling trunk axis; simple, short-petiolate; stipulate.
Leaf blades 30-69 x 1 1-19 mm at first node; 30-82 x 13-17 mm at
nodes 2-15, to 195 x 52 mm on saplings; straight; narrowly obovate
to narrowly elliptic at nodes 1-15 and on sapling; apex acuminate or
long-acuminate, rarely acute at first node; bases cuneate to narrowly
acute; margins entire. Blades green, papery at early nodes becoming
subcoriaceous on larger saplings; sometimes moderately pusticulate
above and below on later leaves, the pustules pale above; the blade
nearly glabrous when fully expanded, the midvein below often
sparsely short-pubescent, but moderately to densely pubescent on
the blade and midvein below and along margins when expanding
leaves of seedlings are < 10 mm long. Hairs simple, basifixed, < 0.25
mm long, clear to white. Venation pinnate, brochidodromous. Midvein
moderate-sized; straight; flush to slightly raised above, prominently
raised below. Secondary veins thin, 14-22 at first node, to 22 at later
nodes; alternate; mostly '/4 to V3 length of blade, longest centrally;
mostly hooked or arched, diverging at a moderately acute angle;
looped, obtusely joined; slightly raised above and below.
Intersecondary veins infrequent. Tertiary veins reticulate in inter-
costal area, looped in marginal area. Petioles 1 mm long at first node,
to 2 mm long at nodes 2-15, to 7 mm on saplings; shallowly
channelled, with erect narrow margins above. Stipules paired; 1 .0-
1.2 mm long at first node, to 1.7 mm long at nodes 2-15; simple,
narrowly triangular; the inner margin and base continuing above the
petiole and apparently terminating near the lateral bud; the outer
margin and base usually terminating abruptly, but sometimes form-
ing a low interpetiolar ridge with the stipule of the opposite leaf as
the stem thickens at some early nodes; persistent; often sparsely
short-pubescent toward the apex.
Vochysia sp.
Fig. 2E-G.
COLLECTIONS. Panama. Colon Province: Santa Rita Ridge,
Garwood 135 (seedlings with seed fragment); Chiriqui Province:
Fortuna, Hampshire & Whitefoord 269 (BM, seedlings); 15 seed-
lings to 9 cm tall examined.
TAXONOMIC NOTE. Specific identification is not yet possible, as
Garwood 135 had only a seed fragment attached and Hampshire &
Whitefoord 269 lacked seeds altogether. In addition to Vochysia
ferruginea, V. allenii Standl. & L.O. Williams, V. jefensis A.
Robyns, and V. speciosa Warm, have been reported from eastern
Panama and V. hondurensis (- V. guatemalensis) from western
Panama (D'Arcy, 1987). As immature leaves of both collections
have T-shaped hairs > 0.25 mm long on the lower surfaces (see
below), it is not V. guatemalensis, which has simple basifixed hairs
< 0.25 mm long (see above). The T-shaped hairs on the first leaves
ally it with V. ferruginea, but the cotyledons are about 1.5-2 times
larger. Thus, the Chiriqui collection is a new species record for
western Panama. As the cotyledons from the Chiriqui collection
are consistently larger than those from the Colon collection (Fig.
3), two species might be represented. Given their similarity and the
paucity of material, they are described together below, with the few
differences noted.
Seedlings
Seedlings phanero-epigeal with photosynthetic cotyledons.
ROOTS. Primary root 45-70 mm long, 1 mm wide, circular in t.s.,
slightly sinuous, medium brown.
STEMS. Hypocotyl 50-86 x = 1 mm long and wide, ± square to
rectangular in t.s., 4-ridged to 4-winged, each wing < 0.25 mm tall
and continuing at apex into lower margin of petiole; glabrous.
Epicotyl (immature) 4-ridged or 4-winged, each wing < 0.25 mm
tall; glabrous. Buds: Cotyledonary buds hidden in axils (Garwood
1 35) or 0.75 mm tall and densely pubescent (Hampshire & Whitefoord
269).
COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, stipu-
late, emergent from seed, spreading apart, persistent. Blades 14-21
x 22-40 mm (14-17 x 25-32 mm in Garwood 135, 19-21 x 34-^0
mm in Hampshire & Whitefoord 269); straight, flat, depressed
obovate, with right side (viewed from lower or abaxial surface)
usually larger than or sometimes equal to left side; apex truncate,
emarginate, or cordate, sometimes obliquely so; base rounded,
truncate or emarginate, sometimes unequal; margins entire. Blades
green, glabrous. Venation palmate, 3-nerved; acro-brochidodromous,
basal and perfect. Primary veins 3, moderate-sized; central primary
vein forked, with straight stem2/3 to3A distance to apex and branches
recurved; lateral primary veins basal, about 3/4 length of blade,
mostly curved, diverging at narrowly acute angle, looped, joining
branches of central vein to form ± small obovate or triangular
intercostal area, the pair of intercostal areas usually spanning about
40-50% of the width of the cotyledon. Secondary veins moderate-
sized basally becoming fine apically; 10-16, 0-2 along midvein,
6-9 along lateral primaries, 3-5 along branches of midvein; mostly
marginal; '/3 to nearly full length of blade, longest basally; mostly
arched to hooked, diverging at narrowly acute angles basally to
perpendicularly apically; looped marginally, acutely to perpendicu-
larly joined; flush above, moderately raised below. Tertiary veins
reticulate within intercostal areas, looped in marginal areas. Petioles
about 2 mm long, transversely oblong in t.s., 2-ridged above, with
ridges continuing into the lateral primary veins; 4-ridged along
margin, with lower ridges continuing into hypocotyl ridges and
upper ridges terminating in the stipules; glabrous. Stipules usually
fused and interpetiolar, sometimes separate and paired; composed of
two structures, both variable in shape and development. The lower
part a pair of ± conical projections between the petioles near the base
of the upper marginal ridge of each petiole, sometimes absent or
bifid, mostly 0.25-0.5 mm long, usually perpendicular to the stem,
sometimes gland-like apically. The upper part usually a flattened
scale-like outgrowth between and just above the petioles, initially
folded over the terminal bud but becoming erect and visible as the
epicotyl expands, ± transversely oblong to depressed ovate, 0.5-
0.75 x 0.75-1.0 mm tall, 3- to 5-lobed and gland-like at the apex,
sometimes divided.
LEAVES (only immature leaves at first node present). Opposite,
decussate on trunk axis; simple, short-petiolate. Blades moderately
pubescent below and along margins when expanding blades are < 5
mm long, sparsely pubescent when half-expanded. Hairs 0.25-
0.625 mm long, T-shaped, sessile, pale orange-browa Venation
pinnate, brochidodromous.
Key to known seedlings of Vochysia in Panama and
Central America
1 . T-shaped hairs present on lower surface of young leaves at first and early
nodes
2. Cotyledons 13-23 mm wide and 10-15 mm long
V. ferruginea
N.C. GARWOOD
2. Cotyledons 22-40 mm wide and 14-21 mm long V. sp.
Simple basifixed hairs present on lower surface of young leaves at first
and early nodes or hairs absent; cotyledons 25-37 mm wide and 1 5-26
mm long V. guatemalensis
DISCUSSION
Vochysiaceae is a moderate-size, primarily neotropical family di-
vided into two tribes. Vochysieae includes Vochysia (100 species),
Callisthene (12 species), Salvertia (1 species), and Qualea sensu
lato (77 species, including Ruizterania); Erismeae includes Erisma
(16 species) and Erismadelphus (2 species), the only African taxa
(Stafleu, 1948-1954; Keay & Stafleu, 1953; Marcano-Berti, 1969;
Kawasaki, in press). The monotypic Euphronia (= Lightia), some-
times included in Vochysiaceae (Lleras, 1976; Cronquist, 1981) or
segregated as the Euphroniaceae (Marcano-Berti, 1989, 1990), is
not closely related to Vochysiaceae according to recent molecular
studies (Litt et al., 1995) and will not be further discussed.
Fruit
There are two basic types of fruit in Vochysiaceae (Warming, 1 875;
Stafleu, 1948- 1954; Keay & Stafleu, 1953). The dehiscent trilocular
capsules of Vochysieae, derived from superior ovaries, contain
winged seeds. The indehiscent unilocular fruits of Erismeae, derived
from inferior to semi-inferior ovaries, have persistent enlarged
winged calyx lobes. Within Vochysieae, fruits of Callisthene have a
thick persistent central column and a fragile exfoliating exocarp not
found in the other genera. Several of the 12 ovules per locule mature
into seeds in Qualea (Stafleu, 1953); one or more of the few ovules
per locule mature in Callisthene, (Stafleu, 1952). In Vochysia and
Salvertia, however, only one of the two ovules in each locule usually
develops into a mature seed (Warming, 1875; Stafleu 1948; Standley
&Steyermark, 1949;Robyns, 1967a;Gaos, 1978;Moreira&Arnaez,
1994; this study), although two mature according to Flores (1993a,
b).
Fruits of Vochysia ferruginea are typical of the genus (Stafleu,
1948), being 3-locular loculicidal capsules, oblong or obovate in
outline and 3-angled. Variation in size, shape, and surface texture of
fruits in Vochysia is not well-documented: fruits of many species are
still unknown (Warming, 1875; Stafleu, 1948; but see van Roosmalen,
1985). The gross morphology of fruits has been little studied. Stafleu
( 1 948) described the pericarp of the genus as consisting of a black or
dark bluish exocarp 'strongly adhering to the shining golden-yellow
endocarp' , an observation repeated by later authors (Robyns, 1 967a;
van Roosmalen, 1985), but Flores (1993a, b) noted the presence of
a well-developed mesocarp in both V. ferruginea and V? guatemalensis.
In V. ferruginea, there is a brown to black exocarp, a light to medium
brown spongy to woody mesocarp, thickest near the centre of the
fruit, and a pale crustaceous endocarp of even thickness around each
locule (Fig. 1C). The mesocarp is penetrated by longitudinal resin
canals adjacent to the locules (Fig. 1C) and is also well-vascularized
(Flores 1993a, b). In mature fruit, the endocarp is as well-developed
around empty locules as around those filled with mature seeds,
indicating that development of each locule is not dependent on the
growth of fertilized ovules within it. The locules complete develop-
ment before the seeds and determine the shape of the seed wing
(Boesewinkel & Venturelli, 1987).
Seed
Seeds are winged in the Vochysieae. The wings are large and
unilateral in Vochysia, Qualea, and Salvertia, but small and circum-
ferential in Callisthene, and are formed from compacted testal hairs
in Vochysia and Salvertia, but from extensions of the testa in Qualea
and Callisthene (Warming, 1875; Stafleu, 1948-1953; Boesewinkel
& Venturelli, 1987). The seeds of V. ferruginea are typical of the
genus (Warming, 1875). The wing is on the anti-hilar side of the
seed, and the micropyle extends along the seminiferous area and
terminates near the hilum, as in V. pygmaea Bong, and Salvertia
(Boesewinkel & Venturelli, 1987). In contrast, the wing of Qualea is
on the hilar side of the seed, such that the micropyle extends along
the anti-hilar margin of the wing and opens at the apex of the wing
away from the hilum (Boesewinkel & Venturelli, 1987). The final
shape of the wing in Vochysia is determined by the shape of the
locule: the unicellular hairs grow upward from the testa to fill and
take their shape from the empty locule (Boesewinkel & Venturelli,
1987). Hence, much of the variation in size and shape of seeds
among species in Vochysia probably arises from differences in the
size and shape of the mature locules.
There has been little work on seed anatomy of Vochysiaceae
(Corner, 1976; Boesewinkel & Venturelli, 1987). Stafleu (1948,
1 953) described the testa of Vochysia as brown, thin, and chartaceous,
and the tegmen as very thin, paper white, and adhering to the testa,
and the testa of Qualea as chartaceous. The mature seed coat of V.
pygmaea, however, consists only of the outer layers of the testa,
because the inner layers of the testa are crushed and the tegmen
resorbed during development (Boesewinkel & Venturelli, 1987).
The seed coat also appeared as one undifferentiated layer at maturity
in V. ferruginea (this study), but Flores (1993a, b) described a dark
exotesta distinct from the dead cell layers and air spaces of the
mesotesta and endotesta and remnants of the tegmen in V. ferruginea
and V. guatemalensis. In contrast, the mature seed coat of Qualea
densiflora Warm, and Callisthene retains both the inner layers of the
testa and tegmen (Boesewinkel & Venturelli, 1987). The seed coat of
Erisma and Erismadelphus is thick and contains a large branched
vascular bundle (Boesewinkel & Venturelli, 1987). The vascular
bundle of V. ferruginea, hitherto undescribed in Vochysieae, is
unbranched and terminates at the tip of the seminiferous area. The
hilum of V. ferruginea (7-10 mm long) is longer than in V. pygmaea
(3-4 mm long), and stretches over a greater proportion of the length
of the winged seed (35-40% versus about 20%). Due to the long
hilum, the seed type of V. ferruginea is described as hilar-anatropous.
Endosperm is absent in mature seeds of Vochysiaceae (Bentham
& Hooker, 1867; Baillon, 1874; Warming, 1875; Petersen, 1896;
Stafleu, 1948-1953; Standley & Steyermark, 1949; Robyns, 1967a;
Gaos, 1978; Boesewinkel & Venturelli, 1987; Flores, 1993o, b).
None was seen in Vochysia ferruginea. Occasional reports that
endosperm is rarely well-developed (Cronquist, 1981; Niembro
Rocas, 1989) do not appear well-founded. A thin, whitish aril,
adhering to the testa, was reported in V. guatemalensis (Gaos, 1978)
and in the family (Niembro Rocas, 1989): this appears to be a
mistranslation of Stafleu's (1948) description of the tegmen (see
above).
Embryo
In all genera, the embryo of the mature seed is straight and the
plumule and radicle poorly developed (Warming, 1875; Stafleu,
1948-1954). In Vochysia ferruginea, the embryonic axis is well-
developed; the hypocotyl is relatively long compared to the poorly
developed radicle, compressed, 4-winged, and annulate at the collet.
The winged hypocotyl and annulate collet persist into the seedling
stage, confirming the identification of the structures in the seed. In
Vochysieae, the embryonic cotyledons are generally thin, folia-
ceous, and folded in various ways (see below), whereas, in Erismeae,
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
they are thick, fleshy, and not or scarcely folded (Baillon, 1874;
Warming, 1875; Boesewinkel & Venturelli, 1987; Kawasaki, in
press). The cotyledons of Erismadelphm are plano-convex and
slightly convolute at the margins (Boesewinkel & Venturelli, 1987),
while those of Erisma are narrowly elongate and semi-cylindrical
(Baillon, 1874), plano-convex (Boesewinkel & Venturelli, 1987), or
flat (Stafleu, 1954).
The embryonic cotyledons have been described as unequal in
Vochysia (Warming, 1875; Stafleu, 1948) and Salvertia (Stafleu,
1 948), although no comparable information was given for Qualea or
Callisthene. Unequal could mean either that each cotyledon is
asymmetrical or that the two cotyledons are heterocotylar, differing
in shape or size. Cotyledons are asymmetric in two species of
Vochysia and one species of Qualea but symmetric in one species of
Vochysia and Salvertia (Table 1 ), based on the illustrations of the
unfolded embryo in Warming ( 1 875). In V.ferruginea, each cotyle-
don is usually asymmetric, with the right side (viewed from the
lower abaxial surface) larger, at least basally. The base of the smaller
side tends to be rounded, and that of the larger side cordate, as in the
expanded cotyledon of the seedling (Fig. U). The embryonic coty-
ledons of V. ferruginea, when unfolded, and the fully expanded
cotyledons of the seedling (Fig. 1 H, P), are isocotylar. The larger
side of one cotyledon opposes the smaller side of the other on the
embryo and seedling, hence, the two cotyledons are identical in
shape, not mirror-images.
Most monographic and floristic works indicate that the embry-
onic cotyledons of Vochysiaceae are folded, but give insufficient
detail to characterize the complex pattern of folding. It is usually
unclear which component of folding is described: ptyxis, the folding
of an individual organ such as a leaf in the bud or cotyledon in the
seed, or vernation, the relationship of the immature organs to each
other (Bell, 1991). Some terms, such as convolute, are used for both
ptyxis and vernation but have different meanings (Bell, 1991).
Definitions of terms vary considerably among authors.
In Vochysia ferruginea, each cotyledon is folded four times
around the longitudinal axis of the embryo (Fig. 1E-G). The folds
(f,-f4) are parallel to the radicle but not parallel to the midvein of
the cotyledon (Fig. IF). The margin of the larger side of the
cotyledon (the right side as viewed from the lower abaxial surface)
is on the exterior of the embryo, while the margin of the smaller
side is in the interior. The folded smaller side is completely en-
closed within the folded larger side. The inner margin is narrowly
bent back with the adaxial surfaces appressed at the first fold (f,).
Because of the oblique folding of the cotyledon, the base of the
larger side of each asymmetric cotyledon is folded around the
hypocotyl, nearly obscuring it (Fig. IF). Except for the com-
pressed nature of the embryo and distinct lines of folding, ptyxis is
essentially supervolute (often called convolute) but oblique. It is
not conduplicate, although the cotyledon appears to be folded
around the midvein in cross-sections through the middle of the
embryo (Fig. 1G). Vernation is essentially convolute sensu Bell
(1991), although unusually compressed. Each cotyledon partially
overlaps the other, with inner margins opposite near the centre of
the embryo and outer margins opposite on the exterior (Fig. 1G).
Confusingly, vernation is not convolute sensu Radford et al.
(1974), because one rolled or supervolute cotyledon is not com-
pletely enclosed within a second rolled cotyledon.
Convolute vernation and supervolute ptyxis are also found in
Vochysia pygmaea, V. tucanorum Mart., and Salvertia convallari-
dorea A. St.-Hil., as seen in transverse sections of the embryo
(Warming, 1875: figs 16-11, 17-11; Boesewinkel & Venturelli, 1987:
fig. 10B). The embryos are less compressed and angular than in V.
ferruginea, and the inner margins of the cotyledons overlap rather
than being folded back individually. Embryos of V. guatemalensis, V.
obscura Warm., V. tucanorum, and 5. convallaridorea (Warming,
1875: figs 13, 1611, 1711; Flores, 1993a: fig. 9) are externally similar
to that of V. ferruginea, and the position of the outer cotyledon
margin is also consistent with convolute vernation. Ptyxis is prob-
ably oblique is most species because the hypocotyl is hidden, as in V.
ferruginea, but may differ in V. obscura, given that the hypocotyl is
well-exposed in spite of the deeply lobed cotyledon base. Although
folding appears somewhat angular in section because the seed is
compressed, cotyledons are not plicate in the sense of fan-like. Thus,
there is one basic pattern of vernation and ptyxis for cotyledons of
Vochysia and Salvertia, with minor variations, whether cotyledons
are asymmetric or not.
Embryonic cotyledons of the genus Vochysia have previously
been described as convolute or rolled (Bentham & Hooker, 1867;
Petersen, 1896; Flores, \993a, b), spirally convolute (Baillon, 1874),
obliquely spirally convolute (Warming, 1875), spirally folded
(Boesewinkel & Venturelli, 1987), and spirolobal (Stafleu, 1948).
The cotyledons of the species discussed above are not spirolobal, a
term now used for bent embryos with incumbent once-folded coty-
ledons (Duke, 1969;Radfordetal., 1974). The remaining descriptions
are not inconsistent with convolute vernation and/or obliquely
supervolute (= convolute) ptyxis, but do not completely characterize
ptyxis and vernation.
In Qualea, embryonic cotyledons have been described as convo-
lute or rolled (Petersen, 1896; Boesewinkel & Venturelli, 1987) or
convolute and plicate (Warming, 1875; Stafleu, 1953). The cotyle-
dons of Q. grandiflora Mart. (Warming, 1875: fig. 8-II) are more
complexly folded than in Vochysia, although the complete pattern
cannot be resolved from the embryo and its cross-section. Supervolute
ptyxis and convolute vernation, as in Vochysia, can be eliminated,
however, and folding may be irregular but is not fan-like. The
exposed margins of the cotyledon are very uneven, almost lobed. In
contrast, the exposed cotyledon margins of Q. multiflora Mart.
(Warming, 1875: fig. 7-II) are unlobed and parallel, similar to
Vochysia except that they are close together on the exterior of the
embryo. There appears to have been some folding around the
longitudinal axis through the radicle, but the specific pattern is again
unknown. In both species of Qualea, the folded cotyledons nearly
obscure the embryonic axis. Thus, ptyxis and vernation appear very
different in these two Qualea species compared to the homogeneous
pattern observed in Vochysia.
Embryonic cotyledons in Callisthene have been described as
plicate and/or convolute (Warming, 1875; Petersen, 1896; Stafleu,
1952; Boesewinkel & Venturelli, 1987). In C. major Mart. (Warm-
ing, 1875: fig. 3-II), the embryo is straight and compressed. Each
cotyledon is irregularly folded, appearing more wrinkled (corru-
gate) than plicate on the surface; each margin is involute, folding
back with the adaxial surfaces touching. The folded cotyledons are
pushed together for most of their length and width, but the folds are
only rarely synchronous between the two. The involute margin of the
first cotyledon is exposed at the edge of the embryo and overlaps the
involute but hidden margin of the second, whereas the exposed
involute margin of the second cotyledon abuts, rather than overlaps,
the hidden margin of the first. No standard terms describe this
pattern of folding, but corrugate-involute ptyxis and semi-convolute
or valvate-convolute vernation might be used. In contrast to other
genera in Vochysieae, the embryonic axis is fully exposed and the
embryo appears almost spathulate, except for the wrinkled slightly
overlapping cotyledons.
Embryos of Vochysia, Qualea, and Callisthene are rich in lipids
and poor in starch, those of Erisma are poor in lipids and starch, and
those of Erismadelphus are rich in lipids and starch (Boesewinkel &
10
Venturelli, 1987). The seeds of V. hondurensis are rich in protein, as
well as lipids (Blanche et al., 1991).
Seedling
There is little published information on seedlings and saplings of
Vochysiaceae. Only Vochysia duquei Pilg., V. guatemalensis, and V.
ferruginea (del Amo, 1979; Ricardi et al., 1987; Flores, 19930. b;
this study) have been described in detail. Limited data are available
for ten additional species and some information can be inferred from
the embryos of five species (Tables 1-2). In Erismeae, cotyledons of
Erismadelphus are cryptocotylar, hypogeal, and food-storing. Em-
bryonic cotyledons ofErisma are also fleshy and food-storing, but it
is not known whether these are cryptocotylar or phanerocotylar, or
epigeal or hypogeal, after germination. No other information on
seedlings is available for Erismeae.
In Vochysieae, cotyledons are all phanerocotylar, epigeal, and
foliaceous (Table 1 ). Where known, they persist after the first leaves
are produced, and presumably continue to photosynthesize. In
Vochysia thyrsoidea Pohl, they are the major photosynthetic organ
20-
10-
15 20
Length (mm)
Fig. 3 Relationship of cotyledon length and width in Vochysiaceae.
Vochysia species: circles, V. ferruginea (Panama: Garwood 1630;
Garwood 2005; Foster s.n.; Costa Rica: Flores, 19936); squares, V.
guatemalensis (Belize: Garwood 4031; Monro 868-871, Monro et al.
M4162X; del Amo, 1979; Costa Rica: Flores, 1993a); diamonds, V. sp.
(Panama: Hampshire & Whitefoord 269); octagons, V. sp. (Panama:
Garwood 135); inverted triangles, V duquei (Venezuela: Ricardi et al.,
1987); open triangle, V. surinamensis (Guyana: Polak, 1992); triangle
with black circle, V. sp. (French Guiana: Oldeman & Fundter, 1986);
triangle with white circle, V thyrsoidea (Brazil: Rizzini, 1965). Other
genera (stars): C, Callisthene major (Brazil: Labouriau et al., 1964); Q,
Qualea grandiflora (Brazil: Thomazini et al., 1973). Shading of
symbols indicates country of origin: black, Panama; cross-hatching,
Belize; horizontal hatching, Costa Rica; vertical hatching, Mexico;
white. South America. Dashed lines represent width to length ratios of
2.0 (upper line) and 1.3 (lower line). Data are measurements of all
seedlings in each collection, values cited in references, or my estimates
from illustrations in references where values are not given. Overlapping
points have been off-set by 0.5-1 .0 mm to improve readability.
N.C. GARWOOD
for many months (Rizzini, 1965). Cotyledons range from about 1 to
3 cm long and 1 to 5 cm wide (Fig. 3). They are smallest in
Callisthene major and largest in an unidentified species of Vochysia
(Fig. 3). Within Vochysia, those of V. ferruginea are the smallest (Fig.
3). Cotyledons are mostly 1.3 to 2.0 times wider than long in
Vochysia, Salvertia, and Callisthene, but only 0.8 to 1.3 times wider
than long in Qualea, and broadest above the middle in Vochysia and
Salvertia. They are broadest around the middle in Callisthene and
Qualea (Table 1; Fig. 3). They are asymmetric or symmetric in
Vochysia (Fig. U and 2 A, E-F), as in the embryos, but this may be
variable within some species or collections (Table l).The apex of Q.
grandiflora was short-acuminate, that of C. major ± obcordate, and
those of Vochysia truncate, rounded, retuse or rarely obcordate (Fig.
U and 2A, E-F; Table 1 ). Venation can only be assessed in Vochysia,
but the pattern is unusual. The midvein forks toward the apex,
whether the apex is rounded or lobed, and two strong lateral prima-
ries diverge from the base: in most species, the lateral primaries join
with the recurved branches of the midvein to form triangular to
obovate intercostal areas (Fig. U and 2A, E-F; Table 1). Smaller
veins form one or more concentric series of loops around the central
intercostal areas.
The cotyledonary stipules in the three species of Vochysia de-
scribed in this study are unusual two-parted structures (Fig. 1K-M;
Fig. 2C-D, G). The lower part usually consists of one to three narrow
projections between the petioles, usually perpendicular to the
hypocotyl, situated at the base of the upper (adaxial) surface of the
petiole where the upper ridge or wing of the petiole terminates. The
upper part consists of a flattened interpetiolar scale-like outgrowth
just above the petioles, initially folded over the terminal bud but
becoming erect and visible as the epicotyl expands, completely or
partially divided or torn into two to five parts but rarely unlobed. The
apex of the lower and/or upper parts often appears gland-like.
Neither part appears to be fused over the petioles, and the axillary
buds are often visible above the cotyledonary petiole. Although
there was some intraspecific variation, the stipules of V. ferruginea
usually had one lower projection and a lobed upper part and those of
V. guatemalensis two lower projections and two separate upper
parts. The reduced stipules of V. guatemalensis have also been
described as 2-3 trichomes of a glandular nature (Flores, 1993a), but
the thick 3-lobed pair of structures illustrated (fig. 7e) appear rather
large and fleshy for trichomes and may be a different form of the
variable structures described in this study. Two-parted cotyledonary
stipules similar to those of Vochysia have not been seen in seedlings
of more than 600 dicotyledonous species in 88 families surveyed
from Panama (Garwood, in prep.), although the fleshy cotyledonary
stipules of Lafoensia (Lythraceae) are similar to the lower stipular
parts of Vochysia.
Leaves are opposite at the early nodes in Vochysia, Qualea, and
Callisthene (Table 2). In Vochysia, the pair of leaves at the first node
is decussate with respect to the cotyledons and leaves at the second
node (Table 2). In Q. grandiflora, the first pair of fully developed
leaves is parallel to the cotyledons but decussate to the second pair
of leaves (Table 2); a pair of 'prophylls' between the cotyledon and
first pair of leaves (Thomazini et al., 1973) might be vestiges of the
'missing' pair of decussate leaves or homologous to the interpetiolar
cotyledonary stipules of Vochysia. Leaves remain opposite into the
adult stage in V. ferruginea, but become 3- then 4-whorled in
V. guatemalensis sometime in the sapling stage. The initial axis of C.
major appears to be plagiotropic, with opposite distichous leaves
smaller than the cotyledons, but this will need confirmation because
it is difficult to interpret details in the published photograph of the
seedlings (Labouriau et al., 1964: fig. 26).
Stem characters are known only from Vochysia. The collet, the
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
11
Table 1 Cotyledon characters in seedlings and embryos of Vochysiaceae.
Genus
Species
Source
Emer-
gence
Posi-
tion
Devel-
Persistence
Width:Length
Ratio
Shape
Base
Apex Symmetry Venation Stipules
opment node months
Vochysia
V. densiflora Spruce
18*
-
-
F
-
1.7
2
5-7
8
A
V duquei Pilg.
15
P
E
Fr
2 >1
1.6-1.9
2-3,6
4-6
3-5
S 7,2 +
V.ferruginea
6
P
E
F
3-8 >9
1.3-1.9
2
3-6
6-7
A 1-3 f+f,g
Mart.
10
-
E
-
-
-
_
_
_
_ _ _
4b
P
E
F
2
1.6
2
6
5
8^4 7 f
V. guatemalensis
6
P
E
F
3
1.2-1.8
2
3-6
3-5
A 1-3 s+s,g
Donn. Sm.
10
-
E
-
- _
_
_
_
_
_ _ _
4a
P
E
F
/
1.2-1.3
2
6
4
A 7,4 s,g
1,5
P
E
F
6
1.5
2
4-5
4
A 1,2
V. obscura Warm.
18*
-
-
F
- -
1.4
2
7
4-5
S
V surinamensis Stafleu
14
P
E
n
6
2.7
1
—
3-4
— — _
V. thyrsoidea Pohl
13
-
E
-
= 12
-
-
-
-
_ _ _
16
P
E
Fr
1 >12
= 7-2
7
2
4
_ _ _
V. tucanorum Mart.
18*
-
-
F
- -
7.6
2
6-7
3^t
A
V. sp. (Brazil)
2
P
E
F
- -
-
1
_
-
— _ _
V. sp. (Fr. Guiana)
11
P
E
F
-
2.7
2
4
5
5 2,5
V. sp. (Panama)
6
P
E
F
1
1.4-2.1
2
3-5
4-6
S,A 1-3 s+f,g
Salvertia
S. convallaridorea
18*
-
-
F
_ -
1.6
2
2-3
4-5
S
A. St.-Hil.
Qualea
Q. grandiflora
12
P
E
F
2 >6
-
-
-
-
_ _ _
Mart.
3
P
E
F
_ 4
-
-
—
_
_ _ _
17
P
E
F
2 >6
1.1
5
3^f
7
Al P
Q. multiflora Mart.
18*
-
-
F
-
0.8-1.3
4
3
9
A
Callisthene
C. major Mart.
8
P
E
Fl
1-3?
1.2-1.5
3
-
5-6
_ _ _
18*
-
-
F
-
1.2-1.6
-
-
-
- - -
Erisma
E. spp.
7*
-
-
R
-
-
-
-
-
-
Erismadelphus
E. exsul Mildbr.
9
C
H
R
-
-
-
-
-
_ _ _
E. sp.
9
C
H
R
-
-
-
-
-
Source: 1, del Amo 1979; 2, Duarte 1979; 3, Felippe & Dale 1990; 4a,b, Flores 1993a, b; 5, Gaos 1978; 6, Garwood, this study; 7, Kawasaki, in press; 8. Labouriau et al. 1964:
9, Miquel 1987; 10, Moreira & Arnaez 1994; 11, Oldeman & Fundter, 1986; 12, Paulilo et al. 1993; 13, Paviani & Haridasan 1988; 14, Polak 1992; 15, Ricardi et al. 1987; 16,
Rizzini 1965; 17, Thomazini et al. 1973; 18, Warming 1875. *, all characters based only on the embryonic cotyledon in the mature seed.
Emergence: P, phanerocotylar; C, cryptocotylar.
Position: E, epigeal; H, hypogeal.
Development: F, foliaceous; R, fleshy, presumably with stored reserves; r, somewhat fleshy or succulent, with reserves or stored water.
Persistence: measured as i) the last recorded leaf node to produce leaves while the cotyledons are still present (e.g. node=3. cotyledons still present when leaves produced at
node 3) or ii) the length of time [months] that cotyledons persist after germination. These measures underestimate persistence when only the youngest seedling stages have
been studied.
Width:Length Ratio: width divided by length.
Shape: 1, obtriangular to shallowly obtriangular, including 'triangular'; 2, depressed obovate; 3, transversely oblong; 4, very widely obovate; 5, transversely widely elliptic to
transversely widely oblong; 6 transversely reniform, rectangular-reniform.
Base & apex: 1, short-acuminate; 2, acute; 3, rounded; 4, truncate; 5, retuse to emarginate; 6, cordate, obcordate, 2-lobed; 7, 2-cleft or -parted; 8, 3-lobed; 9, irregular. When
blade is asymmetric, the base and/or apex may be unequal on the two sides (e.g. half-cordate, half-rounded) or oblique (e.g. obliquely truncate).
Symmetry of blade: A, asymmetrical; S, symmetrical.
Venation: 1, branches of forked central primary vein join lateral primaries forming a pair of obovate to triangular intercostal areas; 2, palmate or 3-nerved; 3, basal acro-
brochidodromous; 4, basal actinodromous; 5, branches of central primary vein forked; lateral primaries each form a closed loop apically, terminating near but not joined
directly to the tips of the central vein.
Stipules: s, usually separate; f, usually fused and interpetiolar; f+f, f+s, and s+s, two structures (fused or separate) present at cotyledonary node (first is lower structure, second
is upper structure); g, gland-like; +, present; P 'prophylls' present above cotyledons.
-, Information not available.
Characters in italics were inferred or estimated from illustrations in the citations, not from the text of the original author. Where authors identified the cotyledons as foliaceous,
it was assumed that they were also phanero-epigeal.
junction of the radicle and hypocotyl, is annulate in at least three
species (Table 2; Fig. IF). The hypocotyl, epicotyl, and later
internodes are usually square, with ridges or wings on each corner
(Table 2). Ridges on the hypocotyl continue without interruption
into the ridges on the lower margins of the cotyledonary petioles in
V.ferruginea and V. guatemalensis (Figs IK, 2G; Flores, 1993a, b).
In contrast, the wings or ridges on the epicotyl and later internodes
terminate in small narrowly triangular to linear stipules at the top of
each internode (Fig. 1O, S; Table 2), with the inner margin and base
of the stipules apparently extending above the petiole but not
attached to it. The winged stems do not originate directly from
decurrent leaf bases, as sometimes suggested, because the attenuate
base of the blade narrows into a medial pair of short thin ridges on
the upper (adaxial) surface of the petiole, interrupting the lamina
12
N.C. GARWOOD
Table 2 Root, stem, and leaf characters in seedlings of Vochysiaceae.
Genus
Species
Source
Collet
Hypocotyl
Epicotyl
Internodes
Phyllotaxis
Leaf
stipules
Nodes 1-2
Later nodes
Vochysia
V duquei
15
a
s
s
0
-,v
+
V. ferruginea
6
a
s,w
s,w
s,w o,d
o,d
+,s
4b
c
s
-
o,d
o,d
-
V. guatemalensis
6
a
s,w
s,w
s,w o,d
o,w
+,s
4a
c
s,w
s,w
o,d
o,w
-
1,5
c
c,w
s,w
s,w o,d
0
+,i
V. surinamensis
14
-
-
-
S,W 0
o,v
-
V. thyrsoidea
13
c
t
-
-
-
-
V. sp. (Panama)
6
-
s,w
w
0
-
-
Qualea
Q. grandiflora
17
-
-
-
o,p
-
—
Callisthene
C. major
8
-
o
a?
Sources and authorities for names: see Table 1.
Collet (junction of hypocotyl and primary root): a, annulate ridge present; c, constriction, abrupt change, or other feature present, but nature of structure not discernable from
illustration.
Hypocotyl. Epicotyl, and Internode: cross-sectional shape of hypocotyl, epicotyl, and early internodes (about first ten), respectively: s, square to rectangular; c, circular; t,
tuberous and circular, at least when older; w, 4 wings or ridges present.
Phyllotaxis:
Nodes 1-2: o, leaves at first node opposite; d, pair of leaves at first node decussate to both cotyledons and pair of leaves at second node; p, pair of leaves at first node parallel to
cotyledons and decussate to pair of leaves at second node.
Later nodes: o, leaves opposite and decussate (at least to node 5, sometimes reported up to node 20); d, leaves opposite and decussate on larger seedlings, saplings, and adults;
w, leaves whorled on larger seedlings or saplings (and adults), with three or four leaves per node, size of juveniles at time of transition from opposite to whorled leaves
unknown; v, juvenile leaves unknown, but adult leaves whorled; a, axis becoming plagiotropic with pairs of leaves distichous.
Leaf stipules: +, small, narrowly triangular to linear stipules present from the first node; i, stipules intrapetiolar; s, stipules separate.
-, Information not available.
Characters in italics were inferred or estimated from illustrations in the citations, not from the text of the original author.
between the blade and wings. It is unclear, however, whether the
petiolar ridges are joined to the lateral extensions of the stipules.
Striking ontogenetic changes are seen in hair morphology in
Vochysia ferruginea (this study). The epicotyl and first leaves are
only sparsely pubescent below, but stems and leaves at higher
nodes become progressively more densely pubescent. Hairs on
early internodes and lower leaf surfaces are nearly sessile,
appressed, and T-shaped, with straight arms of equal length; those
on later leaves and stems are short-stalked, ascending to erect, 2-
branched, with the longer arm curved to coiled and about eight
times longer than the shorter arm. Hairs on the adult leaves are
similar, except the branches are even more unequal in length
(Garwood, pers. obs.)- On seedlings to adults of V. ferruginea,
hairs on the midrib above are always simple. In Vochysia sp.
(Garwood 135 and Hampshire & Whitefoord 269), hairs on the
epicotyl and eophylls were similar to those on V ferruginea. In
contrast, hairs on the young expanding leaves of V guatemalensis
were simple and basifixed, but these were usually sparse or absent
on mature leaves (Flores, 1993a; this study). All parts of V duquei
are glabrous (Ricardi et al., 1987).
Seedlings from the Brazilian cerrado have a number of special
morphological adaptations to the dry cerrado environment (Labouriau
et al., 1964; Rizzini, 1965; Paviani & Haridasan, 1988). Vochysia
thyrsoidea has ± fleshy succulent cotyledons, a thick watery
hypocotyl which is mostly buried, minute eophylls which remain
hidden between the cotyledons, greater growth of the taproot rather
than shoot in the first few months, and a swollen subterranean organ
which forms from the hypocotyl and upper part of the taproot in the
first two years. Qualea grandiflora also forms a swollen under-
ground organ.
Phylogeny
There have been no phylogenetic analyses of relationships within
Vochysiaceae. Since the completion of Stafleu's ( 1 948- 1 954) mono-
graph more than 40 years ago, the numbers of collections and
species described have increased, but major systematic work has
been limited to elevating one section of Qualea to Ruizterania
(Marcano-Berti, 1969) and monographing Erisma (Kawasaki, in
press). Fruit and seed characters have long been important in delim-
iting genera within the family, but the utility of seedling characters
has been untested because seedlings have not been available for
study. The potential of the new seedling and seed characters de-
scribed above to contribute to phylogenetic and systematic studies is
discussed here. If such traits look promising, data on more species
should be collected.
Phanero-epigeal seedlings are reported only from Vochysieae,
and crypto-hypogeal seedlings from Erismeae, following the current
tribal division. In Vochysieae, shape characters of the foliaceous
cotyledon fell into three groups corresponding to Qualea, Callis-
thene, and Vochysia/Salvertia, although some variation occurred in
all groups. Symmetry of the cotyledons varied most in Vochysia, and
vernation and ptyxis of the embryonic cotyledons inQualea. Several
traits generally uncommon in dicotyledon seedlings occur together
in Vochysia, including unusual cotyledonary stipules and
cotyledonary venation, annulate collet, and winged hypocotyls and
early internodes, but these have not been assessed in many species or
other genera. Supra-cotyledonary prophylls, also unusual in dicoty-
ledon seedlings, were reported in one species of Qualea. The
existence of considerable variation in cotyledon characters, and the
presence of many unusual seedling characters in this small sample of
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
13
about 20 species (Tables 1-2), suggests further study of other taxa
would be profitable.
Ontogenetic changes from the seedling to adult stages can also be
important phylogenetic characters. In Vochysia, leaves of adults are
either opposite or 3- or 4-verticillate, but the first seedling leaves of
all species described are opposite (Table 2). In V. guatemalensis,
leaves remain opposite at early seedling nodes, but gradually be-
come 3- then 4-verticillate in larger saplings (Flores, 1993o).
Although presence of opposite seedling leaves in all taxa would not
be phylogenetically informative, the timing of the switch from
opposite to verticillate leaves may provide a useful character. Adult
leaves are also verticillate in Salvertia and some species of Erisma,
but opposite in other taxa (Stafleu, 1948-1954).
Pronounced ontogenetic changes in hair morphology were ob-
served in Vochysia ferruginea, but not in V. guatemalensis. Because
studies of ontogenetic changes of trichomes on leaves of seedlings
and adults have helped identify groups of species within the large
and complex genera of Solanum and Rhododendron (Seithe 1979,
1980; Seithe & Anderson, 1982), further study would be warranted.
Unfortunately, the morphology of hairs in Vochysia is poorly known.
They have been described as simple or sometimes unicellular and 2-
branched (Warming, 1875; Metcalfe & Chalk, 1950); hairs have
arms of equal length and short stalks in at least one species
(Shanmukha Rao & Vijayalakshmi Sarma, 1992). Stafleu (1952)
indicated that simple and biramose hairs occurred in Vochysieae, but
did not specifically describe those of Vochysia (Stafleu, 1948),
although frequent references to 'peculiar indumentum' or 'charac-
teristic indumentum' suggests that their morphology is unusual.
An examination of adult specimens of Vochysia at BM indicates
that species with 2-branched or simple hairs are both common
(Table 3). The distribution of simple and branched hairs is clearly not
random with respect to the sections of Vochysia delimited by Stafleu
(1948). Branched hairs are especially abundant in subsection
Ferrugineae, which contains V. ferruginea. There was considerable
variation among species in the morphology of the branched hairs of
the adult stage, ranging from nearly sessile appressed T-shaped hairs
with straight branches of equal length, to long-stalked erect 2-
branched hairs with curled arms of equal or unequal length (Garwood,
pers. obs.). In some species, the smaller arm was so reduced in
length that the hair could easily be misinterpreted as simple, sug-
gesting that at least some of the simple hairs in Vochysia might be
derived from much-reduced branched hairs, as occurs in Malpighi-
aceae (Anderson, 1990). The range of hair morphology from the
seedling to adult stages in V. ferruginea spans nearly the entire range
of branched hairs seen in the genus. Therefore, examining the
ontogeny of branched hairs might reveal similarities in the early
stages that would be phylogenetically informative in comparison
with other species of Vochysia, or other genera. Hairs are also simple
or 2-branched in Qualea (Stafleu, 1953), but stellate in Erisma
(Warming, 1875; Stafleu, 1954), and simple in Callisthene (Stafleu,
1952), Salvertia (Garwood, pers. obs.), and Erismadelphus
(Garwood, pers. obs.).
Seed, fruit, and seedling characters also bear on the higher order
relationships of the family. Vochysiaceae has usually been included
in the Polygalales with Polygalaceae, Trigoniaceae, Malpighiaceae,
Xanthophyllaceae, Tremandaceae, and Krameriaceae (Dahlgren,
1980;Takhtajan, 1980; Cronquist, 1981). Boesewinkel & Venturelli
(1987) used anatomy of the seed coat to reassess the position of the
family in Polygalales. They concluded that only Polygalaceae and
Xanthophyllaceae should be retained in the Polygalales, and that the
relationship of Vochysiaceae and the remaining families needed
further study. Reviewing numerous floral and vegetative characters
in Malpighiaceae, Anderson (1990) decided that Malpighiaceae was
most similar to Vochysiaceae and Trigoniaceae and least similar to
Polygalaceae and Krameriaceae. The presence of 2-branched uni-
cellular hairs in Vochysiaceae, Trigoniaceae, and Malpighiaceae,
and their absence in Polygalaceae, also supports this grouping
(Shanmukha Rao & Vijayalakshmi Sarma, 1992).
Recent molecular analyses of the angiosperms using the
chloroplast gene rbcL also supported the break-up of the Polygalales,
but not the grouping of Vochysiaceae with Malpighiaceae and
Trigoniaceae (Chase et al., 1993). Instead, Vochysiaceae, repre-
sented by Qualea, fell within the Myrtales. A more detailed study of
Myrtales using the same gene confirmed the Myrtalean affinities of
Vochysiaceae, represented by Vochysia, Erisma, and Qualea (Conti
et al., 1996). Vochysiaceae was sister group to the Myrtaceae and
these, with Heteropyxidaceae and Psiloxylacecae, formed the sister
clade to that containing Melastomataceae, Memecylaceae, and other
families. Work with chloroplast gene matK also supports placement
of Vochysiaceae in Myrtales, but it falls between Myrtaceae (with
Heteropyxidaceae and Psiloxylacecae) and Melastomataceae (with
Memecylaceae) (P. Wilson, pers. comm.).
Previous systematic treatments based on morphology have never
included Vochysiaceae in the Myrtales (Conti et al., 1996), regard-
Table 3 Distribution of simple and branched hairs in Vochysia.
Section
Subsection
Number of species
Described
K,, ^'i:,!! .11 /IG/1Q\
Examined
Type of hair
by Matleu (iy4o)
for study
Simple
Branched
Unknown
Vochysiella
Decorticanthes
10
6
6
0
0
Calophylloideae
10
7
6
0
1
Ciliantha
Micranthae
11
9
8
0
1
Lutescentes
23
13
6
4
3
Discolores
3
2
0
2
0
Chrysophyllae
1
1
1
0
0
Megalanthae
5
3
3
0
0
Ferrugineae
31
22
1
18
3
Pachyantha
-
3
0
—
—
~
Type of hair: simple, only simple unbranched hairs present on leaves, stems, and reproductive parts; branched, some or all of hairs present branched, including T-shaped hairs;
unknown, specimens glabrous or hairs too small or dense to characterize. Leaves, stems, and inflorescences of one to several specimens of each species at BM monographed by
Stafleu (1948) were examined under a 40x dissecting microscope.
14
ing this relationship as so remote that the presence of two of the
most definitive Myrtalean characters, vestured pits in the second-
ary xylem and bicollateral vascular bundles in the primary stem,
was not considered significant (van Vliet & Baas, 1984). In retro-
spect, many vegetative characters of Vochysiaceae are equally at
home in Myrtales (sensu Dahlgren & Thorne, 1984) as in
Polygalales, including opposite or verticillate leaves with entire
margins, brochidodromous venation, and stipules. The highly
zygomorphic flowers with one fertile stamen are, however, unu-
sual in the order (Conti et al., 1996). When a thorough
morphological analysis of the relationship of Vochysiaceae within
Myrtales is undertaken, several seedling and seed characters de-
scribed for Vochysiaceae deserve further study.
Vochysiaceae share other vegetative characters with Myrtales,
including the occasional presence of 2-branched unicellular or
tufted hairs in Combretaceae, Lythraceae, and Myrtaceae
(Metcalfe & Chalk, 1950). Cotyledonary stipules, generally un-
common in dicotyledons, occur as rudimentary glandular
structures in some Lythraceae, Onagraceae, and Myrtaceae, and
as 2- to 3-lobed structures or glandular processes near the basal
margins of the blade in other groups of Myrtaceae (Weberling,
1984). The latter type might be homologous to the unusual
cotyledonary stipules of Vochysia (Flores, 1993«; this study),
which are composed of an upper lobed part and lower possibly
glandular part.
Wings are well-developed on adult stems in some Lythraceae
and Penaeaceae, and often develop auricles at the apex, which are
distinct from the rudimentary stipules found closer to the petioles
(Weberling, 1984). In Lythraceae, auricled wings are already
present at the seedling stage in Lafoensia (Garwood, in prep.).
Wings are well-developed on seedlings of Vochysia, although usu-
ally reduced to ribs on adults (Stafleu, 1948; Garwood, pers.
obs.). As the small stipules of Vochysia occur at the apex of the
wings, they may be homologous to the auricles at the apex of the
wings of these Myrtales, rather than the small, often dissected,
stipules characteristic of Myrtales (Weberling, 1984). If so,
Vochysia may lack true stipules. Herbarium specimens of V.
ferrugineae and V. guatemalensis seedlings were carefully exam-
ined for the presence of dissected or rudimentary stipules over the
petioles, but none were seen (Garwood, pers. obs.). The occa-
sional presence of a minute shrivelled structure, however, suggests
that actively growing seedlings should be examined to verify
whether minute caducous stipules are present but hidden by the
small erect expanding leaves. In seedlings of Myrtaceae, auricles
are lacking on the winged epicotyl and later internodes of Psidium
and Calycolpus and on the winged hypocotyl of Myrcia, and
wings are lacking in Eugenia, but all have rudimentary gland-like
stipules (Garwood, in prep.).
Seeds of Vochysiaceae and Myrtales generally lack endosperm
and arils and have straight or slightly curved embryos (Cronquist,
1981; Dahlgren & Thorne, 1984). A fibrous exotegmen and an
endotestal crystal layer are found in Qualea and some Lythraceae,
Combretaceae, and Onagraceae, but the exotegmen lacks fibres and
is crushed in Vochysia and some Melastomataceae and Myrtaceae
(Corner, 1976; Dahlgren &Thorne, 1984; Boesewinkel &Venturelli,
1987). The embryonic cotyledons are variously folded in
Vochysiaceae and many Myrtales, with folding frequently described
as 'spiral' in Vochysia as well as Combretaceae and Oliniaceae
(Dahlgren & Thorne, 1984). As brief descriptions of vernation and
ptyxis in Vochysiaceae are generally incomplete or incorrect (see
above), further detailed studies should be carried out throughout the
order.
N.C. GARWOOD
Ecology
Seed germination has been the most frequently studied aspect of the
seed and seedling ecology of Vochysiaceae. Germination is usually
rapid, occurring 1 week to 3 months after seed dispersal or sowing in
species of Vochysia, Salvertia, and Qualea from both cerrado and
rainforest (Rizzini, 1965; Thomazini et al., 1973; Macedo, 1977;
Duarte, 1979; Sabatier, 1985; Paviani & Haridasan, 1988; de Godoy
& Felippe, 1992a; Moreira & Arnaez, 1994; Flores 1993a, b; this
study; M. Bustamante, pers. comm.; P.E. Oliveira, pers. comm.; P.
Wilson, pers. comm.). Chemical or mechanical scarification of the
seed coat, or its complete removal, may hasten germination in Q.
grandiflora (Felippe, 1990; Felippe & Dale, 1990), but is not
required (M. Bustamante, pers. comm.; P.E. Oliveira, pers. comm.;
P. Wilson, pers. comm.). Germination is not inhibited by dark or
shaded conditions in Q. retusa Spruce ex Warm., Q. grandiflora, or
V.ferruginea (Macedo, 1977; Felippe, 1990; this study), although it
can be somewhat depressed in direct sunlight (Flores, 19936).
Recently dispersed seeds of V. maxima Ducke buried under the leaf
litter either lost viability or had germinated within 2 months (Viana,
1990). Thus, there is no indication of any type of seed dormancy in
the family, and seeds should form transient, rather than persistent,
seed banks under natural conditions.
Under artificial conditions, seeds of Qualea cordata Spreng.,
Vochysia guatemalensis, and V.ferruginea retained high viability for
at least 2-3 months when stored at room temperature and humidity
(de Godoy & Felippe, 1992a; Flores, 1993a, b), in spite of a
relatively high initial moisture content in Vochysia (23-32%, fresh
weight basis). Reducing the moisture content extended viability of
the apparently orthodox seeds, although they were intolerant of
chilling (Flores, \993a). The potential to store seed should promote
the ongoing silvicultural development of the family.
Seeds of all species are wind-dispersed, as indicated by the
morphology of seeds of Vochysieae and indehiscent fruits of
Erismeae, except Erisma calcaratum (Link) Warm, from the Ama-
zonian varzea, which is water-dispersed (Stafleu, 1954). In tropical
forests, seeds of Vochysia maxima are dispersed near the parent,
mostly < 30 m from the trunk (Viana, 1990), seedlings of V. maxima
and V. ferruginea persist in the shaded understorey (Viana, 1990;
Garwood, pers. obs.), but growth is fast only in treefall gaps and
larger openings (Viana, 1990; Boucher et al., 1994). In plantations,
seedlings of V. guatemalensis and V.ferruginea can reach 2-3 m tall
within three years (Flores 1993a, b). Thus, tall canopy Vochysia
appear to be gap opportunists, regenerating from a persistent seed-
ling bank following disturbances throughout the year. Regeneration
of Qualea may follow a different pattern. Following a severe hurri-
cane, most of the severely defoliated adults of Q. paraensis Ducke
survived but there was no regeneration from seedlings or saplings; in
contrast, all the equally abundant and severely defoliated adults of V.
ferruginea died, but there was extensive regeneration from seedlings
and saplings (Boucher et al., 1994). If this pattern is typical, species
of Vochysia may be more amenable to sustainable management or
plantation development than Qualea.
In the drier and more open cerrado, seedlings of Qualea
grandiflora, Q. cordata, and Vochysia thyrsoidea appear to grow
inherently slowly, with neither extra moisture nor nutrients pro-
moting rapid growth (Rizzini, 1965; de Godoy & Felippe, 19926;
Paulilo et al., 1993; Felippe & Dale, 1990). Although the basis for
the slow growth is not understood, many species develop special
morphological adaptations to the cerrado environment, such as
large subterranean storage organs and thick succulent cotyledons
(Labouriau et al., 1964; Rizzini, 1965; Paviani & Haridasan,
1988).
SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE
15
ACKNOWLEDGEMENTS. I am especially grateful to Margaret Tebbs for
illustrations, Simon Thornton-Wood for cross-sectional diagrams of seeds,
Ruth Dempsey for composition of the figures, and Eduardo Sierra for
technical assistance in Panama. I also thank C.J. Humphries for encourage-
ment and advice; S. Knapp, D.M. Williams and an anonymous reviewer for
helpful comments on the manuscript; M. Beasley for bibliographic assist-
ance; INRENARE for permission to work in Panama; M.L. Kawasaki, M.
Bustamante, P.E. Oliveira, and P.O. Wilson for sharing work in press,
unpublished data, and observations; and the many people on BCI and at STRI
who helped the Project in various ways but are too numerous to thank here
individually. The Seedling Flora Project has been generously supported by
the National Science Foundation (BSR-85-17395), the Natural Environment
Research Council (GR3-695 1 ), the Smithsonian Tropical Research Institute,
The Natural History Museum, and the Linnean Society of London.
REFERENCES
del Amo, S. 1 979. Clave para plantulas y estados juveniles de especies primaries de una
selva alta perennifolia en Veracruz, Mexico. Biotica 4: 59-108.
Anderson, W.R. 1990. The origin of the Malpighiaceae-the evidence from morphol-
ogy. Memoirs of the New York Botanical Garden 64: 210-224.
Kit illi in, H.E. 1874. Vochysiaceae. Histoire des plantes 5: 93-104.
Bell, A. 1991. An illustrated guide to flowering plant morphology. Oxford.
Bentham, G. & Hooker, J.D. 1867. Vochysiaceae. Genera plantarum 1(3): 975-978.
Blanche, C.A., Dodges, J.D., Gomes, A.E. & Gonzalez, E. 1991. Seed chemistry of
the tropical tree Vochysia hondurensis Sprague. Forest Science 37: 949-952.
Boesewinkel, F.D. & Bouman, F. 1984. The seed: structure. In B.M. John (Ed.),
Embryology of angiosperms'. 567-610. Berlin.
& Venturelli, M. 1987. Ovule and seed structure in Vochysiaceae. Botanische
Jahrbiicher fitr Systematik, Pflanzengeschichte und Pflazengeographie 108: 547-
566.
Boucher, D.H., Vandermeer, J., Mallona, M.A., Zamora, N. & Perfecto, I. 1994.
Resistance and resilience in a directly regenerating rainforest: Nicaraguan trees of
the Vochysiaceae after Hurricane Joan. Forest Ecology and Management 68: 1 27-
136.
Butterfield, R.P. & Fisher, R.F. 1994. Untapped potential: native species for reforesta-
tion. Journal of Forestry 92: 37^3.
Chase, M.W., Soltis, D.E., Olmstead, R.G., Morgan, D., Les, D.H., Mishler, B.D.,
Duvall, M.R., Price, R.A., Hills, H.G., Qiu, Y.-L., Kron, K.A., Rettig, J.H., Conti,
E., Palmer, J.D., Manhart, J.R., Sytsma, K.J., Michaels, H.J., Kress, W.J., Karol,
K.G., Clark, W.D., Hedren, M., Gaut, B.S., Jansen, R.K., Kim, Ki-J., Wimpee,
C.F., Smith, J.F., Furnier, G.R., Strauss, S.H., Xiang, Q.-Y., Plunkett, G.M.,
Soltis, P.S., Swensen, S.M., Williams, S.E., Gadek, P.A., Quinn, C.J., Eguiarte,
L.E., Goldenberg, E., Learn, G.H., Jr., Graham, S.W., Barrett, S.C.H.,
Dayanandan, S. & Albert, V.A. 1993. Phylogenetics of seed plants: an analysis of
nucleotide sequences from the plastid gene rbcL. Annals of the Missouri Botanical
Garden 80: 528-580.
Condit, R., Hubbell, S.P. & Foster, R.B. 1993. Identifying fast-growing native trees
from the Neotropics using data from a large, permanent census plot. Forest Ecology
and Management 62: 123-143.
Conti, E., Litt, A. & Sytsma, K.J. 1996. Circumscription of Myrtales and their
relationships to other rosids: evidence from rbcL sequence data. American Journal
of Botany 83: 221-233.
Corner, E.J.H. 1976. The seeds of dicotyledons. Cambridge.
Croat, T.B. 1978. Flora of Barm Colorado Island. Stanford.
Cronquist, A. 1981. An integrated system of classification of flowering plants. New
York.
Dahlgren, R.M.T. 1980. A revised system of classification of angiosperms. Botanical
Journal of the Linnean Society 80: 91-124.
& Thorne, R.F. 1984. The order Myrtales: circumscription, variation, and
relationships. Annals of the Missouri Botanical Garden 71: 633-699.
D'Arcy, W.G. 1987. Flora of Panama. Checklist and index. Parts I and II. St Louis.
(Monographs in systematic botany from the Missouri Botanical Garden 17-18)
Duarte, A.P. 1979. Contribui?ao ao conhecimento da germina?ao das sementes nas
essencias mais usuais. Rodriguesia 31: 59-68.
Duke, J.A. 1969. On tropical tree seedlings. I. Seeds, seedlings, systems, and system-
atics. Annals of the Missouri Botanical Garden 56: 125-161.
Felippe, G.M. 1990. Qualea grandiflora: the seed and its germination. Revista
Brasileira de Botanica 13: 33-37.
& Dale, J.E. 1990. The effects of phosphate supply on growth of plants from the
Brasilian cerrado: experiments with seedlings of the annual weed, Bidens gardneri
Baker (Compositae) and the tree, Qualea grandiflora (Mart.) (Vochysiaceae).
Oecologia 82: 81-86.
Finegan, B. 1992. The management potential of neotropical secondary lowland rain
forest. Forest Ecology and Management 47: 295-32 1 .
Flores, E.M. I993a. Vochysia guatemalensis, chanco bianco- white yemeri. Arboles y
semillas del Neotropico - Trees and seeds from the Neotropics 2: 1-27.
— 1993/7. Vochysia ferruginea, mayo colorado-red yemeri. Arboles y semillas del
Neotropico - Trees and seeds from the Neotropics 2: 29-52.
Gaos, G. 1978. Vochysiaceae. In A. G6mez-Pompa (Ed.), Flora of Veracruz 4: 1-5.
Garwood, N.C. 1994. Morphology and ecology of seedlings, fruit and seeds of
Panama: Bixaceae and Cochlospermaceae. Bulletin of the Natural History Museum,
London (Botany) 24: 161-171.
1995. Studies in Annonaceae. XX. Morphology and ecology of seedlings, fruits
and seeds of selected Panamanian species. Botanische Jahrbiicher fur Systematik,
Pflanzengeschichte und Pflazengeographie 117: 1-152.
in prep. Illustrated guide to seedlings of Barm Colorado Island and adjacent
Panama.
& Humphries, C.J. 1993. Seedling diversity in the Neotropics. NERCNews (No.
27, October): 20-23.
de Godoy, S.M.A. & Felippe, G.M.I 992a. Qualea cordata: a semente e sua germinac, ao.
Revista Brasileira de Botanica 15: 17-21.
1992/7. Crescimento inicial de Qualea cordata, uma arvore dos cerrados. Revista
Brasileira de Botanica 15: 23-30.
Kawasaki, M.L. in press. Systematics of Erisma (Vochysiaceae). Memoirs of the New
York Botanical Garden.
Keay, R.W.J. & Stafleu, F.A. 1953. Erismadelphus. Ada Botanica Neerlandica 1:
594-599.
Labouriau, L.G., Marques Valio, I.F. & Heringer, E.P. 1964. Sobre o sistema
reproductive de plantas dos Cerrados-I. Anais da Academia Brasileira de Ciencias
36: 449-464.
Litt, A. J., Conti, E., Chase, M.W. & Sytsma, KJ. 1995. The systematic position of
Euphmnia (Polygalales) based on molecular and morphological evidence. American
Journal of Botany 82(6, Suppl.): 146. (Abstract 423)
Lleras, E. 1976. Revision and taxonomic position of the genus Euphmnia Martius ex
Martius & Zuccarini. Acta Amazonica 6: 43-47.
Macedo, M. 1977. Dispersao de plantas lenhosas de uma Campina Amazonica. Acta
Amazonica 7(Suppl.): 1-69.
Marcano-Berti, L. 1969. Un nuevo genero de las Vochysiaceae. Pittieria 2: 3-29.
— 1989. Euphroniaceae: una nueva familia. Pittieria 18: 15-17.
— 1990. Descripcion y tipificacion de la nueva familia Euphroniaceae. Emstia 57: 5-
7.
Metcalfe, C.R. & Chalk, L. 1950. Anatomy of the dicotyledons. 1. Oxford.
Miquel, S. 1987. Morphologic fonctionnelle de plantules d'especes forestieres du
Gabon. Bulletin du Museum National d' Histoire Naturelle, 4e serie, Section B,
Adansonia9: 101-121.
Moreira G., I. & Arnaez S., E. 1994. Morfologfa de las estructuras reproductoras y
germinacion de nueve especies forestales nativas de Costa Rica. Revista de Biologia
Tropical (San Jose) 42(Suppl. 2): 73-82.
Niembro Rocas, A. 1989. Semillas de plantas lenosas: morfologia comparada.
Mexico, D.F.
Oldeman, R.A.A. & Fundter, J.M. 1986. On diptero-vochysian tree strategies,
Compte-rendu du Colloque International 1'Arbre, Montpellier 9-14 septembre
1985. Naturalia monspeliensia numero hors serie: 191-208.
Paulilo, M.T.S., Felippe, G.M. & Dale, J.E. 1993. Crescimento inicial de Qualea
grandiflora. Revista Brasileira de Botanica 16: 37— 46.
Paviani, T.I. & Haridasan, M. 1988. Tuberosdidade em Vochysia thyrsoidea Pohl
(Vochysiaceae). Ciencia e Cultura, S. Paulo 40: 998-1003.
Petersen, O.G. 1 896. Vochysiaceae. In A. Engler & K. Prantl (Eds), Die naturlichen
Pflanzenfamilien 3(4): 312-319.
Polak, A.M. 1992. Major timber trees of Guyana: afield guide. Wageningen.
Radford, A.E., Dickison, W.C., Massey, J.I.R. & Bell, C.R. 1974. Vascular plant
systematics. New York.
Ricardi, M., Hernandez, C. & Torres, F. 1 987. Morfologia de plantulas de drboles de
los basques del estado Merida. Merida.
Rizzini, C.T. 1965. Experimental studies on seedling development of cerrado woody
plants. Annals of the Missouri Botanical Garden 52: 410-426.
Robyns, A. 1967a. Vochysiaceae. In R.E. Woodson Jr. & R.W. Schery (Eds), Flora of
Panama. Annals of the Missouri Botanical Garden 54: 1-7.
1 967 b. Vochysia jefensis A. Robyns, sp. nov. (Vochysiaceae). Annals of the
Missouri Botanical Garden 54: 188-189.
van Roosmalen, M.G.M. 1985. Fruits of the Guianan flora. Utrecht.
Sabatier, D. 1985. Saisonnalite et determinisme du pic de fructification en foret
guyanaise. Revue d'Ecologie (La Terre et La Vie) 40: 289-320.
Seithe, A. 1979. Hair types as taxonomic characters in Solanum. In J.G. Hawkes, R.N.
Lester & A.D. Skelding (Eds), The biology and taxonomy of the Solanaceae: 307-
319. London.
1980. Rhododendron hairs and taxonomy. In J.L. Luteyn & M.E. O'Brien (Eds),
Contributions towards a classification of Rhododendron: 89-115. New York.
& Anderson, GJ. 1982. Hair morphology and the relationships of species in
Solanum sect. Basarthrum. Plant Svstematics and Evolution 139: 229-256.
16
N.C. GARWOOD
Shanmukha Rao, S.R. & Vijayalakshmi Sarma. 1992. Morphology of 2-armed
trichomes in relation to taxonomy: Malpighiales. Feddes Repertorium 103: 559-
565.
Stafleu, F.A. 1948. A monograph of theVochysiaceae. Recueildes trauvaux botaniques
neerlandais 41: 397-540.
1952. A monograph of the Vochysiaceae. II. Callisthene. Acta Botanica
Neerlandica 1: 222-242.
1 953. A monograph of the Vochysiaceae. III. Qualea. Acta Botanica Neerlandica
2: 144-217.
1954. A monograph of the Vochysiaceae. IV. Erisma. Acta Botanica Neerlandica
3: 459-^80.
M.I IK lk->. P.C. & Steyermark, J.A. 1949. Vochysiaceae, Flora of Guatemala: Part VI.
Fieldiana: Botany 24: 2-5.
Takhtajan, A.L. 1980. Outline of the classification of flowering plants (Magnoliophyta).
Botanical Review 46: 225-359.
Thomazini, L.I., Massa, C.S. & Piccolo, A.L.G. 1973. Qualea grandiflora Mart.,
considerac,5es sobre a semente, a germinagao a plantula. Revista de Agricultura,
Piraciaba Brazil 48: 101-106.
Viana, V.M. 1990. Seed and seedling availability as a basis for management of natural
forest regeneration. In A.B. Anderson (Ed.). Alternatives to deforestation: steps
toward sustainable use of the Amazon rainforest: 99-1 15. New York.
van Vliet, G.J.C.M. & Baas, P. 1984. Woody anatomy and classification of the
Myrtales. Annals of the Missouri Botanical Garden 71: 783-800.
de Vogel, E.F. 1 980. Seedlings of dicotyledons: structure, development, types; descrip-
tions of 150 woody Malesian taxa. Wageningen.
Warming, E. 1875. Vochysiaceae. In K.F.P. von Martius & A.W. Eichler (Eds), Flora
brasiliensis 13(2): 17-116.
Weberling, F. 1984. Stipular structures. In R. Dahlgren & R.F. Thorne (Eds), The order
Myrtales: circumscription, variation, and relations. Annals of the Missouri Botanical
Garden 71: 641-644.
Bull. not. Hist. Mus. Land. (Bot.) 28(1): 17^0
Issued 25 June 1998
A revision of the genus Mandragora
(Solanaceae)
STEFAN UNGRICHT* SANDRA KNAPP AND JOHN R. PRESS
Department of Botany, Tne~Natural History Museum, Cromwell Road, London SW7 5BD
* Present address: Waldmatt 6, CH-5242 Birr, Switzerland
CONTENTS
Introduction
Mythological and medicinal history
Taxonomic history
Materials and methods
Material examined
Taxonomic concepts
Morphometrics
Cladistics
Results and discussion
Species delimitations using morphometric analyses
Phylogeny
Biogeography
Taxonomic treatment
Mandragora L
Key to the species of Mandragora
1. Mandragora officinarum L
2. Mandragora turcomanica Mizg
3. Mandragora caulescens C.B. Clarke
References
Exsiccatae
Taxonomic index ...
17
18
18
19
19
20
21
22
22
22
26
26
29
29
30
30
33
34
36
38
40
SYNOPSIS. The Old World genus Mandragora L. (Solanaceae) is revised for the first time across its entire geographical range.
The introduction reviews the extensive mythological and medicinal as well as the taxonomic history of the genus. On
morphological and phenological grounds three geographically widely disjunct species can be distinguished: the Mediterranean
M. officinarum L., the narrowly local Turkmenian endemic M. turcomanica Mizg. and the Sino-Himalayan M caulescens C.B.
Clarke. The generic monophyly of Mandragora L. as traditionally circumscribed is supported by cladistic analysis of
morphological data. The ecological and historical phytogeography of the genus is discussed and alternative biogeographical
scenarios are evaluated. Finally, a concise taxonomic treatment of the taxa is provided, based on the evidence of the preceeding
analyses.
INTRODUCTION
The nightshade family (Solanaceae) is a cosmopolitan but predomi-
nantly tropical group and is especially well represented in the
Neotropics. The family is of considerable economic importance.
Food plants include potato (Solanum tuberosum L.), tomato (Sola-
num lycopersicum L. (=Lycopersicon esculentum Mill.)), pepper
(Capsicum annuum L.) and aubergine (Solanum melongena L.j.
Another species of great commercial value is tobacco (Nicotiana
tabacum LJ. The family also contains ornamental plants such as
garden petunias (species of Petunia Juss.) and the Chinese lantern
(Phy sails alkekengi L.j. Many members of the family are poisonous,
but several such as deadly nightshade (Atropa belladonna L.),
henbane (Hyoscyamus niger L.) and thorn-apple (Datura stramo-
nium L.) provide drugs traditionally used in medicine. The subjects
of this study, the mandrakes (Mandragora L.), belong to this latter
group of pharmaceutically interesting Solanaceae.
The long history of mythology and medicinal use of the mandrake
combined with the variable morphology and phenology have led to
considerable confusion in the classification of Mandragora. The
nomenclatural situation is, considering the small size of the genus,
amazingly chaotic. In particular the Mediterranean Mandragora is
burdened with synonyms, and some authors (Mill, in litt.) have
expressed doubts about the placement of the Himalayan mandrake
in Mandragora at all. This study represents the first taxonomic
revision of the genus over its entire range. We address the following
main questions in this study. Firstly, which of the described Man-
dragora taxa are justifiable if the whole range of the morphological
variation is considered? Secondly, is the genus as currently delim-
ited (including the Himalayan mandrake, M. caulescens)
monophyletic? Finally, does the phylogeny, combined with the
distribution patterns of the taxa, support a particular historical
biogeographical scenario and do the ecological requirements of the
defined taxa impinge upon this?
© The Natural History Museum, 1998
18
Mythological and medicinal history
The origin of the myth surrounding the mandrake root is hard to
trace. It is clear, however, that the mandrake was a focus of supersti-
tion and belief in magic and demons for many centuries. The
sometimes vaguely humanoid shape of the root and its potent,
psychotropically active alkaloids are probably responsible for the
long history of myths. However, the properties ascribed to the plant
as a whole, and especially to its root, are highly ambiguous and the
first references in Ancient Egypt and in the Bible remain controver-
sial. Ornaments on the casket and throne from the tomb of
Tutankhamun seem to depict mandrakes (Hepper, 1990; Roland,
199 1 ). Mandragora is not known to be native in Egypt but may have
been cultivated there. The Biblical 'Dudaim' is often said to be
identical with Mandragora (Moldenke & Moldenke, 1952; Fein-
brun-Dothan, 1978; Fleisher & Fleisher, 1994). Dudaim is mentioned
in the Song of Solomon (7: 13) and in another passage from the Old
Testament (Genesis 30: 14-18), according to which it was used in
relation to fertility. The 'Moly ' of Homer in his Odyssey is presumed
to be the mandrake, as is the 'Baraa' of Flavius Josephus' Jewish
Wars, where it is described as shining in the dark (sic) and is
attributed exorcistic powers. Pliny in his Natural History and
Dioscorides in De Materia Medica wrote that the mandrake was
known as 'Circaeon' after Circe, the mythical sorceress who turned
men into sexually supercharged swine (Marzell, 1975; Roland 1991;
Fleisher & Fleisher, 1 994). The Ancient Greeks and Romans consid-
ered the mandrake so potent and valuable as a narcotic and restorative
plant that the collectors of its roots had to obey rigid ceremonial
rules which were described by Theophrastus and Pliny (Randolph,
1905).
In Medieval Europe the plant was known as 'Alraune' in Germany
and 'Main de gloire' in France. Carved to emphasize the anthropo-
morphic male or female character of the root, it was popular as
aphrodisiac, charm against sterility and love talisman alike. But the
plant was soon connected with witchcraft; for example, Joan of Arc,
who was tortured and burnt at the stake in 143 1 as a heretic, was also
accused of the witches' crime of possessing a mandrake (Thompson,
1934). It was believed that the plant grew under gallows and that the
root owed its human shape to urine or sperm of unjustly hanged men.
The gathering of the root was considered to be highly dangerous and
dogs had to perform the supposedly lethal procedure of pulling out
the plants, whereupon the roots would shriek so 'that living mortals
hearing them run mad': Shakespeare, Romeo and Juliet Act IV
(Randolph, 1905; Moldenke & Moldenke, 1952; Marzell, 1975;
Roland, 1991; Talalaj et al, 1992; Fleisher & Fleisher, 1994).
Perhaps the most interesting aspect of the mythology of the man-
drake is convergent myths in the eastern area of the genus in the
disjunct region of Central Asia, the Himalayas and China where
Mandragora has a similarly long history of application and tradi-
tional use. For instance in Sikkim, the roots of Mandragora
caulescens, called 'Lakshmana', were used in magical rites and in
folk medicine (Mehra, 1979); and in Central Asia the mandrake,
called 'Khaoma-soma', was considered a holy plant (Khlopin,
1979).
The mythology of the mandrake and its use in traditional medi-
cine are hard to keep apart. In a wide range of applications it was
used as an aphrodisiac, hypnotic, emetic and purgative, sedative and
narcotic, for example as a pain-killer at childbirth as well as to
relieve crucifixion agony (Duke, 1985; Talalaj et al., 1992); in fact,
according to Heiser (1969), it has been questioned that Christ died
on the Cross at all, suggesting that the vinegar supplied to him had
contained the drug, inducing merely a deathlike trance. The second-
ary metabolites of major pharmaceutical importance are chiefly
S. UNGRICHT, S. KNAPP AND J.R. PRESS
tropane alkaloids and their esters. Staub (1962), Hegnauer (1973),
Romeike (1978), Evans (1979) and Jackson & Berry (1979) list
alkaloids of Mandragora and other Solanaceae. Tropane alkaloids
occur mainly in the Solanaceae, but they are not entirely restricted to
this family; other tropane alkaloids of importance include cocaine,
found in the family Erythroxylaceae. However, esters with tropic or
related acids of alkamines derived from tropane do seem to be
restricted to the Solanaceae (Romeike, 1978).
The most important alkaloid components of Mandragora are:
atropine, apoatropine, belladonnine, cuscohygrine, hyoscyamine,
hyoscyamine-N-oxide, norhyoscyamine, scopolamine, 3oc-tigloyl-
oxytropane, 3oc,6p-ditigloyloxytropane, and 6p-hydroxyhyoscy-
amine. The root, stem, leaves, fruits and seeds of Mandragora all
contain a different alkaloid profile. However, the root is considered
to be the site of synthesis of the tropane alkaloids (Romeike, 1978)
and is the most poisonous part of Mandragora, due to the high
concentration therein. The total content of alkaloids in the root is
around 0.4% (Kessler, 1951; Maugini, 1959; Duke, 1985), hyo-
scyamine being the most important constituent (Duke, 1985). In
contrast, the berries are said to be sweet and edible in moderation
(Viney, 1994). The synthetic pathways leading to the alkaloids in
different taxa of the Solanaceae have been proposed to be homolo-
gous to each other (Tetenyi, 1987) and to be a linear or spiralling
phyletic sequence. Tropane alkaloids are effective as analgesics,
anaesthetics, antispasmodics, and are used to increase the circula-
tion, reduce secretions and dilate pupils. Scopolamine is a hypnotic
(Duke, 1985) and achieved a notorious reputation as a 'truth-drug' in
police and secret service investigations (Heiser, 1969). 'Quishen', a
Chinese herbal medicine consisting of dried roots of M. caulescens,
is used in Yunnan and Xizang as a ginseng substitute for all kinds of
physical weaknesses (Xiao & He, 1983). In allopathic western
medicine, however, the mandrake has largely lost its importance as
valued source of alkaloids, although the plant itself is still sur-
rounded by myth.
Taxonomic history
The classification of Mandragora
MEDITERRANEAN PLANTS. As mentioned above, considering the
small size of the genus, there has been a fair amount of confusion in
respect to the classification of Mandragora. The long taxonomic
history, especially of the Mediterranean Mandragora species, led to
a large number of often imprecisely defined species and subspecies
or varieties. As a consequence, there is a plethora of names, and their
somewhat indiscriminate use renders the meaning and reliability of
identifications of specimens in herbaria, as well as in the literature,
highly problematic. Here we present the taxonomic history of the
Mediterranean part of the genus in periods, with examples of major
taxonomic works and their treatment of the genus, in order to outline
the main taxonomic trends through the centuries.
Pre-Linnaean: The Ancients distinguished two species that they
called the female or black mandrake (Mandragorasfcemina), and the
male or white mandrake (Mandragoras mas). These plants were
described by both Dioscorides and Pliny (cf. Randolph, 1905 for
translations), and this division was widely adopted, for instance by
Parkinson (1629) and by Jean Bauhin et al. (1651) as late as the
seventeenth century. The male mandrake corresponds to a spring
flowering entity; the female mandrake represents an autumn flower-
ing one. Caspar Bauhin (1623) and Tournefort (1719), on the other
hand, each recognized three 'species'. Both authors listed Man-
dragora fructu rotundo and M. /lore subcaeruleo, purpurascente.
MANDRAGORA
19
Bauhin's third species was M.fructu pyri while Tournefort's was M.
flore subcaeruleo, foliis minoribus, fructu globoso).
Linnaeus: In hisHortus cliff ortianus, Linnaeus ( 1 738) considered
these variants as mere forms of a single species. In the first edition of
Species plantarum (1753), he accordingly recognized only one
species that he named Mandragora officinarum. Later, however, he
changed his mind with respect to the distinctness of this taxon and
placed it within his genus Atropa, calling it Atropa mandragora
(Linnaeus, 1759, 1762).
Post-Linnaean: Linnaeus's idea of a single species did not last
very long. The nineteenth and the first half of the twentieth centuries
were characterized by an increasingly elaborate division of the
genus (Table 1 ). Heldreich ( 1 886) additionally described an infertile
hybrid between two vernal entities: Mandragora vernalis Bertol.
and M. haussknechtii Heldr., called M. x hybrida Haussk. & Heldr.
Post-1950: In recent decades, however, there has been a trend
towards reversing this ever greater subdivision of the genus. In fact,
since the 1950s there have been a number of proposals to re-adopt
the two traditional, pre-Linnaean taxa only (Tercinet, 1950; Hawkes,
19726; Jackson & Berry, 1979). This viewpoint was adopted in
Flora Europaea (Hawkes, 1972a), using the names Mandragora
officinarum L. for a vernal entity and M. autumnalis Bertol. for an
autumnal one. However, M. officinarum L. might be considered an
ambiguous name since it was formerly used for an autumnal entity
as well (Greuter & Rechinger, 1967; see Table 1). Hawkes (19726)
rejected this reservation on the grounds of the priority, the wide use
and the generic type status of the Linnaean epithet officinarum.
Knapp in Jarvis et al. (1993) finally lectotypified M. officinarum L.
with a specimen in the Bursar herbarium, to which no phenological
data are attached.
SlNO-HiMALAYAN PLANTS. In contrast to the Mediterranean plants,
the classification of the disjunct Central and East Asian part of the
genus was never as contentious and did not arouse much comment,
in part reflecting its far younger taxonomic history. Mandragora
caulescens was described by C.B. Clarke (1883) in Hooker's The
flora of British India (Hooker fil. and Thomson labelled the speci-
mens 'Scopolia humilis ' but this name was never published). Anisodus
caulescens (C.B. Clarke) Diels and A. mariae Pascher, as well as
Mairella yunnanensis H. Lev., were placed in synonymy with
Mandragora caulescens in the Flora of China (Zhang et al., 1994).
Grubov (1970) described a new species from Xizang, China which
he named Mandragora tibetica, and Kuang Ko-zen & Lu An-ming
(1978) described a new species found in Qinghai and Xizang which
they named Mandragora chinghaiensis. In the Flora of China
(Zhang et al., 1994) the latter two taxa were provisionally put into
synonymy with M. caulescens until a more detailed revision could
Table 1 Post-Linnaean classifications of the increasingly subdivided
Mediterranean taxa of Mandragora.
Classification type Vernal species
Autumnal species
1 VERNAL, 1 AUTUMNAL TAXON
Sprengel (1825) M. vernalis Bertol.
1 VERNAL, 2 AUTUMNAL TAXA
Bertoloni ( 1 835), M. vernalis Bertol.
Dunal (1852)
2 VERNAL, 2 AUTUMNAL TAXA
Heldreich ( 1 886) M. vernalis Bertol.
M. haussknechtii Heldr.
3 VERNAL, 2 AUTUMNAL TAXA
Vierhapper (1915) M. mas Garsault
M. hispanica Vierh.
M. haussknechtii Heldr.
M. autumnalis Bertol.
M. officinarum L.
M. microcarpa Bertol.
M. autumnalis Spreng.
M. microcarpa Bertol.
M. autumnalis Bertol.
M. foemina Garsault
establish their distinctiveness. Mandragora shebbeard C.E.C.
Fischer, described from a single specimen from Xizang (Fischer,
1934), was transferred to the monotypic solanaceous genus
Przewalskia by Grubov (1970). One attempt to subdivide Man-
dragora caulescens was made by Grierson & Long (1978) who split
it into four subspecies (subsp. caulescens, subsp. flavida Grierson
& Long, subsp. purpurascens Grierson & Long and subsp. brevi-
calyx Grierson & Long) differentiated on the basis of corolla colour
and various morphometric characters, especially of flower parts.
TURKMENIAN PLANTS. Finally, Mandragora turcomanica Mizg.
was described in 1942 from a small area (the Shevlan 'pocket',
Mizgireva, 1955) near Kara Kala in the southwestern Kopet Dag in
Turkmenistan. Only a few collections of this restricted endemic
exist, and these are all to be found in herbaria of the former Soviet
Union. Its status as a distinct taxon has never really been questioned,
but only because so few specimens exist for comparison. Mizgireva
( 1 955) worked for 1 2 years on the detailed anatomy and morphology
of this species and her revision remains the best source for data on
characters and their variation.
The position of Mandragora within the Solanaceae
More than 200 years after the original outline by Jussieu (1789), the
higher level classification of Solanaceae is far from being settled,
and the placement of Mandragora in the various proposed classifi-
cations is just as controversial. In the following review, the most
widely followed classifications of the Solanaceae are presented
together with the positions of Mandragora and the other genera used
in our cladistic analysis: Anisodus Link, Atropa L., Lycium L. and
Nicotiana L.
In his artificial sexual system, Linnaeus (1753) placed Man-
dragora within his group Pentandria Monogyna, together with
Atropa, Lycium and Nicotiana. Jussieu (1789) defined the 'order'
Solaneae with Mandragora in group II (fructus baccatus) alongside
Atropa and Lycium. Nicotiana on the other hand was placed in group
I (fructus capsularis). In the nineteenth century, classifications be-
came more elaborate and tribes and subtribes were established in
order to approximate a more 'natural' classification. In Table 2, the
three most widely adopted classifications of the Solanaceae in this
period (Dunal, 1852;Bentham, 1 876; Wettstein, 1895) are presented
together with the more recent one of Baehni (1946).
The last few decades have seen the analysis of new types of data,
such as DNA sequences (Olmstead & Sweere, 1994) and secondary
plant metabolites (Tetenyi, 1987), as well as application of cladistic
methodology to morphological data (Hoare & Knapp, 1997). This
has led to new hypotheses concerning relationships within the
Solanaceae, as well as the placement of Mandragora. However,
opinion still differs widely on whether Mandragora should be
placed within the tribe Hyoscyameae (Hoare & Knapp, 1997; =
Atropeae sensu Tetenyi, 1987) or in an isolated, basal monogeneric
tribe Mandragoreae (Olmstead & Sweere, 1994; Hunziker, 1995;
Olmstead et al., in press).
MATERIALS AND METHODS
Material examined
This revision is based exclusively on herbarium specimens. The
material examined includes the Mandragora specimens in the her-
baria of The Natural History Museum of London (BM) and Kew (K)
together with specimens borrowed from the following herbaria:
20 S. UNGRICHT, S. KNAPP AND J.R. PRESS
Table 2 Tribal classification of the Solanaceae and the position of Mandragora and the outgroup genera Anisodus, Atropa, Lycium and Nicotiana.
Author
Classification
Genera of this study
Dunal(1852)
Nolaneae
Nolanineae
Grabowskieae
Solaneae
Triguereae
Solaneae Solanineae
Atropineae
Mandragora
Atropa
Lycineae
Lycium
Datureae
Hyoscyameae
Anisodus (Scopolia)
Nicotianeae
Nicotiana
Retzieae
Fabianeae
Metternichieae
Cestreae
Bentham(1876)
Solaneae
Atropeae
Mandragora
Atropa
Lycium
Hyoscyameae
Anisodus (Scopolia)
Cestrineae
Nicotiana
Salpiglossidae
Wettstein(1895)
Nicandreae
Solaneae
Lyciinae
Atropa
Lycium
Hyoscyaminae
Anisodus (Scopolia)
Solaninae
Mandragorinae
Mandragora
Daturae
Cestreae
Cestrinae
Goetzeinae
Nicotianinae
Nicotiana
Salpiglossideae
Baehni(1946)
Solaneae
Solaninae
Sarachinae
Margaranthinae
Physalidinae
lochrominae
Goetzeinae
Discopodiinae
Atropeae
Atropinae
Mandragora
Atropa
Lycium
Markeinae
Hyoscyaminae
Anisodus (Scopolia)
Parabouchetiinae
Anthocercideae
Nicotianeae
Nicotianinae
Nicotiana
Daturinae
Nicandrinae
Salpiglossideae
Berkeley (UC, JEPS), Edinburgh (E), Kunming (KUN), Palermo
(PAL) and Reading (RNG). The collections of the Museum National
d'Histoire Naturelle in Paris (P) were examined on site during a
research visit. Regrettably it was not possible to obtain any speci-
mens, including the types, of Mandragora turcomanica and M.
tibetica from St. Petersburg (LE) or M. chinghaiensis from Xining
(HNWP).
Material with the same collection number or specimens collected
at the same locality and at the same date were unified and treated as
one sample. Unlocalized material or specimens with illegible,
doutbful or very imprecise collection locality and collector data
were excluded, cultivated material being the exception. Only collec-
tions with precise locality data are listed in the taxonomic account of
the species. A complete listing of the specimens examined can be
found in the Exsiccatae.
Taxonomic concepts
The adopted taxonomic methodology in this study of herbarium
material is necessarily largely pattern-oriented. If process-based
terms like 'population' or 'hybrid' are used, these properties are only
informally applied, i.e. inferred rather than rigidly defined, let alone
experimentally circumscribed.
The adopted species concept is operational, i.e. the ability to
distinguish between sets of specimens serves as the criterion for the
delimitation of the terminal taxa (species). Cladistic analysis results
in an estimate of these relationship between the species and the
delimitation of higher level taxa, i.e. monophyletic groups and their
hierarchical relationships. The requirement for a species description
is thus at least one distinguishing qualitative, or discontinuous
quantitative, diagnostic character state (monothetic species) or a
unique set of character states (polythetic species) (cf. Nelson &
MANDRAGORA
Platnick, 1 981; Nixon & Wheeler, 1990). Morphological and pheno-
logical variation plus ecological and geographical information are
the data that were collected from herbarium specimens and their
labels respectively.
Morphometrics
Specimen sampling
The inherent weakness of the traditional herbarium record of a taxon
for taxonomic studies is well known (Du Rietz, 1930; Anderson &
Turrill, 1935; Anderson, 1941). For species that are well represented
in herbaria, one can obtain a more or less accurate record of the
general range of variation. Seldom or never, however, is it possible
to calculate frequencies of the different variants and the resolution of
the classification is limited by the lack of a random sample.
Plant variation also occurs on different hierarchical levels (cf.
Bateman, 1989), which can become confused and overlap. In Man-
dragora the confounding of within-group and between-group
variation is most pronounced in the Sino-Himalayan complex, due
to the secondary growth of the plants during the late flowering and
the fruiting period (ontogenetic variation, see Fig. 9, also Hoare &
Knapp, 1997: fig. 5d). Combining all samples to a pooled matrix
easily leads to a confusion of the two sources of variation which can
seriously compromise interpretation of many applications of
morphometric analyses (Gibson et al, 1984). Avoiding within-
group variation, for example by calculating the mean or only
considering the largest or smallest example of a particular character
is not a solution, but merely reduces the total amount of variation
encountered in a taxon, leading ultimately to a simplistic taxon
delimitation. In the taxonomic process, characters and their variabil-
ity necessarily have priority over all other considerations since taxa
are attributed to characters — not the other way around (characters all
attributed to individuals). It follows that the basal unit of investiga-
tion, the specimen, must be evaluated in respect to its variability.
Therefore, maximal as well as minimal values of the flowering part
measurements of each specimen were assessed, rather than solely
mean or maximal values.
Character sampling
A genus with a long taxonomic history is unlikely to yield any 'new'
gross morphological characters which distinguish species. This is
especially true when working with inadequate herbarium material
(i.e. incomplete with much missing data and/or very old and often in
poor condition). The emphasis in this study was thus much more on
evaluating the traditional and obvious characters in respect to their
ability consistently to delimit proposed taxa. The analysis was
therefore carried out using the following data.
GEOGRAPHY. The entire distributional range of Mandragora was
divided into 20 geographic areas. These are (1) Cultivated speci-
mens, typically from botanic gardens north of the Alps, (2) North
Africa, (3) Iberia, (4) northern Italy and Dalmatia, (5) peninsular
Italy with Sardinia and Sicily, (6) mainland Greece, (7) Aegean
islands, (8) Asia Minor, (9) Cyprus, (10) Levant, (11) Kopet Dag,
(12) Nepal, (13) Sikkim, (14) Bhutan, (15) Arunchal Pradesch
(Assam), (16) north Myanmar (Burma), (17) Yunnan, (18) Sichuan,
(19) Qinghai and (20) Xizang.
PHENOLOGY. Flowering and fruiting dates were recorded, result-
ing in the cumulative reproductive period.
MORPHOLOGY. Corolla length, calyx length, fruit size, fruiting
calyx length, leaf length, leaf length/width ratio. The corolla colour
21
of herbarium material is not reliably determinable and was thus only
recorded if it was stated on the labels. The shape of the berry
(globose or ovoid) was excluded as a character since it is not
accessible through pressed herbarium material and is only rarely
stated on the labels. Berry shape in the Solanaceae is quite variable
and this variation is occasionally a developmental sequence (Bohs,
1994).
Data analysis
Univariate and multivariate analyses were applied as a descriptive,
explorative tool to study the joint relationships of the morphological,
phenological and geographical variables of the specimens at hand.
As mentioned above, since the examined specimens are not a
random sample, the result of inferential data analysis with signifi-
cance testing would be invalid. A cluster analysis or a cladistic
analysis of the data at population level, on the other hand, was
rejected on the grounds that these methods assume hierarchical
relationships, an assumption that is not justified. Phenotypic charac-
ters may be distributed clinally or in a reticulate manner, but are
unlikely to be strictly hierarchical (Crisp & Weston, 1993). If
hierarchical structure exists, it would suggest that the populations
have already undergone speciation and have to be treated as terminal
taxa in the phylogenetic analysis. The explorative data analysis was
performed on SYSTAT for the Macintosh Version 5.2 (Wilkinson,
1992).
UNIVARIATE ANALYSES. Phenological data are circular (Fisher,
1993). The reproductive period was thus analysed in a polar coordi-
nate system. In order to dampen peak values, which might merely
reflect a particularly high collecting effort, the collection proportion
of each month was converted to its logarithm.
MULTIVARIATE ANALYSES. Ordination techniques reduce a data
matrix of distances (similarities) among the samples to one or a few
dimensions. These types of techniques seem to be most appropriate
under a nonhierarchical model of infraspecific variation. Principal
component analysis (PCA) is the method most commonly applied in
multivariate systematics studies (James & McCulloch, 1990). It
aims to show the eventual existence of distinguishable groups with
discontinuous variation without a priori postulates of groups such as
populations. Since data sets with missing data cannot be processed
using PCA, only the most commonly represented characters of the
herbarium specimens (see above and Table 3) were analysed. Fortu-
nately, these are also the characters that are most commonly applied
to differentiate between the traditionally recognized Mediterranean
taxa. In constrast to many other analyses of specimen-based data,
within-collection variation was accounted for by recording and
analysing both minimal and maximal values of the flowering part
lengths. Only fully developed and open flowers were considered and
measured. The length/width ratio of the leaves was not included in
the PCA analysis on the grounds that ratios are often not linearly
related to the length measurements (Miles & Ricklefs, 1 984). All the
variables have been standardized (normalized) by the standard
Table 3 Variables used in the principal component analysis of the
Mediterranean complex.
Length measurements
Variable 1
Corolla length max.
mm
Variable 2
Corolla length min.
mm
Variable 3
Calyx length max.
mm
Variable 4
Calyx length min.
mm
Variable 5
Leaf length max.
cm
Phenology
Variable 6
Flowering time
month
22
deviation to obtain a variance of one and a mean of zero by
performing the analysis on a correlation matrix.
Due to a lack of available specimens from Asia of the taxa
Mandragora caulescens subsp. brevicalyx, M. chinghaiensis, M.
tibetica and M. turcomanica, the discussion of variability among
these taxa (and its implications on their delimitation) has to remain
qualitative.
Cladistics
Cladistic analysis is a parsimony-based search for hierarchical
arrangements or patterns of terminal units (in most cases species).
The premises are accordingly that there is a hierarchical structure of
the taxa and therefore of the characters used to define them, and that
the level of homoplasy in the chosen characters is low enough not to
infer convergent groups. As a strictly numerical procedure, there is
no need for additional models or assumptions relating to biological
processes, such as reproduction or descent (Davis & Nixon, 1992).
The resulting cladogram therefore represents a synapomorphy
scheme, merely suggesting closeness of relationship due to recency
of shared common ancestry. It might be congruent with the actual
phylogenetic tree, featuring the ancestors and descendants; how-
ever, there are many possible phylogenetic trees compatible with
one cladogram (Eldredge, 1979). The cladogram was rooted by
means of outgroup comparison.
A phylogeny of the tribe Hyoscyameae based mainly on morpho-
logical characters was recently published by Hoare & Knapp (1997)
suggesting that Mandragora, as traditionally defined, is mono-
phyletic. To test the monophyly of Mandragora as a whole, we
included in this analysis material of species representing two other
genera of the Hyoscyameae, Anisodus luridus Link and Atropa
belladonna (see Table 4). The three recognized terminal taxa of
Mandragora represent the ingroup (M. caulescens, M. officinarum,
Table 4 Taxa used in the cladistic analysis.
Nicotiana glauca Graham
Lycium chinense Mill.
Anisodus luridus Link
Atropa belladonna L.
Mandragora officinarum L.
M. turcomanica Mizg.
M. caulescens C.B. Clarke
S. UNGRICHT, S. KNAPP AND J.R. PRESS
Table 6 Data matrix used in the HENNIG86 analysis
Taxon
Character
1 I
1234567890 1
Nicotiana glauca
Lycium chinense
Anisodus luridus
Atropa belladonna
Mandragora officinarum
M. turcomanica
M. caulescens
00200010000
0020000001 0
11201 111110
10201 10001 1
11011 10001 0
11011 10101 ?
11111 10111 1
M. turcomanica). The outgroup, Nicotiana glauca Graham, was
chosen based upon previous cladistic analyses of chloroplast DNA
data sets (Olmstead & Palmer, 1992; Olmstead & Sweere, 1994;
Olmstead et al., in press). Lycium chinense Mill, was included as an
additional taxon (but not defined as an outgroup, see Tables 2 and 4).
Although acknowledging that there is no such thing as an a priori
'perfect cladistic character' (Thiele, 1993), the suggestion of Chappill
(1989) and Stevens (1991) not to include purely morphometric,
quantitative data in the cladistic analysis was followed. This limita-
tion is feasible in view of the minimal size of the ingroup and the
already established analysis of an extensive character set including
quantitative character states (Hoare & Knapp, 1997). The characters
used in the present work (Table 5) and their states are largely self-
explanatory. The data matrix used in the analysis is presented in
Table 6.
Cladistic analyses were undertaken withHENNlG86 (Farris, 1988)
using the ie option (implicit enumeration) with all characters
unordered, thus avoiding any character state change assumptions.
The generated cladogram statistics include length (L), the number of
character state changes on the tree, the ensemble consistency index
(CI) which provides a measure of character fit on the cladogram
(amount of homoplasy), and the ensemble retention index (RI)
which expresses the fraction of similarities on the cladogram inter-
preted as synapomorphy (Farris, 1989). Both of these indices have
an optimal value of 1 .
RESULTS AND DISCUSSION
Table 5 Character set used in the cladistic analysis.
Habit
1 . Woody plant (shrub or tree) 0; herb 1
Root
2. Tap-root: enlarged tap-root absent 0; present 1
Stem
3. Axis: condensed 0; first condensed, later elongated 1 ; elongated 2
Leaves
4. At anthesis: leaves clustered in a rosette 0; leaves alternate 1
Inflorescence
5. Type: numerous flowers in panicle 0; l(-3) axillary flowers 1
Corolla
6. Shape: tubular-infundibuliform 0; campanulate-urceolate 1
Fruit
7. Type: berry 0; capsule 16
Fruiting calyx
8. Not enlarged in fruit 0; markedly enlarged in fruit 1
9. Without prominent ribs 0; with prominent ribs 1
Seeds
10. Embryo: not curved 0; curved circularly or in a spiral 1
1 1 . Testal cells deep 0; shallow 1
Species delimitation using morphometric analyses
For the Mediterranean taxa a number of specific hypotheses were
tested using the above described statistical analyses.
Univariate analysis of reproductive period. — Are there two
distinct vernal and autumnal taxa ?
By adopting the two species Mandragora vernalis and M. autumn-
alis, Sprengel ( 1 825) argued against Linnaeus's unified Mandragora
officinarum concept. Even in later, more elaborate classifications the
division into these two major groups remained important (Table 1)
since it was obvious that two groups with widely differing flowering
periods would hardly be able to interbreed due to the seasonal
isolation. So, is there evidence in the compiled Mediterranean data
set that there are two phenologically distinct species? Figure 1
shows the distribution of the collecting dates (in flower or in fruit) as
an indicator of the reproductive period.
It is evident that there is actually only one, very extensive period
of reproductive activity from autumn to spring. In fact only during
the hottest summer months does the cycle break. If conditions are
MANDRAGORA
23
Mar
Jun
Feb
Jan
Dec
Sep
Nov
Fig. 1 Relative distribution of the vegetative period of the Mediterranean
complex (Mandragora officinaruiri) in the course of the year
(logarithimized monthly fractions: for analysis see text).
favourable, the same individual can flower twice a year, in autumn
and then again in spring, as can be seen from annotated herbarium
material, especially from cultivated material in botanical gardens.
The key factor is probably sufficient rainfall; for instance, sudden
and widespread flowering after exceptionally heavy autumn rains
was recorded from Tunisia (label data from Tomkinson 72). A more
detailed discussion pertaining to the influence of the climate and the
summer dormancy is given in the section on ecological biogeo-
graphy below.
Multivariate analysis of morphometric characters. - Are
there distinct clusters within the Mediterranean mandrakes
that are recognizable from gross morphology?
The different eigenvector values of the PCA (Table 8) indicate the
relative contribution of the variables. The first three axes (see Table
7) explain 92.2% of the variation if length measurements are consid-
ered only (Fig. 2a), and 87. 1% of the variation if the analysis was run
including the phenological variable of flowering time (Fig. 2b). In
both cases it is obvious that no clear-cut clusters can be distin-
guished. Cryptic species may exist within the Mediterranean region
but are likely to be revealed only if extensive, field-based studies
over the entire range, as well as transplant experiments, are under-
taken (Clausen et al, 1940). The information currently available
does not justify recognition of more than a single species of Mediter-
ranean mandrake.
Other characters traditionally applied in keys but not analysed
quantitatively here include the following:
Corolla colour. Hawkes (1972b) considered the elaborate split-
ting of the vernal and autumnal groups as unjustified and recognized
only the white-flowered vernal plants from northern Italy and Dal-
matia as distinct from the remaining Mediterranean group. Restricted
to only a small part of the total Mediterranean range, the white-
flowered plant has often been considered as the most distinctive
group (Heldreich, 1886; Vierhapper, 1915). The diagnostic value of
this character which has been extensively used in past and current
Table 7 Principal components (PC) of the Mediterranean complex. A:
Length measurements only. B: Length measurements and flowering
period combined.
Component
Eigenvalue
% of variance explained
A PC 1
3.248
64.97
PC 2
0.977
19.54
PC 3
0.383
7.66
B PC 1
3.328
55.47
PC2
1.410
23.50
PC 3
0.490
8.17
Table 8 Eigenvectors of the three most important axes (component
loadings).
Variable
PC 1
PC 2
PC 3
A
1
Corolla length max.
0.885
0.023
0.131
2
Corolla length min.
0.875
0.011
0.435
3
Calyx length max.
0.898
0.114
-0.391
4
Calyx length min.
0.924
0.065
-0.147
5
Leaf length max.
-0.197
0.979
0.047
B
1
Corolla length max.
0.882
0.067
-0.160
2
Corolla length min.
0.851
0.189
0.042
3
Calyx length max.
0.897
0.123
0.066
4
Calyx length min.
0.905
0.156
0.175
5
Leaf length max.
-0.181
0.865
-0.453
6
Flowering time
-0.413
0.763
0.471
keys is minimal, however, if one considers that flowers are very
variable in colour even within individuals (Jury et al. 12238 from
Morocco, Jury & Jury 13111 from Spain).
Fruiting calyx length: The relative length of the accrescent fruit-
ing calyx to the berry (longer: M. autumnalis or shorter: M.
officinarum) has also been proposed as a diagnostic character
(Hawkes, 19726). Again, it seems that within-group variation is just
as high, considering for instance a specimen housed in P of a
cultivated plant which features two berries that are clearly shorter
than the calyx, but three others which are longer.
Leaf shape: This is perhaps the most variable character of all. One
can get at least an idea of the variability if the length to width ratios
of mature leaves are compared. The variability within an individual
ranges from 1.7:1-4:1 (in Reverchon s.n.) to 4.6:1-10.5:1 (in
Welwitsch 159). The total variability within the whole set of exam-
ined specimens was found to be between 1.5:1-10.5:1.
Any distinction between Mandragora officinarum and M.
autumnalis or any other splitting of the Mediterranean Mandragora
on morphological or phenological grounds would be intentionally
giving formal status to divergent ends of clines. Thus overlapping
intermediates would be difficult if not impossible to assign to either
of the taxa in any other way than arbitrarily. Extensive overlap and
putative hybrids must be considered if aiming at expressing the
whole range of variability. This confirms Meikle's (1985) doubts
concerning the validity of the different proposed Mediterranean
species. Although certain plants of particular populations are quite
different in appearance from some plants of other populations, when
the total pattern of variation over the entire range is considered, no
diagnosable taxa within the Mediterranean complex of Mandragora
are discernable. Further morphological support for the unification of
the Mediterranean Mandragora comes from the anatomy of the tap
roots which has been shown to be identical in vernal and autumnal
taxa and in their chemical composition which is also similar (Jackson
& Berry, 1979).
For Mandragora turcomanica, a statistically valid morphometric
analysis is not feasible with the available literature data only. The
24
S. UNGRICHT, S. KNAPP AND J.R. PRESS
/.•
•/ •
••
I
•2
f 1
•2-10 1
FACTOR(2)
•1 0 1
FACTORO)
-3
I
T I T
J I I I
-2
-1
0 1 2 3 -4 -3 -2 -1 0 1 2 3
FACTORC2) FACTOR(3)
Fig. 2 Principal component analysis of the Mediterranean complex (Mandragora officinarum). Top pair: a. Bottom pair b. Left: PC 1,2. Right: PC 2,3.
range of the Turkmenian plants is so small that variability at popula-
tion level is considered equivalent with the variability over the whole
range. Mizgireva ( 1 955) meticulously documented teratogenic forms
of flowers and fruits of M. turcomanica, but did not cite field
collected vouchers for these forms. She also documented huge
variability in the shape and size of tap roots, and described the
growth of the plant in detail, from seed germination to fruiting.
Mandragora turcomanica is accepted as a distinct species on the
grounds of the evidence given in Mizgireva (1942, 1955, 1978) and
Lincevskij (1955). The whole plant is much larger than M.
officinarum. The lower leaves (± 90 x 60 cm) are especially large
resulting in a rosette with a diameter of 150-180 cm. Mandragora
turcomanica blooms in both the autumn and the spring (Mizgireva,
1955, see taxonomic treatment), with different individuals in the
population exhibiting one or the other flowering time. Geographi-
cally the species is isolated, occuring only in a narrowly restricted
range, and within that range it is only found in localized regions
(Mizgireva, 1955). It is also widely disjunct, being some 1500 km
from the closest Mediterranean Mandragora population in the Near
East and about 2500 km from the nearest Himalayan population in
western Nepal.
In common with the Mediterranean mandrake, the Himalayan
plants make up a complex with considerable variation in size, shape
and colour of the flowers and leaves. The following questions are
addressed:
MANDRAGORA
Is the recognition of the four infraspecific taxa sensu
Grierson & Long within Mandragora caulescens sensible?
After checking herbarium material from almost the entire range of
Mandragora caulescens (north-western Sichuan and southern
Qinghai excluded), Grierson & Long (1978) were well aware of the
extreme variability of the morphological characters they used to
describe their four subspecies. The characters used were corolla
colour, calyx and corolla lengths and overall plant size. As with
Mandragora officinarum, flower colour, according to observations
of the fresh material given on herbarium labels, is variable within
individuals as well as within local populations (e.g. Alden et al.
1 185: 'flowers yellow to purplish-brown', Long et al. 266: 'corolla
yellow, purple at base, inside and out', and Polunin 631: 'petals
yellow with purple veins'). Colours probably merely reflect to what
extent the yellow pigments are masked by purple ones. Grierson &
Long (1978) were unable to find any diagnostic morphometric
discontinuities. Their proposed classification is based on the rule
that due to its type status, the holotype of M. caulescens, collected in
Sikkim and representing an intermediate between the extremes of
the 'typical' small, yellow-flowered plants and the large, purple-
flowered ones, must retain the name M. caulescens (and must thus
be given the subspecific epithet caulescens). They therefore split the
complex not into two but into three subspecies representing the two
extreme forms (subsp. flavida and subsp. purpurascens) and the
intermediates (subsp. caulescens), and additionally described a
fourth subspecies with a shorter calyx (subsp. brevicalyx) based on
only two collections from Xizang. As a consequence, their classifi-
cation is based on non-diagnostic, largely overlapping characters,
rendering M. caulescens subsp. caulescens as a 'basket' taxon for
any intermediates. Indeed, it might be hypothesized that there are at
least two different species present within this complex with frequent
25
introgression of the sympatric populations. However, the many
putative 'hybrids' linking them and resulting in an extensive overlap
of their features make them unsuitable as terminal taxa for the
cladistic analysis. Furthermore, a restrictive taxonomic concept as
described earlier does not allow for the recognition of infraspecific
taxa if their characters and their ranges are extensively overlapping.
Are Mandragora tibetica and M. chinghaiensis distinct
species or are they both conspecific with M. caulescens?
As in the case of the Turkmenian mandrake, the unavailability of the
types and a reasonable number of specimens renders the data
somewhat meagre, so the original descriptions by Grubov ( 1 970) for
Mandragora tibetica and by Kuang & Lu ( 1 978) for M. chinghaiensis
must provide much of the necessary information. These descrip-
tions, together with their accompanying keys, which distinguish the
newly described species from M. caulescens, recognize the small,
yellow-flowered plants as distinct from the larger, purple-flowered
ones. Both new taxa are therefore covered by the range of variability
given in Grierson & Long's ( 1978) subdivision of M. caulescens into
four subspecies, which are defined only by overlapping differential
characters and, as such, are not acceptable as distinct taxa. Indeed, a
specimen in P, Polunin et al. 878, is annotated asM. chinghaiensis by
Lu An-ming and is also an isotype of M. caulescens subsp. flavida.
Some doubts must remain concerning M. tibetica and M. caulescens
subsp. brevicalyx that together may represent a single distinct taxon
with a very short calyx. An unusual specimen from western China
(Wilson 4195 (K), no precise locality given) bears a single flower
with a conspicuously narrow corolla. A note by Grierson and Long
on the herbarium sheet suggests that they considered this specimen
not conspecific or even congeneric with M. caulescens. Without
further material, however, proposing a new taxon would be just as
Fig. 3 The single most parsimonious cladogram from the HENNIG86 analysis. The characters are discussed in the text, and characters states are shown in
Table 5. For characters marked on the branches of the cladogram: single lines indicate unambiguous synapomorphies, parallel lines parallelisms and
crosses reversals (homoplasy).
26
S. UNGRICHT, S. KNAPP AND J.R. PRESS
speculative as disregarding the flower as a mere artefact of prepara-
tion. The synonymy presented in the taxonomic treatment here,
especially in respect to the undercollected plants from central and
northeastern Xizang, is therefore provisional and subject to confir-
mation once more material of both flowering and fruiting plants
becomes available. Only then will it be possible to judge whether or
not there exist distinct taxa not included within the range of variation
of M. caulescens.
Mandragora is a genus with pronounced variability, both within
and between individuals and therefore necessarily within the pro-
posed polymorphic species. Furthermore, discordant variation in the
applied gross morphological characters impedes a workable subdi-
vision based on a whole set of differential characters, rather than just
one (apparently non-existent) diagnostic character.
Phylogeny
The cladistic analysis resulted in a single most parsimonious
cladogram of length = 16, CI = 75 and RI = 73 (Fig. 3). The genus
Mandragora defined in the traditional sense is monophyletic in our
analysis: the species share synapomorphies in characters 3 (a con-
densed axis), 4 (leaves clustered in a rosette) and 9 (fruiting calyx
with prominent ribs). If the inaperturate pollen known from M.
caulescens and M. officinarum (see Hoare & Knapp, 1997 for
details) is also found in M. turcomanica, this too would be a
synapomorphy of the genus.
Mandragora officinarum and M. turcomanica are more closely
related to each other than to M. caulescens. They share reversals in
characters 3 (a condensed axis) and 9 (a fruiting calyx without
prominent ribs), but no unambiguous synapomorphies. The peculiar
life-cycle, i.e. the mode of development of the leaves and flowers, is
evidence for a shared common ancestry of the Mediterranean-
Turkmenian clade. Mizgireva (1955) also suggested this in her
revision of the Turkmenian species. In order to determine more
clearly the relationships among the three species of Mandragora, a
different, probably field-based set of characters is necessary. Here,
however, we are concerned more with the delimitation of terminal
taxa, especially among the Mediterranean plants, and whether or not
the genus as currently defined is monophyletic.
A comparison between alternative hypotheses concerning the
position of Mandragora based on morphological, chemical and
molecular (chloroplast DNA) characters is given in Figure 4. The
most pronounced discrepancy concerning the position of Man-
dragora is that the molecular data set (Fig. 4a) deviates from the
morphological (Fig. 4c) and chemical ones (Fig. 4b) by attributing
this genus a very independent position due to extremely divergent
and autapomorphic cpDNA sequences. This is reflected in the
proposal of a monogeneric tribe Mandragoreae (Olmstead et al., in
press). Morphological and chemical evidence, on the other hand,
suggests a placement within the tribe Hyoscyameae (Hoare &
Knapp, 1997; = AtropeaesenswTetenyi, 1987). The present study is
far too limited in its scope to allow any conclusions concerning
higher level classification within the Solanaceae. However, it is clear
that our limited data set supports the inclusion of Mandragora
within the Hyoscyameae (synapomorphies in characters 1, 5 and 6)
but, as mentioned above, a much larger and more field-based data set
will be needed to address this problem. Mandragora is an anoma-
lous genus in the family, both morphologically and in terms of
molecules, but this should not prevent us from eventually identifying
its true affinities and relatives given adequate data.
Biogeography
The biogeography of an exclusively Laurasian genus within the
Solanaceae is of interest because the family is largely Gondwanan in
distribution. It is useful to approach biogeography from two differ-
ent angles. Ecological biogeography considers a short temporal
scale in evaluating ecological factors relevant for present-day distri-
butions of taxa. Historical biogeography, on the other hand, is
concerned with the change of distributions and the potential causes
thereof on a long temporal scale. We attempt here to summarize
some of the most important data with respect to the geographical
distribution of Mandragora, both now and in the past, and pinpoint
major gaps in knowledge.
Ecological biogeography
The present approximate distribution of the genus is given in Figure
5. Apart from the herbarium material examined, the sources pre-
sented in Table 9 were used. Most comprehensive are the distribution
data in Heldreich (1886) and Vierhapper (1915). Mandragora does
not appear to penetrate into the Arabian Peninsula, being absent
ABC
Fig. 4 Differing hypotheses of the relationships of Mandragora within the Solanaceae. a: Olmstead et al., in press, b: Tetenyi, 1987, c: Hoare & Knapp, 1997.
MANDRAGORA
27
20
Fig. 5 Distribution of the genus Mandragom. 1) M. officinarum, 2) M. turcomanica, and 3) M. caulescens.
even in the most northern parts of Saudi Arabia (Migahid, 1989) as
well as from Mesopotamia (Rechinger, 1964; Mouterde, 1979). The
genus was naturalized in Malta in the nineteenth century but is now
presumably extinct there (Borg, 1927; Maugini, 1959). Mandra-
gora does not appear in works covering France and Corsica (Fournier,
1990), Albania (Demiri, 1983), Egypt (Tackholm, 1974; Boulos,
1995) or Libya (Siddiqi, 1978) and is presumed absent from these
areas. Likewise, the genus is absent from the Macaronesian archi-
pelagos of the Canaries, Madeira and the Azores (not recorded either
by Hansen & Sunding, 1993; or Press & Short, 1994).
Mandragora turcomanica is found only in the Kopet Dag range of
southern Turkmenistan. It has not yet been reported from the Iranian
part of these mountains (Rechinger, 1972). In eastern Asia, Man-
dragora (M. caulescens) occurs in the central and eastern Himalaya
Table 9 Sources used for determining the overall distribution of the
genus Mandragora.
Geographical area
Reference
Tunisia
Algeria
Morocco
Portugal
Spain (incl.
Balearic Is.)
Italy (incl. Sicily,
Sardinia)
Former Yugoslavia
Greece (incl. Aegean Is.)
Turkey
Cyprus
Syria
Lebanon
Israel
Jordan
Turkmenistan
Nepal
India (Sikkim,
Arunachal Pradesh)
Bhutan
Myanmar
China
Bouquet (1952), Pettier- Alapetite (1981)
Quezel& Santa (1963)
Jahandiez & Maire (1934)
Amaral Franco (1984)
Lazaro e Ibiza (1907), Bonafe Barcelo (1980),
Valdesetal. (1987)
Maugini (1959), Pignatti (1982),
Cellinese et al. (1994)
Hayek &Markgraf( 1931)
Halacsy (1902), Hayek & Markgraf (1931)
Baytop(1978)
Meikle (1 985), Viney ( 1994)
Post (1933), Feinbrun-Dothan (1978),
Mouterde (1979)
Post (1933), Feinbrun-Dothan (1978),
Mouterde (1979)
Post (1933), Feinbrun-Dothan (1978)
Post (1933), Feinbrun-Dothan (1978)
Pojarkova (1955), Kurbanov ( 1 994)
Grierson& Long (1978)
Hara (1966), Grierson & Long (1978), Deb(1979)
Hara (1971), Grierson & Long (1978)
Grierson & Long (1978)
Grierson & Long (1978), Zhang et al. (1994).
Zhang &Lu( 1994)
and southwestern China, but there are no records from the western
extensions of these mountain ranges (Karakorum, Pamir, Hindu
Kush and Tien Shan).
Apart from historical factors, dealt with below, the principal
factor determining the geographical distribution of taxa on a large
scale is typically climate. Although one might disagree on the
relative importance of means and extreme values of various climatic
variables, the prime importance of climate in general has long been
accepted (Cain, 1944).
A convenient way of depicting and comparing different climates
is by means of standardized diagrams. The purpose of the so-called
ecological climate-diagrams is to portray the climate of a locality in
a readily comparable way and visually to present the most important
climatic factors affecting the growth of plants (Walter et al., 1975).
By choosing a scale at which 10°C correspond to 20 mm precipita-
tion, a relatively wet (dark) and relatively dry (light) season can be
identified on the diagrams (Fig. 6). These climate diagrams make it
evident that the range of Mandragora comprises not only quantita-
tively different climates but qualitatively different climate types.
The amount and especially the main season of rainfall is particularly
significant.
The Mediterranean climate is characterized by mild, rainy winters
and hot, dry summers. Although the total amount of rainfall for other
locations for Mandragora officinarum may differ somewhat from
that presented here (Iraklio, on the northern coast of Crete, Fig. 6),
the general aspect of a winter rainfall regime remains the same all
over the Mediterranean basin, even extending towards Afghanistan
and the Hindu Kush. The typical vegetation consists of adapted
sclerophyllous vegetation, analogous to that found in the same
climate type in California, Chile, the Cape region of South Africa
and southwestern Australia.
The Kopet Dag is a Transcaspian mountain range surrounded by
the Irano-Turanian desert. The climate there represents a continen-
tal, dry version of the Mediterranean type with much lower rainfall
(Fig. 6: Kisil-Arvat)) and an extended summer arid period. During
this dry summer, Mandragora turcomanica exhibits a long period of
xerothermic dormancy (Kurbanov, 1994). Mizgireva (1955) sug-
gested that M. turcomanica is much more cold-tolerant than its
mediterranean congeners. The typical vegetation in the Kopet Dag
consists of broadleafed deciduous forest in the moister microhabitats
and open shrubland in drier areas.
The eastern part of the Mandragora range in the Sino-Himalayan
28
S. UNGRICHT, S. KNAPP AND J.R. PRESS
Irakiio 27 m alt.
Gangtok 1725m alt.
IN
so
M
4*
20
0
Kisil-Arvat 101 m alt.
100
m
M
40
20
0
30
20
10
0
Gyantse 3996 m alt.
A
Fig. 6 Climate diagrams of four representative weather stations in three
different climate regimes. Given are the rainfall and temperature curves,
resulting in wet (dark shading) and dry (light shading) seasons. Based
on Walter et al. 1975.
region is characterized by a completely different rainfall regime
altogether. The influence of the monsoon renders not the winter but
the summer the season of the highest rainfall. The amount of
precipitation during the monsoon period, however, does vary enor-
mously. The Himalayan chain functions as a rain screen resulting in
high rainfall on the southern slope of the mountain range (Fig. 6:
Gangtok) as well as in the southwestern Chinese mountains of
Yunnan and Sichuan, and much lower rainfall in the inner regions of
Xizang (Fig. 6: Gyantse). Even at high altitudes, winters are very
mild in the high rainfall areas (see climate-diagram of Gangtok,
about 1700 m above sea level) and snow is scarce. Generally
speaking, the eastern Himalaya and southeastern Xizang, Sichuan
and Yunnan enjoy a much more humid and milder climate than the
western Himalaya and central Xizang. The locality data of the
examined specimens suggest that the distributional boundary of the
Himalayan mandrake towards the central Xizang highland lies
within the 200-400 mm annual precipitation strip north of the main
ranges. Due to the immense altitudinal gradient, the vegetation is
strikingly zonal and ranges from subtropical forests to alpine scrub
and meadows. The upper forest limit lies between about 3600 and
4200 m, the upper tree limit between 4200 and 4500 m. The belt in
between is characterized mainly by a succession of various Rhodo-
dendron species. This is the typical habitat of Mandragora caule-
scens. The permanent snow line lies at about 4800 m in the south and
5800 m in the north towards central Xizang (Walter, 1962).
Other factors affecting plant distribution, albeit on a smaller and
less important scale, include soil type and competition. Man dragora
officinarum especially grows in ruderal habitats, on clayey and
nitrogen-rich soils. So far, it is not clear whether M. caulescens
occurs on acidic (granitic) or base-rich (limestone) soils in the
subalpine and alpine zone of the Himalaya. It cannot be ruled out
that there might be two different, vicariant ecological groups as is
often the case in the Alps, for example in Rhododendron (Walter,
1966). The type of climate and soil typically only indirectly deter-
mine the occurrence of a species in a certain locality, in that they
affect the fitness of that species in relation to others (Walter, 1977).
Both Mandragora officinarum and M. turcomanica seem more or
less restricted to low-competition localities, representing typical r-
species (good colonizers but poor competitors).
Historical biogeography
The atemporal cladogram of a group together with historical evi-
dence, namely paleoclimatic and paleogeographical data allow for
the formulation of historical scenarios. A scenario is a model of the
basic features (including space and time) of the evolutionary history
of a group which can serve to stimulate directions of further investi-
gation. However, the essentially speculative character of such a
model renders it more like a tool than a serious scientific statement
(Eldredge, 1979) and ad hoc assumptions are inevitable (Humphries
& Parenti, 1986). In attempting to construct scenarios for the
historical biogeography of any group, plate movement, eustasy
(worldwide changes of sea level) and climate changes are consid-
ered the major factors to be considered (Hallam, 1981). In the case
of the Solanaceae, virtually no fossil record is available (PaleoNet
Listserver, http://www.ucmp.berkeley.edu/Paleonet/), thus histori-
cal hypotheses concerning Mandragora are inevitably founded on
extant taxa only.
The Mediterranean region presents a very complex tectonic jig-
saw puzzle, hard to resolve adequately (Hallam, 1981). It seems
clear, however, that during most of the Tertiary, summer was the
rainy season and only before the Pleistocene was there a shift
towards the current pattern of rainfall in winter. The plants had to
adapt and reduce activity during the dry summer months (Walter,
1977). The northern Mediterranean localities for Mandragora in
northern Italy and on the Dalmatian coast are very patchy, resulting
in a highly fragmented range there, and the precise status of these
populations is uncertain. We have been unable to discover any
modern collections from these regions (Cellinese et al., 1994) and
some, or all, of the populations may no longer exist. Vierhapper
(1915) maintained that these plants are Tertiary relics, the scattered
remnants of a formerly larger, continuous range, that have been
unable to adapt to the changing climate and still exhibit the vernal
phenology type. Relict status based on Tertiary processes, however,
is normally attributed to groups of far wider disjunct ranges than is
the case here. Hawkes ( 1972a) based his definition of M. officinarum
L. on these northern Italian and Dalmatian populations only, using
corolla colour and phenology. On the evidence of our multivariate
analyses, Hawkes's position is untenable, since the plants fall within
the range of variation exhibited by the rest of the Mediterranean
plants. It could be argued that the different corolla colour (white) of
the northern Italian and Dalmatian plants, coupled with their dis-
crete distribution is sufficient to recognize them at least at subspecific
rank. We have already pointed out that the corolla colour is only an
MANDRAGORA
29
apparent distinction, with similar colours occurring in populations
outside northern Italy and Dalmatia. The somewhat isolated distri-
bution alone is insufficient reason to recognize the Italian and
Dalmatian plants as a separate taxon.
The close relationship between the Mediterranean and the
Turkmenian mandrake as suggested by the cladistic analysis is
reflected in a linked history of the regions particularly during the
Tertiary. It can be proposed that in this period, after the Tethys Sea
receded and the middle Asian mountains became exposed during
their simultaneous uplift, a vicariance event of the Mediterranean
and Turkmenian ancestors took place. Both extant species show a
comparable adaptation to the aridization and change from a summer
to a winter rainfall regime. In other words, Mandragora turcomanica
can be seen as an example of a plant that has been evolving in
isolation in Turkmenistan since the Tertiary from ancient Tethyan
stock (Kurbanov, 1994). It is of some interest to note that there is a
highly congruent pattern within Atropa where the disjunct distribu-
tion of the Turkmenian A tropa komarovii Blin. & Schal., consists of
a very narrow range south of the river Sumbar only a few miles away
from the Mandragora turcomanica population north of the river
(Kurbanov, 1994).
The Sino-Himalayan area is of particular interest because of its
position at the transition between the Palearctic and the Oriental
floristic realms. The period of the collision of the Indian sub-
continent with Asia is contentious but probably occurred in the
Paleogene approximately 50 million years ago (Scotese et al., 1988)
with the penetration of the Eurasian border continuing during the
early Neogene (c. 20 millions years ago) (Dercourt et al., 1986). The
connection between Tethys and the Mediterranean, on the other
hand, probably did not close until the mid-Miocene (c. 15 million
years ago) (Hallam, 1994). Climate changes during the Paleogene
were severe. Temperatures increased up to the early Eocene (c. 55
million years ago) followed by a global cooling which marks the
passage from a warm domain of the Mesozoic and early Palaeogene
to the cool domain thereafter (Hallam, 1994). According to Rud-
diman et al. (1989) much of the uplift in the Xizang plateau region
occurred only in the last 5 to 10 million years (Plio-Pleistocene
uplift). It must be assumed that the glaciation of the high altitude
regions of the Sino-Himalayan area during the Pleistocene (Ice-
Ages) was more or less complete, posing the question whether
refugia further south and at lower altitudes might have served as
speciation centres.
Schuster (1976) argued that the Indian plate served as a vehicle for
the dissemination of Gonwanan taxa into Laurasia. But at the same
time he expected that due to the migration through several climatic
zones the Indian plate probably arrived with a rather depleted and
altered flora and therefore only remnants of the original Gondwanan
flora are recognizable in India today. If angiosperms were indeed on
the Indian plate in early Cretaceous time, they could probably not be
assigned to presently recognizable families. On the basis of a
proposed late Triassic (c. 210 million years ago) start of the Indian
plate migration, serious doubt is cast on the possibility that early
Solanaceae could have been 'on board'. The main objection is that
the plate drift simply occurred too early, before modern groups were
in existence. However, other data (Smith et al., 1994) suggest that
India did not break free from Gondwana until the late Jurassic to
early Cretaceous (130-140 million years) or even near the Creta-
ceous-Tertiary boundary only 60-65 million years ago (Hallam,
1994). Lu & Zhang (1986) proposed southwestern China as a later,
secondary speciation centre of the whole tribe Hyoscyameae on the
grounds of its high species-richness.
A different biogeographical scenario proposes a migration from
Gondwana to the Mediterranean basin. Migration routes that are
closed today were wide open in the Mesozoic. Indeed, massive and
relatively unimpeded migrations from Gondwana to Laurasia might
have been feasible until the late Cretaceous (Schuster, 1976). Con-
trary to the first scenario, the mandrake ancestors would therefore
have spread eastwards from the Protomediterranean to the develop-
ing eastern Asian mountain ranges where the massive uplift possibly
resulted in a vicariance event. The Mediterranean populations on the
other hand would have had to adapt to the increasing aridization in
the western part of the continent during the late Tertiary (Weinert,
1972) and thus slowly changed their mode of development from a
summer flowering to a autumn- winter-spring flowering rhythm.
Proskuryakova & Belyanina (1985) claim that the slow succession
of the development phases, particularly of the germination, the long
growth and flowering period and the large size of the plant speak for
the great age of the Mediterranean group. They draw the conclusion
that the Mediterranean section must be older, representing a Terti-
ary relic, than the apparently highly adapted Himalayan mandrake.
The Himalayan mandrake, in fact, might have arisen from now-
extinct ancestors as late as the Quaternary glaciations.
To conclude, the massive timespan covered in the Tertiary, the
many fluctuations in climate and transgressions of the seas and the
current very inaccurate timing of even the major events render
definitive statements or falsifications concerning the above scenar-
ios and the history of Mandragora hardly possible. In this context
the distinction between a cladogram (what is more closely related?)
and the actual phylogenetic tree (what is the ancestor?) must be
stressed. The cladogram (Fig. 3) suggesting a more recent common
ancestry of Mandragora officinarum and M. turcomanica is con-
gruent with only one of the many possible trees equally resulting in
this particular cladogram, which therefore represents a far less
informative (more general) mode of expression. Thus, it does not
allow for statements concerning the ancestor and does not imply that
Mandragora caulescens is the older taxon. Nevertheless, it can be
proposed that the major events affecting speciation and present
distribution of Mandragora in the Mediterranean-Turkmenian area
were probably the receding of the Tethys and the aridization of the
climate in the Tertiary. In the Sino-Himalayan area, on the other
hand, the collision of the Indian plate and the subsequent uplift of the
Himalayan range, as well as the later glaciation periods during the
Quaternary, are probably most important.
TAXONOMIC TREATMENT
Mandragora L., Sp. pi. 1: 181 (1753). Type species: M.
officinarum L.
Perennial herbs. Stout tap-root, often branched, sometimes vaguely
anthropomorphic. Stem very short or with secondary growth during
flowering and fruiting, sometimes branched. Leaves simple, nor-
mally entire, occasionally toothed. Inflorescence of typically solitary
axillary, pedicellate flowers. Floral envelope actinomorphic. Calyx
5(6)-lobed, persistent, enlarged in fruit. Corolla 5(6)-lobed. Sta-
mens 5(6), included, inserted in lower half of corolla-tube; filaments
filiform, pubescent at base; anthers dorsifixed, dehiscing longitudi-
nally. Ovary 2-locular; ovules numerous; style elongate; stigma
capitate or somewhat bilobed. Fruit a fleshy, many seeded berry,
globose or ovoid, borne beneath the leaves on nodding pedicel or
lying on the ground. Seeds compressed; embryo curved.
FLORAL FORMULA. K(5) C(5) A5 G(2)
DIVERSITY AND DISTRIBUTION. Three disjunct species: Mediterra-
nean region, central Asia, Sino-Himalayan region.
30
PHENOLOGY AND HABITAT. All three Mandragora species are
geophytes with a conspicuous tap-root which is the perennating
organ of these herbs, linking the annual vegetative periods. The
Turkmenian mandrake with its prolonged dry season dormancy has
been referred to as an ephemeroid (Proskuryakova & Belyanina,
1985), its rhythm of development more closely related to the Medi-
terranean than to the Himalayan climatic cycle. Propagation is
usually by seeds or vegetatively by underground buds on the tap-
root. The species grow in open woodland, olive groves, grassland,
fallow land, waysides, railway embankments, ruins, in rock crev-
ices, grassy and stony mountain slopes and screes. The altitudinal
range is 0-1200 and 3000-4900 m.
ETHNOBOTANY. Once important medicinal plants with a long tra-
dition in mythology and superstition. All parts of the plant contain
potent narcotic alkaloids.
ETYMOLOGY. Origin uncertain, possibly from the Greek |0,otv-
Spcryopaa, original meaning controversial; alternatively from the
Sanskrit mandros, meaning sleep and agora, meaning object or
substance (Thompson, 1934).
Key to the species of Mandragora
1 Young leaves arranged in a cluster on top of a stem with scales, later
along the secondarily elongated, sometimes branched stem; corolla
yellow to dark purple; style not exceeding stamens. Sino-Himalayan.
Alt. 3000-4900 m 3. M. caulescens
Leaves permanently arranged in a rosette, the stem not elongating;
corolla white to pale blue or violet; style exceeding stamens 2
2 Mature lower leaves in a large rosette of 1 50- 1 80 cm; berries 40-60 mm
in diameter. Turkmenian. Alt. 500-700 m 2. M. turcomanica
Mature lower leaves in a rosette of less than 1 00 cm; berries less than 40
mm in diameter. Mediterranean. Alt. 0-1200 m ... 1. M. officinarum
1. Mandragora officinarum L., Sp. pi. 1: 181 (1753). Type: Herb.
Burser IX: 26 (UPS-lectotype, designated by Knapp in Jarvis et
al. (1993), microfiche-BM!).
Figs 7, 8.
= Atropa mandragora L., Syst. nal 10th ed.: 933 (1759), nom.
illeg. (Art. 52.1).
= Mandragora foemina Garsault, Fig. pi. med. 3: 221, t. 363 p.p.
(1764).
= Mandragora mas Garsault, Fig. pi. med. 3: 221, t. 363 p.p.
(1764).
= Mandragora acaulis Gaertn., Fruct. sem. pi. 2: 236, t. 131
(1791).
= Atropa humilis Salisb., Prodr. stirp. Chap. Allerton: 132 (1796).
= Atropa acaulis Stokes, Bot. mat. med. 1: 396 (1812).
= Mandragora autumnalis Bertol., Elench. plant, viv.: 6 (1820).
= Mandragora vernalis Bertol., Virid. Bonon. veg.: 6 (1824).
= Mandragora praecox Sweet, Brit.fl. gard. 2: t. 198 (1827).
= Mandragora neglecta G. Don ex Loudon, Hon. brit.: 71 (1830).
= Mandragora microcarpa Bertol., Comm. Mandrag.: 12, t. 3
(1835).
= Mandragora haussknechtii Heldr. in Mitt. Geogr. Ges. Jena. 4:
77, app. (1886).
= Mandragora x hybrida Hausskn. & Heldr. in Mitt. Geogr. Ges.
Jena. 4:77(1886).
= Mandragora hispanica Vierh. in Osterr. Bot. Z. 65: 132-133
(1915).
ILLUSTRATIONS. Wettstein (1895: fig. 12M-O); Feinbrun-Dothan
S. UNGRICHT, S. KNAPP AND J.R. PRESS
(1978: fig. 278); Pignatti (1982: figs on p. 518); Hoare & Knapp
( 1997: fig. 5c).
Plant virtually stemless. Root stout, often branched, very long.
Leaves arranged in a rosette, leaf shape and size very variable, but
usually elliptic to obovate, max. 45 cm long, length/width ratio 1 .5-
10, variously pubescent. Flowers borne in leaf axils. Pedicels very
variable in length, max. 15 cm. Calyx 6-28 mm long, lobed '/2-2/3 of
the way to the base, lobes 3-15 mm, acuminate, only slightly
accrescent in fruit. Corolla 12-65 mm long, campanulate, greenish
white to pale blue or violet, lobed l/2 to nearly to the base, the lobes
6-60 mm. Stamens adnate to lower half of corolla; filaments 7-15
mm long; anthers 2.5^.0 mm long, yellow to brown, sometimes
pale blue. Style 8-20 mm, exceeding stamens; stigma capitate. Fruit
5^0 mm in diameter, globose to ellipsoid, glossy yellow to orange
when ripe, juicy, edible. Seeds 2.5-6.0 mm long, reniform, the
surface reticulate, yellow to light brown.
HABITAT. Open woodland, olive groves, fallow land, waysides,
railway embankments, ruins, crevices. Altitudinal range 0-1200 m.
VERNACULAR NAME. Mediterranean mandrake.
PHENOLOGY. Flowering September to April. Fruiting November
to June.
CONSERVATION STATUS. Populations are scattered throughout the
species range. Northern Italian populations are vulnerable (Cellinese
et al., 1994) as they are apparently known only from old herbarium
records and the present occurrence and abundance of these
populations are unclear.
DISTRIBUTION. Circum-Mediterranean. (Tunisia, Algeria, Mo-
rocco, Portugal, Spain, Italy, former Yugoslavia, Greece, Cyprus,
Turkey, Syria, Lebanon, Israel, Jordan).
SELECTED SPECIMENS EXAMINED. TUISIA. Cap Bon, roadsides and
orchards, 0-150 m, 6 October 1969, Tomkinson, M.J. 72, 72A (BM). Nabel,
60 m, 14 October 1907, Gandoger, M. 84 (K).
MOROCCO. Guercif, 144 km E. of Fes near 'La Kazbah', 38 km
before the town of Guercif, 34°16'N, 3°45'W, 510 m, disturbed gravelly
area by road, 3 November 1994, Jury, S.L. et al. 15434 (RNG); Gharb, c. 4
km NNE of Souk-El-Arbaa-du-Rharb on road to Ksar-El-Kebir, by bridge
and salt works, 34°43'N, 6°00'W, 30 m, pasture between fields at side of
salt pans, 7 November 1994, Jury, S.L. et al. 15701 (RNG); Zerhoun, c. 3
km NW of Moulay Idriss and 33 km from Meknes along road to Sidi
Kacem and Tanger, 34°04'N, 5°33'W, 390 m, ruins of Roman city of
Volubilis, 1 November 1994, Jury, S.L et al. 15398 (RNG); Prerif Central,
N. of Fes, on the road to Ouezzane, 34°22'N, 5°09'W, 210 m, roadside
ruderal community, 24 October 1993, Jury, S.L et al. 12634 (RNG); Haul
Ouerrha, c. 50 km N. of Fes on the road between Fes-el-Bali and Ourtzarh,
34°33'N, 5°02'W, 120 m, in olive plantation on steep N-facing slope with
Zizyphus lotus, 24 October 1993, Jury, S.L et al. 12653 (RNG); Temara, 10
km S. of Rabat, 0-30 m, sandy roadside, 14-20 October 1981, Lewalle, J.
10054 (BM, RNG); 5 km W. of Oued, Beht on Rabat-Meknes road,
33°52'N, 5°58'W, 400 m, 27 September 1991, Gardner, M.F. et al. 4885 (E,
RNG); Loukkos, SW of Chefchaouen, on road to Ksar-el-Kebir, 35°1'N,
5°45'W, 120 m, in cultivated fields of wheat, 21 October 1993, Jury, S.L. et
al. 12542 (RNG); Loukkos, c. 1 km E. of Larache on road to Asilah, c. 200
m from Oued Loukkos, 35°12'N, 6°08'W, 5 m, at side of road by saltpans,
28 October 1994, Jury, S.L et al. 15237 (RNG), Tanga, just N. of airport,
road to Cap Spartel, 35°45'N, 5°55'W, 35 m, fallow area at side of ploughed
field with Urginea maritima, 16 October 1993, Jury, S.L. et al. 12217
(RNG), Tanga, E. on road to Sebta (Ceuta), Oued Dliane, 35°50'N, 5°39'W,
5 m, nitrophilous areas on fallow cultivated land on heavy clay alluvial
plain, 17 October 1993, Jury, S.L. et al. 12238 (RNG); Allal-Tazi, close to
the river Sedou, 1 November 1965, Talon, A. 1630 (BM).
SPAIN. Baleares, Palma, between S. Sardina and S. Bernardo, 22 Septem-
ber 1913, Bianor & Sabasien 1877 (BM, E, P, RNG); Province Cadiz,
MANDRAGORA
Fig. 7 Mandragora officinarum. a) Flowering, cultivated at RBG Kew, b) Fruiting, cultivated at RBG Kew.
32
S. UNGRICHT, S. KNAPP AND J.R. PRESS
MANDRAGORA
33
Chiclana, on clayey soil by the river, 10 January 1 855, Bourgeau, E. 2304 (E,
K, P);Torredonjimeno, 600 m, 10 December 1986, Espinosa, A. 870252 (K);
alrededores de Cordoba, Cuesta del Espino, 2 December 1967, Galiano, E.F.
& Valdes, B. 820 (E, RNG); Province Malaga, C 341 from Campillos to
Ronda, 36°52'N, 5°00'W, 525 m, disturbed ground near new road cutting, 27
October 1990, Gardner, M.F. & Knees, S.G. 4769 (RNG); Province Cadiz, 23
km from Jerez de la Frontera, 36°32'N, 5°59'W, 1 30 m, ruderal area by road
with abundant Silybum marianum below ruined castle of El Pedroso, 22
December 1993, Jury, S.L. & Jury, L.C. 13111 (BM, RNG); Province Ja6n,
Torredonjimeno, between Porcuna andTorredonjimeno, near Pilar de Moya,
500 m, roadside on chalky soil, 3 November 1979, Ladero, M. et al. 10605
(RNG); Malaga, near Churiana, 3 December 1957, Partridge, F. 2 (BM);
Campamento, 9 November 1911, Wolley-Dod, A.M. 22 (BM).
ITALY. Sicily, Province Palermo, Madonie, near Polizzi, 950 m, rocky
limestone slopes, 1 8 August 1964, Davis, P.M. 40134 (E); Sicily, Trapani, 200
m, steep slopes, 25 August 1964, Davis, P.M. 40182 (RNG); Sicily, Province
Siracusa, Augusta, October-November 1911, Vaccari, A. 2326 (BM, K).
FORMER YUGOSLAVIA. Hercegovinae, Doljnja Glavska, 6 May
1908, Reiser, C. s.n. (K); Silieski, near Ragusa, 500 m, 29 April 1909,
Maly, K. s.n. (K).
GREECE. Near Corinth, 30 m, waste grounds and waysides, April 1931,
Atchley, S.C. 819 (K); Crete, Frangokastello, 29 March 1973, Barclay, C.
3004 (K); Crete, Canea, 1 December 1943, Bickerich, G. 15324 (BM, K);
Crete, Kolokithia, December 1977, Bowen, H.J.M. 694 (RNG); Crete, Agios
Nikolaos, December 1983, Bowen, H.J.M. 3088 (RNG); Peloponnese, 5 km
from Pilos on the road from Kalamata, 250 m, in cultivated fields near stream,
12 November 1973, Brickell, C. & Mathew, B. 8088 (K); Crete, Canea,
frequent in hills and ditches throughout Crete up to 300 m, 1 January 1940,
Davis, P.H. 1100(E,K);Cyclades,Amorgos, lOApril 1940, Davis, P.M. 1438
(K); Dodecanese, Kos, Asfendion, 300-600 m, rocky slopes, 27 March 1965,
Davis, P.H. 40478 (K); Crete, near Candia, Hagios-Deka, December 1913,
Duffour, C. 1501 (P); Cyclades, Kea, 30 April 1961, Gathorne-Hardy, E. 14
(E); Dodecanese, Tilos, near chapel of Ayios Nicolaos c. 180 m above
Livadia, 26 April 1962, Gathorne-Hardy, E. 139 (E); Crete, near Sitia, 150-
200 m, on clayey soil, 30 October 1966, Greuter, W. 7821 (E); Near Corinth,
Examillia, 8 February 1931, Guiol, F. 1731 (BM); Crete, Canea, on waste-
land, 10 March 1883, Reverchon, E. 117 (BM, E, K, P, PAL); Dodecanese,
Kos, by track near Asclepeion, 23 March 1985, Townsend, C.C. 85/11 (K);
Dodecanese, Rhodes, 9 February 1914, Vaccari, A. 1141 (K); Crete, Nomos
Lasithiou, Monastiraki, roadside, 5 April 1980, Walree, A.M. 22226 (BM).
CYPRUS. Kyrenia, fallow olive grove, 13 February 1949, Casey, B.C.
221 (K).
TURKEY. Province Mugla, Kemer-Kestep, 50 m, edge of fields, 29
March 1956, Davis, P.H. & Polunin, O. 25469 (BM, E, K); Side, 22 February
1966, Baytop, A. et al. 8504 (E).
SYRIA. Baniyas, 300 m, foot of hill, beneath tree, 11 March 1945,
Norris, F.H. s.n. (BM).
ISRAEL. Jerusalem, fields, 800 m, 20 February 1908, Meyers, F. 4377
(E); Mount Carmel, upper Nahal Nesher, 2 km S. of Nesher, 32°45'N,
35°31'E, 280 m, terra rossa on hard limestone, 6 April 1989, Danin, A. &
Knees, S.G. 1709 (RNG); Mount Gerizim, near Nablus, fields, 700 m, 22
December 1910, Meyers, F.S. & Dinsmore, J.E. 6377 (E); District Haifa, near
Waldheim (Alloney Aba), open places in Quercus ithaburensis forest, 28
March 1942, Davis, P.H. 4166 (E, K).
The confusion surrounding specific epithets for the taxon we are
here calling Mandragora officinarum is incredible. This morass has
arisen for two basic reasons, firstly orthographic errors, and sec-
ondly, the confusion over the phenological status of type specimens.
Orthographic errors made by early authors persist through floristic
works, adding and compounding any ambiguities that may have
arisen. A name often used in synonymy (or as a valid name) of M.
officinarum is Mandragora officinalis Mill., said to be validly
published in 1768 (see Jackson & Berry, 1979: 508, footnote).
However, in the eighth edition of The gardener's dictionary (Miller,
1768), where Miller used Linnaean epithets for the first time, Miller
actually used the epithet officinarum ('MANDRAGORA
(officinarum)'), notofficinalis. In the ninth edition (Miller & Martyn,
1807), the taxon was classified as Atropa mandragora, following
Systema naturae (Linneaus, 1759) and the second edition of Species
plantarum (Linnaeus, 1762). Mandragora officinalis was cited in
synonymy, with reference made to the illustrations produced to
compliment the earlier editions (Miller, 1760). These illustrations
bear no specific epithets, thus the 1807 reference to the epithet
officinalis is almost certainly an orthographic error for the 1768 use
of Linneaus's epithet officinarum, which was not cited in synonymy
in the ninth edition (Miller & Martyn, 1807). Thus the name
Mandragora officinalis Mill, has persisted in the literature, but has
no real nomenclatural standing.
Similarly, Bertoloni (1820) attributed the epithet officinalis (see
below) to Willdenow, explicitly citing 'W. En.\ Willdenow (1809),
in his Enumeratio plantarum, attributed the epithet officinalis to
DeCandolle (in Lamarck & DeCandolle, 1805), where it is again
attributed, in error, to Miller's Gardener's dictionary. Thus it is
apparent that the numerous occurences of the epithet officinalis,
attributed to many authors, are traceable to a single orthographic
error from Miller's 1767 edition of the Gardener's dictionary. Even
if the epithet officinalis had nomenclatural standing, it would be
invalid under Article 52.1 (see above), as it is homotypic with
Mandragora officinarum L.: in DeCandolle (in Lamarck & DeCan-
dolle, 1805) and Willdenow (1809), Atropa mandragora L. is cited
in synonymy.
Assumptions as to the phenological status of type specimens has
also led to confusion about the correct application of epithets (e.g.
Jackson & Berry, 1979). Bertoloni (1820, 1824, 1835) contributed
greatly to the confusion surrounding names for the Mediterranean
mandrake. In a list of plants cultivated in the Botanic Gardens at
Bologna (Bertoloni, 1820), he included two species of mandrakes:
Mandragora autumnalis, which he coined as a new name, and M.
officinalis, which he attributed to Willdenow. In a later work (Berto-
loni, 1824) he designated two different taxa: M. vernalis and M.
officinarum, explicitly citing Linnaeus as the source of the latter
epithet. He put M. officinalis in synonymy with M. vernalis and put
M. autumnalis in synonymy with M. officinarum. In Commentarius
de Mandragoris (Bertoloni, 1835), he revised the genus and recog-
nized three species: M. vernalis, M. officinarum andM. microcarpa.
In synonymy he left out both of the previously used epithets,
autumnalis and officinalis, which he may have considered to be
superfluous.
The lectotype of Mandragora officinarum, selected from amongst
the elements used by Linneaus in his description, bears no pheno-
logical information at all. The specimen in Burser's herbarium is
labelled 'Misnia, Lusatia', indicating it came from Germany. It is
probable therefore that it is from a botanical garden, and since plants
of Mandragora in northern European gardens usually bloom in
spring, it has been assumed that this one did as well. This is not
founded in fact, thus rendering much of the debate as to which name
is the vernal or autumnal taxon irrelevant.
2. Mandragora turcomanica Mizg. in Trudy Turkmen. Fil. Akad.
Nauk SSSR.Ashkabad2: 165 (1942). Type: Turkmenistan, west-
ern Kopet Dag, Kara Kala region, southern foot of Mt Syunt,
Schevlan valley, on scree slopes, 26 November 1938, O.G.
Mizgireva, M. Nastacalicz & G. Nastacalicz s.n. (ASH-holotype;
LE-isotype(?)).
ILLUSTRATIONS. Mizgireva (1942: figs 1, 2; 1955: figs 2-7, 9, 14,
15, 18, 19, 24); Belyanina (1982: figs 1, 2).
Plant virtually stemless. Leaves arranged in a rosette: lower leaves c.
90 x 60 cm, length/width ratio c. 1 .3-1 .5, broadly elliptical or ovate,
34
S. UNGRICHT, S. KNAPP AND J.R. PRESS
usually with large, irregular teeth in upper half, upper leaves smaller,
oblong-ovate or broadly lanceolate, usually without large teeth, on
both sides sparsely hairy, the hairs mostly arranged along veins.
Flowers borne l(-3) in leaf axils, occasionally the pedicels up to 2-
6-flowered. Pedicels 2-3 cm long, 7-18 cm in fruit. Calyx 15-20
mm long, lobes 10-15 mm, acuminate, accrescent in fruit. Corolla
20-25 mm long, campanulate, violet or purple, with three narrow
white stripes at base, the lobes c. 15 mm. Stamens c. 10 mm long,
attached to lower half of corolla; filaments 6-7 mm, densely pubes-
cent at the base and in the lower Vr, anthers c. 4 mm long, pale blue.
Style exceeding stamens; stigma capitate, slightly bilobed, green.
Fruit 40-60 mm in diameter, glossy yellow to orange when ripe,
pulpy, juicy, smelling of melons, edible. Seeds 4-5 x 6-7 mm,
reniform, yellow to light brown.
HABITAT. Shrubland in ravines and on stony mountain slopes,
especially among Paliurus spina-christi Mill. (Rhamnaceae).
Altitudinal range 500-700 m.
VERNACULAR NAME. Turkmenian mandrake.
PHENOLOGY. Flowering October to March. Fruiting until June.
Summer dormancy until autumn rains begin. Xerothermic
ephemeroid.
CONSERVATION STATUS. Only small populations in a restricted
area within the Syunt-Khassardagh Reserve, and in the valleys of
Chokhadgh, Shevlan, Altybai, Ekechinar, Sarymsakli and Dagdanli
are known (Kurbanov, 1994). According to Mizgireva (1978) the
population in the Syunt-Khassardagh reserve consists of fewer than
a thousand plants.
DISTRIBUTION. Southwestern Kopet Dag. (Turkmenistan, ?Iran).
SELECTED SPECIMENS EXAMINED. No specimens were available
for examination: the data here are drawn exclusively from the
literature.
In the original article in which this species was described (Mizgireva,
1942) the type is cited as being in Ashkabad (ASH). Subsequently,
in the Flora of the USSR (Lincevskij, 1955), the type was said to be
in Leningrad (now St. Petersburg, LE) and this citation has been
followed in other floristic works (Schonbeck-Temesy, 1972). Since
we have been unable to obtain specimens from either herbarium, it
is uncertain whether or not the holotype remains in ASH or has been
transfered to LE, thus we have cited its location as in the original
publication.
Mizgireva's (1955) detailed study of this species documented the
flowering times (Mizgireva, 1955: fig. 16) of individual plants and
clearly shows that some individuals flower in autumn (October and
November) while others flower in spring (February and early March).
This is the same situation as that in Mandragora officinarum, and is
markedly different from that found in much of the rest of the flora of
Kopet Dag (Mizgireva, 1978). Mizgireva (1955) stated that the
differences between her species, M. turcomanica, and the Mediter-
ranean mandrakes (separated by her into M. officinarum and M.
autumnalis) were largely size of plant, colour of anthers and relative
sizes of calyx and corolla. It is clear from her work that these
characteristics are consistent throughout the range of wild and
greenhouse grown material she examined, but since she cites no
specimens, nor have we been able to examine any of the known
specimens of M. turcomanica, an in-depth analysis of these charac-
ters relative to the range of variation found in M. officinarum awaits
future study.
3. Mandragora caulescens C.B. Clarke, Fl. Brit. Ind. 4: 242
(1883). Type: India, Sikkim, Lachen, Hooker s.n. (Kl-lectotype,
designated here by R. Mill).
Fig. 9.
= Anisodus mariae Pascher in Feddes Repert. Spec. Nov. 7: 227
(1909).
= Mairella yunnanensis H. Lev., Cat. pi. Yun-Nan: 199 (1916).
= Anisodus caulescens (C.B. Clarke) Diels in Feddes Repert. Spec.
Nov.Beih. 12:480(1922).
= Mandragora tibetica Grubov in Rast. Tsentral. Azii. 5: 108-109
(1970).
= Mandragora caulescens subsp. brevicalyx Grierson & D.G. Long
in Notes Roy. Bot. Card. Edinburgh 36: 144 (1978).
= Mandragora caulescens subsp. flavida Grierson & D.G. Long in
Notes Roy. Bot. Card. Edinburgh 36: 143 (1978).
= Mandragora caulescent subsp. purpurascens Grierson & D.G.
Long in Notes Roy. Bot. Card. Edinburgh 36: 143 (1978).
= Mandragora chinghaiensis Kuang & A.M. Lu, Fl. reip. pop. Sin.
67: 159 (1978).
ILLUSTRATIONS. Grubov (1970: pi. IV, fig. 2; pi. V, fig. 2); Kuang
& Lu (1978: fig. 36, 1-6). Hoare & Knapp (1997: fig. 5d).
Root stout, often branched. Stem present, max. 60 cm long. Leaves
arranged in a cluster on top of the young stem above scale leaves,
later along the secondary growth of the stem, max. 30 cm long,
length/width ratio 1.9-5.5, oblanceolate or spathulate, glabrous to
densely pubescent beneath, especially along the veins. Flowers
borne in leaf axils, often nodding, pleasantly scented. Pedicels 5-10
cm long. Calyx 5-30 mm long, often nearly as long as corolla, lobed
c. l/4 of the way to the base, accrescent in fruit. Corolla 5-30 mm
long, campanulate to cup-shaped, yellow to purple, the venation
usually darker, lobed about 1A of the way to the base, the lobes 2-10
mm. Stamens adnate in lower half of corolla; filaments 3-10 mm;
anthers 1-5 mm long, white to yellow. Style 3-6 mm long, not
exceeding the stamens; stigma capitate, somewhat bilobed. Ovary
2.5-5 mm in diameter, globose. Fruit 10-25 mm in diameter,
globose, pale greenish white or greenish yellow. Seeds 1.5-3.0 mm
long, numerous, reniform, the surface pitted, yellow to light brown.
HABITAT. Grassy and stony slopes, open moorland, pastures and
screes. Subalpine and alpine zone, especially among Rhododendron
shrubs. Altitudinal range 3000-4900 m.
VERNACULAR NAMES. Himalayan mandrake. China: qie shen.
PHENOLOGY. Flowering April to September. Fruiting August to
October.
CONSERVATION STATUS. Locally common. Not threatened.
DISTRIBUTION. Sino-Himalayan. (Nepal, India, Bhutan, Myanmar,
China).
SELECTED SPECIMENS EXAMINED. NEPAL. Topke Goia, 27°39'N,
87°35'E, 3810 m, in short grassland, 3 July 1971, Beer, LW. 8264 (BM);
Tarakot, 28°53'N, 82°59'E, 3900 m, 25 May 1969, Dobremez, J.F. 175 (BM);
Jangla Banyang, wet place, 3800 m, 5 June 1973, Einarsson, L. et al. 430
(BM); Ganesh Himalaya, Shior Khola, 4270 m, bouldery alpine meadow,
Gardner, P.C. 479 (BM); Topke Gola-Tasagon, 3600-4350 m, 17 August
1977, Ohashi, H. et al. 775165 (BM); Langtang Valley, 28°14'N, 85°32'E,
4420 m, among rocks in turf, June 1949, Polunin, O. 63 1 (BM); 5 miles E. of
Timure, oak wood, 41 10 m, 3 July 1949, Polunin, O. 780 (BM); pass NW of
Jumla, 6 miles Pansae Dara, 3050 m, growing in loose soil recently free of
snow, 30 April 1952, Polunin, O. et al. 878 (BM); Dozam Schola, near
Simikot, 3510 m, grass slopes, 2 June 1952, Polunin, O. et al. 4250 (BM);
near Jumla, 3810 m, open grassy slopes, 14 July 1952, Polunin, O. et al.
MANDRAGORA
35
X S
\U*iin I
I
36
S. UNGRICHT, S. KNAPP AND J.R. PRESS
4668 (BM); Arun-Tamur watershed, Topke Gola, 3960 m, 11 May 1956,
Stainton, J.D.A. 254 (BM); Balangra Pass, W. of Tibrikot, 29°06'N, 82°41'E,
3810 m, 28 May 1963, Stainton, J.D.A. 4263 (BM); Rolwaling, 27°55'N,
86°23'E, 3810 m, beneath small shrubs, 25 June 1964, Stainton, J.D.A. 4676
(BM);Khunde-Khumjung,3820m,6Junel952,Z//n/w^r/nann,A. 688 (BM).
INDIA. Sikkim: Changu, 3960 m, in crevices of rocks, 28 June 1913,
Cooper, R.E. 80 (E); Gnatong, 3960 m, peaty marsh, 31 August 1913,
Cooper, R.E. 759 (E); Jongri, 4000-4200 m, 22 May 1960, Hara, H. et al.
443 (BM, K); Tungu, 3660-3960 m, 23 July 1849, Hooker, J.D. s.n. (K);
Jongri, 4420 m, campsite towards Yakhut amongst Juniperus scrubs, 29
May 1990, Kirkpatrick, G. 53 (E); District Darjeeling, Phalut, 3350-3660
m, 30 May 1902, Lace, J.H. 2266 (E); West District, Bikbari, Choktsering
Chhu valley; 27°30'53"N, 88°08'28"E, 3950 m, amongst scrub, 12 July
1992, Long, D.G. et al. 266 (E); Chulong, 4570 m, 12 September 1912,
Rhomoo, L 210 (E); Gamothang, 3960 m, 12 August 1913, Rhomoo, L.
1059 (E); Giagong, 3960 m, 6 September 191 1, Ribu & Rhomoo 5492 (BM,
E, K); Phalut, 3350-3660 m, 27 May 1911, Smith, W.W. 4593 (E); Jongri,
4000 m, in shelter of dwarf Rhododendron and Juniperus, 23 June 1983,
Starling, B.N. et al. 92 (K); Tangshing campsite, 4000 m, open grazed areas
among Rhododendron and around campsite, 24 June 1983, Starling, B.N. et
al. 106 (K); summit of Sandakphu, 3660 m, small dense tufts amongst
grass, 18 May 1881, Watt, G. 5373 (E, K). Arunachal Pradesh (Assam):
Orka La, Bhutan frontier, 3960 m, on stony slopes, scattered, 7 June 1938,
Kingdon Ward, E 13840 (BM).
BHUTAN. Paco Chu, 4110m, loose sandy crevice, 15 May 1966, Bowes
Lyon, S. 3 156 (BM); Phajudin Timpu, 3960 m, 5 August 1914, Cooper, R.E.
3232 (BM, E); Tare-La area, 3660 m, 19 May 1938, Gould, B.J. 109 (K);
Laum Thang, 3900 m, in alpine herbage with moderate humidity, 18 May
1967, Hara, H. et al. 1 1973 (BM); Western Bhutan, 3510 m, among boulders
above tree line, 17 June 1933, Ludlow, F. & Sheriff, G. 107 (BM); Byasu La,
4110m, growing among stones on open hillside, 18 May 1937, Ludlow, F. &
Sherriff, G. 3073 (BM); Cheli La, 3810 m in open moorland, 4 May 1949,
Ludlow, F. et al. 16139 (BM); Pangotang, 3960 m, open hillside above fir
zone, 27 May 1949, Ludlow, Eel al. 18986(BM); Kantanang,Tsampa,4110
m, among shrubs on steep hillside, 3 June 1949, Ludlow, Eetal. 19040(BM);
Shingbe, Me La, 3350 m, growing amongst other herbs and small shrubs, 16
May 1949, Ludlow, F. et al. 20645 (BM, E).
MYANMAR (Burma). Chenochi Pass, 3600 m, 3 July 1920, Farrer, J.R.
1694 (E).
CHINA. Yunnan: Diqing Prefecture, E. slope of Bai Ma Shan, 28°19'N,
99°05'E, 3895 m, among stones in Abies forest, 9 June 1993, Alden, B. et al.
1021 (E); Diqing Prefecture, Zhongdian County, above Napa Hai, N. of
Zhongdian, 27°55'N, 99°34'E, 4000 m, forested ridge, snow-bed vegetation
and dwarf Rhododendron scrub, 12 June 1993, Alden, B. et al. 1185 (E);
western flank of the Lichiang range, 27°20'N, 3050-3350 m, open alpine
pasture, June 1910, Forrest, G. 5999 (E, K); Lichiang range, 27°25'N,
3660-3960 m, July 1913, Forrest, G. 10474 (BM, E, K); western slopes of
Likiang snow range, Yangtze watershed, 30 May-6 June 1922, Rock, J.F.
4211 (BM); Mo-ting shan, ME of Atuntze, eastern slopes of Yangtze-
Mekong divide, 3660 m, along trail, June 1932, Rock, J.F. 22771 (E); slopes
of Mt Gyi-na lo-ko, the second peak of the Yu-lung shan, 3660 m, in alpine
meadows, April-May 1932, Rock, J.F. 25005 (BM, E); Chungtien, Chiren,
2200 m, grass slope, 17 May 1937, Yu, T.T. 11327 (KUN); Mekong-
Salween divide, Chingpoh, 3800 m, alpine pasture lands, 14 June 1938, Yu,
T.T. 19078 (E); Upper Kiukiang valley, Clulung Lungtsahmuru, 3700 m,
mountain slope, open and grassy place, 9 August 1938, Yu, T.T. 19843 (E,
KUN). Sichuan: Tien-chuan-hsien, 3500 m, 14 June 1936, Chu, K.L. 2791
(BM); mountains E. of Yungning, 27°48'N, 101° E, 3660 m, open stony
pasture, June 1922, Forrest, G. 21407 (BM, E, K); mountains around Muli,
28°12'N, 101° E, 3660 m, July 1930, Forrest, G. 28416 (BM, E); Liuku-
liangdse, 27°48'N, 3700-4200 m, 18 May 1914, Handel-Mazetti, H. von
2349 (K); Minya Konka snow range, S. of Tatsienlu, 4450 m, in meadows,
July 1929, Rock, J.F. 17530 (E); Mu-li-ka-bu, 3600 m, mountain slope,
grassy place, 18 May 1937, Yu, T.T. 5496 (KUN). Qinghai: Dari (Darlag)
Xian, just N. of Manzhang, along the Manzhang He, between Dari and
Banma, 33°17'51"N, 100°25'55"E, 4000 m, on moist rodent disturbed
slope, 12 August 1993, Ho, T.N. et al. 1 172 (E); Chindu Xian, Xiwu Xiang,
E. of pass on road between Xiwu and the Szechwan border, 33°09'14"N,
97°32'35"E, 4250 m. damp slope with extensive pika warrens, 1 1 Septem-
ber 1996, Ho, T.N. et al. 3063 (BM). Xizang: Dotha, 3960 m, 20 June 1945,
Bar & Kirat Ram 20511 (K); Mt. Demula, Yiwei County, 4100 m, 23
August 1973, Chinese Tibet Expedition 1973 1214 (KUN); District
Chengtang, Dingjie County, Ladang to Xingeng, 3800-4000 m, 9 June
1975, Chinese Tibet Expedition 1975 5679 (KUN); Londre Pass, Tsarong,
Mekong-Salween divide, 28°14'N, 98°40'E, 3960 m, July 1921, Forrest, G.
19630 (UC/JEPS, P, K); Tha Chu valley, 3660 m, in clumps, alpine region,
in stony pastures and on grassy slopes, 11 July 1950, Kingdon Ward, F.
19613 (BM); N. of Sanga Choling, 28°41'N, 93°02'E, 4110 m, on open
grassy bank, 14 May 1936, Ludlow, E & Sherriff, G. 1590 (BM); hills S. of
Lhasa, 4420 m, ground left bare by melting snow, 6 June 1942, Ludlow, F.
& Sherriff, G. 8676 (BM); hills S. of Lhasa, 4720 m, herb growing under
boulders, 21 August 1942, Ludlow, F. & Sherriff, G. 9014 (BM); Pome,
above Showa Dzong, 29°55'N, 95°25'E, 3350 m, open hillsides midst grass,
1 1 June 1947, Ludlow, F. et al. 13149 (BM, E); Sobhe La, near Tongyuk
Dzong, Pome, 30°07'N, 94°54'E, 3510 m, 21 May 1947, Ludlow, F. et al.
13750 (BM, E); Province Kongbo, Nyima La, 29°38'N, 94°52'E, 3350 m,
on dry ground under trees, 21 May 1947, Ludlow, F. et al. 15051 (BM, E);
Lo La, Pachakshiri, 28°49'N, 93°59'E, 3810 m, in openings of Rhododen-
dron forest, mostly above Abies zone, 15 May 1938, Ludlow, F. et al. 3776
(BM); Province Kongbo, south side of Lusha Chu, 29°20'N, 94°35'E, 3810
m, in Lonicera and Berberis scrub, 13 June 1938, Ludlow, F. et al. 4591 A
(BM, E); valley above Tse, Tsangpo valley, 29°23'N, 94°22'E, 3350 m, in
grassy banks under Rhododendron, Ludlow, F. et al. 4591 (BM).
The status and identity of the previously proposed subspecies of
Mandragora caulescens (Grierson & Long, 1978) are discussed in
the section on the species delimitation (p. 25). It is clear from our
studies that M. caulescens is extremely variable and that a detailed
field-based study over the entire species range is necessary.
ACKNOWLEDGEMENTS. This study was undertaken as partial fulfilment of
the requirements for an M.Sc. in Advanced Methods in Taxonomy and
Biodiversity at Imperial College, London and The Natural History Museum,
London undertaken by S. Ungricht and supervised by S. Knapp and J.R.
Press. The work has profited from discussions with many members of the
staff at The Natural History Museum, particularly S. Cafferty, N.C. Garwood,
CJ. Humphries, C.E. Jarvis, C. Moncrieff, N.K.B. Robson, K. Sidwell, A.R.
Vickery, D.M.Williams, and C.Whitefoord. We are grateful to Z. Frankiel, K.
Rottcher and Zhan Ren-bin for their translation of the crucial Russian and
Chinese papers respectively. Furthermore, we are indebted to M. Gilbert and
R.R. Mill for their sharing of personal observations and to N. Turland for a
thoughtful and detailed review. Thanks are also due to R. Baldini in Florence
and the directors and staff of the following herbaria who made their collec-
tions available: E, JEPS, K, KUN, P, PAL, RNG and UC. This study has been
supported by a grant from the Park Fund of the Botany Department of The
Natural History Museum, London.
REFERENCES
\ iimi-iil Franco, J. do 1984. Nova flora de Portugal 2. Lisbon.
Anderson, E. 1941. The technique and use of mass collections in plant taxonomy.
Annals of the Missouri Botanical Garden 28: 287-292.
& Ttirrill, W.B. 1935. Biometrical studies on herbarium material. Nature 136:
986.
Baehni, C. 1946. L'ouverture du bouton chez les fleurs de Solanees. Candollea 10:
400-492.
Bateman, R.M. 1989. On measuring Marsh-Orchids — morphometric procedure, taxo-
nomic objectivity and Marsh-Orchid systematics. Watsonia 17: 449-462.
Bauhin, C. 1623. Pinax theatri botanici. Basel.
Bauhin, J., Cherler, J.H. & Chabrey, D. 1650-1651. Historia plantarum. 1-3.
Yverdon.
Baytop, A. 1978 ['1979']. Mandragora. In P.H. Davis (Ed.), Flora of Turkey and the
East Aegean Islands 6: 450—45 1 .
Belyanina, N.B. 1982. K anatomo-morfologicheskoi strukture vegetativnykh organov
mandragory turkmenskoi. Byulleten Glavnogo Botanicheskogo Sada (Moskow) 124:
63-69.
MANDRAGORA
37
Bentham, G. 1876. Solanaceae. In G. Bentham & J.D. Hooker (Eds), Genera plantarum
2: 882-913.
Bertoloni, A. 1 820. Elenchus plantarum vivarum. Bologna.
1824. Viridarii Bononensis vegetabilia. Bologna.
1835. Commentarius de mandragoris. Bologna.
Bohs, L. 1994. Cyphomandra (Solanaceae). Flora Neotropica 63: 1-175.
Bonafe Barcelo, F. 1980. Flora de Mallorca 4. Mallorca.
Borg, J. 1927. Descriptive flora of the Maltese Islands including the ferns and
flowering plants. Malta.
Boulos, L. 1995. Flora of Egypt checklist. Cairo.
Bouquet, J. 1952. La Mandragore en Afrique du Nord. Bulletin de la Societe des
Sciences Naturelles de Tunisie 5: 29-44.
Cain, S.A. 1944. Foundations of plant geography. New York.
Cellinese, N., Jarvis, C.E., Press, J.R., Short, M.J. & Viciani, D. (Eds) 1994.
Threatened plants of Italy: interim report. London.
Chappill, J.A. 1989. Quantitative characters in phylogenetic analysis. Cladistics 5:
217-234.
Clarke, C.B. 1 883. Mandragora. In J.D. Hooker, The flora of British India 4: 241-242.
Clausen, J., Keck, D.D. & Heisey, W.M. 1940. Experimental studies on the nature of
species. I. The effect of varied environments on western North American plants.
Publications of the Carnegie Institute of Washington 520: 1-452.
Crisp, M.D. & Weston, P.H. 1993. Geographic and ontogenetic variation in morphol-
ogy of Australian Waratahs (Telopea: Proteaceae). Systematic Biology 42: 49-76.
Davis, J.I. & Nixon, K.C. 1992. Populations, genetic variation, and the delimitation of
phylogenetic species. Systematic Biology 41: 421-435.
Deb, D.B. 1979. Solanaceae in India. In J.G. Hawkes, R.N. Lester & A.D. Skelding
(Eds), The biology and taxonomy of Solanaceae: 87-1 12. London.
Demiri, M. 1983. Flora ekskursioniste e Shqiperise. Tirana.
Dercourt, J., Zonenshain, L.P., Ricou, L.-E., Kazmin, V.G., Le Pichon, X., Knipper,
A.L., Grandjacquet, C., Sbortshikov, I.M., Geyssant, J., Lepvrier, C., Pechersky,
D.H., Boulin, J., Sibuet, J.-C., Savostin, L.A., Sorokhtin, O., Westphal, M.,
Bazhenov, M.L., Lauer, J.P. & Biju-Duval, B. 1986. Geological evolution of the
Tethys belt from the Atlantic to the Pamir since the Lias. Tectonophysics 123: 241-
315.
Duke, J.A. 1985. Handbook of medicinal herbs. Boca Raton.
Dunal, M.F. 1852. Solanaceae. In A.P. de Candolle (Ed.), Prodromus systematis
naturalis regni vegetabilis 13(1): 546-556.
Du Rietz, G.E. 1930. The fundamental units of biological taxonomy. Svensk Botanisk
Tidskrift 24: 333^28.
Eldredge, N. 1979. Cladism and common sense. In 3. Cracraft & N. Eldredge (Eds),
Phylogenetic analysis and palaeontology: 165-198. New York.
Evans, W.C. 1979. Tropane alkaloids of the Solanaceae. In J.G. Hawkes, R.N. Lester
& A.D. Skelding (Eds), The biology and taxonomy of Solanaceae: 241-254.
London.
Farris, J.S. 1988. //£AW/G 86: version 1.5. Published by the author, Jamaica Plains, NY.
1989. The retention index and the rescaled consistency index. Cladistics 5: 417-
419.
Feinbrun-Dothan, N. 1978. Mandragora. In N. Feinbrun-Dothan, Flora Palestina 3:
167.
Fischer, C.E.C. 1934. Mandragora shebbearei. Bulletin of miscellaneous information,
Royal Botanic Gardens, Kew, England 1934: 260-261.
Fisher, N.I. 1993. Statistical analysis of circular data. Cambridge.
Fleisher, A. & Fleisher, Z. 1 994. The fragrance of biblical mandrake. Economic Botany
48: 243-251.
Fournier, P. 1990. Les quatre flares de France. Paris.
Gibson, A.R., Baker, A.J. & Moeed, A. 1984. Morphometric variation in introduced
populations of the Common Myna (Acridotheres tristis): an application of the
jacknife to principal component analysis. Systematic Zoology 33: 408-421.
Greuter, W. & Rechinger, K.H. 1967. Flora der Insel Kythera. Boissiera 13: 1-206.
Grierson, A.J.C. & Long, D.G. 1978. Notes relating to the flora of Bhutan: I. Notes
from the Royal Botanic Garden, Edinburgh 36: 139-150.
Grubov, V.I. 1970. Mandragora. In V.I. Grubov (Ed.), Plantae asiae centralis 5: 108-
109.
Halacsy, E. de 1902. Conspectus florae graecae 2. Leipzig.
Hallam, A. 1981. Relative importance of plate movements, eustasy, and climate in
controlling major biogeographical changes since the early Mesozoic. In G. Nelson &
D.E. Rosen (Eds), Vwariance biogeography—a critique: 303-340. New York.
1994. An outline ofphanerozoic biogeography. Oxford.
Hansen, A. & Sunding, P. 1993. Flora of Macaronesia: checklist of vascular plants.
4th ed. Oslo.
Hara, H. 1966. The flora of eastern Himalaya. Tokyo.
1971. The flora of eastern Himalaya: second report. Bulletin of the University
Museum of the University of Tokyo 2.
Hawkes, J.G. 1972«. Mandragora. In T.G. Tutin, V.H. Heywood, N.A. Burges, D.M.
Moore, D.H. Valentine, S.M. Walters & D.A. Webb (Eds), Flora Europaea 3: 199-
200.
1972fo. Solanaceae. In V.H. Heywood (Ed.), Flora Europaea. Notulae Systematicae,
13. Botanical Journal of the Linnean Society 65: 356-357.
Hayek, A. & Markgraf, F. 1931. Prodromus florae peninsulae balcanicae2. Dahlem.
Hegnauer, R. 1973. Chemotaxonomie der Pflanzen 6. Basel.
Heiser, C.B. 1969. Nightshades: the paradoxical plants. San Francisco.
Heldreich, T. von 1 886. Bemerkungen iiber die Gattung Mandragora. Mitteilungen der
Geographischen Gesellschaft Jena 4: 75-80.
Hepper, F.N. 1990. Pharaoh's flowers: the botanical treasures of Tutankhamun.
London.
Hoare, A.L. & Knapp, S. 1 997. A phylogenetic conspectus of the tribe Hyoscyameae
(Solanaceae). Bulletin of the Natural History Museum London (Botany Series) 27:
11-29.
Humphries, CJ. & Parent!, L.R. 1986. Cladistic biogeography. Oxford.
Hunziker, A.T. 1995. Studies on Solanaceae XXXVIII. Miscellaneous novelties on the
taxonomy of Solanaceae. Lorentzia 8: 5-8.
Jackson, B.P. & Berry, M.I. 1979. Mandragora — taxonomy and chemistry of the
European species. In J.G. Hawkes, R.N. Lester & A.D. Skelding (Eds), The biology
and taxonomy of Solanaceae: 505-512. London.
Jahandiez, E. & Mai re, R. 1934. Catalogue des plantes du Maroc. Algiers.
James, F.C. & McCulloch, C.E. 1990. Multivariate analysis in ecology and systemat-
ics: Panacea or Pandora's box? Annual Review of Ecology and Systematics 21:
129-166.
Jarvis, C.E., Barrie, F.R., Allan, D.M. & Reveal, J.L. 1 993. A list of Linnaean generic
names and their types. Regnum Vegetabile 127: 1-100.
Jussieu, A.-L. de 1789. Genera plantarum. Paris.
Kessler, J. 1951. Uber einen Vergiftungsfall durch Radix Mandragorae. Berichte der
Schweizerischen Botanischen Gesellschaft 61: 405—409.
Khlopin, I.N. 1979. Mandragora turkmenskaya v istorii narodov vostoka. Izvestija
Akademii Nauk Turkmenskoj SSR, Biologicheskikh Nauk 1979(1): 70-75.
Kuang Ko-zen & Lu An-ming. 1978. Mandragora. In Ko-zen Kuang & An-ming Lu
(Eds), Flora Reipublicae Popularis Sinicae 67(2): 137-141, 159-160.
Kurbanov, D. 1994. Flora of Kopetdagh. In V. Fet & K.I. Atamuradov (Eds), Biogeog-
raphy and ecology of Turkmenistan: 105-126. Dordrecht.
Lamarck, J.B.A.P.M. de & DeCandolle, A.P. de 1805. Flore francaise 3. Paris.
Lazaro e Ibiza, B. 1907. Compendia de la flora Espahola 2. Madrid.
Lincevskij, LA. 1955. Mandragora. In K. Shishkin & E.G. Bobov (Eds), Flora URSS
22: 75-77.
Linnaeus, C. 1738. Hortus cliff ortianus. Amsterdam.
1753. Species plantarum. Stockholm.
1759. Systema naturae. 10th ed. Stockholm.
1762. Species plantarum. 2nd ed., 1. Stockholm.
Lu An-ming & Zhang Zhi-yu. 1986. Studies of the subtribe Hyoscyaminae in China.
In W.G. D'Arcy (Ed.), Solanaceae: biology and systematic s: 56-85. New York.
Marzell, H. 1975. Solanaceae. In G. Hegi, Illustrierte Flora von Mitteleuropa 5(4):
2548-2625.
Maugini, E. 1959. Ricerche sul genere Mandragora. Nuovo Giomale Botanico
Italiano e Bolletino delta Societa Botanica Italiana (n. s.) 66: 34-60.
Mehra, K.L. 1979. Ethnobotany of Old World Solanaceae. In J.G. Hawkes, R.N. Lester
& A.D. Skelding (Eds), The biology and taxonomy of Solanaceae: 1 6 1 - 1 70. London.
Meikle, R.D. 1985. Mandragora. In R.D. Meikle, Flora of Cyprus 2: 1 188-1 190.
Migahid, A.M. 1989. Flora of Saudi Arabia. 3rd ed. Riyadh.
Miles, D.B. & Ricklefs, R.E. 1984. The correlation between ecology and morphology
in deciduous forest passerine birds. Ecology 65: 1629-1640.
Miller, P. 1760. Figures of plants. London.
1768. The gardener's dictionary, 8th ed. London.
& T. Martyn. 1807. The gardner's and botanist's dictionary, 9th ed. London.
Mizgireva, O.F. 1942. A new species of Mandragora in Turkmenia. Trudy Turkmens-
kogo Filiala Akademii Nauk SSSR. Ashkabad. 2: 165.
1 955 . Mandragora turkmenskaya (Mandragora turcomanica).Problemy botaniki,
Moskva-Leningrad, Izd-Vo., Akademii Nauk SSSR 2: 165-205.
1978. Mandragora turkmenskaya (Mandragora turkcomanica Mizgir.). Izvestija
Akademii Nauk Turkmenskoj SSR, Biologicheskikh Nauk 1978(4): 54-55.
Moldenke, H.N. & Moldenke, A.L. 1952. Plants of the Bible. New York.
Mouterde, P. 1 979. Mandragora. In P. Mouterde, Nouvelleflore du Liban et de la Syrie
3: 196.
Nelson, G. & Platnick, N.I. 1981. Systematics and biogeography: cladistics and
vicariance. New York.
Nixon, K.C. & Wheeler, Q.D. 1990. An amplification of the phylogenetic species
concept. Cladistics 6: 211-223.
Olmstead, R.C. & Palmer, J.D. 1992. A chloroplast DNA phylogeny of the Solanaceae:
subfamilial relationships and the character evolution. Annals of the Missouri Botani-
cal Garden 79: 346-360.
& Sweere, J.A. 1994. Combining data in phylogenetic Systematics: an empirical
approach using three molecular data sets in the Solanaceae. Systematic Biology 43:
467-481.
Spangler, R.E., Bohs, L. & Palmer, J.D. [In press]. Phylogeny and
provisional classification of the Solanaceae based on chloroplast DNA In D.E.
Symon & M. Nee (Eds), Solanaceae IV. Kew.
Parkinson, J. 1629. Parodist in sole paradisus terrestris. [Reprinted 1904 in London]
Pignatti, S. 1982. Mandragora. In S. Pignatti, Flora d'ltalia 2: 518-519.
38
S. UNGRICHT, S. KNAPP AND J.R. PRESS
Pojarkova, A.I. 1955. Solanaceae. In K. Shishkin & E. G. Bobov (Eds), Flora URSS
22: 1-117. [Translated 1994]
Post, G.E. 1933. Flora of Syria, Palestine and Sinai. 2nd ed. Beirut.
Pottier-Alapetite, G. 1981. Flore de la Tunisie. Tunis.
Press, J.R. & Short, MJ. (Eds), Flora of Madeira. London.
Proskuryakova, G.M. & Belyanina, N.B. 1985. K sistematike roda Mandragora i ego
istorii. Biologicheskie Nauki (Moskow) 2(254): 76-86.
Quezel, P. & Santa, S. 1963. Nouvelle flore de I'Algerie. Paris.
Randolph, C.B. 1905. The Mandragora of the Ancients in folk-lore and medicine.
Proceedings of the American Academy of Arts and Sciences 40: 487-537.
Rechinger, K.H. 1964. Flora of lowland Iraq. Weinheim.
1972. Mandragora. In E. Schonbeck-Temesy (Ed.), Flora Iranica 100: 43-44.
Roland, J.-D. 1991. La mandragore: le mythe d'une racine, la racine d'un mythe.
Annales des Sciences Naturelle, Botanique 11: 49-81.
Romeike, A. 1978. Tropane alkaloids — occurence and systematic importance in
angiosperms. Botaniska Notiser 131: 85-96.
K IK Id i ma 11. W.F., Prell, W.L. & Raymo, M.E. 1989. History of late Cenozoic uplift in
south-east Asia and the American southwest: rationale for general circulation
modelling experiments. Journal of Geophysical Research 94: 18379-18391.
Schuster, R.M. 1976. Plate tectonics and its bearing on the geographical origin and
dispersal of angiosperms. In C.B. Beck (Ed.), Origin and early evolution of
angiosperms: 48-138. New York.
Scotese, C.R., Gahagan, L.M. & Larson, R.L. 1988. Plate tectonic reconstructions of
the cretaceous and cenozoic ocean basins. Tectonophysics 155: 27-48.
Siddiqi, M. 1978. Solanaceae. In S.M.H. Jafri & A. El-Gadi (Eds), Flora of Libya 62:
1-38.
Smith, A.G., Smith, D.G. & Funnell, B.M. 1994. Atlas of Mesozoic and Cenozoic
coastlines. Cambridge.
Sprengel, K. (Ed.) 1825. Systema vegetabilium. 16th ed. Gottingen.
Staub, H. 1962. Uber die chemischen Bestandteile der Mandragorawurzel. Helvetica
Chimica Acta 45: 2297-2305.
Stevens, P.F. 1991. Character states, morphological variation, and phylogenetic analy-
sis: a review. Systematic Botany 16: 553-583.
Tackholm, V. 1974. Students 'flora of Egypt. Cairo.
Talalaj, S., Talalaj, D. & Talalaj, J. 1992. The strangest plants in the world. London.
Tercinet, L. 1950. Mandragore — Qui-es-tu? Paris.
Tetenyi, P. 1987. A chemotaxonomic classification of the Solanaceae. Annals of the
Missouri Botanical Garden 74: 600-608.
Thiele, K. 1993. The holy grail of the perfect character: the cladistic treatment of
morphometric data. Cladistics 9: 275-304.
Thompson, C.J.S. 1934. The mystic mandrake. London.
Tournefort, J.P. de 1719. Institutiones rei herbariae. Paris.
Valdes, B., Talavera, S. & Fernandez-Galiano, E. (Eds). 1987. Flora vascular de
Andalucia occidental 2. Barcelona.
Vierhapper, F. 1 9 1 5. Beitrage zur Kenntnis der Flora Gretas. Osterreichische Botanische
Zeitschrift 65: 124-138.
Viney, D.E. 1994. An illustrated flora of north Cyprus. Konigstein.
Walter, H. 1 962, Die Vegetationszonen der Erde in okologischer Betrachtung 1, 2. Jena.
1966. Vegetation of the earth. London.
1977. Vegetationszonen und Klima. 3rd ed. Stuttgart.
Harnickell, E. & Mueller-Dombois, D. 1975. Climate-diagram maps. Berlin.
Weinert, E. 1972. Zur Taxonomie und Chorologie der Gattung Scopolia Jacq. Feddes
Repertorium 82: 617-628.
Wettstein, R. von 1895 [printed in 1891]. In A. Engler & K. Prantl (Eds), Die
natiirlichen Pflanzenfamilien 4(3b): 4-38.
Wilkinson, L. 1992. SYSTAT. Version 5.2. Evanston.
Willdenow, C.L von 1809. Enumeratio plantarum. Berlin.
Xiao Pei-gen & He Li-yi. 1983. Ethnopharmacologic investigation on tropane-contain-
ing drugs in Chinese Solanaceous plants. Journal of Ethnopharmacology 8: 1-18.
Zhang Zhi-yun & Lu An-ming. 1994. Mandragora. In Wang Wen-tsai (Editor-in-
chief), Vascular plants of the Hengduan mountains 2: 1754-1755.
& D'Arcy, W.G. 1994. Mandragora. In Zheng-yi Wu & P.H. Raven (Co-
chairs of the editorial committee), Flora of China 17: 329.
EXSICCATAE
Arranged by collector, with collection numbers (s.n.: collections
without collection number), species and herbaria.
Al-Eisawi, D. & Al-Jallad, W. 2221, M. officinarum (BM)
Alden, B., Alexander, J.C.M., Long, D.G., McBeath, R.J.D., Noltie, H.J. &
Watson, M.E 1021, M. caulescens (E); 1185, M. caulescens (E)
Atchley, S.C. s.n., M. officinarum (BM); 315, M. officinarum (K); 819, M.
officinarum (K)
Atherton, G.E. 793, M. officinarum (K); 812, M. officinarum (K); 998 M.
officinarum (K)
Baker, E.W. 149, M. officinarum (BM)
Ball, J. s.n., M. officinarum (E)
Barclay, C. 3004, M. officinarum (K)
Bauer, K. & Spitzenberger, F. 53, M. officinarum (BM)
Baytop, A., Tanker, N. & Sezik, E. 8504, M. officinarum (E)
Beer, L.W. 8264, M. caulescens (BM)
Bianor & Sabasien 1877 , M. officinarum (BM, E, P, RNG)
Bickerich, G. 15324, M. officinarum (BM, K)
Biro, L. s.n., M. officinarum (K)
Bar & Kirat Ram 20481, M. caulescens (K); 2051 1, M. caulescens (K)
Bornmiiller, J. s.n., M. officinarum (K)
Bourgeau, E. s.n., M. officinarum (K, P), 2304, M. officinarum (E, K, P)
Bowen, H.J.M. 694, M. officinarum (RNG); 3088, M. officinarum (RNG)
Bowes Lyon, S. 3156, M. caulescens (BM); 3239, M. caulescens (BM)
Brickell, C. & Mathew, B. 8088, M. officinarum (K)
Carr, J.W. s.n., M. officinarum (RNG)
Casey, E. C. 22 1 , M. officinarum (K)
Chaboisseau, C. 575, M. officinarum (BM)
Chatterjee, D. 75, M. caulescens (BM)
Chinese Tibet Expedition 1973 1214, M. caulescens (KUN)
Chinese Tibet Expedition 1975 5679, M. caulescens (KUN)
Chu, K.L. 2791, M. caulescens (BM)
Clarke, C.B. 34977, M. caulescens (K); 34983 A, M. caulescens (BM);
35704, M. caulescens (K)
Clarke, E.D. s.n., M. officinarum (BM)
Coincy, M.A. de s.n., M. officinarum (P)
Cooper, R.E. 80, M. caulescens (E); 759, M. caulescens (E); 3232, M.
caulescens (BM, E)
Danin, A. & Knees, S.G. 1709, M. officinarum (RNG)
Davis, P.H. 1 100, M. officinarum (E, K); 1438, M. officinarum (K); 2108,
M. officinarum (E, K); 4166, M. officinarum (E, K); 40134, M.
officinarum (E); 40182, M. officinarum (RNG); 40478, M. officinarum
(K)
Davis, P.H. & Polunin, O. 25469, M. officinarum (BM, E, K)
Delavay, J.M. s.n., M. caulescens (P); 263, M. caulescens (P); 1026, M.
caulescens (P)
Dhwoj, L 513, M. caulescens (BM, E)
Dinsmore, J.E. B 377, M. officinarum (E)
Dobremez, J.F. 175, M. caulescens (BM)
Duff our, C. 1501, M. officinarum (P)
Economides, S. 1055, M. officinarum (K)
Einarsson, L, Skarby, L. & Wetterhall, B. 430, M. caulescens (BM)
Espinosa, A. 870252, M. officinarum (K)
Ex Herb. Postian. apud Colleg. Syriens. Protest. 952, M. officinarum (BM)
Farrer, J.R. 1694, M. caulescens (E)
Fay, J.M. 815, M. officinarum (K); 864, M. officinarum (K)
Forbes s.n., M. officinarum (UC/JEPS)
Forrest, G. 569, M. caulescens (E); 5999, M. caulescens (E, K); 10474, M.
caulescens (BM, E, K); 19630, M. caulescens (UC/JEPS, P, K); 21407,
M. caulescens (BM, E, K); 26951, M. caulescens (P); 28416, M.
caulescens (BM, E)
Galiano, E.F. & Valdes, B. 820, M. officinarum (E, RNG)
Gammie, G.A. s.n., M. caulescens (BM, E)
Gandoger, M. s.n., M. officinarum (K); 84, M. officinarum (K); 8400, M.
officinarum (K)
Gardner, M.E, Jury, S.L. & Rejdali, M. 4885, M. officinarum (E, RNG)
Gardner, M.F. & Knees, S.G. 4769, M. officinarum (RNG)
Gardner, PC. 479, M. caulescens (BM)
Garnett, C.S. 33/5, M. officinarum (BM); 158/7, M. officinarum (BM)
Gathorne-Hardy, E. 14, M. officinarum (E); 139, M. officinarum (E); 194,
M. officinarum (E)
Girgenti s.n., M. officinarum (BM)
Gould, B.J. 109, M. caulescens (K)
Greuter, W. 7821, M. officinarum (E)
Guiol, F. 1731, M. officinarum (BM)
Haig, E. s.n., M. officinarum (BM)
Handel-Mazetti, H. von 2349, M. caulescens (K)
MANDRAGORA
39
Hara, H., Kanai, H., Murata, G. Ohashi, H., Tanaka, O. & Yamazaki, T.
1 1973, M. caulescent (BM)
Hara, H., Kanai, H., Murata, G., Togashi, M. & Tuyama, T. 443, M.
caulescens (BM, K)
Hardy, E. s.n., M. officinarum (BM)
Hart, H.C. s.n., M. officinarum (BM)
Hay, J.H. 1707, M. officinarum (K); H. 2484, M. officinarum (K)
Helbaek, H. 74, M. officinarum (K)
Heldreich, T. von s.n., M. officinarum (BM, E, K, P); 257 [1860], M.
officinarum (BM); 257 [1857], M. officinarum (P); 257 [1859], M.
officinarum (K); 2817 [1853], M. officinarum (P); 2817 [1860], M.
officinarum (K, P)
Higgins, E.K. s.n., M. officinarum (BM)
Ho, T.N., Bartholomew, B. & Gilbert, M. 1 172, M. caulescens (E)
Ho, T.N., Bartholomew, B., Watson, M. & Gilbert, M. 3063, M. caulescens
(BM)
Hooker, J.D. s.n., M. caulescens (E, K-lectotype, P)
Huguenin s.n., M. officinarum (BM)
/wry, S.L., Achhal, A. & Mouks, H. & Upson, T.M. 12634, M. officinarum
(RNG); 12653, M. officinarum (RNG)
Jury, S.L., Aitlafkih, M., Hedderson, T. & Kahouadji, A. 15237, M.
officinarum (RNG)
Jury, S.L., Aitlafkih, M., Hedderson, T. & Rutherford, R. W. 15398, M.
officinarum (RNG); 15434, M. officinarum (RNG); 15701, M.
officinarum (RNG)
Jury, S.L. & Jury, L.C. 13111, M. officinarum (BM, RNG)
Jury, S.L, Rejdali, M., Taleb, A. & Upson, T.M. 12542, M. officinarum
(RNG)
Jury, S.L, Taleb, A. & Upson, T.M. 12217, M. officinarum (RNG); 12238,
M. officinarum (RNG)
Jussieu, A. de s.n., M. officinarum (P)
King, G. s.n., M. caulescens (E); 4368, M. caulescens (P)
Kingdon Ward, F. 13840, M. caulescens (BM); 19613, M. caulescens (BM)
Kirkpatrick, G. 53, M. caulescens (E)
Lacaita, C.C. s.n., M. officinarum (BM); s.n., M. caulescens (BM)
Lace, J.H. 2266, M. caulescens (E)
Ladero, M., Lopez Guadalupe, M., Molero, J. & Perez Raya, F. 10605, M.
officinarum (RNG)
Lange, J. s.n., M. officinarum (K, P)
Laukkonen, P. 55, M. officinarum (K)
Le Testu, G. s.n., M. officinarum (BM)
Lewalle, J. 10054, M. officinarum (BM, RNG)
Lojacono-Pojero, M. s.n., M. officinarum (BM)
Long, D.G., McBeath, R.J.D., Noltie, H.J. & Watson, M.F. 266, M.
caulescens (E)
Lowne, B.T. s.n., M. officinarum (BM)
Ludlow, F.& Sheriff, G. 107, M. caulescens (BM); 1590, M. caulescens
(BM); 3073, M. caulescens (BM); 8676, M. caulescens (BM); 9014, M.
caulescens (BM)
LHd/ovv, E, S/zernjft G. cfe E///O?, //.//. 13149, M. caulescens (BM, E);
13750, M. caulescens (BM, E); 15051, M. caulescens (BM, E)
LwoVow, E, Sherriff, G. & Hicks, J.H. 16139, M. caulescens (BM); 18986,
M. caulescens (BM); 19040, M. caulescens, (BM); 20645, M.
caulescens (BM, E)
Ludlow, F., Sherriff, G. & Taylor, G. 3776, M. caulescens (BM); 4591 A,
M. caulescens (BM, E); 4591, M. caulescens (BM)
Maty, K s.n., M. officinarum (K)
Manberg, T. s.n., A/, caulescens (K)
Mavzomoustakis, G.A. 8, M. officinarum (BM)
Me Laren '5 collectors AC 67, M. caulescens (E, P)
Meyers, E 4377, M. officinarum (E)
Meyers, F.S. & Dinsmore, J.E. 6377, M. officinarum (E); B 6377, M.
officinarum (K)
M;7/, y.S. s.n., M. officinarum (K)
Moreau, W.M. 62, M. officinarum (K)
Norris, F.H. s.n., M. officinarum (BM)
, H., ATanai, H., 0/z6a, //. c£ rafm/z/, K 775165, M. caulescens
(BM)
Olivier & Bruguiere s.n., M. officinarum (P)
Orphanides, T.G. 75, M. officinarum (BM, E, K, P)
Pa/ne, /A. s.n., M. officinarum (K)
Pantling, R. s.n., M. caulescens (K); 46387, M. caulescens (K)
Partridge, F. 2, M. officinarum (BM)
Pasta/; 7. <6 Va/des, B. s.n., M. officinarum (RNG)
Perraudiere, H. de la s.n., M. officinarum (P)
Pette;; E s.n., M. officinarum (BM)
Pichler s.n., M. officinarum (BM, E, K)
Pignant s.n., M. officinarum (K)
P/fard, C.-y. 1757, M. officinarum (K)
Polunin, 0. 631, M. caulescens (BM); 780, M. caulescens (BM); 6616, M.
officinarum (K)
Po/Mrtw, O., Sytes, W/?. <fe Williams, L.H.J. 878, M. caulescens (BM, P);
4250, M. caulescens (BM); 4668, M. caulescens (BM); 4696, M.
caulescens (BM)
Pratt, A.E. 755, M. caulescens (BM, K, P)
Probyn, P.M. 49, M. officinarum (K)
Raulin, V. s.n., M. officinarum (P); 302, M. officinarum (P)
Rechinger, K.H. 90, M. officinarum (BM); 3630, M. officinarum (BM, K)
Reiser, C. s.n., M. officinarum (K)
Reverchon, E. s.n., M. officinarum (BM, E, P, K); 1 17, M. officinarum
(BM, E, K, P, PAL); 469 [1888], M. officinarum (BM, P); 469 [1890],
M. officinarum (P)
Rhomoo, L 210, M. caulescens (E); 1059, M. caulescens (E)
/?/£« <fe Rhomoo 5492, M. caulescens (BM, E, K)
flocfc, y.E 3597, M. caulescens (E); 3985, M. caulescens (P); 421 1, M.
caulescens (BM); 17530, M. caulescens (E); 22771, M. caulescens (E);
24750, M. caulescens (BM, E); 25005, M. caulescens (BM, E)
s, //. s.n., M. officinarum (BM)
s, H. 266, M. officinarum (E, K, P)
Schneider, C. 3501, M. caulescens (E, K)
Sintenis & Rigo 8, M. officinarum (K, PAL); 8a, M. officinarum (K)
Smtt/z, WW 4593, M. caulescens (E)
S0H/j<?', y.-A. 849, M. caulescens (P); 1252, M. caulescens (P); 2102, M.
caulescens (P)
Stainton, J.D.A. 254, M. caulescens (BM); 4263, M. caulescens (BM);
4676, M. caulescens (BM)
Starling, B.N., Upward, EM., Brickell, C.D. & Mathew, B. 92, M.
caulescens (K); 106, M. caulescens (K)
Syngrassides, A. 1464, M. officinarum (K)
, A. 38, M. officinarum (RNG)
, A. 1630, M. officinarum (BM)
Thomas, P. s.n., M. officinarum (K)
TOman, //. W 23, M. caulescens (K)
Todaro, A. s.n., M. officinarum (P)
Tbdd, E. s.n., M. officinarum (BM)
Tomkinson, M.J. 72, 72A, M. officinarum (BM)
Townsend, C.C. 85/1 1, M. officinarum (K)
Tbze/; //.E s.n., M. officinarum (K)
Treffewy, A. W 1 1, M. officinarum (K); 20, M. officinarum (K)
Vaccarz, A. 1 141, M. officinarum (K); 2326, M. officinarum (BM, K)
Virantonio s.n., M. officinarum (BM)
Wa/ree, A.M. 22226, M. officinarum (BM)
WaM, £.113, M. caulescens (K)
Watt, G. 5373, M. caulescens (E, K); 5436, M. caulescens (E, K)
Wfe/wttscfc, EM.y. s.n., M. officinarum (P); 159, M. officinarum (BM); 337,
M. officinarum (P)
White, M.F. s.n., M. officinarum (BM)
Williams, LH.J. 700, M. caulescens (BM)
W/son, E.//. 4192, M. caulescens (BM, K, P); 4194, M. caulescens (BM,
K, P); 4195, M. caulescens (K)
WW/ey-ZW, A.//. 22, M. officinarum (BM)
Ku, rr 5496, M. caulescens (KUN); 1 1327, M. caulescens (KUN); 19078,
M. caulescens (E); 19843, M. caulescens (E, KUN)
Zimmermann, A. 688, M. caulescens (BM)
40
S. UNGRICHT, S. KNAPP AND J.R. PRESS
TAXONOMIC INDEX
Accepted names are given in roman, synonyms in italics. An asterisk denotes
a figure.
Anisodus caulescens (C.B. Clarke) Diels 34
A. luridus Link 22
A. mariae Pascher 34
Atropa acaulis Stokes 30
A. belladonna L. 30
A. humilis Salisb. 30
A. mandragora L. 30
Lycium chinense Mill. 22
Mairella yunnanensis H. LeY 34
Mandragora acaulis Gaertn. 30
M. autumnalis Bertol. 30
M. caulescens C.B. Clarke 34, 35*
M. caulescens subsp. brevicalyx Grierson & D.G. Long 34
M. caulescens C.B. Clarke subsp. caulescens 34
M. caulescens subsp. flavida Grierson & D.G. Long 34
M. caulescens subsp. purpurascens Grierson & D.G. Long 34
M. chinghaiensis Kuang & A.M. Lu 34
M. foemina Garsault 30
M. haussknechtii Heldr. 30
M. hispanica F. Vierh. 30
M. x hybrida Hausskn. & Heldr. 30
M. mas Garsault 30
M. microcarpa Bertol. 30
M. neglecta G. Don ex Loudon 30
M. officinarum L. 30, 31*. 32*
M. praecox Sweet 30
M. shebbearei C.E.C. Fisch. 19
M. tibetica Grubov 34
M. turcomanica Mizg. 33
M. vemalis Bertol. 30
Nicotiana glauca Graham 22
Przewalskia Grubov 19
Bull. nat. Hist. Mus. Land. (Bot.) 28(1): 41-66 Issued 25 June 1998
The pteridophytes of Sao Tome and Principe
(Gulf of Guinea)
ESTRELA FIGUEIREDO^ ,_
Centra de Botdnica, Institute de Investigacao Cientifica Tropical, R. da Junqueira 86, 1300 Lisboa, Portugal
CONTENTS
Introduction 41
Diversity 42
Conservation 42
Collections 42
Historical collections 42
Recent collections 42
Catalogue 43
Acrostichaceae 43
Adiantaceae 43
Aspidiaceae 45
Aspleniaceae 48
Athyriaceae 50
Blechnaceae 51
Cyatheaceae 51
Davalliaceae 51
Dennstaedtiaceae 51
Elaphoglossaceae 52
Gleicheniaceae 53
Grammitaceae 53
Hemionitidaceae 54
Hymenophyllaceae 54
Lomariopsidaceae 56
Loxogrammaceae 56
Lycopodiaceae 5"
Marattiaceae 57
Nephrolepidaceae 5£
Oleandraceae 58
Ophioglossaceae 55
Polypodiaceae 5>
Psilotaceae ""
Pteridaceae 60
Selaginellaceae 61
sr\
Sinopteridaceae DZ
f\)
Thelypteridaceae
Vittariaceae "-
Doubtful records
References
64
Systematic index
SYNOPSIS. A catalogue of the pteridophytes of Sao Tome and Principe is presented, with 156 taxa, 12 of which are endemic to
the islands. Recent synonymy for the area is included. All collections examined are cited.
- ' --- The pteridophytes of the neighbouring island of Bioko (Fernando
INTRODUCTION _ _ _ Po) have been recently revised by Benl (1978, 1980, 1982, 1988,
1 99 1 ) who provided descriptions and keys to the taxa. The lack of an
The most recent account of the pteridophytes of Sao Tome and updated checklist for Sao Tome and Principe and the difficulty in
Principe (Gulf of Guinea) was produced by Alston in 1944. Since compiling the dispersed information on this group of plants have
then several nomenclatural changes have taken place and new precluded its inclusion in recent floristic and biodiversity studies
records have been added to the flora of the islands (e.g. Alston 1958, (Figueiredo, 1994).
The incorporation of several recent collections from Sao Tome
© The Natural History Museum, 1998
42
E. FIGUEIREDO
and Principe at LISC, has revealed further new records and new
areas of distribution for some taxa, and provided the basis for a
revision of Alston's (1944) account.
DIVERSITY
There are 153 indigenous taxa of pteridophytes in Sao Tome and
Principe occurring from sea level (e.g. Acrostichum aureum L.,
found in swamps and mangroves) to the highest peaks of the islands
(e.g. Grammitis spp.), and belonging to 28 families and 60 genera.
Three taxa were probably introduced, and became naturalized,
Pityrogramma calomelanos (L.) Link (Alston, 1944), Adiantum
raddianum C. Presl from the Neotropics (Alston, 1956a), and Pteris
vittata L. (Alston 1958, 1959).
One hundred and thirty-seven taxa occur in Sao Tome and 75
occur in Principe; 12 taxa are endemic to the islands. In Bioko, the
pteridophyte richness is higher than in either Sao Tome or Principe,
with 208 taxa in 31 families (Benl, 1978). However, only four taxa
(1.9%) are endemic to Bioko, while eight taxa are endemic to Sao
Tome (5.8%) and three are endemic to Principe (4%).
This difference in the percentage of endemics is less striking than
the difference found for the angiosperms. In that group, Bioko has
3.6% endemic taxa, Sao Tome has 13.4%, and Principe has 8.2%
(Figueiredo, 1994).
Asplenium is the genus of pteridophytes with the largest number
of taxa in Sao Tome and Principe. Twenty-four taxa are represented,
of which three are endemic.
CONSERVATION
Twenty-five taxa collected in Sao Tome and Principe in the nine-
teenth century have not been collected since, even though extensive
collecting took place in the 1950s and also in the 1990s. Although
some of these taxa, represented in the herbarium by a single collec-
tion, may be doubtful records, others were collected more than once
by different collectors. The absence of these taxa amongst the
collections made this century may indicate their rarity or extinction.
For example, Microlepia speluncae (L.) T. Moore was collected in
Sao Tome by Don (s.l.), Quintas (Ilheu das Rolas), and Welwitsch
(Monte Cafe). Although Monte Cafe is a well collected area, visited
by most botanists, this species has not been found during the last 100
years. Elsewhere in Africa, M. speluncae is rare and sporadic,
occurring at low to medium altitudes.
Sixteen taxa are represented in herbaria by a single collection,
either from Sao Tome or from Principe (10 from Sao Tome and 6
from Principe), which may indicate their rarity or even extinction.
When Principe alone is considered, 27 of the total of 75 taxa have
been collected only once. In this case, mistaken recordings or
undercollecting are also a possibility. Several species are repre-
sented in herbaria by a single collection made by Barter, who visited
Principe when stationed at Bioko, and therefore it is possible that
there were some errors when the specimens were labelled. Principe
remains poorly collected and recent collections of pteridophytes
from that island were not available to confirm the presence of these
taxa.
Seven taxa are apparently restricted to the Pico of Sao Tome
[Asplenium friesiorum C. Chr., Ctenitis cirrhosa (Schum.) Ching,
Grammitis molleri (Baker) Schelpe, G. tomensis Schelpe, Huperzia
ophioglossoides (Lam.) Rothm., Lycopodium clavatum L., and
Xiphopteris oosora (Baker) Alston] and five taxa are confined to the
Pico of Principe [Grammitis nigrocincta Alston, Hymenophyllum
polyanthos var. mildbraedii (Brause ex Brause & Hieron.) Schelpe,
Lomariopsis warneckei (Hieron.) Alston, Selaginella squarrosa
Baker, and Triplophyllum jenseniae (C. Chr.) Holttum]. Most of
these are high altitude pteridophytes which are also distributed on
other African mountains. Two of these taxa, Grammitis molleri and
G. tomensis, are very rare and endemic to the Pico of Sao Tome.
Lycopodium clavatum is cosmopolitan but, in the tropics, it is
confined to mountains.
Of the twelve taxa endemic to Sao Tome and Principe, two are
apparently very rare, having been collected only once: Selaginella
monodii Alston and Sphaerostephanos elatus subsp. thomensis
Holttum.
COLLECTIONS
Historical collections
Exell (1944) gives a lively account of the early collectors in Sao
Tome and Principe. The majority of the earliest collections of
pteridophytes from Sao Tome and Principe were made by Don (in
1822), Curror (in 1839), Welwitsch (in 1853), Barter (in 1858),
and Mann (in 1861). These collections are deposited at K (Curror,
Barter, and Mann), BM (Don and Welwitsch), and LISU
(Welwitsch).
In the late nineteenth century extensive collecting was made by
the Portuguese collectors Moller, Quintas, and Newton. The origi-
nal set of these collections is kept at COI, but duplicates were sent
to several European herbaria. The duplicates were distributed as
Flora Africanae Exsiccata (F.A.E.}, a series in which different
collections of the same taxon were grouped and distributed with
the same number. For this reason, most F.A.E. labels refer to
several localities and different collectors After examining the
original sets of the Moller, Quintas, and Newton collections, Exell
(1944: 383-385) tried to link originals with duplicates. However,
it seems that, for the pteridophytes, in most cases the original
collections were scarsely labelled, lacking the information needed
to link them to the duplicates distributed. Many F.A.E. duplicates
sent from COI to K had a second label referring to a single
collector. The numbering on these collector labels is unique to K
specimens, as the original sets at COI are mostly unnumbered.
Although the second label usually refers to a single collector and
locality there is no certainty as to the correspondence between the
specimen and the data. The information consisting of more than
one locality, found on some F.A.E. and original set labels, is
worthless and misleading because it is not possible to relate it to
the specimens it is attached to. This information should be omitted
when those collections are cited.
Together with the F.A.E. collections, the collections made by
Chevalier in 1905 and Exell in 1932-33, the first sets of which are
deposited, respectively, at P and BM, comprise the bulk of the
material examined to produce Alston's account of the pteridophytes
in Exell's Catalogue (1944).
Recent collections
Only a few collections were made during the twenty years that
followed Exell's expedition in 1932-33. The majority of recent
collections were made during the years 1954-1973, and in the
1990s.
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
43
The first set of the collections made by Rozeira in Sao Tome and
Principe in 1954 is deposited at PO. However, several collections of
pteridophytes cited in the literature (Alston, 1958) could not be
located in that herbarium. A set of duplicates of Rozeira's collection
is apparently deposited at BM (Exell & Rozeira, 1958) but could not
be found in 1997. Some duplicates (with the name of the collector
missing) exist at K. Based on Rozeira's collections, new records for
the islands were published by Alston (1958) in the proceedings of
the 'Conferencia Internacional dos Africanistas Ocidentais', which
took place in Sao Tome in 1956.
During 1956 several botanists collected in Sao Tome and
Principe. As part of a Calypso expedition which took place in
1956-57, Rose made several collections in the two islands, in-
cluding living material, which were deposited at P. In the same
year, Monod and Thorold made important collections in Sao Tome
and Principe, which are deposited at BM. These collections were
studied by Alston (1958, 1959) and resulted in several new records
being added to the flora of the islands. Lains e Silva also collected
in Sao Tome in 1956. Although his collections have been cited in
the literature, several numbers could not be found either at LISJC
or LISU. In 1968-1973, Espirito Santo collected in both islands.
The original sets of his collections are kept at LISJC and LISC,
with some duplicates at BM and COI.
Recent collections of pteridophytes from Sao Tome examined
during this study were mainly made by Pinto Basto in 1990, Matos
and Matos & Van Essche in 1994, and Lejoly in 1994-1995, and are
deposited at LISC. Recent collections were also made by Paiva
(COI), but these were not available for examination at the time of
this study.
CATALOGUE
The catalogue includes all the taxa of pteridophytes known to occur
in Sao Tome and Principe, arranged in alphabetical order, with
synonyms and references to the literature for the area published
since 1944. Synonyms already cited in Alston's (1944) work are
omitted.
All the collections examined are cited in this catalogue. Collec-
tions cited in the literature are also included. Some of these specimens
were not located (indicated in the text with 'not found'), and others
were not available for study. Some of the collections cited but not
seen by Alston ( 1 944), probably at B, were not examined during this
study; these are marked with a question mark. It is possible that a
future study of the unseen material may reveal misidentifications by
previous authors.
For each taxon, the collections are cited by alphabetical order of
collectors, but collections from the same locality are grouped.
Collections without locality are placed at the end of the specimens
cited. These include F.A.E. and Moller collections with more than
one locality on the labels. F.A.E. duplicates at K are cited under their
F.A.E. number and reference is made to other labels and numbers if
they exist.
Most of the localities provided by the collectors consist of the
name of a plantation ('ro9a') or a river. For this reason, distribution
maps based on these collections would be very inaccurate. Maps
with 'ro9as' and other collecting localities mentioned in the text are
provided, to allow for an estimate of the distribution of the taxa (Figs
1 and 2).
Data on habit, habitat, and ecology are lacking for the vast
majority of the collections examined.
Acrostichaceae
Acrostichum L.
Acrostichum aureum L., Sp. pi. 2: 1069 (1753); Alston in Exell,
Cat. Vase. PL S. Tome: 85 (1944); Benl inActa Bot. Barcinon. 38:
22(1988).
SAO TOME. Porto Alegre, Chevalier 14194bis (P!), 14200 (P!), salt
marshes, Espirito Santo 5131 (LISC!, LISJC!); Ilheu das Rolas, F.A.E.
(Quintas)69 (BM!, LISU!), Quintass.n. (COI!); PortoAlegre, Praia Jal<£, Rio
Malanza, swamp, Matos 7663 (LISC!); Perseveran?a, Rose 538 (P!).
PRINCIPE. Santo Antonio, salt marshes, Espirito Santo 5170 (LISC!,
LISJC!), mangrove, Monod 12191 (BM!), mangrove, Rose 425 (P!); Rio
Papagaio, swampy border, Quintas 1 (BM!, COI!); Praia General Fonseca,
Rozeira 430 (PO!); s.l., Welwitsch 15 (BM!, LISU!).
DISTRIBUTION. Pantropical.
Adiantaceae
Adiantum L.
Adiantum mettenii Kuhn, Filic. afr.: 65 (1868) nomen; in Jahrb.
Konigl. Bot. Gart. Berl 1: 338 (1881).
Adiantum soboliferum sensu auct. non Wall, ex Hook., Sp.fil. 2: 13,
t. 74A (1858); Alston in Exell, Cat. Vase. PL S. Tome: 82 (1944).
SAO TOME. Monte Forte, 50 m, Quintas s.n. (COI!, K!); s.l., Don s.n.
(BM!).
DISTRIBUTION. Tropical Africa and Madagascar, extending to In-
dia and Philippines.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Adiantum philippense L., Sp. pi. 2: 1094 (1753); Alston in Exell,
Cat. Vase. PL S. Tome: 82 (1944); Alston in Exell, Suppl. Cat.
Vase. PL S. Tome: 1 (1956); Benl in Acta Bot. Barcinon. 32: 26
(1980).
SAO TOME. Casal Pires, waterfall, 200 m, Espirito Santo 1 10 (BM!, COI!,
LISJC!); Boa Entrada, Chevalier 14518 (P); Rio do Ouro, 1 10 m, Quintas
1249 (BM!, COI!, LISJC!); Agua Coimbra, 100 m, Quintas s.n. (COI!),
Moller & Quintas s.n. (LISU!); Diogo Vaz, Rozeira 518 (PO!); Santa
Catarina, between main house and Lemba, Rozeira 840 (PO!).
DISTRIBUTION. Palaeotropical.
Adiantum raddianum C. Presl, Tent, pterid.: 158 (1836); Alston in
Exell, Suppl. Cat. Vase. PL S. Tome: 1 (1956); Alston in Exell &
Rozeira in Conf. Int. Afr. Oc. 3: 79 (1958).
SAO TOME. Quinta da Graca, 600 m, Espirito Santo 124 (BM!, COI!,
LISJC!); Pouso Alto, 490 m, Espirito Santo 4278 (LISC!, LISJC!); Monte
Cafe, Sao Carlos, secondary forest, Espirito Santo 5048 (LISC!, LISJC!);
Sao Nicolau, waterfall, on wet rocks, Espirito Santo 5178 (LISC!, LISJC!);
Saudade, 900 m, Exell 399 (BM!, COI!); Ponta Figo, on the way to Morro
Vilela, Lains e Silva 357 (LISU!); between Pico and Monte Castro, 1200-
1800 m, mountain rainforest, Lejoly 95/13 (LISC!); Monte Cafe, 1010 m,
Monod 1 1694 (BM!, COI!), Oliveira & Noronha s.n. (LISU!); NW of Pico,
1000-1200 m, Monod 12263 (BM!, COI!); Agua Ize\ common, Pinto Basto
199 (LISC!); Tras-os-Montes, Calvario, Rozeira 582 (PO!).
PRINCIPE. Esperansa, near Ribeira Camarao, Rozeira 354 (PO!).
DISTRIBUTION. Cultivated American species, escaped and natural-
ized.
44
E. FIGUEIREDO
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
Adiantum vogelii Mett. ex Keyserl. in Mem. Acad. Imp. Sci. Saint
Petersbourg ser. 7, 22(2): 8, 3 1 ( 1 875);Alston in Exell, Cat. Vase.
PI. S. Tome: 83 (1944);Tardieu-Blot inAubrev., Fl. Cameroun 3:
147 (1964); Benl in Acta Bot. Barcinon. 32: 26 (1980).
PRINCIPE. Santo Antonio, Monod 12072 (BM!, COI!); Berimbau, Newton
18 (BM!, COI!, K!), Newton s.n. (K!); Cimalo, Quintas 36 (BM!, COI!);
Esperan?a, between Montalegre and Maquina, Rozeira 378 (PO!); s.l., Souza
s.n. (COI!).
DISTRIBUTION. W. tropical Africa to Angola, Zanzibar.
Aspidiaceae (=Dryopteridaceae)
Ctenitis (C. Chr.) C. Chr.
Ctenitis cirrhosa (Schum.) Ching in Sunyatsenia 5: 250 (1940);
Benl in Acta Bot. Barcinon. 40: 41 (1991).
Dryopteris crinobulbon (Hook.) C. Chr., Index filic., Suppl. 3: 84
(1934); Alston in Exell, Cat. Vase. PL S. Tome: 61 (1944).
SAO TOME. Pico, 1850 m, Mann s.n. (B, K!-type of Dryopteris
crinobulbon).
DISTRIBUTION. Tropical and SE Africa, Madagascar, Comoros
and Mascarene Is.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Ctenitis lanuginosa (Willd. ex Kaulf.) Copel., Gen. fit.: 124 (1947);
Benl in Acta Bot. Barcinon. 40: 41 (1991).
Dryopteris lanuginosa (Willd. ex Kaulf.) C. Chr., Index filic.: 273
(1905); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944).
SAO TOME. S.I., Mann s.n. (K-not found).
DISTRIBUTION. Tropical and subtropical Africa, Madagascar,
Comoros, Seychelles and Mascarene Is.
45
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Didymochlaena Desv.
Didymochlaena truncatula (Sw.) J. Sm. in J. Bot. (Hooker) 4: 196
(1841); Alston in Exell, Cat. Vase. PL S. Tome: 66 (1944);
Tardieu-Blot in Aubrev., Fl. Cameroun 3: 254 (1964); Benl in
Acta Bot. Barcinon. 40: 35 (1991).
SAO TOME. Rio Contador, 1270-1350 m, F.A.E. (Moller) 36 (BM!,
LISU!);Trds-os-Montes, Gama s.n. (CO\!);Mollers.n. (COI!); Macambrani,
1270 m, Moller s.n. (COI!); Sela do Camelo, near Lemba, 200 m, Rozeira
3450 (PO!); Monte Cafe, 760-850 m, Welwitsch 61 (BM!, K!, LISU!).
DISTRIBUTION. Pantropical.
Dryopteris Adans.
Dryopteris pentheri (Krasser) C. Chr., Index filic.: 284 (1905);
Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 447
(1959); Benl in Acta Bot. Barcinon. 40: 36 (1991).
Dryopteris oligantha sensu auct. non (Desv.) C. Chr., Index filic.
Suppl. 3: 93 (1934); Alston in Exell, Cat. Vase. PL S. Tome: 62
(1944).
SAO TOME. Pico, Chevalier 13651 (P), 14290 (P), Moller s.n. (COI-not
found, P!); between Monte Cafe and Pico, Chevalier 14580 (P); Macambrara,
Ere// 131 (BM), 1 35 (BM);Tras-os-Montes, Gama s.n. (COI-not found), 800
m, Quintas 1286 (BM!, COI-not found), Rozeira 554 (K!, PO!); between
Pico and Monte Castro, 6 km S. of Ponta Figo, mountain rainforest, 1200-
1800 m, Lejoly 95/9 (LISC!); between Monte Quinas and Quinas Alias,
Monod 12007 (BM!); Monte Cafe, 1010 m, Monod 1 1696 (BM!); Lagoa
Amelia, 1300 m, Quintas 1291 (BM!, COI-not found), Rose 199 (P!); Sao
Nicolau, near waterfall, Rozeira 182 (K!, PO!); Santa Catarina, Lemba,
Rozeira 856 (K!); s.l., 1850 m, Mann (K!), F.A.E. (Moller) 39 (BM!, LISU!),
Moller s.n. (COI-not found), Quintas 10 (K!).
DISTRIBUTION. Tropical and southern Africa.
Fig. 1 Sao Tome
1 . Agua Cardoso-2B
2. Agua Coimbra-3A
3. Agua Grande-3B-4B-4A
4. Agua Ize (Ro?a)-4C
5. Agua Pinhao-3B
6. Almas-4B
7. Antonio Soares-4B
8. Bacelar-2D
9. Batepa-4B
10. Bemposta (Ro9a)-3B
ll.Benfica(Roca)-3B
12. Binda(Ro?a)-lC
13. Blu-blu-4B
14. Boa Entrada (Roca)-4A
15. Bom Sucesso (Ro?a)-3B
16. Bombaim (Roca)-3C
17. Cabumbe-2C
18. Caixao Grande-4B
19. Calvario-2B
20. Cantagalo (Ro?a)-4C
21.CaoPequeno-2D
22. Cascata-2B
23. Chami90-3B
24. Colonia Acoreana
(Roca)-4C
25. Cruzeiro (Roca)-3C
26. Diana (Ro?a)-4B
27. Diogo Nunes-4A
28. Diogo Vaz (Roca)-2B
29. Dona Amelia (Roca)-2B
30. Dona Augusta (Roca)-3D
3 1 . Dona Eugenia (Roca)-3D
32. Ermelinda (Roca)-2D
33. Estacao Sousa-2B
34. Famosa-4B
35. Formoso-3C
36. Guaiaquil (Ro?a)-3C
37. Guarda (Ro?a)-3A
38. Ilheu das Rolas-2E-2F
39. Java (Roca)-3B
40. Jou (Roca)-2D
41 . Lagoa Amelia-SB
42. Lemba (Roca)- IB
43. Macambrara (Roca)-3B
44. Maria Fernandes-3C
45. Milagrosa (Roca)-4B
46. Monte Cafe (Roca)-3B
47. Monte Castro-2B
48. Monte Forte (Ro9a)-2A
49. Monte Mario (Roc.a)-2E
50. Monte Quinas-2B
5 1 . Monte Rosa (Roca)-2D
52. Monte Verde-2D
53. Morro Cantagalo-3C
54. Morro Vigia -3B
55. Morro Vilela-2B
56. Nova Ceilao (Ro9a)-3B
57. Nova Moca (Ro9a)-3B
58. Novo Brasil (Ro9a)-2E
59. Novo Destine (Ro9a)-3B
60. Paga Fogo (Ro9a)-2B
61. Pedroma (Ro9a)-4B
62. Perseveran9a (Ro9a)-3D
63. Pico-2B
64. Pinheira (Ro9a)-4B
65. Ponta Figo (Ro9a)-2A
66. Porto Alegre (Ro9a)-2E
68. Praia Jale-2E
69. Praia Melao (Ro9a)-5B
70. Praia Pantufo-5B
71.Quija-2D
72. Quinas Altas -2B
73. Quinta da Gra9a (Roca)-3B
74. RibeiraMo9a-lB-2B
75. Ribeira Peixe-3D
76. Rio Abade-3C-4C
77. Rio Abade (bridge) -4C
78. Rio Campos-2C
79. Rio Caue-2D
80. Rio Contador-2A-2B
8 1 . Rio do Ouro (Ro9a)-3 A
82. Rio Malanza-2E
89. Rodia (Ro9a)-3B
90. Santa Catarina (Ro9a)-lB
9 I.Santa Irene (Ro9a)-2C
92. Santa Margarida (Ro9a)-3B
93. Santelmo (Ro9a)-3C
94. Sao Carlos (Ro9a)-3B
95. Sao Joao dos Angolares-3D
96. Sao Miguel (Ro9a)-lD
97. Sao Nicolau (Ro9a)-3B
98. Sao Pedro-3B
99. Sao Tome -4A
100. Sao Vicente-4B
101.Saudade(Ro9a)-3B
102. Sela do Camelo-2C
103. Tras-os-Montes (Ro9a)-3B
83. Rio Manuel Jorge-3B-4B-5B 104. Trindade-4B
84. Rio Manuel Jorge
(waterfall )-3B
85. Rio Miranda Guedes-3C-
3D
86. Rio Paga Fogo-lB-2B
87. Rio Xufexufe-2D
88. Ro9a Nova (Ro9a)-3B
105. Vale Carmo (Ro9a)-3D
106. Vieira Machado (Roca)-3C
107. Vila Aida (Ro9a)-3D
108. Vila Conceicao (Roca)-2E
109. Vila Jose (Ro9a)-3D
110. Zagaia-2C
111. Zampalma (Ro9a)-3B.
67. Pouso Alto (Ro9a)-3B
The following collecting localities could not be located: Agua Telha (Diogo Vaz), Agua Telha (near Ribeira Mo9a), Casa do Pico (1935 m alt.), Casal Pires
Correia, Monte Figo, Magodinho, Morro Pempem (Tras-os-Montes), Quingombo, Santa Maria, Rio Santo Antonio.
46
E. FIGUEIREDO
Fig. 2 Principe
1 . Barriga Branca- 1 C
2. Bela Vista (Ro?a)- IB
3. Berimbau-2B
4. Cajamanga- 1 B
5. Cimalo-2B
6. Dois Irmaos-2C
7. Esperanca (Ro9a)-lB
8. Fundao-lB
9. General Fonseca-2C
10. Infante D. Henrique
(Rosa)- 1C
H.Lapa(Ro9a)-lB
12. Mamas- 1C
13. Maquina-lB
14. Mencorne-lC
15. Montalegre (Roca)-lB
16. Morro do Leste-lB
17. Neves Ferreira (Rosa)- 1C
18. Oque Caspar (Rosa)- IB
19. 6queNazare-lB
20. Oque Pipi-lB
21.Pico-lC
22. Pico Papagaio-lB
23. Praia da Lapa-lB
24. Praia das Burras-2A
25. Precipfcio-2A
26. RibeiraBibi-lC
27. Ribeira Cambungo-lC-2C
28. Ribeira Chimboto-lC
29. Ribeira Fria (Ro?a)-2B
30. Ribeira Jambere-lB-2B
31.RioPapagaio-lB-2B
32. Santo Antonio-2B
33. Sao Joao (Rosa)-2B
34. Sundi(Ro?a)-lA
35.TerreiroVelho-2B.
The following collecting localities could not be located: Porto Real, Ribeira Camarao and Rio Bambu-Porco.
Lastreopsis Ching
Lastreopsis aff . currori (Mett. ex Kuhn) Tindale in Victoria Natu-
ralist 73: 184 (1957); Tindale in Contr. New South Wales Natl.
Herb. 3: 338(1965).
Aspidium nigritianum Mett. ex Kuhn, Filic. afr.: 138 (1868).
Dryopteris nigritiana (Mett. ex Kuhn) Kuntze, Revis. gen. pi. 2: 813
(1891); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944).
Ctenitis nigritiana (Mett. ex Kuhn) Alston in Bol. Soc. Brot., ser. 2,
30: 12(1956).
PRINCIPE. Rio Bambu-Porco, 60 m, Navel 138 (LISU!), 141 B (not found);
Pico, Rozeira 2607 (PO!); s.l., Barter 1906 (K-type of Aspidium nigritianum,
not found).
Tectaria Cav.
Tectaria angelicifolia (Schum.) Copel. in Philipp. J. Sci., ser. C,
Bot. 2: 410 (1907); Alston in Exell, Cat. Vase. PI. S. Tome: 66
(1944); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 1 (1956);
Benl in Acta Bot. Barcinon. 40: 55 (1991).
SAO TOME. Sao Vicente, 200 m, Espirito Santo 42 (BM!, COI!, LISJC!);
Bombaim, Monod 11886 (BM!); between Sao Tome and Monte Cafe,
Welwitsch 62 (BM!, LISU!); s.l., F.A.E. (Moller & Quintas) 43 p.p. (BM!,
COI!, K also Moller 32!, LISU!), 550 m, Welwitsch s.n. (K!).
PRINCIPE. S.l., Welwitsch 12b (BM!, LISU!).
DISTRIBUTION. W. and Central tropical Africa
Tectaria camerooniana (Hook.) Alston in / Bot. 77: 288 (1939);
Alston in Exell, Cat. Vase. PL S. Tome: 67 (1944);Tardieu-Blot in
Aubrev., Fl. Cameroun 3: 29 1 ( 1 964); Benl in Acta Bot. Barcinon.
40:56(1991).
SAO TOME. Macambrara to Zampalma, 900 m, ground fern, Exell 293
(BM!), 294 (BM!, COI!); between Bom Sucesso and LagoaAmelia, primary
forest, 1350-1420 m,Matos & Van Esschel296 (LISC!); above Monte Cafe,
1010 m, Monod 1 1693 (BM!); between Sao Pedro and LagoaAmelia, 1050
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
m, Monod 11711 (BM!); Calvario, 1200 m, Monod 11815 (BM!), 11817
(BM!); Lagoa Amelia, Rose 232 (P!); s.l., F.A.E. (Moller) 44 (BM!, COI!,
LISJC!, LISU!, P!).
DISTRIBUTION. W. and Central tropical Africa.
Tectaria fernandensis (Baker) C. Chr., Index filic., Suppl. 3: 179
(1934);Alston in Exell, Cat. Vase. PL S. Tome: 67 (1944); Benl in
Acta Bot. Bardnon. 40: 57 (1991).
SAO TOME. Boa Entrada, Chevalier 14360 (P!), 14517 (P!); SW region,
Chevalier 14591 (P!), 14617 (P!); Sao Miguel, 50 m, Espirito Santo 4742
(LISJC!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain
rainforest, 1200-1800 m, Lejoly 95/10 (LISC!); Sao Miguel, Monte Verde,
secondary forest, 10-1 50 m,Lejoly95/%4 (LISC!); Pedroma,Mocquerys 1 257
126(P!);SaoJoaodosAngolares,80m,A'eH'fons.n.(COI!),50m,2M/n/a5S.n.
(COI!); Agualze, common, Pinto Basto2\ 1 (LISC!); Perseveranca,/?o.se537
(P!); s.l., Don s.n. (BM!), F.A.E. (Moller & Quintas) 43 p.p. (LISJC!).
PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350
m, ground fern, Exell 564 (BM!, COI!), 565 (BM!, COI!); Pico Papagaio, 600
m, ground fern, Exell 684 (BM!); s.l., Barter 1907 (K!), Welwitsch 12 (BM!,
Ki.LISU!).
DISTRIBUTION. W. and Central tropical Africa.
Triplophyllum Holttum
Triplophyllum buchholzii (Kuhn) Holttum in Kew Bull. 41: 251
(1986).
Tectaria buchholzii (Kuhn) Copel. in Philipp. J. Sci. 38: 138 (1929)
'buckholzii'; Alston in Exell, Cat. Vase. PL S. Tome: 67 (1944).
Ctenitis buchholzii (Kuhn) Alston in Bol. Soc. Brot., ser. 2, 30: 1 1
(1956); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A,
21: 448 (1959).
PRINCIPE. Pico, 500-600 m, Monod 12119 (BM!), 800 m, Rozeira 886
(PO!), 91 1 (PO!), 2617 (PO!); Rio Bambu-Porco, 60 m, Navel 139B (LISU!);
Infante D. Henrique, 250 m, Rozeira 2300 (PO !); Infante D. Henrique, old path
to Ribeira Bibi, Rozeira 2391 (PO!); Mencorne, 200 m, Rozeira 241 1 (PO!).
DISTRIBUTION. Ivory Coast to Zaire.
Triplophyllum fraternum (Mett. ex Kuhn) Holttum in Kew Bull.
41:253(1986).
Aspidium fraternum Mett. ex Kuhn, Filic. afr.: 132 (1868).
Ctenitisfraterna(Meti.)Tardieu inNotul. Syst. (Paris) 14: 342(1953).
var. elongatum (Hook.) Holttum in Kew Bull. 41: 254 (1986).
Nephrodium subquinquefidum (P. Beauv.) Hook., Sp. fil. 4: 130
(1862) var. elongatum Hook., Sp.fil. 4: 130 (1862).
PRINCIPE. S.l., Mam 1861 (KMectotype of Nephrodium subquinque-
fidum var. elongatum).
DISTRIBUTION. Endemic.
NOTES. The following collections are unlocalized but may have
been collected in Principe, since there are no further collections from
the continent: Niger Exped., Barter s.n. (K!), s.l., Curror s.n. (K!).
Based on the same type specimen, Pichi-Sermolli (1985) consid-
ered this fern to be a new species (Ctenitis attenuata Pic. Serm.) and
later (Pichi Sermolli, 1991) transferred it to Triplophyllum with the
name T. attenuatum (Pic. Serm.) Pic. Serm.
This taxon has not been recorded in the islands since the nine-
teenth century.
Triplophyllum jenseniae (C. Chr.) Holttum in Kew. Bull. 41: 253
(1986).
47
PRINCIPE. Pico, 850 m, Rozeira 1021 (PO!).
DISTRIBUTION. W. and Central tropical Africa.
Triplophyllum principis Holttum in Kew Bull. 41: 246 (1986).
Dryopteris variabilis (Hook.)Alston in Exell, Cat. Vase. PL S. Tome:
64 (1944) var. barteri (Hook.) Alston in Exell, Cat. Vase. PL S.
Tome: 64(1944).
PRINCIPE. Oque Pipi, secondary forest, 300 m, ground fem, Exell 546
(BM!, COI!); between Oque Pipi and Morro do Leste, secondary forest, 350
m, ground fern, Exell 562 (BM!, COI!); above Infante D. Henrique, second-
ary forest, 300 m, ground fern, Exell 618 (BM!, COI!), 624 (BM!); Rio
Bambu-Porco, 60 m, Navel 14 la (LISU!); Oque Nazare, 500 m, Newton s.n.
(COI!); Infante D. Henrique, Rose 402 (P!), Rozeira 2370 (PO!); Neves
Ferreira, Rozeira 2204A (PO!), 2205 (PO!); between Barriga Branca and
Mamas, Rozeira 2552 (PO!); Mencorne, Rozeira 4299 (PO!); s.l., Barter
1927 (BM!, K!-holotype), Wilde et al. 371 (WAG, cited by Holttum, I.e.).
DISTRIBUTION. Endemic.
Triplophyllum protensum (Afzel. ex Sw.) Holttum in Kew Bull.
41: 247 (1986); Benl in Acta Bot. Bardnon. 40: 45 (1991).
Aspidium subquinquefidum P. Beauv., Fl. Oware 1: 34, t. 19 (1805).
Dryopteris protensa (Afzel. ex Sw.) C. Chr., Indexfilic.: 286 (1905);
Alston in Exell, Cat. Vase. PL S. Tome: 63 (1944).
SAO TOME. Bemposta, 600 m, Moller s.n. (BM!, COI!).
PRINCIPE. SW Esperan9a, 200 m, Exell 67 1 a (BM ! ); Sao Joao, Newton s.n.
(COI!); Infante D. Henrique, old path to Ribeira Bibi, Rozeira 2390A (PO!);
Pico Papagaio, Welwitsch 16 (BM!, LISU!); s.l., Souza s.n. (COI!).
DISTRIBUTION. Widespread in W. and Central tropical Africa.
NOTE. Pichi Sermolli (1991) considered that the type of
Triplophyllum protensum was a hybrid and described this taxon as
a new species, T. heudelotii Pic. Serm. He considered
Triplophyllum subquinquefidum (P. Beauv.) Pic. Serm. to be a
distinct species. Of these two taxa only the latter was recorded for
S. Tome and Principe.
Triplophyllum securidiforme (Hook.) Holttum in Kew Bull. 41:
242 (1986); Benl in Acta Bot. Bardnon. 40: 43 (1991).
Dryopteris securidiformis (Hook.) C. Chr., Indexfilic.: 291 (1905).
D. securidiformis var. nana Bonap., Not. pterid. 14: 211 (1923);
Alston in Exell, Cat. Vase. PL S. Tome: 64 (1944).
Triplophyllum securidiforme var. nanum (Bonap.) Holttum in Kew
Bull. 41:243(1986).
SAO TOME. Macambrara, primary forest, 1050-1200 m, Exell 103 (BM!),
151 (BM!); Monte Cafe, 800 m, Moller s.n. (COI!); between Monte Quinas
and Quinas Altas, Monod 12013 (BM!); Sao Joao dos Angolares, 200 m,
Quintas s.n. (COI!); Ribeira Moca, Rozeira 21 19 (PO!); between Ermelinda
and Cabumbe, Rozeira 2819 (PO!), 2823 (PO!); Cantagalo. 400 m, Rozeira
3152 (PO!); Sela do Camelo, 200 m, Rozeira 3451 (PO!); Formoso, Rozeira
3537 (PO!); near Rio Miranda Guedes, Rozeira 3639 (PO!); Vieira Machado,
400 m, Rozeira 3666 (PO!); s.l., F.A.E (Moller & Quintas) 40 (BM!, COI!, K
also Quintas 44!, LISU!, P!).
PRINCIPE. Rio Bambu-Porco, 60 m, Navel 1 39 A (LISU !), 1 4 1 C (not found);
Infante D. Henrique, Rose 409 (P!), 414 (P!); Neves Ferreira, Rozeira 2201
(PO!); Ribeira Camarao, Rozeira 22 19(PO!), 2220(PO!), Rozeira 2268 (PO!);
between Ribeira Jambere and Ribeira Fria, Rozeira 2237 (PO!); near Ribeira
Cambungo, Rozeira 2239 (PO !); Infante D. Henrique, old path to Ribeira Bibi,
Rozeira 2305 (PO!), 2326 (PO!); Infante D. Henrique, Rozeira 2374 (PO!);
Mencorne, Rozeira 2408 (PO !); between Barriga Branca and Mamas, Rozeira
2562 (PO!); s.l., Barter 1929 (P!), Mann s.n. (K-not found).
DISTRIBUTION. W. and Central tropical Africa.
48
Aspleniaceae
Asplenium L.
Asplenium aethiopicum (Burm.f.) Bech. in Candollea 6:22(1 935);
Alston in Exell, Cat. Vase. PL S. Tome: 74 (1944); Benl in Acta
Bot. Barcinon. 40: 24 (1991).
SAO TOME. Pico, primary forest, 1987 m, epiphyte, Matos & Van Essche
7381 (LISC!); Lagoa Amelia, 1430 m, epiphyte, Matos 7495 (LISC!);
between Estasao Sousa and Pico, primary forest, 1700 m, epiphyte, Matos
7554 (LISC!); Macambrara, 1270 m, Moller s.n. (COI!); Lagoa Amelia to
Sao Pedro, 1200 m, Quintas 1392 (BM!, COI!); Monte Cafe, Quintas 1392
(LISJC!); s.l., F.A.E. {Moller) 31 (BM!, K!, LISU!), Moller s.n. (COI!).
DISTRIBUTION. Pantropical and subtropical.
Asplenium africanum Desv. in Ges. Naturf. Freunde Berlin Mag.
Neuesten Entdeck. Gesammten Naturk. 5: 322 (1811); Alston in
Exell, Cat Vase. PI. S. Tome: 74 (1944); Benl in Acta Bot.
Barcmon. 40:6(1991).
SAO TOME. Ilheu das Rolas, Greeffs.n. (B); near Sao Tome, 8 m, Moller
s.n. (COI!); Agua Ize, frequent epiphyte, Pinto Basto2\S (LISC!); s.l., Don
s.n. (BM!), F.A.E. (Moller) 21 (BM!, LISU!, P!).
PRINCIPE. Above Neves Ferreira, 200-350 m, epiphyte, Exell 650 (BM!,
COI!); s.l., Newton (not found at COI).
DISTRIBUTION. W., Central, and E. Africa.
Asplenium anisophyllum Kunze in Linnaea 10: 511 (1836).
Asplenium geppii Carruth., Cat. Afr. PI. 2: 269 (1901); Alston in
Exell, Cat. Vase. PL S. Tome: 78 (1944).
SAO TOME. Near Sao Pedro, 1280 m, F.A.E. (Moller) 26 (BM!, K also
Moller 23 !, LISU!); between Pico and Monte Castro, 6 km S. of Ponta Figo,
mountain forest, 1200-1800 m, Lejoly 95/18 (LISC!); NW of Pico, 1350-
1600 m, Monod 12226 (BM!); Lagoa Amelia, Quintas 1385 (BM!, COI!),
Rose 206 (P!).
DISTRIBUTION. W. tropical Africa.
Asplenium barteri Hook., Sec. cent, ferns: t. 75 (1860); Alston in
Exell, Cat. Vase. PL S. Tome: 75 (1944);Tardieu-Blot inAubrev.,
Fl. Cameroun 3: 192 (1964); Benl in Acta Bot. Barcmon. 40: 13
(1991).
SAO TOME. Sao Miguel, Chevalier 1449 Ibis (P-not found); SW region,
Chevalier 14618 (P!); Porto Alegre, Chevalier 14619bis (P!); Saudade and
Pico, F.A.E. (Moller) 25 p.p. (BM!); Sao Joao dos Angolares, 80 m, Newton
s.n. (BM!, COI!); Cruzeiro, 300 m, Thorold 2065 (BM!); s.l., Matos s.n.
(LISC!).
PRINCIPE. Esperanca, plantation, 100 m, on rocks, Exell 677 (BM!, COI!);
W. of Pico, above Lapa, below 250 m, Monod 12100 (BM!); Pico, near Lapa,
below 250 m, Monod 12103 (BM!); Sao Joao, 450 m, Newton 28 (BM!,
COI!); Lapa (Oeste), Quintas 25 (BM!, COI!); s.l., Barter 1909 (K!), 1910
(K!).
DISTRIBUTION. Widespread in tropical Africa.
Asplenium biafranum Alston & F. Ballard in Hooker's Icon. pi. 34:
t. 3367 (1938); Alston in Exell, Cat. Vase. PL S. Tome: 75 (1944);
Tardieu-Blot in Aubrev., FL Cameroun 3: 200 (1964); Benl in
Acta Bot. Barcinon. 40: 23 (1991).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14543 (P!);
Macambrara, primary forest, 1050-1200 m, Exell 154 (BM!); Pico, primary
forest, 1900-2020 m, Lejoly 94/581 (LISC!); between Lagoa Amelia and
Chamico, Matos 7528 (LISC!); Santa Maria, 1350 m, Moller s.n. (COI!);
E. FIGUEIREDO
Lagoa Amelia, 1400 m, Monod 11747 (BM!); Calvario, 1560 m, Monod
1 1 834 (B M ! ) ; between Casa do Pico and Pico, 1 950 m, Monod 1 1 958 (BM ! ).
DISTRIBUTION. Benin, Bioko, Nigeria, Cameroun.
Asplenium cuneatum Lam., Encycl. 2: 309 (1786); Alston in Exell,
Cat. Vase. PL S. Tome: 75 (1944); Tardieu-Blot in Aubrev., FL
Cameroun 3: 219(1964).
SAO TOME. Tras-os-Montes, Morro Pempem, 1200 m, Quintas 1387
(BM!, COI!).
DISTRIBUTION. W. tropical Africa, tropical America, Polynesia.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Asplenium currorii Hook., Sp. fil. 3: 82 (1860); Alston in Exell,
Cat. Vase. PL S. Tome: 75 (1944); Alston in Exell in Bull. Inst.
Franc. Afrique Noire ser. A, 21: 445 (1959); Tardieu-Blot in
Aubrev., FL Cameroun 3: 180 (1964).
SAO TOME. Macambrara, primary forest, 1050-1200 m, epiphyte, Exell
172 (BM!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain
forest, 1200-1800 m, Lejoly 95/29 (LISC!); between Bom Sucesso and
Lagoa Amelia, primary forest, 1350-1420 m, Matos & Van Essche 7291
(LISC!); Moller s.n. (COI!); Lagoa Amelia, 1500 m, Monod 1 1722 (BM!);
s.l., 550 m, Welwitsch 64 (BM!, LISU!).
PRINCIPE. Pico Papagaio, Welwitsch 24a (BM!); s.l., Barter 1900 (K-not
found), Mann 1861 (K!), Welwitsch 24 (BM!, K-not found, LISU!).
DISTRIBUTION. W. tropical Africa.
Asplenium dregeanum Kunze in Linnaea 10:517(1 836); Alston in
Exell, Cat. Vase. PL S. Tome: 76 (1944); Alston in Exell, Suppl
Cat. Vase. PL S. Tome: 1 (1956).
subsp. brachypterum (Kunze ex Houlston & T. Moore) Pic. Serm.
in Bull. Jard. Bot. Belg. 55: 130 (1985); Benl in Acta Bot. Barci-
non. 40: 28 (1991).
SAO TOME. Monte Cafe, 1200 m, epiphyte, Espirito Santo 164 (BM!,
COI!, LISJC!); Macambrara, primary forest, 1050-1200 m, Exell 96 (BM!),
126 (BM!, COI!); Lagoa Amelia, 1200-1400 m, Lains e Silva 83 (LISU!),
Matos 7493 (LISC!), 1400 m, Monod 1 1741 (BM!), 1400 m, Quintas 1349
(BM!, COI!), Rose 224 (P!), 240 (P!), Rozeira 154 (PO!), 1 161 (PO!); Pico,
primary forest, 1900-2020 m, epiphyte, Lejoly 94/580 (LISC!), 1950 m,
Moller s.n. (COI!); between Bom Sucesso and LagoaAmelia, primary forest,
1350-1400 m, epiphyte, Matos & Van Essche 7287 (LISC!); between Sao
Pedro and Lagoa Amelia, 1050 m, Monod 11705 (BM!, COI!); Calvario,
1400 m, Monod 11852 (BM!); s.l., F.A.E. (Moller) 33 (BM!, COI!, K!,
LISJC!, LISU!), Mann s.n. (K!), Moller s.n. (COI!).
DISTRIBUTION. Zaire, Rwanda, Burundi.
Asplenium emarginatum P. Beauv., Fl. Oware 2: 6, t. 61 (1808);
Alston in Exell, Cat. Vase. PL S. Tome: 76 ( 1944);Alston in Exell,
Suppl. Cat. Vase. PL S. Tome: 1 (1956); Tardieu-Blot inAubrev.,
Fl. Cameroun 3: 192 (1964); Benl in Acta Bot. Barcinon. 40: 10
(1991).
SAO TOME. Sao Vicente, 200 m, common, Esptrito Santo 35 (BM!, COI!,
LISC!, LISJC!); Monte Cafe, 800 m, Moller 349 (BM!, COI!); Santa
Margarida, Antonio Soares, /toze/ra 1225 (PO!); s.l., Don. s.n. (BM!),800m,
Quintas 8 (K!).
PRINCIPE? (Beauvois, I.e.).
DISTRIBUTION. Widespread in tropical Africa.
Asplenium erectum Bory ex Willd., Sp. pi. 5: 328 (1810).
van usambarense (Hieron.) Schelpe in Bol. Soc. Brot., ser. 2, 41:
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
49
207 (1967); Schelpe in Exell & Launert, Fl. Zamb., Pteridophyta:
176 (1970); Benl in Acta Bot. Barcinon. 40: 16 (1991).
Asplenium quintasii Gand. in Bull. Soc. Bot. France 66: 305 (1919);
Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944);Tardieu-Blot in
Aubrev., Fl. Cameroun 3: 198 (1964).
SAO TOME. Monte Caf6, Chevalier 13673 (BM!, P!); Pico, 1850 m, Exell
345a (BM!), Newton s.n. (BM!, COI!), 1650 m, Quintas 1342 (BM!, COI!;
type of Asplenium quintasii); between Pico and Monte Castro, 6 km S. of
Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/19 (LISC!); between
Pico and Ponta Figo, primary forest, 1600-1700 m, epiphyte, Matos & Van
Essche 7389 (LISC!); Calvario, 1450 m, Monod 1 1826 (BM!).
DISTRIBUTION. Widespread in tropical Africa.
Asplenium eurysorum Hieron. in Engl. in Bot. Jahrb. Syst. 46: 364
(191 1); Alston in Exell, Cat. Vase. PI. S. Tome: 76 (1944).
SAO TOME. Pico, Chevalier 13659 (P!), primary forest, 1987 m, epiphyte,
Matos & Van Essche 7382 (LISC!); Calvario, 920 m, Espirito Santo 5070
(LISC!, LISJC!); between Bacelar and Quija, secondary forest, 160 m,
Espirito Santo 5139 (LISC!, LISJC!); Macambrara, primary forest, 1050-
1200 m, ground fern, Exell 132 (BM!), 1385 m, Moller s.n. (COI!); Ribeira
Peixe, Lains e Silva 301 (LISU!); between Pico and Monte Castro, 6 km S. of
Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/1 (LISC!); Santa
Maria, 1260 m, Moller s.n. (COI!); between Sao Pedro and Lagoa Amelia,
1050 m, Monod 11714 (BM!); between Monte Quinas and Quinas Alias,
Monod 12008 (BM!); s.l., F.A.E. (Moller) 28 (BM!, LISJC!, LISU!, P!).
PRINCIPE. Oque Nazare, Newton 27 (BM!, COI!, P!); Pico Papagaio,
Welwitsch 14 (BM!, K!, LISU!); s.l., Barter 1896b (K!).
DISTRIBUTION. Endemic.
Asplenium exhaustum (H. Christ) Alston in Exell, Cat. Vase. PL S.
Tome: 76, f. 2 (1944); Alston in Exell in Bull. Inst. Franc. Afrique
Noire ser. A, 21: 445 (1959).
SAO TOME. Pico, Chevalier 14587 (P!-type); Cruzeiro, 300 m, Thorold
2072 (BM!); s.l., Seabra s.n. (BM!, COI!, LISU!).
DISTRIBUTION. Endemic.
Asplenium formosum Willd., Sp. pi. 5: 329 (1810); Alston in Exell
& Rozeira in Conf. Int. Afr. Oc. 3: 79 (1958).
SAO TOME. Diogo Vaz, near Agua Telha, Rozeira 5 1 8a (PO).
DISTRIBUTION. Pantropical.
Asplenium friesiorumC. Chr. inNotizbl. Bot. Gart. Berlin-Dahlem
9: 181 (1924); Benl in Acta Bot. Barcinon. 40: 22 (1991).
SAO TOME. Pico, 1950 m, F.A.E. (Moller) 27 (BM!, K!, LISU!, P!);
between Casa do Pico and Pico, 1950 m, Monod 1 1957 (BM!).
DISTRIBUTION. Widespread in tropical Africa.
Asplenium hypomelas Kuhn, Filic. afr.: 104 (1868); Alston in
Exell, Cat Vase. PL S. Tome: 78 (1944); Benl in Acta Bot.
Barcinon. 40: 31 (1991).
SAO TOME. Porto Alegre, Chevalier 13658 (P!); between Monte Cafe and
Pico, Chevalier 14579 (P!); Lagoa Amelia, 1400 m, Espirito Santo 4069
(COI!, LISJC!), 1400 m, Monod 1 1735 (BM!, COI!), Rose 233 (P!), Rozeira
1 129 (PO!); Calvario, secondary forest, 920 m, Espirito Santo 5074 (LISC!,
LISJC!), 1200 m,Monod 11813 (BM\),Rose 150 (P!); Macambrara, primary
forest, 1050-1200 m, ground fern, Exell 120 (BM!, COI!), 1200-1350 m,
ground fern, 235a (BM!, COI!);Trds-os-Montes, Gamas.n. (COI!); between
Pico and Monte Castro, 6 km S. of Ponta Figo, mountain forest, 1200-1800
m, Lejoly 95/16 (LISC!); between Bom Sucesso and Lagoa Amelia, primary
forest, 1350-1420 m,Matos& Van £wc/je 7295 (LISC!); Rio Contador, 1400
m, Moller s.n. (COI!); W. of Pico, 1225 m, Monod 11996 (BM!); Pico,
Newton s.n. (BM!, COI!); Sao Pedro, Quintas s.n. (COI!); s.l., F.A.E. (Moller
& Quintas) \\(BM\, COI!, K also Moller 10!, LISJC!, LISU!, P\),Mann s.n.
(K!), Quintas s.n. (LISU!), Souza s.n. (COI!).
DISTRIBUTION. Widespread in tropical Africa, extending to S. and
SE Africa.
Asplenium inaequilaterale Bory ex Willd., Sp. pi. 5: 322 (1810);
Alston in Exell, Cat. Vase. PL S. Tome: 78 (1944); Benl in Acta
Bot. Barcinon. 40: 17 (1991).
SAO TOME. Ribeira Peixe, Lains e Silva 253 (LISU!), 254 (LISU!); Lagoa
Amelia, 1430 m, Matos 7492 (LISC!); between Bom Sucesso and
Macambrara, Matos 7605 (LISC!); Saudade, 750 m, Moller s.n. (COI!);
Calvdrio, Monod 1 1856 (BM!); s.l., F.A.E. (Moller) 25 p.p. (BM!, LISJC!,
LISU!), Moller 25 (BM!, COI!).
DISTRIBUTION. Widespread in tropical Africa, extending to S. and
SE Africa and Madagascar.
Asplenium lividum Mett. ex Kuhn in Linnaea 36: 100 (1869).
Asplenium protensum sensu auct. non Schrad. in Gott. Gel. Anz.
[1818]: 916 (1818); Alston in Exell, Cat. Vase. PL S. Tome: 79
(1944).
SAO TOME. Macambrara to Zampalma, 900 m, ground fem, Exell 296
(BM!, COI!); Agua Pinhao, 850 m, Quintas 1283 (BM!, COI!).
DISTRIBUTION. E. and S. Africa, and S. America.
Asplenium longicauda Hook., Sec. cent, ferns: t. 69 (1860); Alston
in Exell, Cat Vase. PL S. Tome: 78 (1944); Tardieu-Blot in
Aubrev., FL Cameroun 3: 1 86 ( 1 964); Benl in Acta Bot. Barcinon.
40:9(1991).
PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350
m, ground fern, Exell 571 (BM!, COI!); Rio Bambu-Porco, 60 m, Navel 144
(LISU!, P!); s.l., Barter 1900 (K!-syntype).
DISTRIBUTION. W. tropical Africa, Madagascar.
Asplenium megalura Hieron. in Brause & Hieron. in Mildbr., Wiss.
Erg. Deut. Zentr.-Afr. Exped., Bot. 2: 17(1910); Benl inActaBot.
Barcinon. 40: 23 (1991).
var. molleri (Hieron. )Tardieu inMem. Inst. Franc. Afrique Noire 28:
190(1953).
Asplenium molleri Hieron. in Engl. in Bot. Jahrb. Syst. 46: 371
(191 1); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944).
SAO TOME. Lagoa Amelia, 1400 m, epiphyte, Espirito Santo 4068 (COI!,
LISJC!), crater rim, 1350 m, epiphyte, Exell 202 (BM!), 1430 m, rare
epiphyte, Matos 7496 (LISC!), 1400 m, Monod 1 1779 (BM!); Macambrara,
primary forest, 1200 m, epiphyte, Exell 228 (BM!); Bom Sucesso, 1075 m,
F.A.E. (Moller) 29 (BM!, COI!, K!-type, LISU!); between Bom Sucesso and
Lagoa Amelia, secondary forest, 1 150-1300 m, Matos & Van Essche 7310
(LISC!); Calvario, 1400 m, Monod 1 1854 (BM!); W. of Pico, 870-1225 m,
Monod 1 1905 (BM!); s.l., Matos s.n. (LISC!).
DISTRIBUTION. The species is widespread in tropical Africa. The
variety is endemic to Sao Tome.
Asplenium nigritianum Hook., Sp.fil. 3: 223 (1860) et Sec. cent,
ferns: t. 44 (1860); Alston in Exell, Cat. Vase. PL S. Tome: 79
(1944); Alston in Bol. Soc. Brot., ser. 2, 30: 10 (1956).
SAO TOME. S.l., Mann 1 107 (K-not found).
PRINCIPE. Lapa (Leste), Quintas s.n. (BM!, COI!); s.l., Barter 1895 (BM!,
K-not found), (probably Lapa), Quintas B (K!).
50
DISTRIBUTION. Tropical Africa.
NOTES. As shown by Alston ( 1956£), it was wrongly assumed that
the type collection came from Bioko. In fact, Asplenium nigritianum
does not seem to occur in Bioko.
Alston (19566) referred to the type collection as Barter 225,
explaining that Hooker mistook the date, 1 859, for the number and
then miscopied it as 1 895 (in K specimen) or 1 898 (in BM speci-
men). Since the specimen at K was not found, it was not possible to
check its number. The collections of pteridophytes made by Barter
which I examined range from no. 1886 to 1928, so it is likely that
these figures refer to the number and not the date of collection. The
type collection should be referred to as Barter 1895.
The species is poorly represented in herbaria. Neither the speci-
men Barter 1895 (K) nor Mann 1107 (K) could be located. In a
handwritten document kept at BM, reference is made to a duplicate
of the Mann collection kept at B.
This taxon has not been recorded in the islands since the nine-
teenth century.
Asplenium paucijugum F. Ballard in Hooker's Icon. pi. 33: t. 3287
(1935); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944); Benl in
Acta Bot. Barcinon. 40: 8 (1991).
Asplenium variabile Hook., Sp. fil. 3: 93, t. 185 (1860) var.
paucijugum (F. Ballard) Alston in Bol. Soc. Brot., ser. 2, 30: 7
(1956); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 184 (1964).
SAO TOME. Monte Cafe, 800 m, F.A.E. (Moller) 24 (BM!, COI!, LISJC!,
LISU!); LagoaAmelia, primary forest, 1400-1420 m, epiphyte, Matos & Van
Essche 7410 (LISC!); NW of Pico, 1000-1350 m, Monod 12216 (BM!); s.l.,
800 m, Moller 19 (K!), 550 m, Welwitsch 63 (BM!, LISU!).
PRINCIPE. Sundi, Newton 29 (BM!, COI!).
DISTRIBUTION. W., Central, and E. tropical Africa, Madagascar.
Asplenium sandersonii Hook., Sp.fil. 3: 147, t.179 (1860); Alston
in Exell, Cat. Vase. PL S. Tome: 80 (1944); Benl in Acta Bot.
Barcinon. 40: 27 (1991).
SAO TOME. Macambrara, primary forest, 1050-1200 m, epiphyte, Exell
144 (BM!, COI!); Sao Miguel, between Xufexufe and Sao Miguel, 400 m,
epiphyte, Matos 7658 (LISC!); Pico, 1950 m, Moller s.n. (COI!); Bombaim,
Rose 156 (P!); DonaAugusta, Dona Eugenia, epiphyte, Rozeira 1096 (PO!);
LagoaAmelia, 1400 m, epiphyte, Sergio s.n. (LISU!); s.l., F.A.E. (Moller) 23
(BM!, K also Moller 21 !, LISU!), Mann (not found), Moller s.n. (COI!).
DISTRIBUTION. Widespread in tropical and subtropical Africa.
Asplenium unilaterale Lam., Encycl. 2: 305 (1786); Alston in
Exell, Cat Vase. PL S. Tome: 80 (1944); Benl in Acta Bot.
Barcinon. 40: 15 (1991).
SAO TOME. Pico, 1300-1900 m, EA.E. (Moller) 30 (K-not found, LISU!);
Pico, 1900 m, Moller s.n. (COI!); Rio Contador, 1300 m, Moller s.n. (COI!).
PRINCIPE. S.L, Barter 1917 (K!).
DISTRIBUTION. Palaeotropical.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Asplenium variabile Hook., Sp.fil. 3: 93, t. 185 (1860); Alston in
Exell, Cat Vase. PI. S. Tome: 80 (1944); Benl in Acta Bot.
Barcinon. 40: 7 (1991).
SAO TOME. 16 Grande, Sao Joao dos Angolares, 50 m, EA.E. (Quintas) 22
(K also Quintas 20!, LISU!); between Xufexufe and Sao Miguel, 400 m, epi-
phyte, Matos 7657 (LISC!); Sao Joao dos Angolares, 50 m, Quintas s.n.
(COI!).
E. FIGUEIREDO
PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350
m, on rocks, Exell 567 (BM!, COI!); W. of Pico, above Lapa, below 250 m,
Monod 12099 (BM!); Rio Bambu-Porco, 60 m, Navel 140 (LISU!, P!); Pico,
Thorold 2085 (BM!); s.l., Mann s.n. (K-not found).
DISTRIBUTION. W. tropical Africa.
NOTE. Alston (1944) cited the collection Barter 1910 (BM, K). At
BM, only the specimen Barter s.n. (Niger Expedition) was found.
Barter 1910 (K) is Asplenium barteri.
Athyriaceae
Athyrium Roth
Athyrium newtonii (Baker) Diels in Engl. & Prantl, Nat.
Pflanzenfam. 1(4): 224 (1899); Alston in Exell, Cat. Vase. PL S.
Tome: 73 (1944).
SAO TOME. Pico, 1650 m, Quintas 1393 (BM!, COI!, LISJC!), 1650 m,
Quintas s.n. (COI!); s.l., Newton 2 (K!-type), Newton s.n. (COI!), (probably
Pico), 1650 m, Quintas 9 (K!).
DISTRIBUTION. Also in Uganda and Tanzania, above 1000 m.
NOTES. Newton 88 (K) cited by Alston (I.e.) is most probably a
mistake for Quintas 9 (the number 88 after the collector's name is
the date).
This taxon has not been recorded in the islands since the nine-
teenth century.
Diplazium Sw.
Diplazium arborescens (Bory) Sw., Syn.fiL: 92 (1806); Alston in
Exell, Cat. Vase. PL S. Tome: 73 (1944).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14290 (P!); Pico,
Chevalier s.n. (P!); Macambrara, primary forest, 1050-1200 m, Exell 128
(BM!, COI!);Tras-os-Montes, Gama s.n. (COI!); between LagoaAmelia and
Chamico, primary forest, 1400 m, Matos 7526 (LISC!); Monte Cafe, Moller
s.n. (COI!); LagoaAmelia, 1350-1380 m, Monod 1 1772 (BM!, COI!), Rose
234 (P!), Rozeira 1139 (K!, PO!); Cantagalo, 400 m, Rozeira 3157 (PO!);
Formoso, Rozeira 3236 (PO!); Nova Ceilao, Rozeira 4905 (PO!); s.l., EA.E.
(Moller) 34 (BM!, COI!, K also Moller 28!, LISJC!, LISU!, P!), Mann 621
(P!), 1200 m, Mann s.n. (K!), Moller s.n. (COI!), Rozeira 2846 (PO!).
DISTRIBUTION. Also in Madagascar, Comoros, and Mascarene Is.
Diplazium proliferum (Lam.) Thouars, Esquisse fl. Tristan
d'Acugna: 35 (1804); Alston in Exell, Cat. Vase. PL S. Tome: 73
(1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 237 (1964).
Callipteris prolifera (Lam.) Bory, Voy. ties Afrique 1: 283 (1804);
Benl in Acta Bot. Barcinon. 38: 37 (1988).
SAO TOME. Between Bom Sucesso and Lagoa Amelia, Almeida s.n.
(LISC!); Boa Entrada, Chevalier 14333 (P!), 14361 (P!) 14361bis (not
found); Ribeira Peixe, S. of Cabumbe, secondary forest, 350 m, Espirito
Santo 3916 (COI!, LISJC!); Morro Vigia, 800 m, Espirito Santo 4253
(LISC!, LISJC!); Monte Rosa, secondary forest, 50 m, Espirito Santo 4748
(LISC!, LISJC !);Tras-os-Montes, Lains e Silva 229 (LISU!); between Monte
Castro and Ponta Figo, secondary forest, 500-1070 m,Lejoly95/45 (LISC!);
Dona Amelia, Santa Catarina, near Agua Cardoso, Matos 7593 (LISC!);
Pedroma, Mocquerys 89/90 (P!), 131/132 (P!); Nova Moca, near Monte
Cafe, 90 m, Moller s.n. (COI!); Pinheira, Moller s.n. (COI!); Rodia, 540 m,
Moller s.n. (COI!); Calvario, 1200 m, Monod 11816 (BM!); waterfall near
Sao Nicolau, Monod 12037 (BM!, COI!), Rozeira 571 (PO!); Cruzeiro,
Rozeira 765 (PO!); between Milagrosa and Rio Abade, near Formoso,
Rozeira 3188 (PO!); on the way from Santa Irene to the river, Rozeira 3319
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
(PO!); Sela do Camelo, near Lembd, 300-700 m, Rozeira 3459 (PO!); Monte
Cafe, 760 m, Welwitsch 60 (BM!, K!, LISU!); s.l., Don 49 (BM!), F.A.E.
(Moller) 35 (BM!, COI!, LISJC!, LISU!, P!).
PRINCIPE. Between Oque Pipi and Morro de Leste, secondary forest, 350
m, ground fern, Exell 563 (BM!, COI!); Rio Bambu-Porco, 60 m, Navel 137
(LISU!, P!); Infante D. Henrique, Rose 413 (P!); Infante D. Henrique, near
Ribeiro Chimboto, Rozeira 3 1 2 (PO ! ); Lapa, way to Pico, 300 m, Rozeira 9 1 5
(PO!); near Ribeira Camarao, Rozeira 2213 (PO!); Bela Vista, near main
house, Rozeira 3916 (PO!).
DISTRIBUTION. Palaeotropical.
Blechnaceae
Blechnum L.
Blechnum attenuatum (Sw.) Mett., Fit. hort. hot. Lips.: (A, t. 3, figs
1-6 (1856); Alston in Exell, Cat. Vase. PL S. Tome: 80 (1944);
Tardieu-Blot in Aubrev., Fl. Cameroun 3: 295 (1964); Benl in
Acta Bot. Barcinon. 38: 63 (1988).
SAO TOME. Macambrara to Zampalma, primary forest, 800-1050 m, Exell
437 (BM!, COI!), 438 (BM!, COI!); between Casa do Pico and Pico, 1 250 m,
Mono</11960(BM!).
DISTRIBUTION. Tropical and S. Africa, Madagascar.
Cyatheaceae
Cyathea Sm.
Cyathea camerooniana Hook, in Hook. & Baker, Syn. fil: 21
(1865); Alston in Exell, Cat. Vase. PL S. Tome: 60 (1944).
var. currorii Holttum in Kew Bull. 36: 480 (1981).
Alsophila camerooniana var. camerooniana sensu auct. non (Hook.)
R.M. Tryon in Contr. Gray Herb. 200: 30 (1970); Benl in Acta
Bot. Barcinon. 31: 26 (1978).
PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350
m, Exell 556 (BM!, COI!); Rio Bambu-Porco, 60 m, Navel 136 (COI!, K!,
LISU!, P!, WAG); Infante D. Henrique, Rose 400 (P!); Infante D. Henrique,
near Ribeiro Chimboto, Rozeira 325 (PO!); s.l., Strickland s.n. (BM!, COI-
not found).
DISTRIBUTION. Apparently this variety occurs only in Principe,
where the type (W. Africa, S. of Tropics, Curror s.n. (K!)) was
probably collected.
Cyathea manniana Hook, in Hook. & Baker, Syn. fil.: 21 (1865);
Alston in Exell, Cat. Vase. PL S. Tome: 60 (1944); Holttum in
Kew Bull. 36:472(1981).
Alsophila manniana (Hook.) R.M. Tryon in Contr. Gray Herb. 200:
30 (1970); Benl in Acta Bot. Barcinon. 31: 27 (1978).
SAO TOME. Lagoa Amelia, 1400 m, Espirito Santo 5035 (LISC!, LISJC!),
1380 m, Monod 11763 (BM!), Quintas s.n. (COI!), Rozeira 1150 (PO!);
Macambrara, primary forest, 1050-1200 m, Exell 244 (BM!); Pico, primary
forest, 1900-2020 m, Lejoly 94/577 (LISC!), 1850 m, Moller s.n. (COI!);
between Quinas Alias and Monte Quinas, Monod 12021 (BM!); between
Casa do Pico and Pico, 1950 m, Monod 12242ter (BM!).
DISTRIBUTION. Widespread in tropical Africa.
Cyathea welwitschii Hook, in Hook. & Baker, Syn. fil.: 21 (1865);
Alston in Exell, Cat. Vase. PL S. Tome: 60 (1944); Holttum in
Kew Bull. 36:475(1981).
51
Alsophila welwitschii (Hook.) R.M. Tryon in Contr. Gray Herb. 200:
31 (1970).
SAO TOME. Macambrara, primary forest, 1050-1200 m, Exell \ 1 1 (BM!,
COI!), 233 (BM!, COI!); Bom Sucesso, 1200 m, F.A.E. (Moller) 2 (K!,
LISU!, P!), 1200 m, Moller s.n. (BM!, COI!); Pico, 1500-1850 m, F.A.E.
(Moller) 3 (K!, LISU!), 1850 m, Moller s.n. (COI!); Tris-os-Montes, Gama
s.n. (COI!), 900-1200 m, Henriques s.n. (P!); between Quinas Alias and
Monte Quinas, Monod 12010 (BM!); Monte Cafe\ 600 m, Welwitsch 66
(BM!, K!, LISU!); s.l., Mannl 41/1 (P!), Mann 1 104 (KMectotype).
DISTRIBUTION. Endemic.
Davalliaceae
Davallia Sm.
Davallia chaerophylloides (Poir.) Steud., Nomencl. hot. 2: 146
(1824); Alston in Exell, Cat. Vase. PI. S. Tome: 72 (1944); Alston
in Exell, Suppl. Cat. Vase. PL S. Tome: 1 ( 1 956); Benl in Acta Bot.
Barcinon. 33: 27 (1982).
SAOTOME. Sao Vicente, 200 m, epiphyte, Espirito Santo 107 (BM!, COI!,
LISJC!); Benfica, 350 m, epiphyte, F.A.E.(Moller) 9 (BM!, LISU!); Benfica,
near Trindade, 350 m, Moller s.n. (COI!); Rio do Ouro, 350 m, Moller s.n.
(COI !); near Diana, epiphyte, Rozeira 1 2 1 0 (PO !); s.l., 460-600 m, Welwitsch
56 (BM!, K!, LISU!).
Widespread in tropical and S. Africa and Mascarene
DISTRIBUTION.
Is.
NOTE. Nooteboom ( 1 994) synonymizedDava///a chaerophylloides
with D. denticulata (Burm.f.) Mett. ex Kuhn var. denticulata.
Humata Cav.
Humata repens (L.f.) Diels in Engl. & Prantl, Nat. Pflanzenfam.
1(4): 209 (1899); Alston in Exell, Cat. Vase. PL S. Tome: 71
(1944).
SAO TOME. Sao Miguel, Newton s.n. (P!).
DISTRIBUTION. Palaeotropical.
NOTES. Nooteboom (1994) included Humata in Davallia [D.
repens (L.f.) Kuhn].
This taxon has not been recorded in the islands since the nine-
teenth century.
Dennstaedtiaceae (incl. Hypolepidaceae)
Blotiella R.M. Tryon
Blotiella currorii (Hook.) R.M. Tryon in Contr. Gray Herb. 191: 99
(1962); Benl in Acta Bot. Barcinon. 38: 29 (1988).
Lonchitis currorii (Hook.) Mett. ex Kuhn in Von der Decken's
Reisen Ost-Afr., Bot. 3: 10 (1879); Alston in Exell, Cat. Vase. PI.
S. Tome: 85 ( 1944); Alston in Exell & Rozeira in Conf. Int.Afr. Oc.
3: 79 (1958); Alston in Exell in Bull. Inst. Franc. Afrique Noire
ser. A, 21: 441 (1959); Tardieu-Blot in Aubrev., Fl. Cameroun 3:
100(1964).
SAO TOME. Nova Moca, 1 100 m, Quintas 6 (K!, LISU!); Lagoa Amelia,
1400 m, Quintas 1262 p.p. (P!), 1400 m, Quintas 1455 (BM!, COI!, K!); s.l.,
Quintas 1287 p.p. (P!).
PRINCIPE. Infante D. Henrique, Lains e Silva 458 (LISU!), 459 (LISU!);
Pico, 500-600 m, Monod 12109 (BM!), Rozeira 913 (PO).
DISTRIBUTION.
bia.
W. tropical Africa, extending to Sudan and Zam-
52
E. FIGUEIREDO
NOTE. Newton s.n. (P!), from Principe, cited but not seen by
Alston (1944), consists of sterile young fronds.
Blotiella glabra (Bory) R.M. Tryon in Contr. Gray Herb. 191: 99
(1962); Benl in Acta Bot. Barcinon. 38: 31 (1988).
Lonchitis glabra Bory, Voy. ties Afrique 1: 321 (1804); Alston in
Exell, Cat. Vase. PL S. Tome: 85 (1944).
Lonchitis gracilis Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 1
(1956); Tardieu-Blot in Aubrev., Fl. Cameroun 3:102 (1964).
SAO TOME. Porto Alegre, Chevalier 14311 (P!); Pico, 1700-1900 m,
F.A.E. (Moller) 13 (BM!, COI!, LISJC!, LISU!, P!), Mann s.n. (K!), 1750-
1900 m, Moller s.n. (COI!); Lagoa Amelia, 1400 m, Quintas 1262 (BM!).
DISTRIBUTION. Tropical and S. Africa, Madagascar and Reunion.
Blotiella mannii (Baker) Pic. Serm. in Webbia 31: 250 (1977); Benl
in Acta Bot. Barcinon. 38: 30 (1988).
Lonchitis mannii (Baker) Alston in Bol. Soc. Brot., ser. 2, 30: 18
(1956); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 80
(1958).
SAO TOME. Macambrara, primary forest, 1050-1200 m, ground fern, Exell
239 (BM!, COI!), Quintas 1262A (BM!); Tras-os-Montes, Quintas 1287
(BM!, COI!): s.l., Quintas 1262 p.p. (COI!, P!).
PRINCIPE. Pico, Rozeira 903 (BM-not found, PO!), 800 m, Rozeira 912
(K!), 918 (K!, PO!), 600 m, Rozeira 1075 (K!).
DISTRIBUTION. W. tropical Africa.
Histiopteris (J.G. Agardh) J. Sm.
Histiopteris incisa (Thunb.) J. Sm., Hist.fil: 295 (1875); Alston in
Exell, Cat Vase. PL S. Tome: 86 (1944); Benl in Acta Bot.
Barcinon. 38: 25 (1988).
SAO TOME. Near Sao Pedro, F.A.E. (Moller) 19 (BM!, LISU!); Macambrara,
1280m, Moller s.n. (COI!).
PRINCIPE. Oque Nazare, Newton s.n. (COI!).
DISTRIBUTION. Pantropical.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Hypolepis Bernh.
Hypolepis sparsisora (Schrad.) Kuhn, Filic. afr.: 120 (1868);
Alston in Exell, Cat. Vase. PL S. Tome: 82 (1944); Benl in Acta
Bot. Barcinon. 38: 27 (1988).
SAO TOME. Macambrara, primary forest, 1050-1200 m, Exell 1 12 (BM!,
COI!), 1200 m, Quintas 4 (COI!-also with no. 1354, K!), 1200 m, Quintas
1354 (BM!, COI!, LISJC!); Tras-os-Montes, Gama s.n. (COI!).
DISTRIBUTION. Tropical and S. Africa, Madagascar, Comoros and
Mascarene Is.
Lonchitis L.
Lonchitis occidentalis Baker in Hook. & Baker, Syn. fil: 128
(1867); Benl in Acta Bot. Barcinon. 38: 34 (1988).
Anisosorus occidentalis (Baker) C. Chr., Cat. pi. mad., pter.: 54
(1932); Alston in Exell, Cat. Vase. PL S. Tome: 85 (1944).
SAO TOME. Macambrara, 1290 m, Moller s.n. (COI!); Sao Joao dos
Angolares, 200 m alt, Quintas s.n. (COI!); s.l., F.A.E (Moller & Quintas) 14
(BM!, K also Moller 1 1 !, LISU!).
DISTRIBUTION. Widespread in tropical and SE Africa, Madagascar.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Microlepia C. Presl
Microlepia speluncae (L.)T. Moore, Index fil.: 93 (1857); Alston in
Exell, Cat Vase. PL S. Tome: 72 (1944); Benl in Acta Bot.
Barcinon. 38: 23(1988).
SAO TOME. Ilheu das Rolas, Quintas s.n. (BM !, COI!); Monte Cafe, 760 m,
Welwitsch 56(2) (BM!, LISU!); s.l., Don s.n. (BM!), F.A.E. (Moller &
Quintas) 10 (K-not found, LISU!), 760 m, Welwitsch 56 (K!, LISU!).
DISTRIBUTION. Palaeotropical, rare and sporadic.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Odontosoria Fee
Odontosoria chinensis (L.) J. Sm. in Seem., Bot. voy. Herald: 430
(1857).
van divaricata H. Christ in J. Bot. (Morot), ser. 2, 2: 23 (1909).
Stenoloma chinense (L.) Bedd., Handb. ferns Brit. Ind.: 70 (1883)
var. divarication (H. Christ) Alston in Exell, Cat Vase. PL S.
Tome: 73 (1944).
Sphenomeris chusana (L.) Copel. in Bernice P. Bishop Mus. Bull.
59: 69 (1929) var. divaricata (H. Christ) Tardieu in Humbert, Fl.
Madag. et Comoros 1, fam. 5: 29 (1958).
Sphenomeris chinensis (L.)Maxonin/ Wash. Acad. Sci. 3: 144(1913)
var. divaricata (H. Christ) Kramer in Blumea 15: 572 (1968).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14309 (P!-type of
the var.); Calvario, 1500 m, ground fern, Exell 321 (BM!, COI!); Pico, 1500-
\900m, F.A.E. (Moller) 12 (BM!, COI!, LISJC!, LISU!), 1900m,Mo//ers.n.
(COI!), Newton s.n. (BM!, COI!); Tras-os-Montes, Gama s.n. (COI!); be-
tween Calvario and Esta?ao Sousa, primary forest, 1500 m, Matos & Van
Essche 7357 (LISC!).
DISTRIBUTION. Occurs also sporadically in tropical Asia and
Oceania, and in Madagascar, Reunion, Mauritius and Seychelles.
NOTE. Kramer (1972) suggested that the collectionsMo//e/; Gama,
Exell, Newton and F.A.E. (Moller), cited above, might represent an
undescribed species of Sphenomeris (= Odontosoria), noting that
more material and field studies are required to resolve the problem.
Pteridium Gled. ex Scop.
Pteridium aquilinum (L.) Kuhn in Von der Decken's Reisen Ost-
Afr., Bot. 3: 1 1 (1879); Alston in Exell, Cat. Vase. PL S. Tome: 86
(1944); Benl in Acta Bot. Barcinon. 38: 35 (1988).
SAO TOME. Boa Entrada, Chevalier 13791 (P); between Macambrara and
Sao Nicolau, secondary woodland, 1050 m, common, Exell 272 (BM!,
COI!); Nova Moca, 800-900 m, F.A.E. (Moller) 17 (LISU!); between Morro
Vigia and Guarda, Rozeira 688 (PO!); Pico, 1900 m, Monod 1 1920 (BM!);
Monte Cafe, 760 m, Welwitsch 59 (BM!, LISU!); s.l., Moller s.n. (COI!),
Welwitsch 57 (LISU!).
PRINCIPE. Cajamanga, Rozeira 633 (PO!); Pico Papagaio, Welwitsch 18
(BM!, LISU!).
DISTRIBUTION. Cosmopolitan.
Elaphoglossaceae
Elaphoglossum Schott ex J. Sm.
Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe in J. S.
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
53
African Bot. 30: 196 (1964); Schelpe in Contr. Bolus Herb. 1: 29
(1969); Benl in Acta Bot. Barcinon. 40: 71 (1991).
SAO TOME. Macambrara, Exell 153 (BM!); Calvario, Monod 11846
(BM !); Pico, Rose 311 (P!).
DISTRIBUTION. Widespread in tropical and subtropical Africa,
Madagascar, Comoros and Reunion.
Elaphoglossum aubertii (Desv.) T. Moore, Index fit. 2: 5 (1857);
Alston in Exell, Cat. Vase. PL S. Tome: 91 (1944); Schelpe in
Contr. Bolus Herb. 1: 32 (1969); Benl in Acta Bot. Barcinon. 40:
75(1991).
SAO TOME. Pico, 1900 m, F.A.E. (Moller) 63 (K also Moller 561, LISU!),
1900 m, Moller s.n. (BM!, COI!); Calvario, 1550 m, Monod 1 1841 (BM!).
DISTRIBUTION. Widespread in tropical and subtropical Africa,
Madagascar, Comoros and Mascarene Is.
Elaphoglossum chevalieri H. Christ in 7. Bot. (Morot), ser. 2, 2: 23
(1909); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 8 (1956);
Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 447
(1959); Schelpe in Contr. Bolus Herb. 1: 33 (1969).
Elaphoglossum clarenceanum sensu auct. non (Baker) C. Chr.,
Index filic.: 304 (1905); Alston in Exell, Cat. Vase. PL S. Tome: 92
(1944).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14292bis (P!-type);
Pico, 1950 m, Moller s.n. (COI!), Rose 298 (P!); Calvario, 1550 m, Monod
1 1837 (BM!); near Casa do Pico, 1930 m, Monod 1 1953 (BM!); E. of Pico,
1700 m, Quintas 1379 (BM !, COI!); s.l., F.A.E. (Moller)64 (LISU!), Henriques
5/84 p.p. (K also labelled F.A.E. (Moller) 64!).
PRINCIPE. Pico, 700-800 m, Monod 12134 (BM!, COI!), above 300 m,
77zoro«2087(BM!).
DISTRIBUTION. W. tropical Africa, SE Africa.
Elaphoglossum isabelense Brause in Brause & Hieron. in Bot.
Jahrb. Syst. 53: 432 (1915); Alston in Exell, Cat. Vase. PL S.
Tome: 92 (1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 297
(1964); Benl in Acta Bot. Barcinon. 40: 73 (1991).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14540 (not found);
Lagoa Amelia, Exell 221 (BM!), Rose 195 (P!); Macambrara, Exell 227
(BM!); Calvario, Monod 1 1823 (BM!); Rio Contador, 1450 m, Quintas 1356
(BM!, P!); s.l., F.A.E. (Moller) 62 (BM!, LISU!, P!), Henriques 5/84 p.p.
(K!).
PRINCIPE. S.l., Barter 1903 (K!).
DISTRIBUTION. W. tropical Africa.
Elaphoglossum salicifolium (Willd. ex Kaulf.) Alston in Exell,
Cat. Vase. PL S. Tome: 92 (1944); Tardieu-Blot in Aubrev., FL
Cameroun 3: 302 (1964); Benl in Acta Bot. Barcinon. 40: 73
(1991).
Elaphoglossum petiolatum (Sw.) Urb., Symb. antill. 4: 61 (1903)
subsp. salicifolium (Willd. ex Kaulf.) Schelpe in Contr. Bolus
Herb. 1: 34(1969).
SAO TOME. Sao Joao dosAngolares, F.A.E. (Quintas) 61 p.p. (BM!); Pico,
Rose 315 (P!).
DISTRIBUTION. Tropical Africa, Madagascar, Comoros, Sey-
chelles and Mascarene Is.
Gleicheniaceae
Dicranopteris Bernh.
Dicranopteris linearis (Burm.f.) Underw. in Bull. Torrey Bot. Club
34: 250 (1907); Benl in Acta Bot. Barcinon. 31: 23 (1978).
Gleichenia linearis (Burm.f.) C.B. Clarke in Trans. Linn. Soc.
London, Bot. 1: 428 (1880); Alston in Exell, Cat. Vase. PL S.
Tome: 93 (1944); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 8
(1956).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14310 (P); Famosa,
400 m, Espirito Santoll (BM!, COI!, LISJC!); between Java and Bombaim,
secondary forest, 520 m, Espirito Santo 501 3 (LISC!, LISJC!); Macambrara
to Zampalma, 1050 m, ground fern, Exell 298 (BM!, COI!); Sao Nicolau, 900
m, Figueiredo & Arriegas 52 (LISC!); Jou, Lains e Silva 331 (LISU!);
between Monte Castro and Ponta Figo, secondary forest, 500-1070 m,Lejoly
95/35 (LISC!); Pico, summit, Mann (K!), 1800 m, Moller s.n. (COI!);
between Calvario and Esta^ao Sousa, primary forest, 1500 m, Matos & Van
Essche 7358 (LISC!); Rio Contador, 1325 m, Moller s.n. (COI!); Calvano,
1 100 m, Monod 11810 (BM!, COI!); Agua Ize, common, Pinto Basto 198
(LISC!); Nova Moca, 750 m, Quintas 1465 (BM!, COI!, K!); between Vale
Carmo and Guaiaquil, Rozeira 18 (PO!); Tras-os-Montes, s.c. s.n. (COI!);
s.l., F.A.E. (Moller) 1 (BM!, COI!, LISJC!, LISU!).
PRINCIPE. Infante D. Henrique, 200 m, common ground fern, Exell 654
(BM!, COI!), 240 m, Lains e Silva 467 (LISU!); Oque Nazare, Newton s.n.
(BM!, COI!); Pico Papagaio, 760 m, Welwitsch 19 (BM!, LISU!).
DISTRIBUTION. Pantropical.
Grammitaceae
Grammitis Sw.
Grammitis ebenina (Maxon) Tardieu in Mem. Inst. Franc. Afrique
Noire 28: 21 1, t. 42, figs 1 1-12 (1953); Schelpe in Contr. Bolus
Herb. 1: 3 (1969).
Polypodium ebeninum Maxon in Bull. Torrey Bot. Club 42: 224
(1915); Alston in Exell, Cat. Vase. PL S. Tome: 88 (1944).
SAO TOME. Cabumbe, 800 m, epiphyte, Newton s.n. (P!).
DISTRIBUTION. Also in St. Helena.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Grammitis molleri (Baker) Schelpe in Bol. Soc. Brot., ser. 2, 40:
162 (1966); Schelpe in Contr. Bolus Herb. 1: 5 (1969).
Polypodium molleri Baker in Henriq. in Bol. Soc. Brot. 4: 154, t. 2,
fig. B (1887); Alston in Exell, Cat. Vase. PL S. Tome: 88 (1944).
SAO TOME. Pico, 1900 m, F.A.E. (Moller) 5\ (BM!, COI!, LISJC!, LISU!,
P!); s.l. (probably Pico), Moller 64 (K-holotype).
DISTRIBUTION. Endemic.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Grammitis nigrocincta Alston in Exell in Bull. Inst. Franc. Afrique
Noire ser. A, 21: 444 (1959); Schelpe in Contr. Bolus Herb. 1: 3
(1969).
Grammitis sp.; Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 80
(1958).
PRINCIPE. Pico, 600 m, Thorold 2074 (BM !), Monod 1 2 1 62 (BM !-holotype,
COl\), Rozeira 921 (PO).
54
E. FIGUEIREDO
DISTRIBUTION. Also in Madagascar.
Grammitis tomensis Schelpe in Contr. Bolus Herb. 1: 6 (1969).
SAO TOME. Pico, 2000 m, Quintas 1 1 (BM!-holotype, COI!); s.l. (prob-
ably Pico), Newton 1 1 p.p. (K).
DISTRIBUTION. Endemic.
NOTES. At BM there is another collection labelled Quintas 1 1
which consists of Pleris tripartita.
This taxon has not been recorded in the islands since the nine-
teenth century.
Xiphopteris Kaulf.
Xiphopteris cultrata (Willd.) Schelpe in Bol. Soc. Brot, ser. 2, 41:
217 (1967); Schelpe in Contr. Bolus Herb. 1: 9 (1969); Benl in
Acta Bot. Barcinon. 33: 22 (1982).
Polypodium cultratum Willd., Sp. pi. 5: 187 (1810) van elasticum
(Bory ex Willd.) Baker in Hook. & Baker, Syn.fil.: 327 (1867);
Alston in Exell, Cat. Vase. PL S. Tome: 87 (1944).
SAO TOME. Macambrara to Zampalma, primary forest, 800-1 050 m, Exell
441 (BM!); between Bom Sucesso and Lagoa Amelia, epiphyte, Matos 7585
(LISC!); Lagoa Amelia, primary forest, 1400-1420 m, epiphyte, Matos &
Van Essche 7412 (LISC!).
DISTRIBUTION. W. and Central Africa and tropical America.
Xiphopteris oosora (Baker) Alston in Bol. Soc. Brot., ser. 2, 30: 26
(1956); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 325 (1964);
Schelpe in Contr. Bolus Herb. 1: 12 (1969); Benl in Acta Bot.
Barcinon. 33: 21 (1982).
Polypodium oosorum Baker in Henriq. in Bol. Soc. Brot. 4: 154, t. 2,
fig. A (1887); Alston in Exell, Cat. Vase. PL S. Tome: 89 (1944).
SAO TOME. Pico, 1950 m, Moller s.n. (BM!, COI!); Pico, 1950 m, F.A.E.
(Moller) 52 (BM!, K?, LISU!, P!); s.l. (probably Pico), Moller 1337 (K-
holotype).
DISTRIBUTION. W. and E. tropical Africa, Madagascar.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Xiphopteris villosissima (Hook.) Alston in Bol. Soc. Brot., ser. 2,
30: 27 (1956); Schelpe in Contr. Bolus Herb. 1: 8 (1969); Benl in
Acta Bot. Barcinon. 33: 23 (1982).
Polypodium villosissimum Hook., Sp. fil. 4: 197 (1862); Alston in
Exell, Cat. Vase. PL S. Tome: 90 (1944).
SAO TOME. S.l., Mann s.n. (K), Newton (not found at COI).
DISTRIBUTION. W., E., and SE tropical Africa.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Hemionitidaceae
Coniogramme Fee
Coniogramme africana Hieron. in Hedwigia 57: 293 (1916); Als-
ton in Exell, Cat. Vase. PL S. Tome: 81 (1944); Tardieu-Blot in
Aubrev., FL Cameroun 3: 132 (1964); Benl in Acta Bot. Barc-
inon. 32: 26 (1980).
SAO TOME. Macambrara to Zampalma, 900 m, ground fern, Exell 292
(BM!); Macambrara, primary forest, 1050-1200 m, ground fern, Exell 421
(BM!, COI!); Pico, 1800 m, Moller s.n. (COI!); W. of Pico, 1225 m, Monod
1 1994 (BM!); NW of Pico, 1225 m, Monod 1 1995 (BM!); Lagoa Amelia,
Rozeira 1 182 (PO!); s.l., F.A.E. (Moller) 58 (BM!, LISU!), Mann s.n. (K!),
Moller s.n. (COI!).
DISTRIBUTION. Widespread in tropical Africa.
Pityrogramma Link
Pityrogramma calomelanos (L.) Link, Handbuch 3: 20 (1833)
'calomelas'; Alston in Exell, Cat. Vase. PL S. Tome: 81 (1944);
Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 442
(1959).
van calomelanos; Benl in Acta Bot. Barcinon. 32: 30 (1980).
SAO TOME. Between Bom Sucesso and Macambrara, Matos 761 1 (LISC!);
Agua Ize, Pinto Basto 215 (LISC!); near Bombaim, Rozeira 499 (PO!).
PRINCIPE. Santo Antonio, Monod 12064 (BM!, COI!); s.l., Barter 1912
(K-not found).
DISTRIBUTION. A weed introduced from tropical America.
Hymenophyllaceae
Hymenophyllum Sm.
Hymenophyllum capillare Desv. in Mem. Soc. Linn. Paris 6: 333
(1827); Alston in Exell, Cat Vase. PL S. Tome: 59 (1944);
Tardieu-Blot inAubrev., Fl. Cameroun 3: 75 (1964); Benl inActa
Bot. Barcinon. 32: 7 (1980).
SAO TOME. Pico, 1 500-2000 m, Moller s.n. (BM!);W. of Pico, 1225-1430
m, Monod 11913 (BM!, COI!); Calvario, Rose 155 (P!); s.l., F.A.E. (Moller
& Quintas) 5 p.p. (K also Moller 4!, P!).
DISTRIBUTION. Widespread in tropical Africa, Mascarene Is. and
Tristan da Cunha.
Hymenophyllum hirsutum (L.) Sw. in J. Bot. (Schrader) 1800(2):
99 (1801), emend. C.V. Morton in Contr. U.S. Nad. Herb. 29:
155, 173 (1947); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 76
(1964); Benl in Acta Bot. Barcinon. 32: 9 (1980).
Hymenophyllum ciliatum Sw. in J. Bot. (Schrader) 1800(2): 100
(1801) var. boryanum (Willd.) Mett. ex Kuhn, Filic. afr.: 39
(1868); Alston in Exell, Cat. Vase. PL S. Tome: 59 (1944).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14292 p.p. (P!);
Calvario, 1500 m, ground fern, Exell 327 (BM!, COI!), 1560 m, Monod
1 1835 (BM!, COI!); Lagoa Amelia, primary forest, 1400-1420 m, epiphyte,
Matos & Van Essche 7414 (LISC!); Pico, 1500-2000 m, Moller s.n. (BM!,
COI!); Bom Sucesso, 1 150 m, Moller & Quintas s.n. (COI!); Binda, Newton
(P-specimen is missing from sheet); Sao Nicolau, 900 m, Quintas s.n.
(LISU!); s.l., F.A.E. (Moller & Quintas) 5 p.p. (BM!, COI!, K!, LISJC!,
LISU!, P!), Quintas s.n. (K!).
DISTRIBUTION. Pantropical and subtropical.
Hymenophyllum polyanthos Sw. in J. Bot. (Schrader) 1800(2):
102(1801).
var. kuhnii (C. Chr.) Schelpe in Bol. Soc. Brot., ser. 2, 40: 156
(1966); Benl in Acta Bot. Barcinon. 32: 6 (1980).
Hymenophyllum kuhnii C. Chr., Index filic.: 363 (1905); Alston in
Exell, Cat. Vase. PL S. Tome: 59 (1944); Alston in Exell & Rozeira
in Conf. Int. Afr. Oc. 3: 78 (1958); Alston in Exell in Bull. Inst.
Franc. Afrique Noire ser. A, 21: 441 (1959); Tardieu-Blot in
Aubrev., FL Cameroun 3: 72 (1964).
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
55
SAO TOME. Between Monte Cafe and Pico, Chevalier 14292 p.p. (P!),
14292ter(BM!, P!), 14551 (P!); SW region, Chevalier 14588 (P!); Calvario,
1 500 m, ground fern, Exell 323 (BM !); between Lagoa Amelia and Change,
primary forest, 1 250 m, epiphyte, Matosl534 (LISC!); Pico, 1950 m, Moller
s.n. (COI!), Rose 316 (P!), 319 (P!), Rozeira 967 (PO!); Tras-os-Montes,
800- 1 200 m, Quintas 1315 (BM !, COI-not found); s.l., F.A.E. (Moller) 4 p.p.
(BM!, COI!, K also Moller 3!, LISJC!, LISU!, P!).
PRINCIPE. W. of Pico, 500-600 m,Monod\2\ 13 (BM!); Pico, 500-600 m,
Monod 121 16 (BM!), 800-900 m, Rozeira 973 (BM-not found, PO), 600 m,
epiphyte, Thorold 2073 A (BM!).
DISTRIBUTION. Widespread in tropical Africa.
Hymenophyllum polyanthos Sw. in J. Bot. (Schroder) 1800(2):
102(1801).
var. mildbraedii (Brause ex Brause & Hieron.) Schelpe in Garcia de
Orta, ser. Bot. 3: 54(1976).
Hymenophyllum mildbraedii (Brause ex Brause & Hieron.) Alston
in Exell, Cat. Vase. PL S. Tome: 60 (1944); Alston in Exell in Bull.
Inst. Franc. Afrique Noire ser. A, 21: 441 (1959).
PRINCIPE. Pico, Rozeira 965 (K!), 600 m, epiphyte, Thorold 2073 p.p.
(BM!), 900 m, Thorold 2084 p.p. (BM!).
DISTRIBUTION. Angola and Annobon.
Hymenophyllum splendidum Bosch in Ned. Kruidk. Arch. 5(3):
192 (1863); Alston in Exell, Cat. Vase. PL S. Tome: 60 (1944);
Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 441
(1959); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 76 (1964);
Benl in Acta Bot. Barcinon. 32: 8 (1980).
SAO TOME. Macambrara, primary forest, 800-1050 m, Exell 439 (BM!,
COI!); s.l., F.A.E. (Moller & Quintas) 5 p.p. (P!).
PRINCIPE. Pico,Rozeira 966 (K!), 600 m, epiphyte, Thorold 2013B (BM!),
900 m, epiphyte, Thorold 2084A (BM!).
DISTRIBUTION. Tropical Africa, extending to Tanzania.
Trichomanes L.
NOTE. Recent work (Dubuisson, 1996) supports the division by
earlier workers of the genus Trichomanes s.l. into several genera.
Although the genus is treated here in its sensu lato, the names of the
taxa in the segregate genera are also provided, when available.
Trichomanes borbonicum Bosch in Ned. Kruidk. Arch. 5(2): 158
(as '185') (1861); Alston in Exell, Cat. Vase. PL S. Tome: 57
(1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 88 (1964);
Benl in Acta Bot. Barcinon. 32: 22 (1980).
[Vandenboschia borbonica (Bosch) G. Kunkel in Nova Hedwigia
6:213(1963)]
SAO TOME. Pico, 1 500-2000 m,Moller s.n. (BM!,COI!);Tras-os-Montes,
1200 m, Quintas 1269A (BM!, COI!).
DISTRIBUTION. Widespread in tropical Africa, SE and S. Africa,
Madagascar and Reunion.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Trichomanes crispiforme Alston in Exell, Cat. Vase. PL S. Tome:
57 (1944); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 78
(1958); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 90 (1964).
SAO TOME. Cabumbe, Newton (P?).
PRINCIPE. Oque Pipi, secondary forest, 300 m, epiphyte, Exell 501 (BM!,
COI!); W. of Pico, 500-600 m, Monod 12123 (BM!), 700-800 m, Monod
12143 (BM!); Pico, Rozeira 905 (BM-not found, PO); s.l., Barter 1918 (K-
not found).
DISTRIBUTION. W. and Central tropical Africa.
Trichomanes digitatum Sw., Syn.fil: 370 (1806); Alston in Exell
in Bull Inst. Franc. Afrique Noire ser. A, 21: 441 (1959).
[Microtrichomanes digitatum (Sw.) Copel. in Philipp. J. Sci. 67:
36(1938)]
SAO TOME. Calvario, 1400 m, Monod 1 1855 (BM!, COI!).
DISTRIBUTION. Also in Madagascar, Mauritius, Reunion, Malaya
and Borneo.
Trichomanes erosum Willd., Sp. pi. 5: 501 (1810); Alston in Exell,
Cat. Vase. PI. S. Tome: 58 (1944); Benl in Acta Bot. Barcinon. 32:
16(1980).
[Microgonium erosum (Willd.) C. Presl, Gefassbiindel Farm: 21
(1847)]
Trichomanes chamaedrys Taton in Bull. Soc. Roy. Bot. Belgique 78:
29, pi. 2, figs K-L (1946); Alston in Exell & Rozeira in Conf. Int.
Afr. Oc. 3:78(1958).
SAO TOME. Porto Alegre, Chevalier 14178 (P); Pico, 1900 m, Moller s.n.
(COI!); Sao Joao dosAngolares, &Om,Newton s.n. (BM!, COI!); Macambrara,
1200 m, epiphyte, Quintas 1269B (BM!, COI!); Bom Sucesso, Quintas s.n.
(BM!, COI!); Ribeira Peixe, Rozeira 205 (BM-not found, PO); s.l., F.A.E.
(Moller & Quintas) 6 (K!, LISU!), Quintas 3 (COI!).
PRINCIPE. S.l., Barter 1919 (K!).
DISTRIBUTION. Tropical Africa.
Trichomanes mannii Hook, ex Hook. & Baker, Syn.fil.: 75 (1867);
Alston in Exell, Cat. Vase. PL S. Tome: 58 (1944); Benl in Acta
Bot. Barcinon. 32: 17 (1980).
[Gonocormus mannii (Hook, ex Hook. & Baker) G. Kunkel in
Nova Hedwigia 6: 212 (1963)]
SAO TOME. Lagoa Amelia, Exell 203 (BM!, COI!); between Sao Pedro and
Lagoa Amelia, 1050 m, Monod 11706 (BM!); Tras-os-Montes, 850 m,
Quintas 1269 (BM!, COI!).
DISTRIBUTION. W., Central and E. Africa, Madagascar and
Comoros.
Trichomanes pyxidiferum L., Sp. pi. 2: 1098 (1753).
[Vandenboschia pyxidifera (L.) Copel. in Philipp. J. Sci. 67: 53
(1938)]
var. melanotrichum (Schltdl.) Schelpe in/ 5. African Bot. 30: 181
(1964); Benl in Acta Bot. Barcinon. 32: 24 (1980).
Trichomanes melanotrichum Sch\td\.,Adumbr.pL: 56 (1832); Alston
in Exell, Cat. Vase. PL S. Tome: 58 (1944).
SAO TOME. Sao Nicolau, 900 m, F.A.E. (Moller) 1 (BM!, K!, LISU!), 950
m, Moller s.n. (COI!); Lagoa Amelia, Rozeira 1084 (K!); s.l. (probably Sao
Nicolau), F.A.E. (Moller) 4 p.p. (P!).
DISTRIBUTION. Widespread in tropical and subtropical Africa.
Trichomanes radicans Sw. in J. Bot. (Schroder) 1800(2): 97
(1801); Benl in Acta Bot. Barcinon. 32: 14 (1980).
[Vandenboschia radicans (Sw.) Copel. in Philipp. J. Sci. 67: 54
(1938)]
Trichomanes giganteum sensu auct. non Bory ex Willd., Sp. pi. 5:
514 (1810); Alston in Exell, Cat. Vase. PL S. Tome: 58 (1944).
SAO TOME. Macambrara, primary forest, 1050- 1200m, ground fern, Exell
56
E. FIGUEIREDO
185 (BM!, COI!), Moller s.n. (COI!); near Sao Tome, 20 m, Mailer s.n.
(COI!); Pico, 1800 m, Moller s.n. (COI!), Newton s.n. (BM!, COI!); Sao
Pedro, Quintas s.n. (COI!); s.l., F.A.E. (Moller & Quintas) 8 (BM-not found,
COI!, LISU!, P!), Mann s.n. (K!).
DISTRIBUTION. W., Central and E. Africa, Madagascar, Comoros
and Reunion.
Trichomanes rigidum Sw., Prodr.: 137 (1788).
[Selenodesmium rigidum (Sw.) Copel. in Philipp. J. Sci. 67: 81
(1938)]
Trichomanes cupressoides Desv. in Mem. Soc. Linn. Paris 6(2): 330
(1827); Alston in Exell, Cat. Vase. PL S. Tome: 57 (1944); Alston
in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 78 (1958).
SAO TOME. Maria Femandes, secondary forest, 275 m, Carvdho 2
(LISC!); Sao Miguel, Monte Verde, secondary forest, 10-150 m, Lejoly 95/
96 (LISC!).
PRINCIPE. Oque Nazare, Newton s.n. (BM!, COI!); Pico, above Neves
Ferreira, Rozeira 650 (BM-not found, PO), 683 (BM-not found, PO); Pico,
Rozeira 907 (BM-not found, PO).
DISTRIBUTION. Widespread in tropical and subtropical Africa,
tropical America.
Lomariopsidaceae
Bolbitis Schott
Bolbitis acrostichoides (Afzel. ex Sw.) Ching in C. Chr., Index filic.
Suppl. 3: 47 (1934); Alston in Exell, Cat. Vase. PI. S. Tome: 68
(1944); Hennipman, Monogr. fern genus Bolbitis: 149 (1977);
Benl in Acta Bot. Barcinon. 40: 61 (1991).
SAO TOME. Near Sao Tome, F.A.E. (Moller) 67 (K also Moller 41!,
LISU!); Monte Cafe, 800 m, Moller s.n. (COI!); Welwitsch 63b (BM!,
LISU!); Sao Pedro, 1 100 m, Quintas s.n. (COI!).
DISTRIBUTION. Tropical Africa, Madagascar.
NOTES. Moller 63 (K), cited by Hennipman (1977), is probably a
mistake for F.A.E. (Moller) 67.
This taxon has not been recorded in the islands since the nine-
teenth century.
Bolbitis auriculata (Lam.) Ching in C. Chr., Index filic. Suppl. 3: 47
(1934) et Alston in/ Bot. 72, Suppl. 2: 3 (1934); Alston in Exell,
Cat. Vase. PL S. Tome: 68 (1944); Hennipman, Monogr. fern genus
Bolbitis: 136 (1977); Benl in Acta Bot. Barcinon. 40: 59 (1991).
SAO TOME. Boa Entrada, Chevalier 14335 (P!); Monte Cafe, Moller s.n.
(COI!), 800 m, F.A.E. (Moller) 66 (BM!, LISU!, P!); Diogo Vaz, 150 m,
Quintas s.n. (BM!, COI!, K!, LISJC!, P!); above Monte Cafe, Welwitsch 58
(BM!, LISU!).
PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350
m, ground fern, Exell 572 (BM!); s.l., Barter 1899 (K-not found).
DISTRIBUTION. Tropical Africa, Madagascar, Mascarenes.
Bolbitis fluviatilis (Hook.) Ching in C. Chr., Index filic., Suppl. 3:
48 (1934); Alston in Exell, Cat. Vase. PL S. Tome: 69 (1944);
Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 78 (1958);
Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 447
(1959); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 316 (1964);
Hennipman, Monogr. fern genus Bolbitis: 142 (1977); Benl in
Acta Bot. Barcinon. 40: 60 (1991).
SAO TOME. Sao Joao dos Angolares, 200 m, F.A.E. (Quintas) 68 (BM!,
COI!, LISU!); Rio Abade, 1300 m, Quintas s.n. (COI!).
PRINCIPE. Pico, 700-800 m, Monod 12137 (BM!); between Lapa and
Pico, Rozeira 893 (PO); s.l., Barter 1897 (K), 1897bis (K!).
DISTRIBUTION. W. tropical Africa.
Lomariopsis Fee
Lomariopsis guineensis (Underw.) Alston in/ Bot. 72, Suppl. 2: 5
(1934); Alston in Exell, Cat Vase. PL S. Tome: 81 (1944);
Tardieu-Blot in Aubrev., FL Cameroun 3: 309 (1964); Benl in
Acta Bot. Barcinon. 40: 64 (1991).
SAO TOME. Ilheu das Rolas, 1 20 m, Quintas 1164 (BM!, COI also labelled
no. 35 from Monte Cafe!), F.A.E. (Quintas) 65 p.p. (BM!, COI!, K!-also
Quintas 35); Monte Cafe, Welwitsch 68 (BM!, LISU!).
PRINCIPE. Esperanc,a, 600 m, Rozeira 892 (K!).
DISTRIBUTION. W. tropical Africa to Angola.
Lomariopsis hederacea Alston in J. Bot. 72, Suppl. 2: 5 (1934);
Alston in Exell, Cat. Vase. PL S. Tome: 81 (1944); Benl in Acta
Bot. Barcinon. 40: 70 (1991).
SAO TOME. Macambrara, 1050-1200 m, epiphyte, Exell 189 (BM!),
primary forest, 1050-1200 m, ground fern, Exell 315 (BM!, COI!); Rio
Contador, 1 300 m, Moller s.n. (COI!); s.l., F.A.E. (Moller & Quintas) 65 p.p.
(BM!, COI!), Moller s.n. (COI!).
DISTRIBUTION. W. tropical Africa to Angola, Central Africa.
NOTE. Barter s.n. (K) from Principe was cited by Alston in the
protologue of the species (paratype). Barter 1 899 (K), also from
Principe, was cited by Alston in 1944. The same collection (Barter
1 899) was cited by Alston ( 1 944) under Bolbitis auriculata. Neither
of these collections was found under Lomariopsis hederacea at K.
Barter 1899 (K) is Bolbitis auriculata. It is possible that both
citations refer to Barter 1902 (K), a specimen of L. warneckei.
Lomariopsis warneckei (Hieron.) Alston in J. Bot. 72, Suppl. 2: 6
(1934); Benl in Acta Bot. Barcinon. 40: 70 (1991).
Lomariopsis nigrescens Holttum in Kew Bull. 1939: 627, figs 13 &
14 (1940); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser.
A, 21: 447 (1959).
Lomariopsis hederacea sensu auct. non Alston in J. Bot. 72, Suppl.
2: 5 (1934); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 79
(1958).
PRINCIPE. Pico, above Neves Ferreira, epiphyte, Rozeira 663 (BM-not
found, PO!); Pico, epiphyte, 77zoroW2086 (BM!); s.l., Barter 1902 p.p. (K!).
DISTRIBUTION. E. and SE Africa and Bioko.
NOTE. Barter 1 902 (K) consists of two herbarium sheets. The one
cited above is apparently a young plant (as noted by Holttum on the
herbarium sheet), with pinnae smaller than in typical Lomariopsis
warneckei. The other sheet contains a larger specimen, referred to L.
hederacea by Holttum, but with much larger and separated pinnae.
Although the distinction between this species and the previous
seems to be clear in material from the continent, some specimens
from Sao Tome and Principe show intermediate features.
Loxogrammaceae
Loxogramme (Blume) C. Presl
Loxogramme lanceolata (Sw.) C. Presl, Tent, pterid.: 215, t. 9, fig.
8 (1836); Alston in Exell, Cat. Vase. PL S. Tome: 90 (1944); Benl
in Acta Bot. Barcinon. 33: 24 (1982).
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
SAO TOME. Macambrara, primary forest, 1200 m, epiphyte, Exell 230
(BM!); between Estacao Sousa and Pico, 1700 m, Matos 7551 (LISC!);
between Bom Sucesso and Lagoa Amelia, primary forest, 1350-1420 m,
Matos & Van Essche 7290 (LISC!); Saudade,780 m, Moller s.n. (COI!); NW
of Pico, 1600-1800 m, Monod 12239 (BM!); Tras-os-Montes, Quintas s.n.
(BM!, COI!); Novo Brasil, Monte Mario, Rozeira 4 (PO!); Cruzeiro, 300 m,
77wrt»W2069(BM!).
PRINCIPE. S.I., Barter 1898 (K!).
DISTRIBUTION. Widespread in tropical and subtropical Africa.
Lycopodiaceae
Huperzia Bernh.
Huperzia mildbraedii (Herter) Pic. Serm. in Webbia 23: 163
(1968); Benl in Acta Bot. Barcinon. 31: 10 (1978).
Lycopodium mildbraedii Herter in Hedwigia 49: 90 (1909); Alston
in Exell, Cat. Vase. PL S. Tome: 95 (1944).
SAO TOME. Lagoa Amelia, epiphyte, Espirito Santo 5032 (LISC!,
LISJC!), primary forest, 1400-1420 m, epiphyte, Matos & Van Essche
7409 (LISC!), 1350 m, epiphyte, Moller s.n. (COI!), 1400 m, Monod 1 1734
(BM!); Calvario, 1500 m, Exell 317 (BM!, COI!), 1550 m, Monod 11832
(BM!); Macambrara, primary forest, 1050-1200 m, epiphyte, Exell 427
(BM!, COI!); Pico, 1700-1900 m, F.A.E. (Moller) 75 (BM!, K!, LISU!, P!),
1850 m, epiphyte, Moller s.n. (COI!), Moller s.n. p.p. (COI!), Rose 323
(P!); Bom Sucesso, 1250 m, epiphyte, Moller s.n. (COI!); Tras-os-Montes,
1100 m, Quintas 1321 p.p. (COI!); s.l., F.A.E. (Moller) 74 p.p. (BM!, COI!,
LISU!), 1 100-1400 m, Moller 76 (K!).
PRINCIPE. Pico Papagaio, 350 m, Exell 702 (BM!, COI!); W. of Pico,
below 250 m, Monod 12097 (BM!); Pico, common up to 900 m, Thorold
2083 (BM!).
DISTRIBUTION. W. tropical Africa.
Huperzia ophioglossoides (Lam.) Rothm. in Feddes Repert. Spec.
Nov. Regni Veg. 54: 62 (1944).
SAO TOME. Pico, primary forest, 1900-2020 m, Lejoly 94/538 (LISC!),
1850m, epiphyte, Moller s.n. p.p. (COll), Rose 328 (P!); s.l. (probably Pico),
F.A.E. (Moller) 74 p.p. (LISU!).
DISTRIBUTION. Tropical, SE and S. Africa, Madagascar and
Mascarene Is.
Huperzia phlegmaria (L.) Rothm. in Feddes Repert. Spec. Nov.
Regni Veg. 54: 62 (1944); Benl in Acta Bot. Barcinon. 31: 9
(1978).
Lycopodium phlegmaria L., Sp. pi. 2: 1101 (1753); Alston in Exell,
Cat. Vase. PL S. Tome: 96 (1944).
SAO TOME. Pico, Chevalier 13660 (P); Lagoa Amelia, Chevalier 14307
(P!); Ribeira Peixe, Vila Irene, epiphyte, Espirito Santo 4601 (LISC!); Sao
Joao dos Angolares, F.A.E. (Quintas) 76 (BM!, K!, LISU!, P!), Quintas s.n.
(COI!); Perseveran?a, Seabra 183b (LISU!).
PRINCIPE. Between Terreiro Velho and Infante D. Henrique, plantation,
100 m, Exell 587 (BM!, COI!); Porto Real, Rose 483 (P!).
DISTRIBUTION. Palaeotropical.
Huperzia verticillata (L.f.) Trevis. in Atti Soc. Ital. Sci. Nat. 17:
248 (1874); Benl in Acta Bot. Barcinon. 31: 8 (1978).
Lycopodium verticillatum L.f., Suppl. pi.: 448 (1782); Alston in
Exell, Cat. Vase. PL S. Tome: 96 (1944); Tardieu-Blot in Aubrev.,
Fl. Cameroun3: 12(1964).
SAO TOME. Bom Sucesso, 1200 m, F.A.E. (Moller) 73 (BM!, COI!, K!,
LISU!, P!); Macambrara, 1 100 m, Moller s.n. (COI!).
57
DISTRIBUTION. Tropical, SE and S. Africa, Comoros and Reunion,
tropical America, and Polynesia.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Huperzia warneckei (Herter ex Nessel) Pic. Serm. in Webbia 23:
163(1968).
Lycopodium warneckei (Herter ex Nessel) Alston in Bol. Soc. Brot.,
ser. 2, 30: 20 (1956); Alston in Exell & Rozeira in Conf. Int. Afr.
Oc. 3:81 (1958).
SAO TOME. Between Sao Miguel and Zagaia, primary forest, 170-400 m,
Matos & Van Essche 7247 (LISC!); Tras-os-Montes, 1 100 m, Quintas 1321
p.p. (COI!); Ribeira Peixe, between Vila Jose and Ermelinda, Rozeira 288
(PO!).
DISTRIBUTION. W. tropical Africa.
Lycopodiella Holub
Lycopodiellacernua(L.) Pic. Serm. inWebbia23: 166 (1968); Benl
in Acta Bot. Barcinon. 31: 1 1 (1978).
Lycopodium cernuum L., Sp. pi. 2: 1103 (1753); Alston in Exell,
Cat. Vase. PL S. Tome: 95 (1944); Sobrinho in Portugaliae Acta
BioL, Ser. B, Sist. 3: 393 (1952).
SAO TOME. Porto Alegre, Chevalier 14196 (P); Sao Nicolau, 1000 m,
Espirito Santo4448 (LISC!, LISJC!),900m,Mo/ters.n. (COI!); Macambrara,
primary forest, 1200 m, Exell 94 (BM!); Santa Maria, 1300 m, Exell 196
(BM!,COI!), 1380m, Moller s.n. (COI!); Lagoa Amelia, 1 300 m, Exell 206
(BM!, COI!), crater swamp, 1300m, Exell 209 (BM!), 1412 m, Matos 7504
(LISC!); Tras-os-Montes, Gama s.n. (COI!); road to Monte Mario, Lains e
Silva279 (LISU!); Pico, primary forest, 1900-200 m,L<yo/y94/546 (LISC!);
Bom Sucesso, secondary forest, 1 150 m, Matos & Van Essche 7459 (LISC!);
Saudade, 950 m, Moller s.n. (COI!); above Monte Cafe, 1050 m, Monod
11679 (BM!); between Cascata and Morro Vilela, 700-1000 m, Monod
12202 (BM!); Santelmo, Oliveira & Noronha 6-B (LISU!); Monte Cafe, in
forest and plantations, Oliveira & Noronha s.n. (LISU!); Tras-os-Montes,
between Calvario and main house, Rozeira 527 (PO!); s.l., Campos 16
(COI!), F.A.E. (Moller) 77 (BM!, COI!, K!, LISU!).
PRINCIPE. Oque Pipi, secondary forest, 300 m, Exell 551 (BM!, COI!);
Oque Caspar, Newton s.n. (COI!); Ribeira Jambere, Rozeira 593 (PO!); Pico
Papagaio, 900 m, Welwitsch 10 (BM!, LISU!).
DISTRIBUTION. Cosmopolitan.
Lycopodium L.
Lycopodium clavatum L., Sp. pi. 2: 1101 (1753); Alston in Exell,
Cat. Vase. PL S. Tome: 95 (1944); Tardieu-Blpt in Aubrev., Fl.
Cameroun 3: 16 (1964); Benl in Acta Bot. Barcinon. 31: 10
(1978).
SAO TOME. Pico, 2000 m, Chevalier 13676 (P!), 2000 m. F.A.E. (Moller)
78 (BM!, COI!, K!, LISU!, P!), 1850 m, Monod 1 1919 (BM!, COI!), Rose
320 (P!).
DISTRIBUTION. Cosmopolitan.
Marattiaceae
Marattia Sw.
Marattia fraxinea Sm., PL icon. ined. 2: t. 48 (1790); Alston in
Exell, Cat Vase. PL S. Tome: 94 (1944); Benl in Acta Bot.
Barcinon. 32: 3 (1980); Pichi Sermolli in Webbia 23: 332 (1969).
Marattia robusta Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 8
(1956).
58
SAO TOME. Monte Cafe, primary forest, 1 200 m, Espirito Santo 191 (BM!-
holotype of Marattia robusta, COI!, LISJC!), 760 m, Welwitsch 70 (BM!,
LISU!); Ribeira Peixe, S. of Cabumbe, secondary forest, Espirito Sanfo3899
(COI!, LISJC!); Macambrara, primary forest, 1050-1200 m,Exell 129 (BM!,
COI!), 1250 m, Moller s.n. (COI!); Lagoa Amelia, 1200-1400 m, very
common, Lains e Silva 88 (LISU!), 88a (LISU!), 1350-1380 m, Monod
\ 1771 (BM!, COI!), Rozeira 1091 (PO!); Sao Miguel, Monte Verde, second-
ary forest, 10-150 m, Lejoly 95/88 (LISC!); between Sao Miguel and Zagaia,
primary forest, 170-400 m, Matos & Van Essche 7239 (LISC!); Bom
Sucesso, 1250 m, Moller s.n. (COI!); between Sao Pedro and Lagoa Amelia,
1050 m, Monod 11717 (BM!); Sao Joao dos Angolares, 150 m, Quintas s.n.
(COI!); Porto Alegre, near Cao Pequeno, Rozeira 5 1 (PO!); between Lagoa
Amelia and Calvario, Rozeira 1171 (PO!); Dona Augusta, Rozeira 1179
(PO); s.l., EA.E. (Moller & Quintas) 71 (BM!, COI!, LISU!).
PRINCIPE. S.I., Barter 1916 (K!).
DISTRIBUTION. Palaeotropical.
NOTE. Pichi Sermolli ( 1 969) considered that Marattia robusta was
a strong form of M. fraxinea and that the plants from Sao Tome
might be recognized at infraspecific level. The collection from
Principe is from a smaller plant (M. fraxinea s.s.).
Nephrolepidaceae
Arthropteris J. Sm.
Arthropteris monocarpa (Cordem.) C. Chr., Cat. pi. mad., pter.: 32
(1932); Alston in Exell, Cat. Vase. PL S. Tome: 69 (1944); Alston
in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 445 (1959);
Benl in Acta Bot. Barcinon. 33: 35 (1982).
SAO TOME:. Pico, Campos 77 (COI!), Chevalier s.n. (P!), primary forest,
1987 m, Matos & Van Essche 7385 (LISC!); Macambrara, virgin forest,
1050-1200 m, Exell 121 (BM!), 1200-1350 m, 122a (BM!); Lagoa Amelia,
crater swamp, 1300 m, ground fern, Exell 210 (BM!), 1430 m, epiphyte,
Matos 7494 (LISC!); between Macambrara and Sao Nicolau, secondary
woodland, 1050 m, Exell 403 (BM!, COI!); Tras-os-Montes, Gama s.n.
(COI!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain
rainforest, 1200-1800 m, Lejoly 95/23 (LISC!); Calvario and surroundings,
primary forest, 1600-1620 m, epiphyte, Matos & Van Essche 7349 (LISC!);
Sao Joao dos Angolares, 80 m, Newton s.n. (BM!, COI!); s.l., Moller s.n.
(COI!), F.A.E. (Moller) 38 p.p. (BM!, LISJC!, LISU!, P!).
PRINCIPE. Porto Real, Rose 450 (P!); Pico, 950 m, Thorold 2082 (BM!).
DISTRIBUTION. Widespread in tropical Africa and Mascarene Is.
Arthropteris orientalis (J.F. Gmel.) Posth. in Recueil Trav. Bot.
Neerl. 21: 218 (1924); Alston in Exell, Cat. Vase. PL S. Tome: 70
(1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 116 (1964);
Benl in Acta Bot. Barcinon. 33: 34 (1982).
SAO TOME. Between Monte Cafe and Pico, Chevalier 14562 (P!); Pico,
Moller s.n. (COI-not found); Cruzeiro, 300 m, Thorold 2068 (BM!); s.l.,
EA.E. (Moller) 38 p.p. (BM!, COI!, LISJC!, P!).
PRINCIPE. Rio Bambu-Porco, 60 m, epiphyte, Navel 143 (P!).
E. FIGUEIREDO
DISTRIBUTION. Tropical Africa, tropical Asia, Philippines.
Nephrolepis Schott
Nephrolepis biserrata (Sw.) Schott, Gen.fiL: ad t. 3 (1834); Alston
in Exell, Cat. Vase. PL S. Tome: 70 ( 1 944); Sobrinho inPortugaliae
Acta BioL, ser. B, Sist. 3: 392 (1952); Benl in Acta Bot. Barcinon.
33:36(1982).
SAO TOME. Boa Entrada, Chevalier 13789 (P); between Monte Caf£ and
Pico, Chevalier 14291bis (P); Sao Miguel, 50 m, not frequent, Espirito Santo
4743 (LISC!, LISJC!); Agua Ize, littoral, ground fern, Exell 482 (BM!,
COI!), common epiphyte, Pinto Basto 214 (LISC!); Rio Contador, 1300 m,
Moller s.n. (COI!); Bombaim, Monod 11887 (BM!); W. of Pico, Monod
1 1895 (BM!); between Quinas Alias and Monte Quinas, 520-750 m, Monod
12021 (BM!), 12022 (BM!); Santelmo, secondary forest, very abundant
epiphyte, Oliveira & Noronha 23-A (LISU!); Sao Nicolau, Dependencia
Roca Nova, road to Tras-os-Montes, Oliveira & Noronha s.n (LISU!); Ilheu
das Rolas, Quintas s.n. (COI!); Cruzeiro, 300 m, Thorold 2063 (BM!); s.l.,
EA.E. (Moller) 46 (BM!, COI!, LISU!), Moller s.n. (COI!), Rattray s.n. (E),
Souza s.n. (COI!).
PRINCIPE. Terreiro Velho, plantation, 200 m, ground fern, Exell 525 (BM !,
COI!), 526 (BM!, COI!); s.l., Rattray s.n. (E), Souza s.n. (COI!), Welwitsch
23 (BM!, K!, LISU!), Welwitsch 23b (BM!, LISU!).
DISTRIBUTION. Pantropical.
Nephrolepis pumicicola F. Ballard inKewBulL [1955]: 467 (1955);
Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 445
(1959); Tardieu-Blot in Aubrev., FL Cameroun 3: 108 (1964);
Benl in Acta Bot. Barcinon. 33: 37 (1982).
SAO TOME. Lagoa Amelia, Chevalier 13650 (P!), 1412 m, Matos 7505
(LISC!), 1380 m, Monod 11759 (BM!); Pico, Chevalier 13659ter (P!);
between Monte Cafe and Pico, Chevalier 14581 (P!); s.l., 1500 m, Mann
1 106 (K!), Mann s.n. (P!).
DISTRIBUTION. Cameroun, Bioko.
Nephrolepis undulata (Afzel. ex Sw.) J. Sm. in Bot. Mag. 72,
Comp.: 37 [as '35'] (1846); Alston in Exell, Cat. Vase. PL S.
Tome: 71 (1944); Benl in Acta Bot. Barcinon. 33: 39 (1982).
SAO TOME. Sao Nicolau, 870 m, Espirito Santo 5053 (LISC!, LISJC!);
Macambrara, coffee plantation, 1200 m, epiphyte, Exell 116 (BM!, COI!),
primary forest, 1050-1200 m, epiphyte, Exell 190a (BM!); Lagoa Amelia,
1300 m, ground fern, Exell 212 (BM!); Tras-os-Montes, Gama s.n. (COI!);
Santa Catarina, Lains e Silva 387 (LISU!); between Monte Castro and Ponta
Figo, secondary forest, 500-1070 m, Lejoly 94/39 (LISC!); Pico, primary
forest, 1900-2020 m, Lejoly 94/540 (LISC !); Nova Moca, 850 m, Moller s.n.
(COI!); Saudade, Moller s.n. (COI!); Monte Cafe, 690 m, Monod 11669
(BM!), 600 m, Thorold 2061 (BM!); Bombaim, Monod 11885 (BM!);
between Monte Quinas and Quinas Alias, Monod 12016 (BM!); NW of Pico,
above Cascata, 1000 m, Monod 12267 (BM!); Agua Ize, Pinto Basto 209
(LISC!); s.l., Brentel s.n. (COI!), EA.E. (Moller) 45 (BM!, LISU!), Mann
(not found).
PRINCIPE. Pico, summit, Rozeira 924 (PO!).
DISTRIBUTION. Widespread in tropical Africa and Mascarene Is. DISTRIBUTION. Widespread in tropical and S. Africa.
Arthropteris palisotii (Desv.) Alston inBol. Soc. Brot., ser. 2, 30: 6
(1956); Benl in Acta Bot. Barcinon. 33: 31 (1982).
Arthropteris obliterata sensu auct. non (R. Br.) J. Sm., Ferns Brit,
for.: 163 (1866); C. Chr., Index filic.: 62 (1905); Alston in Exell,
Cat. Vase. PL S. Tome: 70 (1944).
SAO TOME. Sao Joao dos Angolares, 120 m, EA.E. (Quintas) 47 (BM!,
LISU!), 80 m, Newton s.n. (BM!, COI!), 120 m, Quintas s.n. (COI!); Agua
Pinhao,650m, Quintas 1 454 (BM!, COI!); Monte Cafe, 760m, Welwitsch 55
(BM!, K!,LISU!).
Oleandraceae
Oleandra Cav.
Oleandra distenta Kunze in Bot. Zeitung (Berlin) 9: 347 (1851);
Alston in Exell, Cat. Vase. PL S. Tome: 69 (1944); Benl in Acta
Bot. Barcinon. 33: 28 (1982).
Oleandra annetii sensu auct. non Tardieu in Notul. Syst. (Paris) 14:
332 (1953); Pichi Sermolli in Webbia 20: 763 (1965) p.p. quoad
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
distr. Sao Tome; Benl in Acta Bot. Bardnon. 33: 30 (1982) p.p.
quoad distr. Sao Tome.
SAO TOME. Macambrara, primary forest, 1050-1200 m, epiphyte, Exell
237 (BM!); Nova Moca, 950 m, Matter s.n. (COI!); Bom Sucesso, 1250 m,
Matter s.n. (COI!); Calvario, Espirita Santo 5058 p.p. (LISC!); Sao Joao dos
Angolares, Newton s.n. (BM!, COI!); Tra"s-os-Montes, Quintas s.n. (COI!);
Monte Cafe, Welwitsch 52 (BM!, LISU!); s.l., F.A.E. (Matter & Quintas) 48
(BM!, G, K also Moller 43\, LISU!).
PRINCIPE. S.I., Barter 1928 (K!).
DISTRIBUTION. Widespread in tropical and subtropical Africa and
Mascarene Is.
NOTE. Some collections from Sao Tome previously identified as
Oleandra annetii Tardieu have squarrose rhizomes with abruptly
subulate to gradually tapering scales and for this reason, they are
here referred to O. distenta s.l.
Ophioglossaceae
Ophioglossum L.
Ophioglossum costatum R. Br., Prodr.: 163 (1810); Alston in
Exell, Cat. Vase. PL S. Tome: 94 (1944).
SAO TOME. Between Macambrara and Sao Nicolau, Exell 307 (BM-not
found).
DISTRIBUTION. Widespread in tropical Africa, Asia, and Australa-
sia.
Ophioglossum reticulatum L., Sp. pi. 2: 1063 (1753); Alston in
Exell, Cat Vase. PI. S. Tome: 94 (1944); Benl in Acta Bot.
Bardnon. 33: 3 (1982).
SAO TOME. Near Sao Pedro, 900-1200 m, Chevalier 14532 (P!);
Magodinho, 50 m, Espirito Santo 4929 (LISC!, LISJC!); Macambrara,
primary forest, 1050-1200 m, Exell 444 (BM!); Nova Moca, 800 m, F.A.E.
(Quintas) 72 (BM!, COI!, K!, LISU!, P!), Quintas s.n. (COI!); Diogo Nunes,
Newton s.n. (BM!, COI!, P!); Diogo Vaz, between Paga Fogo and Rio Paga
Fogo, Rozeira 698 (PO!); s.l. (probably Nova Moca), Quintas 62 (K!),
Welwitsch 30 (BM\, LISU!).
DISTRIBUTION. Pantropical and subtropical.
Polypodiaceae
Anapeltis J. Sm.
Anapeltis lycopodioides (L.) J. Sm., Cult, ferns: 6 (1857).
var. owariensis (Desv.) Benl in Acta Bot. Bardnon. 33: 18 (1982).
Polypodium lycopodioides sensu auct. non L., Sp. pi. 2: 1082 (1753);
Alston in Exell, Cat. Vase. PL S. Tome: 88 (1944); Sobrinho in
Portugaliae Acta BioL, Ser. B, Sist. 3: 393 (1952).
Microgramma owariensis (Desv.) Alston in Bol. Soc. Brot., ser. 2,
30: 20 (1956); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3:
80(1958).
SAO TOME. Pico, Chevalier 13639 (P); Porto Alegre, Chevalier 1421 Ibis
(P); Vila Conceicao, 100 m, Matos 7718 (LISC!); Caixao Grande, 200 m,
Moller s.n. (COI!); Trindade, 250 m, Matter s.n. (COI!); near Sao Tome,
Mailer s.n. (COI!); Ribeira Peixe, epiphyte, Oliveira & Noronha 35-A
(LISU!); Agua Ize, epiphyte on plantation, Pinto Basto206 (LISC!); Correia,
50 m, Quintas 1246 (BM!, COI!); Ilheu das Rolas, Quintas s.n. (COI!); Sao
Nicolau, 900 m, Quintas s.n. (LISU!); Ribeira Peixe, between Vila Jose and
Ermelinda, epiphyte, Rozeira 236 (PO!); Diogo Vaz, Rozeira 869 (PO); s.l.,
F.A.E. (Moller & Quintas) 54 (COI!, BM!, LISJC!, LISU!), Seabra s.n.
(COI!, LISU!).
59
PRINCIPE. Terreiro Velho, 200 m, epiphyte, Exell 534 (BM!, COI!); Porto
Real, epiphyte, Lains e Silva 420 (LISU!); Sundi, epiphyte, Monod 12076
(BM!), Thorold 2080 (BM!); Berimbau, Newton 23 (BM!, COI!); Cimalo,
Quintas 43 (BM!, COI!); Infante D. Henrique, at the base of Dois Irmaos,
epiphyte, Rozeira 435 (BM-not found, PO!); Esperan?a, 1 20 m, Rozeira 996
(PO), Thorold2016 (BM!); Santo Antdnio, sea level, epiphyte, Thorold2019
(BM!); s.l., Rattray s.n. (E, K!).
DISTRIBUTION. W. tropical Africa.
Belvisia Mirb.
Belvisia spicata (L.f.) Mirb. ex Copel., Gen.fiL: 192 (1947); Alston
in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 444 (1959);
Tardieu-Blot in Aubrev., FL Cameroun 3: 342 (1964); Benl in
Acta Bot. Bardnon. 33: 12 (1982).
Hymenolepis spicata (L.f.) C. Presl, Epimel. hot.: 159 (1851);
Alston in Exell, Cat. Vase. PL S. Tome: 90 (1944); Sobrinho in
Garcia de Orta 7: 89 (1959).
SAO TOME. Calvario, secondary forest, 920 m, epiphyte, Espirito Santo
5075 (LISJC!); Ribeira Peixe, 140 m, epiphyte, Lains e Silva 248 (LISU!),
257 (LISJC-not found), 284 (LISJC-not found); Tras-os-Montes, 700 m,
Quintas 1322 p.p. (BM!, COI!); Nova Moca, 1 100 m, Quintas s.n. (LISU!);
Ribeira Peixe, between Vila Jose and Ermelinda, Rozeira 272 (PO!); Cruzeiro,
300 m, Thorold2067 (BM!), 300 m, 2071 (BM!); s.l., 900 m, Quintas?, (K!).
DISTRIBUTION. Tropical Africa and Mascarene Is.
Drynaria (Bory) J. Sm.
Drynaria laurentii (H. Christ ex De Wild. & T. Durand) Hieron.
in Engl. & Drude, Veg. Erde 9(2): 57, fig. 54 (1908); Alston in
Exell, Cat. Vase. PL S. Tome: 90 (1944); Benl in Acta Bot.
Bardnon. 33: 10(1982).
SAO TOME. Monte Cafe, Chevalier 14291 (P!), 600 m, Thorold 2062
(BM!), 600-760 m, Welwitsch 69 (BM!, K!, LISU!); Blu-blu, 50 m, Moller
s.n. (COI!); Agua Ize, epiphyte, Pinto Basto2\l (LISC!); Diana, Rozeira 957
(PO!); Perseveranca, Seabra 173 (LISU!); s.l., Don s.n. (BM!), F.A.E.
(Moller) 57 (BM!, K also Moller 52!, LISU!).
PRINCIPE. S.l., Welwitsch 17 (BM!, K!, LISU!).
DISTRIBUTION. Tropical Africa.
Drynaria volkensii Hieron. in Engl. & Drude, Veg. Erde 9(2): 57
(1908) nomen; in Bot. Jahrb. Syst. 46: 393 (1911); Alston in
Exell & Rozeira in Conf. Int. Afr. Oc. 3: 80 (1958); Alston in
Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 444 (1959);
Benl in Acta Bot. Bardnon. 33: 1 1 (1982).
SAO TOME. Nova Moca, 900 m, Moller s.n. (COI!); Cruzeiro, 300 m,
Thorold 2066 (BM!); Agua Ize, 180 m, Thorold 2092 (BM!).
PRINCIPE. Praia da Lapa, epiphyte, Rozeira 1902 (BM-not found, PO!);
s.l., Barter 1921 (K!).
DISTRIBUTION. Widespread in tropical Africa.
Microsorum Link
Microsorum punctatum (L.) Copel. in Univ. Calif. Publ. Bot. 16:
1 1 1 (1929); Benl in Acta Bot. Bardnon. 33: 14 (1982).
Polypodium irioides Poir., Encycl. 5: 513 (1804); Alston in Exell,
Cat. Vase. PL S. Tome: 88 (1944).
SAO TOME. Between Bom Sucesso and Lagoa Amelia, Almeida s.n.
(LISC!); Boa Entrada, Chevalier 13788 (P!), 14375 (not found); Sao Joao
dos Angolares, F.A.E. (Quintas) 55 (LISU!), Quintas s.n. (COI!); Agua Ize",
common epiphyte, Pinto Basto 203 (LISC!); s.l., Mendonca s.n. (P!), Souza
s.n. (BM!, COI!).
60
E. FIGUEIREDO
PRINCIPE. SantoAntonio, sea level, epiphyte, Monod 1 2 1 89 (BM !); Sundi,
150 m, Quintas 62 (COI!); Esperan?a, 120 m, Thorold 2077 (BM!); Lapa,
near sea level, Thorold 2089 (BM!), 2090 (BM!); s.l., Rattray s.n. (E),
Welwitsch 25 (BM\,USU\).
DISTRIBUTION. Palaeotropical.
Phymatosorus Pic. Serm.
Phymatosorus scolopendria (Burm.f.) Pic. Serm. in Webbia 28:
460 (1973); Benl in Acta Bot. Barcinon. 33: 13 (1982).
Polypodium phymatodes L., Mant. pi. 2: 306 ( 1 77 1 ); Alston in Exell,
Cat. Vase. PL S. Tome: 89 (1944); Alston in Exell, Suppl. Cat.
Vase. Pl.S. Tome: 8(1956).
SAO TOME. Boa Entrada, Chevalier 13786 (P); Porto Alegre, Chevalier
14243 (K!); Famosa, 400 m, Espirito Santo 69 (BM!, COI!, LISJC!); Ilheu
das Rolas, plantation, 70 m, epiphyte, Espirito Santo 5 1 10 (LISC!, LISJC!),
Quintas s.n. (COI!); Agua Ize, epiphyte, Exell 480 (BM!, COI!), epiphyte,
Pinto Basto 201 (LISC!); bridge over R. Abade, Figueiredo & Arriegas 34
(LISC!); Rio SantoAntonio, Matos 7646 (LISC!); Praia Pantufo, Moller s.n.
(COI!); nearBatepa, 500 m, Moller s.n. (COI!); Bombaim, epiphyte, Monod
1 1880(BM!);NW of Pico, 1000 m,M0/i<«/12268(BM!, COI!); Quingombo,
100 m, Newton s.n. (COI!); Cruzeiro, 300 m, Thorold 2070 (BM!); Monte
Cafe, 850m, Welwitsch 5\ (BM!, LISU!); s.l, Don s.n. (BM!), F.A.E. (Moller
& Quintas) 56 (BM!, COI!, LISJC!, LISU!), Mendonfa (COI!, K!), Moller
s.n. (COI!), Rozeira 1 (PO!), Seabra s.n. (COI!).
PRINCIPE. Terreiro Velho, plantation, 200 m, ground fern, Exell 523 (BM!,
COI!); Esperanca, 120 m, Thorold2015 (BM!), 2078 (BM!); Lapa, near sea
level, Thorold 2091 (BM!); s.l., Rattray s.n. (E), Welwitsch 9 (BM!, K!,
LISU!), 9bis (K!).
DISTRIBUTION. Palaeotropical.
Platycerium Desv.
Platycerium stemaria (P. Beauv.) Desv. \nMem. Soc. Linn. Paris 6:
213 (1827); Alston in Exell, Cat. Vase. PI. S. Tome: 91 (1944);
Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 8 (1956); Benl in
Acta Bot. Barcinon. 33: 9 (1982); Hennipman & Roos, Monogr.
fern genus Platycerium: 108 (1982).
SAO TOME. Sao Vicente, 200 m, epiphyte, Espirito Santo 24 (BM!, COI!,
LISJC!); near Sao Tome, Chevalier 13726 (P!);Trindade, 250 m, Moller s.n.
(COI!), Quintas s.n. (COI!); Rio do Ouro, 380 m, Moller s.n. (COI!); Diogo
Vaz, epiphyte, Rozeira 829 (PO!); s.l., 250-350 m, F.A.E. (Moller) 70
(LISU!), Mendonfa s.n. (K!, P!), 250 m, Moller 60 (K!).
PRINCIPE. Precipfcio, Monod 12178 (BM!); Praia das Burras, epiphyte,
Rozeira 1029 (PO!); s.l., Barter 1886 (K!), Welwitsch 20 (K-not found,
LISU!), 20b (BM-not found, LISU!).
DISTRIBUTION. W. and Central tropical Africa.
Pleopeltis Humb. & Bonpl. ex Willd.
Pleopeltis excavata (Bory ex Willd.) Sledge in Bull. Br. Mus. (Nat.
Hist.) Bot. 2: 138 (1960); Benl in Acta Bot. Barcinon. 33: 16
(1982).
Polypodium preussii Hieron. in Engl. in Bot. Jahrb. Syst. 46: 386
(1911); Alston in Exell, Cat. Vase. PL S. Tome: 89 (1944).
SAO TOME: Calvario, Espirito Santo 5058 p.p. (LISC!, LISJC!);
Macambrara, primary forest, 1050-1200 m, epiphyte, Exell 113 (BM!);
Lagoa Amelia, Exell 216 (BM-not found), 1412 m, Matos 751 1 (LISC!),
1380 m, Monod 11758 (BM!), 1450 m, Monod 11768 (BM!, COI!);
Macambrara to Zampalma, 900 m, epiphyte, Exell 295 (BM!); Pico, light
woodland, 2000m, epiphyte, Exell 341 (BM!), primary forest, 1 900-2020 m,
Lejoly 94/536 (LISC!), Mann (not found), primary forest, 1987 m, epiphyte,
Matos & Van Essche 7386 (LISC!); Bom Sucesso, 1080 m, F.A.E. (Moller &
Quintas) 53 (BM!, LISU!), Moller s.n. (COI!); near Sao Nicolau waterfall,
800 m, Lains e Silva 129 (LISU!); Casa do Pico, 1930 m, Monod 12250
(BM!);s.l., Souzas.n. (COI!).
DISTRIBUTION. Palaeotropical.
Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. in Berlin. Jahrb.
Pharm. Verbundenen Wiss. 21: 41 (1820).
SAO TOME. Lagoa Amelia, primary forest, 1400-1420 m, epiphyte, Matos
& Van Essche 741 1 (LISC!).
DISTRIBUTION. Pantropical and subtropical
Pyrrosia Mirb.
Pyrrosia lanceolate (L.) Farw. in Amer. Midi. Naturalist 12: 245
(1931); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 80
(1958); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 340 (1964);
Hovenkamp, Monogr. fern genus Pyrrosia: 191 (1986).
Cyclophorus spissus (Bory ex Willd.) Desv. in Ges. Naturf. Freunde
Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 301 (1811)
var. continentalis Hieron. ex Engl. in Engl. & Drude, Veg. Erde 9(2):
55, fig. 53 (1908); Alston in Exell, Cat . Vase. PI. S. Tome:9\ (1944).
SAO TOME. Cruzeiro, Rozeira 690 (BM-not found, PO).
PRINCIPE. Cimalo, Quintas 30 (BM!, COI!, P!, Z); Esperan?a, way to
Montalegre, Rozeira 392 (PO!); Praia da Lapa, Rozeira 1944 (PO!); s.l.,
Keulemans s.n. (L), Quintas A (K!).
DISTRIBUTION. Palaeotropical.
Psilotaceae
Psilotum Sw.
Psilotum nudum (L.) P. Beauv., Prodr. aetheogam.: 106, 112
(1805); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 9 (1956);
Benl in Acta Bot. Barcinon. 31: 21 (1978).
SAO TOME. Praia Melao, Rozeira 57 (PO!); s.l., Don s.n. (BM!).
DISTRIBUTION. Widespread in tropical and subtropical regions
worldwide.
Pteridaceae
Pteris L.
Pteris atrovirens Willd., Sp. pi. 5: 385 (1810); Alston in Exell, Cat.
Vase. PL S. Tome: 83 (1944); Tardieu-Blot in Aubrev., FL Cam-
eroun 3: 166 (1964); Benl in Acta Bot. Barcinon. 38: 16 (1988).
SAO TOME. Boa Entrada, Chevalier 13792 p.p. (P!); Pico, Chevalier 14617
(P!); Tras-os-Montes, Gama s.n. (COI!); between Quinas Altas and Monte
Quinas, Monod 12009 (BM!, COI!); Agua Ize, Pinto Basto 210 (LISC!);
Pedroma, Rozeira 1370 (K!); s.l., Don s.n. (BM!), F.A.E. (Moller) 18 p.p.
(BM!, COI!, LISU!, P!), Moller s.n. p.p. (COI!), Moller s.n. (COI!), Rattray
s.n. (E, K!).
PRINCIPE. Esperanga, Navel 146 (P!); Sundi, Newton 42 (COI!); Pico
Papagaio, Welwitsch 22 (BM!, LISU!); s.l., Jardim (cited by Kuhn, 1868),
Rattray s.n. (E).
DISTRIBUTION. W. tropical Africa.
Pteris burtonii Baker in Ann. Bot. (London) 5: 218 (1891); Alston
in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 442 (1959);
Benl in Acta Bot. Barcinon. 38: 7 (1988).
PRINCIPE. S.l., Fox s.n. (BM!).
DISTRIBUTION. W. and E. tropical Africa.
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
61
NOTES. This taxon has not been recorded in the islands since the
nineteenth century.
The collection Walg 1 1 (WAG), from Sao Tome, was interpreted
as a possible hybrid between Pteris atrovirens and P. burtonii by
Benl(1988: 17).
Pteris dentata Forssk., Fl. aegypt.-arab.: 186 (1775); Alston in
Exell, Cat. Vase. PL S. Tome: 84 (1944).
subsp. flabellata (Thunb.) Runemark in Bot. Not. 115: 190 (1962);
Benl in Acta Bot. Bardnon. 38: 1 1 (1988).
Pteris communata sensu auct. non Kuhn in Von der Decken's Reisen
Ost-Afr., Bot. 3: 20 (1879); Alston in Exell in Bull. Inst. Franc.
Afrique Noire ser. A, 21: 442 (1959).
SAO TOME. Lagoa Amelia, 1400 m, Monod 1 1749 (BM!).
PRINCIPE. Esperanca, 600 m, Rozeira 890 (K!).
DISTRIBUTION. Subtropical Africa, Madagascar, Mascarene Is.,
Ascension, St Helena.
Pteris linearis Poir., Encyd. 5: 723 (1804); Alston in Exell &
Rozeira in Conf. Int. Afr. Oc. 3: 79 (1958); Benl in Acta Bot.
Bardnon. 38: 18(1988).
Pteris biaurita sensu auct. non L., Sp. pi. 2: 1076 (1753); Alston in
Exell, Cat. Vase. PI. S. Tome: 83 (1944).
SAO TOME. Boa Entrada, Chevalier 13792 p.p. (P); Ilheu das Rolas, palm
plantation, Espirito Santo 5 145 (LISC!, LISJC!), Rozeira 74 (BM-not found,
PO!); Bombaim, Rozeira 495 (PO); Santa Catarina, palm plantation, Rozeira
790 (K!); s.l., Don s.n. (BM!), F.A.E. (Moller) 16 (BM!, COI!, LISU!), 900 m,
Moller 14 (K!), Moller s.n. (COI!), Rattray s.n. (E).
PRINCIPE. Oque Nazare, Carvalho s.n. (COI!);Terreiro Velho, plantation,
200 m, ground fern, Exell 529 (BM!, COI!); s.l., Welwitsch 22b (LISU!).
DISTRIBUTION. Palaeotropical.
Pteris paucipinnata Alston in Exell in Bull. Inst. Franf. Afrique
Noire ser. A, 21: 443 (1959); Benl in Acta Bot. Bardnon. 38: 14
(1988).
Pteris catoptera sensu auct. non Kunze in Linnaea 18: 119 (1844);
Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 79 (1958).
Pteris quadriaurita sensu auct. non Retz, Observ. hot. 6: 38 (1791);
Alston in Exell, Cat. Vase. PL S. Tome: 84 (1944) p.p. quoad spec.
Sao Tome.
SAO TOME. Macambrara, primary forest, 1050-1200 m, ground fern, Exell
187 (BM !); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain
rainforest, 1200-1 800 m,Lejoly 95/1 1 (LISC!); Monte Cafe, lQlOm,Monod
11690 (BM!-holotype); Colonia A?oreana, Rozeira 267 (PO); Diogo Vaz,
Rozeira 516 (PO); s.l., 350 m, Welwitsch 54 (BM!, LISU!).
PRINCIPE. Pico Papagaio, 600 m, ground fern, Exell 686 (BM!).
DISTRIBUTION. Bioko.
Pteris pteridioides (Hook.) F. Ballard in Kew Bull. [1937]: 348
(1937); Alston in Exell, Cat Vase. PL S. Tome: 84 (1944);
Tardieu-Blot in Aubrev., Fl. Cameroun 3: 170 (1964); Benl in
Acta Bot. Bardnon. 38: 9 (1988).
SAO TOME. Tras-os-Montes, Gama s.n. (COI!); Macambrara, 1200 m,
Quintas 5 (K!), 1200 m, Quintas 1352 (BM!, COI!, K!, P!).
DISTRIBUTION. Widespread in tropical and subtropical Africa.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Pteris similis Kuhn in Von der Decken's Reisen Ost-Afr., Bot. 3: 21
(1879); Benl in Acta Bot. Bardnon. 38: 18 (1988).
Pteris atrovirens sensu auct. nonWilld.,5/?. pi. 5: 385 (1810);Alston
in Exell, Cat. Vase. PI. S. Tome: 83 (1944) p.p.
SAO TOME. Sao Miguel, Monte Verde, secondary forest, 10-150 m, Lejoly
95/98 (LISC!); Agua Ize, common, Pinto Basto 200 (LISC!); Monte Cafe,
760 m, Welwitsch 53 (BM!, LISU!); s.l., Don s.n. (BM!).
PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350
m, ground fern, Exell 561 (BM!, COI!); Sundi, 150 m, Quintas 45 (BM!,
COI!, K!, P!); Infante D. Henrique, Rose 407 (P!).
DISTRIBUTION. W. tropical Africa to E. Africa.
Pteris togoensis Hieron. in Brause & Hieron. in Engl. in Bot. Jahrb.
Syst. 53: 402 (1915); Benl in Acta Bot. Bardnon. 38: 13 (1988).
PRINCIPE. S.l., Barter 1926 (K!).
DISTRIBUTION. W. tropical Africa to E. Africa.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Pteris tripartita Sw. in J. Bot. (Schroder) 1800(2): 67 (1801);
Alston in Exell, Cat. Vase. PL S. Tome: 84 (1944); Benl in Acta
Bot. Bardnon. 38: 15 (1988).
SAO TOME. Boa Entrada, Chevalier 13720 (P!); Ilheu das Rolas, EA.E.
(Quintas) 20 (BM!, LISU!), Quintas s.n. (COI!); s.l., Don s.n. (BM!); s.l.,
Rattray s.n. (E).
PRINCIPE. Sundi, littoral, Quintas 1 1 (BM!, COI!, K!, P!).
DISTRIBUTION. Pantropical.
Pteris vittata L., Sp. pi. 2: 1074 (1753); Alston in Exell & Rozeira
in Conf. Int. Afr. Oc. 3: 79 (1958); Alston in Exell in Bull. Inst.
Franc. Afrique Noire ser. A, 21: 443 (1959).
SAO TOME. Monte Cafe, 690 m, Monod 1 1671 (BM!), Rozeira s.n. (PO);
Agua Ize, Pinto Basto 216 (LISC!).
DISTRIBUTION. Pantropical. Alston (1958, 1959) suggested that
this might be an escape from cultivation.
Selaginellaceae
Selaginella P. Beauv.
Selaginella cathedrifolia Spring in Mem. Acad. Roy. Sd. Belgique
24: 112 (1849); Alston in Exell, Cat. Vase. PL S. Tome: 96
(1944); Tardieu-Blot in Aubrev., FL Cameroun 3: 30 (1964);
Benl in Acta Bot. Bardnon. 31: 15 (1978).
PRINCIPE. Above Infante D. Henrique, secondary forest, 300 m, ground
fern, Exell 629 (BM!); Pico Papagaio, 600 m, ground fern, Exell 687 (BM!,
COI!); near Lapa, Monod 12094 (BM!); s.l., Barter 1931 (K\),Curror 3 (K!-
type), Mann s.n. (K!).
DISTRIBUTION. W. and Central tropical Africa.
Selaginella mannii Baker in J. Bot. 23: 1 80 ( 1 885); Alston in Exell,
Cat. Vase. PL S. Tome: 96 (1944).
SAO TOME. Between Monte Caf<§ and Pico, Chevalier 14552 (P!);
Macambrara, primary forest, 1050-1200 m, Exell 95 (BM!, COI!), 106
(BM!), 168 (BM!), 1260 m, Moller s.n. (COI!); between Bom Sucesso and
Calvario, primary forest, 1350-1400 m, Matos & Van Essche 7347 (LISC!);
Bom Sucesso, 1 250 m,Moller s.n. (COI!); Pico, 1900m,Afo//ers.n. (COI!);
Rio Contador, 1350 m, Moller s.n. (COI!); Lagoa Amelia, 1400 m, Monod
62
E. FIGUEIREDO
1 1784 (BM!); W. of Pico, 870-1225 m, Monod 1 1904 (BM!); Calvario, Rose
151 (P!); s.l., F.A.E.(Moller) 80 (BM!, COI!, K also Matter 75!, LISU!, P!),
Henriques 4 (K!), 1500 m, Mann 1 108 (K!-type).
PRINCIPE. Pico Papagaio, 600 m, ground fern, Exell 683 (BM!).
DISTRIBUTION. Endemic.
Selaginella molleri Hieron. in Engl. & Prantl, Nat. Pflanzenfam.
1(4): 697 (1901); Alston in Exell, Cat Vase. PL S. Tome: 97
(1944); Benl in Acta Bot. Barcinon. 31: 20 (1978).
SAO TOME. Pico, Chevalier 13668 (P!); Macambrara, primary forest,
1050-1200 m, Exell 180 (BM!, COI!); Bom Sucesso, 1250 m, Moller s.n.
(COI!); Rio Manuel Jorge, waterfall, 800 m, Moller s.n. (COI!); s.l., F.A.E.
(Moller) 79 p.p. (BM type!, COI!, K!, LISU!).
DISTRIBUTION. Liberia, Nigeria, Bioko.
Selaginella molliceps Spring in Mem. Acad. Roy. Sci. Belgique 24:
257 (1849); Alston in Exell, Cat. Vase. PL S. Tome: 97 (1944);
Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 8 1 ( 1 958); Benl
in Acta Bot. Barcinon. 31: 20 (1978).
SA TOME. Between Bacelar and Quija, secondary forest, 160 m, Espirito
Santo 5140 (LISC!, LISJC!); Ribeira Peixe, Cao Grande, primary forest,
Matos 7636 (LISC!); between Sao Miguel and Zagaia, primary forest, 170-
400 m, Matos & Van Essche 7238 (LISC!); Diogo Vaz, Rozeira 879 (PO).
PRINCIPE. Oque Pipi, secondary forest, 300 m, ground fern, Exell 500
(BM!, COI!); Sao Joao, 450 m, Newton s.n. (COI!), Newton 4 (K!); Pico
Papagaio, 900 m, Welwitsch 1 1 (BM!, LISU!); s.l., Curror 2 p.p. (K!-type).
DISTRIBUTION. W. and Central tropical Africa.
Selaginella monodii Alston in Exell in Bull. Inst. Franc. Afrique
Noire ser. A, 21: 440 (1959).
SAO TOME. Blu-blu, Monod 1 1806 (BM!-holotype, COI!).
DISTRIBUTION. Endemic.
Selaginella myosurus (Sw.) Alston in J. Bot. 70: 64 (1932).
PRINCIPE. S.l., Curror 2 p.p. (K!).
DISTRIBUTION. Tropical Africa.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Selaginella squarrosa Baker in J. Bot. 23: 180 (1885); Alston in
Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 440 (1959).
PRINCIPE. Pico, 500-600 m, Monod 12111 (BM!).
DISTRIBUTION. Cameroun, Equatorial Guinea.
Selaginella thomensis Alston in Exell, Cat. Vase. PL S. Tome: 97, f.
3 (1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 40 (1964).
SAO TOME. Between Santa Maria and Zampalma, 1200 m, Exell 261
(BM!, COI!); Macambrara, primary forest, 1050-1200 m, Exell 423 (BM!-
holotype, COI!); road Monte Cafe to Sao Nicolau, Figueiredo & Arriegas
121 (LISC!); Nova Moca, 890 m, Moller s.n. (COI!, K!); Pico, 1950 m,
Moller s.n. (COI!); s.l., F.A.E. (Moller) 79 p.p. (BM!, K!, LISU!).
DISTRIBUTION. Sierra Leone, Nigeria, Cameroun.
Sinopteridaceae
Pellaea Link
Pellaea doniana Hook., Sp.fil. 2: 137, t. 125 fig. A (1858); Alston
in Exell, Cat Vase. PL S. Tome: 82 (1944); Tardieu-Blot in
Aubrev., FL Cameroun 3: 140 (1964); Benl in Acta Bot.
Barcinon. 32: 29 (1980).
SAO TOME. Pico, Don s.n. (BM!-type); Ilheu das Rolas, plantation,
Espirito Santo 5106 (LISC!, LISJC!), F.A.E. (Quintas) 15 (K also Quintas
13!, LISU!), Quintas s.n. (COI!), Rozeira 92 (PO!).
PRINCIPE. Between Esperan9a and Fundao, primary forest, 300-350 m,
Ere// 739 (BM!).
DISTRIBUTION. Widespread in tropical Africa
Thelypteridaceae
Christella H. Lev.
Christella dentata (Forssk.) Brownsey & Jermy in Brit. Fern Gaz.
10: 338 (1973); Benl in Acta Bot. Barcinon. 38: 53 (1988).
Dryopteris dentata (Forssk.) C. Chr. in Kongel. Danske Vtdensk.
Selsk. Naturvidensk. Math. Afh. 8(6): 24 (1920); Alston in Exell,
Cat. Vase. PI. S. Tome: 61 (1944); Alston in Exell, Suppl. Cat.
Vase. PL S. Tome: 7(1956).
SAO TOME. Boa Entrada, Chevalier 13787 (P!), 14336 (P!); Sao Vicente,
200 m, Espirito Santo 41 p.p. (BM!, COI!, LISJC!); Novo Destino, 760 m,
Welwitsch 61 (BM!, K!, LISU!); Monte Cafe, Welwitsch 65 (BM!, LISU!);
s.l., F.A.E. (Moller) 42 p.p. (COI!, BM!), Rattray s.n. (BM!, E).
PRINCIPE. 6queNazare,A^ewtons.n.(BM!,COI!);s.l.,^a«ra>'s.n.(BM!,E).
DISTRIBUTION. Palaeotropical.
Christella hispidula (Decne.) Holttum in Kew Bull. 31: 312 (1976);
Benl in Acta Bot. Barcinon. 38: 51 (1988).
Dryopteris quadrangularis (Fee) Alston in J. Bot. 75: 253 (1937);
Alston in Exell, Cat. Vase. PL S. Tome: 63 (1944).
SAO TOME. Agua Grande, 100 m, Moller s.n. (COI!); Saudade, 720 m,
Moller s.n. (COI!); above Monte Cafe, 1010 m, Monod 11691 (BM!);
between Monte Quinas and Quinas Alias, Monod 12006 (BM!).
DISTRIBUTION. Pantropical.
Cyclosorus Link
Cyclosorus striatus (Schum.) Ching in Bull. Fan Mem. Inst. Biol.
10: 249 (1941); Benl in Acta Bot. Barcinon. 38: 55 (1988).
Dryopteris striata (Schum.) C. Chr., Indexfilic.: 294 (1905); Alston
in Exell, Cat. Vase. PL S. Tome: 64 (1944).
SAO TOME. Lagoa Amelia, Chevalier 13653 (P).
DISTRIBUTION. Tropical Africa.
Metathelypteris (H. Ito) Ching
Metathelypteris fragilis (Baker) Holttum in /. S. African Bot. 40:
127 (1974).
Dryopteris fragilis (Baker) C. Chr., Indexfilic.: 266 (1905); Alston
in Exell, Cat. Vase. PL S. Tome: 62 (1944).
subsp. guineensis Benl in Nova Hedwigia 27: 150 (1976); Benl in
Acta Bot. Barcinon. 38: 43 (1988).
SAO TOME. Macambrara, primary forest, 1050-1200 m, Exell 133 (BM!);
s.l., Newton 1 (K!).
DISTRIBUTION. Bioko.
NOTE. The collection cited by Holttum (I.e.) as Newton s.n. (K ex
COI), 1885, is Newton 1 (K).
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
Pneumatopteris Nakai
Pneumatopteris afra (C. Chr.) Holttum in Blumea 21: 306 (1973);
Benl in Acta Bot. Barcinon. 38: 57 (1988).
Dryopteris oppositifolia sensu auct. non (Hook.) C. Chr., Index
filic.: 281 (1905); Alston in Exell, Cat Vase. PL S. Tome: 63
(1944) p.p. quoad pi. ex Principe.
Cyclosorus afer (H. Christ) Ching in Bull. Fan Mem. Inst. Biol. 10:
242 ( 1 94 1 ); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser.
A, 21:446(1959).
SAO TOME. Sao Vicente, Espirito Santo 41 p.p. (LISC!); Sao Miguel, 50m,
Espirito Santo 4744 (LISC!); between Pico and Monte Castro, 6 km S. of
Ponta Figo, mountain rainforest, 1200-1800 m, Lejoly 95/30 (LISC!);
Pedroma, Mocquerys 39/40 (P!); between Quinas Altas and Monte Quinas,
Monod 12023 (BM!); Agua Ize, Pinto Basto 213 (LISC!); Lagoa Amelia,
Rose 204 (P\), 231 (P!).
PRINCIPE. Terreiro Velho, plantation, 200 m, Exell 530a (BM!, COI!);
between Oque Pipi and Morro do Leste, secondary forest, 350 m, ground
fern, Exell 566 (BM!, COI!); Infante D. Henrique, Rose 398 (P!); s.l., Barter
1924 (K!), Rattray s.n. (E), 300 m, Welwitsch 21 (BM!, K!, LISU!), 21bis
(K!).
DISTRIBUTION. Widespread in W. tropical Africa.
Pneumatopteris oppositifolia (Hook.) Holttum in Blumea 21: 304
(1973); Holttum in/ S. African Bot. 40: 157 (1974); Benl inActa
Bot. Barcinon. 38: 62 (1988).
Dryopteris oppositifolia (Hook.) C. Chr., Index filic.: 281 (1905);
Alston in Exell, Cat. Vase. PI. S. Tome: 63 (1944) p.p. quoad pi. ex
Sao Tome.
Cyclosorus oppositifolius (Hook.)Tardieu inNotul. Syst. (Paris) 14:
346 ( 1 953); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser.
A, 21:445(1959).
SAO TOME. Between Lagoa Amelia and Calvario, primary forest, 150 m,
ground fern, Exell 259 (BM!, COI!); Macambrara to Zampalma, 900 m,
ground fern, Exell 297 (BM!); Tras-os-Montes, Gama s.n. (COI!); Ribeira
Peixe, 140 m, frequent, Lains e Silva 268 (LISU!); Pico, 1500 m, Mann s.n.
(K!-type); Lagoa Amelia, 1400 m, Monod 11748 (BM!); Calvario, 1200 m,
Monod 11814 (BM!); between Quinas Altas and Monte Quinas, Monod
12018 (BM!); s.l., F.A.E. (Moller) 41 p.p. (BM!, COI!, LISU!, P!).
DISTRIBUTION. Bioko, Annobon.
Pneumatopteris venulosa (Kuntze) Holttum in Blumea 21: 315
(1973); Benl in Acta Bot. Barcinon. 38: 59 (1988).
Dryopteris venulosa Kuntze, Revis. gen. pi. 2: 814 (1891); Alston in
Exell, Cat. Vase. PL S. Tome: 66 (1944).
Cyclosorus elatus (Mett. ex Kuhn) Alston in Bol. Soc. Brot., ser. 2,
30: 13 (1956); Alston in Exell in Bull. Inst. Franc. Afrique Noire
ser. A, 21: 446 (1959).
SAO TOME. Boa Entrada, Chevalier 13680 (P!); Macambrara, primary
forest, 1050-1200 m, ground fern, Exell 242 (BM!, COI!); Lagoa Amelia,
1400 m, Monod 1 1750 (BM!); s.l., Don s.n. (BM-not found), F.A.E. (Moller)
42 p.p. (K also Moller 39!, P!).
PRINCIPE. Esperan9a, Navel 145 (LISU!, P!).
DISTRIBUTION. Equatorial Guinea (continent, Bioko and Annobon).
Pseudophegopteris Ching
Pseudophegopteris henriquesii (Baker) Holttum in Blumea 17: 15
(1969); Holttum in J. S. African Bot. 40: 128 (1974).
Polypodium henriquesii Baker in Henriq. in Bol. Soc. Brot. 4: 154,
t.l (1887).
63
Dryopteris henriquesii (Baker) C. Chr., Index filic.: 270 (1905);
Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944).
Thelypteris henriquesii (Baker) Tardieu in Notul. Syst. (Paris) 14:
344(1953).
Macrothelypteris henriquesii (Baker) Pic. Serm. in Webbia 23: 179
(1968).
SAO TOME. Tras-os-Montes, Gama s.n. (COI!); Nova Moca, Moller s.n.
(COI!), 750 m, Quintas 1464 (COI!, LISJC!, P!); Macambrard, 1250 m,
Moller s.n. (COI!); Sao Nicolau, waterfall, Rozeira 162 (PO!); Trds-os-
Montes, Calvario, Rozeira 562 (PO!); s.l., F.A.E. (Moller) 49 (BM!, K!-also
Moller 45, type, LISU!, P!), Moller 1 (BM!).
DISTRIBUTION. Endemic.
Sphaerostephanos J. Sm.
Sphaerostephanos elatus (Bojer) Holttum in J. 5. African Bot. 40:
167(1974).
subsp. thomensis Holttum in J. S. African Bot. 40: 167 (1974).
Thelypteris mauritiana (Fee) C.F. Reed in Phytologia 17: 291
(1968) subsp. thomensis (Holttum) Viane in Bull. Soc. Roy. Bot.
Belg. 118:54(1985).
SAO TOME. S.l., F.A.E. (Moller) 41 p.p. (COI!, P!-type).
DISTRIBUTION. Endemic.
NOTE. This taxon has not been recorded in the islands since the
nineteenth century.
Vittariaceae
Anthrophyum Kaulf.
Anthrophyum immersum (Bory ex Willd.) Mett. ex Hook, in
Hook. & Baker, Syn.fiL: 393 (1868); Alston in Exell in Bull. Inst.
Franc. Afrique Noire ser. A, 21: 442 (1959).
SAO TOME. Monte Verde, secondary forest, 10-150 m, Lejoly 95/86
(LISC!); between Estacao Sousa and Pico, primary forest, 1700 m, rare
epiphyte, Matos 7553 (LISC!); Cruzeiro, 300 m, Thorold 2064 (BM!).
PRINCIPE. Pico, 950 m, Thorold 2081 (BM!).
DISTRIBUTION. Also in Liberia and Mascarene Is.
Anthrophyum mannianum Hook., Sec. cent, ferns: t. 73 (1860);
Alston in Exell, Cat. Vase. PL S. Tome: 87 (1944); Benl in Acta
Bot. Barcinon. 33: 6 (1982).
SAO TOME. Pico, light woodland, 2000 m, epiphyte, Exell 340 (BM!); near
Rio Campos, primary forest, epiphyte, Matos & Van Essche 7441 (LISC!);
between Sao Pedro and Lagoa Amelia, 1250 m, Moller s.n. (COI!);
Macambrara, 1 200 m, epiphyte, Moller s.n. (COI!); Sao JoaodosAngolares,
200 m, Quintas 54A (K!), Quintas s.n. (COI!); Lagoa Amelia, epiphyte,
Rozeira 1097 (PO!); s.l., F.A.E. (Moller & Quintas) 60 (BM!, COI!, K!,
LISJC!, LISU!), 1250 m, Moller 54 (K!).
DISTRIBUTION. Widespread in tropical Africa.
Vittaria Sm.
Vittaria guineensis Desv. in Ges. Naturf. Freunde Berlin Mag.
Neuesten Entdeck. Gesammten Naturk. 5: 325 (1811); Alston in
Exell, Cat. Vase. PL S. Tome: 86 (1944);Tardieu-Blot inAubrev.,
Fl. Cameroun 3: 124 (1964); Benl in Acta Bot. Barcinon. 33: 7
(1982).
SAO TOME. SW region, Chevalier 14598 (P!); Calvario, 970 m, epiphyte,
64
E. FIGUEIREDO
Espi'rito Santo5Q59 (LISC!, LISJC!); between LagoaAmelia and Esperanca,
1300 m, epiphyte, Espi'rito Santo5\56 (LISC!, LISJC!); between Sao Carlos
and Bom Sucesso, secondary forest, Espi'rito Santo 5 1 90 (LISJC!); Sao Joao
dosAngolares, littoral, F.A.E. (Newton) 61 p.p. (BM!), 80 m, F.A.E. (Quintas)
61 p.p. (BM!, COI!, K!-mixed with Barter 1908, LISU!), Newton s.n. p.p.
(COI!, P!); Ilheu das Rolas, Greeffs.n. (B?); between Pico and Monte Figo,
6 km S. of Ponta Figo, mountain rainforest, 1200-1800 m, epiphyte, Lejoly
95/6 (LISC!); W. of Pico, 870-1225 m, Monod 1 1906 (BM!); Macambrara,
1200 m, Quintas s.n. (COI!); Ribeira Peixe, between Vila Aida and main
house, epiphyte, Rozeira 291 (PO!).
PRINCIPE. Pico Papagaio, primary forest, 450 m, epiphyte, Exell 719
(BM!, COI!); Oque Caspar, Newton 25 p.p. (BM!); s.l., Barter 1908 (K!-
mixed with F.A.E. 61).
DISTRIBUTION. W. tropical Africa to Uganda.
Vittaria owariensis Fee, Mem.foug. 3: 2 1 , t. 3, fig. 2 ( 1 852); Alston
in Exell, Cat. Vase. PL S. Tome: 87 (1944).
SAO TOME. Sao Joao dosAngolares, F.A.E. (Newton)6\ p.p. (BM!), F.A.E.
(Quintas) 61 p.p. (BM!, P!), Newton s.n. p.p. (P!), Quintas s.n. (COI!); Vila
Concei9§o, 100 m, epiphyte, Matos 7716 (LISC!); Rio Caue, epiphyte,
Rozeira 30 (PO!); Rio Manuel Jorge, between Pinheira and Almas, epiphyte,
Rozeira 83 (PO!); s.l., Seabra s.n. (COI!).
PRINCIPE. Oque Caspar, Newton 25 p.p. (BM!, COI!); Porto Real, Rose
467 (P!); Infante D. Henrique, at the base of Dois Irmaos, epiphyte, Rozeira
480 (PO!); s.l., Welwitsch 13 (BM-not found, K!, LISU!).
DISTRIBUTION. W. tropical Africa.
DOUBTFUL RECORDS
Asplenium hemitomum Hieron. in Engl. in Bot. Jahrb. Syst. 46:
365 (1911); Alston in Exell, Cat. Vase. PL S. Tome: 78 (1944);
Benl in Acta Bot. Barcinon. 40: 20 (1991).
PRINCIPE ?
NOTE. Alston (I.e.) recorded this species based on the collection
Barter 1896a (K!) from Principe. Barter 1896 is a mixed collec-
tion, containing A. hemitomum and A. eurysorum. Since Barter
also collected A. hemitomum in Bioko it is possible that a specimen
of this species, from Bioko, was attached to the herbarium sheet
with A. eurysorum from Principe.
Ceratopteris cornuta (P. Beauv.) Lepr. in Ann. Sci. Nat. (Paris) 19:
103, t. 4A (1830); Alston in Exell, Cat. Vase. PL S. Tome: 92
(1944).
PRINCIPE ? Barter (Kuhn, 1868).
NOTE. No specimens were seen at BM, K or P. However, at BM
there is a specimen collected by Barter during the Niger Expedition,
unlocalized, which could have been collected in Principe.
Dryopteris manniana (Hook.) C. Chr., Index filic.: 276 (1905);
Benl in Acta Bot. Barcinon. 40: 38 (1991).
SAO TOME ?
NOTE. No specimens were seen.
Lastreopsis nigritiana (Baker) Tindale in Contr. New South Wales
Natl. Herb. 3: 245 (1963);Tardieu-Blot inAubrev., Fl. Cameroun
3: 280 (1964); Benl in Acta Bot. Barcinon. 40: 51 (1991).
Polypodium nigritianum Baker in Hook. & Baker, Syn. fil.: 313
(1867).
PRINCIPE ?
NOTE. No specimens were seen. Although there are references in
the literature to the occurrence of this species in Principe, Tindale
(1965) did not record it for Sao Tome and Principe in his monograph
of the genus.
Lygodium microphyllum (Cav.) R. Br., Prodr.: 162 (1810).
Lygodium scandens sensu auct. non (L.) Sw. in /. Bot. (Schrader)
1800(2): 106 (1801); Alston in Exell, Cat. Vase. PL S. Tome: 93
(1944).
PRINCIPE ? (Barter, 1860).
NOTE. No specimens were seen at BM or K. There are specimens
of Lygodium microphyllum collected by Barter in 1859 during the
Niger Expedition, some of which are unlocalized and could have
been collected on the island. Being a lowland species, L.
microphyllum might be extinct due to the extensive cultivation of
these areas.
Ophioglossum gramineum Willd. in Schrift. Akad. Erfurt: 18, t.l,
f.l (1802); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 46 (1964).
SAO TOME ?
NOTE. No specimens seen.
Trichomanes africanum H. Christ in J. Bot. (Morot), ser. 2, 2: 21
(1909); Benl in Acta Bot. Barcinon. 32: 23 (1980).
[Vandenboschia africana (H. Christ) G. Kunkel in Nova Hedwigia
6:213(1963)]
SAO TOME ?
NOTE. No specimens seen.
ACKNOWLEDGEMENTS. Thanks are due to Fundagao Calouste Gulbenkian
for a research grant. I would also like to thank the directors of BM, K, and P
for providing all the facilities during my stay in their herbaria, and also
Frederic Badre, Gustav Benl, Josephine Camus, Peter Edwards, Elisa
Folhadela, Mary Gibby, Bob Johns, and Alison Paul.
REFERENCES
Alston, A.H.G. 1944. Pteridophyta. In A. Exell, Catalogue of the vascular plants ofS.
Tome (with Principe and Annobon): 57-99. London.
1956a. Pteridophyta. In A. Exell, Supplement to the catalogue of the vascular
plants ofS. Tome (with Principe and Annobon): 7-9. London.
1956/7. New African ferns. Boletim da Sociedade Broteriana, serie 2, 30: 5-27.
1958. Pteridophyta. In A. Exell &A. Rozeira, Aditamento a Flora das ilhas de Sao
Tome e Principe. Conferencia Internacional dos Africanistas Ocidentais 3: 78-81.
1959. Pteridophyta. In A. Exell, Additions to the flora of S. Tome and Principe.
Bulletin de I'lnstitut Franfaise d'Afrique Noire, serie A, 21: 440-448.
Barter, C. 1 860. Letter from Mr Charles Barter, natural history collector to the Niger
Expedition. Journal of the Proceedings of the Linnean Society, Botany. 4: 17-23.
Benl, G. 1 978. The Pteridophyta of Fernando Po I. Acta Botanica Barcinonensia 31: 1-
31.
1980. The Pteridophyta of Fernando Po II. Acta Botanica Barcinonensia 32: 1-
34.
1982. The Pteridophyta of Fernando Po III. Acta Botanica Barcinonensia 33: 1-
46.
1 988. The Pteridophyta of Bioko (Fernando Po) IV. Acta Botanica Barcinonensia
38: 1-69.
199 1 . The Pteridophyta of Bioko (Fernando Po) V. Acta Botanica Barcinonensia
40: 1-106.
Dubuisson, J.-Y. 1996. Evolutionary relationships within the genus Trichomanes
sensu lato (Hymenophyllaceae) based on anatomical and morphological characters
and a comparison with rbcL necleotide sequences: preliminary results. In J. Camus
et al. (Eds) Pteridology in perspective: 285-287. Kew.
PTERIDOPHYTES OF SAO TOME AND PRINCIPE
65
Exell, A. 1944. Catalogue of the vascular plants of S. Tome (with Principe and
Annobon). London.
Exell, A. & Rozeira, A. 1958. Aditamento a flora das ilhas de Sao Tome e Principe.
Conferencia International dos Africanistas Ocidentais 3: 77-91.
Figueiredo, E. 1994. Diversity and endemism of angiosperms in the Gulf of Guinea
islands. Biodiversity and Conservation 3: 785-793.
Kramer, K. 1972. The Lindsaeoid ferns of the Old World-IX Africa and its islands.
Bulletin du Jardin Botanique National de Belgique 42: 305-345.
Kuhn, M. 1 868. Filices africanae. Lipsiae.
Nooteboom, H.P. 1994. Notes on Davalliaceae. II. A revision of the genus Davallia.
Blumea39: 151-214.
Pichi Sermolli, R.E.G. 1969. Adumbratio Florae Aethiopicae. 16. Marattiaceae.
Webbia 23: 329-351.
1985. On the taxonomy and nomenclature of some species of Ctenitis
(Aspidiaceae) from Tropical Africa. Webbia 39: 1-28.
1991. On the taxonomy and nomenclature of some species from Tropical
Africa of the genus Triplophyllum Holttum (Dryopteridaceae). Webbia 45: 117-
135.
Tindale, M.D. 1 965. A monograph of the genus Lastreopsis Ching. Contributions from
the New South Wales National Herbarium 3: 249-339.
SYSTEMATIC INDEX
Accepted names are in roman and synonyms in italic.
Acrostichum aureum L. 42, 43
Adiantum mettenii Kuhn 43
Adiantum philippense L. 43
Adiantum raddianum C. Pres 42, 43
Adiantum soboliferum auct. 43
Adiantum vogelii Mett. ex Keyserl. 45
Alsophila camerooniana van camerooniana auct. 51
Alsophila manniana (Hook.) R.M. Tryon 5 1
Alsophila welwitschil (Hook.) R.M. Tryon 51
Anapeltis lycopodioides van owariensis (Desv.)
Benl 59
Anisosorus occidentalis (Baker) C. Chr. 52
Anthrophyum immersum (Bory ex Willd.) Mett. ex
Hook. 63
Anthrophyum mannianum Hook. 63
Arthropteris monocarpa (Cordem.) C. Chr. 58
Arthropteris obllterata auct. 58
Arthropteris orientalis (J.F. Gmel.) Posth. 58
Arthropteris palisotii (Desv.) Alston 58
Aspidium fraternum Mett. ex Kuhn 47
Aspidium nigritlanum Mett. ex Kuhn 46
Aspidium subquinquefidum P. Beauv. 47
Asplenium aethiopicum (Burm.f.) Bech. 48
Asplenium africanum Desv. 48
Asplenium anisophyllum Kunze 48
Asplenium barteri Hook. 48, 50
Asplenium biafranum Alston & F. Ballard 48
Asplenium cuneatum Lam. 48
Asplenium currorii Hook. 48
Asplenium dregeanum subsp. brachypterum (Kunze
ex Houlston & T. Moore) Pic. Serm. 48
Asplenium emarginatum P. Beauv. 48
Asplenium erectum var. usambarense (Hieron.)
Schelpe 48
Asplenium eurysorum Hieron. 49, 64
Asplenium exhaustum (H. Christ) Alston 49
Asplenium formosum Willd. 49
Asplenium friesiorum C. Chr. 42, 49
Asplenium geppii Carruth. 48
Asplenium hemitomum Hieron. 64
Asplenium hypomelas Kuhn 49
Asplenium inaequilaterale Bory ex Willd. 49
Asplenium lividum Mett. ex Kuhn 49
Asplenium longicauda Hook. 49
Asplenium megalura var. molleri (Hieron.)
Tardieu 49
Asplenium molleri Hieron. 49
Asplenium nigritianum Hook. 49, 50
Asplenium paucijugum F. Ballard 50
Asplenium protensum auct. 49
Asplenium quintasil Gand. 49
Asplenium sandersonii Hook. 50
Asplenium unilaterale Lam. 50
Asplenium variabile Hook. 50
Asplenium variabile var. paucijugum (F. Ballard)
Alston 50
Athyrium newtonii (Baker) Diels 50
Belvisia spicata (L.f.) Mirb. ex Copel. 59
Blechnum attenuatum (Sw.) Mett. 51
Blotiella currorii (Hook.) R.M. Tryon 51
Blotiella glabra (Bory) R.M. Tryon 52
Blotiella mannii (Baker) Pic. Serm. 52
Bolbitis acrostichoides (Afzel. ex Sw.) Ching 56
Bolbitis auriculata (Lam.) Ching 56
Bolbitis fluviatilis (Hook.) Ching 56
Callipteris prolifera (Lam.) Bory 50
Ceratopteris cornuta (P. Beauv.) Lepr. 64
Christella dentata (Forssk.) Brownsey & Jermy 62
Christella hispidula (Decne.) Holttum 62
Coniogramme africana Hieron. 54
Ctenitis attenuata Pic. Serm. 47
Ctenitis buchholzii (Kuhn) Alston 47
Ctenitis cirrhosa (Schum.) Ching 42, 45
Ctenitis fraterna (Mett.) Tardieu 47
Ctenitis lanuginosa (Willd. ex Kaulf.) Copel. 45
Ctenitis nigritiana (Mett. ex Kuhn) Alston 46
Cyathea camerooniana var. currorii Holttum 5 1
Cyathea manniana Hook. 51
Cyathea welwitschii Hook. 5 1
Cyclophorus spissus var. continentalis Hieron. ex
Engl. 60
Cyclosorus afer (H. Christ) Ching 63
Cyclosorus elatus (Mett. ex Kuhn) Alston 63
Cyclosorus oppositifolius (Hook.) Tardieu 63
Cyclosorus striatus (Schum.) Ching 62
Davallia chaerophylloides (Poir.) Steud. 51
Davallia denticulata var. denticulata (Burm.f.) Mett.
ex Kuhn 51
Davallia repens (L.f.) Kuhn 51
Dicranopteris linearis (Burm.f.) Underw. 53
Didymochlaena truncatula (Sw.) J. Sm. 45
Diplazium arborescens (Bory) Sw. 50
Diplazium proliferum (Lam.) Thouars 50
Drynaria laurentii (H. Christ ex De Wild. & T.
Durand) Hieron. 59
Drynaria volkensii Hieron. 59
Dryopteris crinobulbon (Hook.) C. Chr. 45
Dryopteris dentata (Forssk.) C. Chr. 62
Dryopteris fragilis (Baker) C. Chr. 62
Dryopteris henriquesii (Baker) C. Chr. 63
Dryopteris lanuginosa (Willd. ex Kaulf.) C. Chr. 45
Dryopteris manniana (Hook.) C. Chr. 64
Dryopteris nigritiana (Mett. ex Kuhn) Kuntze 46
Dryopteris oligantha auct. 45
Dryopteris oppositifolia auct. 63
Dryopteris oppositifolia (Hook.) C. Chr. 63
Dryopteris pentheri (Krasser) C. Chr. 45
Dryopteris protensa (Afzel. ex Sw.) C. Chr. 47
Dryopteris quadrangularis (Fee) Alston 62
Dryopteris securidiformis (Hook.) C. Chr. 47
Dryopteris securidiformis var. nana Bonap. 47
Dryopteris striata (Schum.) C. Chr. 62
Dryopteris variabilis var. barteri (Hook.) Alston 47
Dryopteris venulosa Kuntze 63
Elaphoglossum acrostichoides (Hook. & Grev.)
Schelpe 52
Elaphoglossum aubertii (Desv.) T. Moore 53
Elaphoglossum chevalieri H. Christ 53
Elaphoglossum clarenceanum auct. 53
Elaphoglossum isabelense Brause 53
Elaphoglossum petiolatum subsp. salicifolium (Willd.
ex Kaulf.) Schelpe 53
Elaphoglossum salicifolium (Willd. ex Kaulf.)
Alston 53
Gleichenia linearis (Burm.f.) C.B. Clarke 53
Gonocormus mannii (Hook, ex Hook. & Baker) G.
Kunkel 55
Grammitis ebenina (Maxon) Tardieu 53
Grammitis molleri (Baker) Schelpe 42, 53
Grammitis nigrocincta Alston 42, 53
Grammitis sp. 53
Grammitis tomensis Schelpe 42, 54
Histiopteris incisa (Thunb.) J. Sm. 52
Humata repens (L.f.) Diels 51
Huperzia mildbraedii (Herter) Pic. Serm. 57
Huperzia ophioglossoides (Lam.) Rothm. 42, 57
Huperzia phlegmaria (L.) Rothm. 57
Huperzia verticillata (L.f.) Trevis. 57
Huperzia warneckei (Herter ex Nessel) Pic. Serm. 57
Hymenolepis spicata (L.f.) C. Presl 59
Hymenophyllum capillare Desv. 54
Hymenophyllum ciliatum var. boryanum (Willd.)
Mett. ex Kuhn 54
Hymenophyllum hirsutum (L.) Sw. 54
Hymenophyllum kuhnii C. Chr. 54
Hymenophyllum mildbraedii (Brause ex Brause &
Hieron.) Alston 55
Hymenophyllum polyanthos var. kuhnii (C. Chr.)
Schelpe 54
Hymenophyllum polyanthos var. mildbraedii (Brause
ex Brause & Hieron.) Schelpe 42, 55
Hymenophyllum splendidum Bosch 55
Hypolepis sparsisora (Schrad.) Kuhn 52
Lastreopsis aff. currori (Mett. ex Kuhn) Tindale 46
Lastreopsis nigritiana (Baker) Tindale 64
Lomariopsis guineensis (Underw.) Alston 56
Lomariopsis hederacea Alston 56
Lomariopsis hederacea auct. 56
Lomariopsis nigrescens Holttum 56
Lomariopsis warneckei (Hieron.) Alston 42, 56
Lonchitis currorii (Hook.) Mett. ex Kuhn 51
Lonchitis glabra Bory 52
Lonchitis gracilis Alston 52
Lonchitis mannii (Baker) Alston 52
Lonchitis occidentalis Baker 52
Loxogramme lanceolata (Sw.) C. Presl 56
Lycopodiella cernua (L.) Pic. Serm. 57
Lycopodium cemuum L. 57
Lycopodium clavatum L. 42, 57
Lycopodium mildbraedii Herter 57
Lycopodium phlegmaria L. 57
Lycopodium verticillatum L.f. 57
Lycopodium warneckei (Herter ex Nessel) Alston 57
Lygodium microphyllum (Cav.) R. Br. 64
Lygodium scandens auct. 64
Macrothelypteris henriquesii (Baker) Pic. Serm. 63
Marattia fraxinea Sm. 57, 58
Marattia robusta Alston 57, 58
Metathelypteris fragilis subsp. guineensis Benl 62
Microgonium erosum (Willd.) C. Presl 55
66
E. FIGUEIREDO
Microgramma owariensis (Desv.) Alston 59
Microlepia speluncae (L.) T. Moore 42, 52
Microsorum punctatum (L.) Copel. 59
Microtrichomanes digitatum (Sw.) Copel. 55
Nephmdium subquinquefidum var. elongatum
Hook. 47
Nephrolepis biserrata (Sw.) Schott 58
Nephrolepis pumicicola F. Ballard 58
Nephrolepis undulata (Afzel. ex Sw.) J. Sm. 58
Odontosoria chinensis var. divaricata H. Christ 52
Oleandra annetii auct. 58
Oleandra annetii Tardieu 59
Oleandra distenta Kunze 58
Ophioglossum costatum R. Br. 59
Ophioglossum gramineum Willd. 64
Ophioglossum reticulatum L. 59
Pellaea doniana Hook. 62
Phymatosorus scolopendria (Burm.f.) Pic. Serm. 60
Pityrogramma calomelanos var. calomelanos (L.)
Link 42, 54
Platycerium stemaria (P. Beauv.) Desv. 60
Pleopeltis excavata (Bory ex Willd.) Sledge 60
Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. 60
Pneumatopteris afra (C. Chr.) Holttum 63
Pneumatopteris oppositifolia (Hook.) Holttum 63
Pneumatopteris venulosa (Kuntze) Holttum 63
Polypodium cultratum var. elasticum (Bory ex Willd.)
Baker 54
Polypodium ebeninum Maxon 53
Pol\podium henriquesii Baker 63
Polypodium irioides Poir. 59
Polypodium lycopodioides auct. 59
Polypodium molleri Baker 53
Polypodium nigritianum Baker 64
Polypodium oosorum Baker 54
Polypodium phymatodes L. 60
Polypodium preussii Hieron. 60
Polypodium villosissimum Hook. 54
Pseudophegopteris henriquesii (Baker) Holttum 63
Psilotum nudum (L.) P. Beauv. 60
Pteridium aquilinum (L.) Kuhn 52
Pteris atrovirens Willd. 60, 61
Pteris atmvirens auct. 61
Pteris biaurita auct. 61
Pteris burtonii Baker 60, 61
Pteris catoptera auct. 61
Pteris communata auct. 6 1
Pteris dentata subsp. flabellata (Thunb.) Runemark 61
Pteris linearis Poir. 61
Pteris paucipinnata Alston 61
Pteris pteridioides (Hook.) F. Ballard 61
Pteris quadriaurita auct. 61
Pteris similis Kuhn 61
Pteris togoensis Hieron. 61
Pteris tripartita Sw. 54, 61
Pteris vittata L. 42, 61
Pyrrosia lanceolata (L.) Farw. 60
Selaginella cathedrifolia Spring 61
Selaginella mannii Baker 61
Selaginella molleri Hieron. 62
Selaginella molliceps Spring 62
Selaginella monodii Alston 42, 62
Selaginella myosurus (Sw.) Alston 62
Selaginella squarrosa Baker 42, 62
Selaginella thomensis Alston 62
Selenodesmium rigidum (Sw.) Copel. 56
Sphaerostephanos elatus subsp. thomensis
Holttum 42, 63
Sphenomeris chinensis var. divaricata (H. Christ)
Kramer 52
Sphenomeris chusana var. divaricata (H. Christ)
Tardieu 52
Stenoloma chinense var. divaricatum (H. Christ)
Alston 52
Tectaria angelicifolia (Schum.) Copel. 46
Tectaria buchholzii (Kuhn) Copel. 47
Tectaria camerooniana (Hook.) Alston 46
Tectaria fernandensis (Baker) C. Chr. 47
Thelypteris henriquesii (Baker) Tardieu 63
Thelypteris mauritiana subsp. thomensis (Holttum)
Viane 63
Trichomanes africanum H. Christ 64
Trichomanes borbonicum Bosch 55
Trichomanes chamaedrys Taton 55
Trichomanes crispiforme Alston 55
Trichomanes cupressoides Desv. 56
Trichomanes digitatum Sw. 55
Trichomanes erosum Willd. 55
Trichomanes giganteum auct. 55
Trichomanes mannii Hook, ex Hook. & Baker 55
Trichomanes melanotrichum Schltdl. 55
Trichomanes pyxidiferum var. melanotrichum
(Schltdl.) Schelpe 55
Trichomanes radicans Sw. 55
Trichomanes rigidum Sw. 56
Triplophyllum attenuatum (Pic. Serm.) Pic. Serm. 47
Triplophyllum buchholzii (Kuhn) Holttum 47
Triplophyllum fraternum var. elongatum (Hook.)
Holttum 47
Triplophyllum heudelotii Pic. Serm. 47
Triplophyllum jenseniae (C. Chr.) Holttum 42, 47
Triplophyllum principis Holttum 47
Triplophyllum protensum (Afzel. ex Sw.) Holttum 47
Triplophyllum securidiforme (Hook.) Holttum 47
Triplophyllum securidiforme var. nanum (Bonap.)
Holttum 47
Triplophyllum subquinquefidum (P. Beauv.) Pic.
Serm. 47
Vandenboschia africana (H. Christ) G. Kunkel 64
Vandenboschia borbonica (Bosch) G. Kunkel 55
Vandenboschia pyxidifera (L.) Copel. 55
Vandenboschia radicans (Sw.) Copel. 55
Vittaria guineensis Desv. 63
Vittaria owariensis Fee 64
Xiphopteris cultrata (Willd.) Schelpe 54
Xiphopteris oosora (Baker) Alston 42, 54
Xiphopteris villosissima (Hook.) Alston 54
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 2
No. 1 New Himalayan species of Pedicularis with special reference
to those from the eastern Himalaya. P.C. Tsoong. 1955. Pp. 1-
34. Facsimile edition. £4.35
No. 2 Mosses of Dominica, British West Indies. E.B. Bartram. 1955.
Pp. 37-49. Mosses of the Ecuadorian Andes collected by P.R.
Bell. E.B. Bartram. 1955. Pp. 51-64. Facsimile edition. £4.00
No. 3 Novitates Himalaicae-1. F. Ludlow & W.T. Stearn. 1956. Pp.
65-81, 8 plates, 1 1 figs. Facsimile edition. £5.25
No. 4 Saxifraga of the Himalaya 1. Section Kabschia. H. Smith.
1958. Pp. 83-129, 14 figs. £9.65
No. 5 The Polypodiaceae and Grammitidaceae of Ceylon. W.A.
Sledge. 1960. Pp. 131-158, 4 figs. Facsimile edition. £3.90
No. 6 Allium and Milula in the central and eastern Himalaya. W.T.
Stearn. 1960. Pp. 159-191, 4 plates, 10 figs. Facsimile edition.
£4.35
No. 7 The identity of Isopyrum aquilegioides. L.G. de Beer & W.T.
Stearn. 1960. Pp. 193-202, 3 figs. Facsimile edition. £3.15
No. 8 On the geographical relationships of the angiosperm flora of
New Guinea. R. Good. 1960. Pp. 203-226, 1 fig. Facsimile
edition. £3.60
No. 9 Saxifraga of the Himalaya II. Some new species. H. Smith.
1960. Pp. 227-260. 9 plates, 17 figs. Facsimile edition. £5.65
No. 10 New species of Taraxacum from the Himalayan region. J.L.
van Soest. 1961. Pp. 261-273, 8 plates. Facsimile edition.
£6.15
No. 1 1 The athyriod ferns of Ceylon. W.A. Sledge. 1962. Pp. 275-323.
3 plates. Facsimile edition. £5.25
No. 12 The genus Epilobium in the Himalayan region. PH. Raven.
1962. Pp. 325-382, 7 plates, 13 figs. Facsimile edition. £5.90
Volume 3
No. 1 A revision of the genera Buchenavia and Ramatuella. A.W.
Exell & C.A. Stace. 1963. Pp. 1-46, 5 figs. Facsimile edition.
£5.15
No. 2 The diatom genus Capartogramma and the identity of
Schizostauron. R. Ross. 1963. Pp. 47-92, 2 plates, 38 figs.
Facsimile edition. £5.15
No. 3 Angiosperms of the Cambridge Annobon Island Expedition.
A.W. Exell. 1963. Pp. 93-1 18, 10 plates. Facsimile edition.
£3.75
No. 4 A revision of the genus Petrorhagia. P.W. Ball & V.H.
Hey wood. 1964. Pp. 1 19-172, 3 plates, 22 figs. Facsimile
edition. £4.90
No. 5 Marine algae of Gough Island. Y.M. Chamberlain. 1965. Pp.
173-232, 4 plates, 80 figs. £12.50
No. 6 The Ceylon species ofAsplenium. W.A. Sledge. 1965. Pp. 233-
277, 1 plate, 3 figs. Facsimile edition. £5.15
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
No. 5
No. 6
No. 7
A synopsis of Jamaican Myrsinaceae. W.T. Stearn. 1969. Pp.
143-178, 8 plates, 25 figs. £4.55
The Jamaican species of Columnea and Alloplectus
(Gesneriaceae). W.T.Stearn. 1969. Pp. 179-236, 8 plates, 29
figs. £6.40
New or little known Himalayan species of Swertia and
Veratrilla (Gentianaceae). H. Smith. 1970. Pp. 237-258, 16
plates, 7 figs. £8.25
A survey of the tropical genera Oplonia and Psilanthele
(Acanthaceae). W.T. Stearn. 1971. Pp. 259-323, 10 plates, 18
figs. £12.40
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. 1977. 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. PS. 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 & PS. 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. PS. Dixon. 1983. Pp.
1-15, 2 figs. £5.55
No. 2 A taxonomic study of the lichen genus Micarea in Europe. B.J.
Coppins. 1983. Pp. 17-214, 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. 291^63, 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 fern 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. Umbraculoides. N.K.B. Robson. 1985.
Pp. 163-325, 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. 131-276, 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. 1^2, 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. 107-233, 15 plates, 31 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.
1 989. 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 Solatium 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. 1-151, 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.
147-152.
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. Pp. 1-34.
Studies on the lichen genus Sticta (Schreber) Ach.: II.
Typification of taxa from Swartz's Prodromus of 1788. D.J.
Galloway. Pp. 35-48.
Seaweeds of the western coast of tropical Africa and adjacent
islands: a critical assessment. IV. Rhodophyta (Florideae) 4.
Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F.
Prud'homme van Reine and W.J. Woelkerling. Pp. 49-90.
Studies on the Cretan flora 3. Additions to the flora of •
Karpathos. N.J. Turland and L. Chilton. Pp. 9 1-100. £43.25
No. 2 Observations on the benthic marine algal flora of South
Georgia: a floristic and ecological analysis. D.M. John, P.J.A.
Pugh and I. Tittley. Pp. 101-1 14.
Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical
species (excluding Australia). D.J. Galloway. Pp. 1 15-160.
Morphology and ecology of seedlings, fruits and seeds of
Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp.
161-172.
A study of Bixa (Bixaceae), with particular reference to the leaf
undersurface indumentum as a diagnostic character. R.E.
Dempsey and N.C. Garwood. Pp. 173-180. £43.40
Volume 25
No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. Pp.
1-94.
William Roxburgh's St Helena plants. Q.C.B. Cronk. 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. Pp. 99-122.
A new species of Odontorrhynchos (Orchidaceae,
Spiranthinae) from Boliva. D.L. Szlachetko. Pp. 123-125.
Linnaeus's interpretation of Prospero Alpino's De plantis
exoticis, with special emphasis on the flora of Crete. N.J.
Turland. Pp. 127-159.
Book review. M.G. Gilbert. P. 1 6 1 . £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. £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.
£43.40
Volume 27
No. 1 Notes on the diatom species Tetracyclus castellum (Ehrenb.)
Grunow with a description of Tetracyclus pseudocastellum nov.
sp. D.M. Williams. Pp. 1-5.
A new species of Calymperes (Musci: Calymperaceae) from
Peninsular Malaysia. L.T. Ellis. Pp. 7-9.
A phylogenetic conspectus of the tribe Hyoscyameae
(Solanaceae). A.L. Hoare and S. Knapp. Pp. 1 1-29.
A revision of Solanum section Ptemidea: Solanaceae. S.
Knapp and T. Helgason. Pp. 3 1-73. 1997. £43.40
No. 2 Systematics of Pogostemon (Labiatae) G.R. Bhatti and M.
Ingrouille. Pp. 77-147. 1997 £43.40
CONTENTS
1 Morphology and ecology of seedlings, fruits and seeds of Panama: Vochysiaceae
Nancy C. Garwood
17 A revision of the genus Mandragora (Solanaceae)
S. Ungricht, S. Knapp and J.R. Press
41 The pteridophytes of Sao Tome and Principe (Gulf of Guinea)
Estrela Figueiredo
tural History Musei
BOTANY SERIES
Vol.28, No. 1, June 1998