Bulletin
British Museum
(Natural History)

VOLUME 20 NUMBER 2 20 DECEMBER 1990

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)
British Museum (Natural History), 1990

ISBN 565 08027 X Botany Series

ISSN 0068 - 2292 Vol 20(2), pp 1 53 - 237

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Bull. Br. Mus. nat. Hist (Bot.) 20(2): 153-168

Issued 20 December 1990

The marine algal flora of Namibia: its
distribution and affinities

BRITISH MUSEUM

[NATURAL IMSIORY)

14J

proem-no

GEORGE W. LAWSON_/ C

23 Sheffield Terrace, London W8 7NQ
RICHARD H. SIMONS

Department of Botany, University of Cape Town, Rondebosch, South Africa 7700

WILLIAM EDWIN ISAAC

20 Spraypoint Road, Blairgowrie, Victoria 3942, Australia

GENERAL

CONTENTS

Introduction 153

The Namibian coast and its environmental conditions 154

Patterns of littoral zonation 154

The phytogeographical status of the Namibian marine algal flora 161

Annotated list of species 164

References 167

SYNOPSIS. The coast of Namibia (South West Africa) which lies partly within and partly outside the tropics on the
south western Atlantic coast of Africa was visited and studied in July 1957. Littoral zonation at three main localities
(Liideritz, Elizabeth Bay, and Swakopmund) is briefly described, and an annotated list is presented of the 136
species of marine algae now known from Namibia, as well as of certain algae identified only to genus.
Phytogeographically, the marine algal flora of Namibia represents a northward extension of the flora of the western
coast of South Africa though somewhat depleted north of the tropic. The boundary between the Namibian flora and
the tropical flora of Angola is a sharp one and occurs at, or just north of, the political border of these two countries.

INTRODUCTION

This paper is a further development of reseach originally
undertaken as part of the survey of South African intertidal
plant ecology then being conducted by Professor William
Edwyn Isaac, of the Department of Botany, University of
Cape Town. Professor Isaac organized the Namibia field
excursion which took place during the southern winter of
1957, in which the participants were Professor W. E. Isaac,
Miss Judith Graves, Mr R. H. Simons, and Dr G. W.
Lawson. Use has already been made of the collections then
assembled in several studies and some of the species identi-
fied have been referred to in publications (e.g. John et al.,
1979; Lawson, 1978; Lawson & Price, 1969; Lawson et al.,
1975; Price et al., 1978, 1986, 1988; Silva, 1959; Simons, 1964,
1966, 1970). It is felt that, despite the time elapsed since the
completion of the field work, and despite the rather limited
nature of some of our ecological observations in the short
time available (which included two successive spring tide
periods), a more comprehensive account here attempted is of
value in view of the very small amount of work hitherto
published on the marine algae of the region and of the current
interest in it (see Hommersand, 1986).

The main object of the excursion was to provide data on
the composition, distribution, and zonation of the marine
algal flora of Namibia (then South West Africa), previously

much under-investigated from those points of view. The
reason for the lack of information is not far to seek. Along the
whole of the coast lies the formidable Namib desert making it
difficult to reach and extremely inhospitable. In fact, over-
land access is only practicable by the roads that have been
built to the few townships along the coast. The main acces-
sible areas are around the towns of Swakopmund (22 40' S)
and Walvis Bay in the centre and Liideritz (26 38' S), some
450 km further south. We also had the opportunity of visiting
Elizabeth ville, a relict town no longer inhabited but acces-
sible by road, a short distance south of Liideritz. Though the
total time available was approximately one month, because of
the great distances to be covered between centres and the
then indifferent nature of the roads, our main field work was
accomplished during about a week when we were based at
Swakopmund and a similar period based at Liideritz. From
the phycological point of view the coast of the Republic of
South Africa is relatively well-known, having been worked
over a considerable number of years (see Seagrief &
Troughton, 1973 and Seagrief, 1984). Isaac, (19376) has given
a description of the intertidal seaweed vegetation of the west
coast of South Africa from Lambert's Bay to the Cape of
Good Hope. He has also described zonation to the south-east
of the Cape of Good Hope (Isaac, 1949). Though the zonation
is similar to that on the west coast, east of Cape Town there
are some differences, especially in the sublittoral fringe. In a
later paper Isaac & Hewitt (1953) pointed out that the region

154

from Cape Point to Cape Agulhas is best regarded as an
extension of the west coast, if False Bay, which possesses
relatively higher sea temperatures, is excepted. It is from the
zoological point of view, however, that the most comprehen-
sive picture of the biogeographical distribution of intertidal
organisms in South Africa has been constructed. Stephenson
and his co-workers were actively engaged on these problems
between 1932 and 1937 and these studies were summarized by
Stephenson in three papers published in 1939, 1944, and
1948. One of the main conclusions that emerged from this
work was that there are three distinct faunas on South
African shores: an essentially sub-tropical fauna on the east
coast, a warm temperate fauna on the south coast, and a cold
temperate fauna on the west coast. There are, of course,
many examples of intergrading and overlapping between
these faunas. Stephenson indicated his belief that the same
situation was substantially true for the algae though it appears
from his diagrams (Stephenson, 1944) that the number of
species present only along the south coast is relatively small.
More recently Bolton (1986) has given a similar but fuller
analysis.

From the point of view of the present study it is the
distribution of the west coast flora that is most interesting. Its
southern limit, as indicated above, is approximately at Cape
Agulhas, but the northern limit is less certain, and the
establishment of that limit was one of the objectives of the
present survey. Stephenson and his co-workers did not pene-
trate into Namibia and their furthest northerly place studied
was the Port Nolloth region, south of the Orange river that
marks the coastal boundary between the Cape Province of
South Africa and Namibia.

Subsequent to our own work in Namibia, however, Penrith
& Kensley (1970a, 19706) and Kensley & Penrith (1980) have
also visited the area with much the same objectives as
ourselves. In their case the interest was mainly zoological and
algae were treated as subsidiary. Kensley & Penrith (1973)
were also able to study rocky shores at Mogamedes (Angola)
and surveys of the marine algae for that same region are
reported by Lawson et al. (1975) and John et al. (1981).
Information from all these sources, as well as from Wynne
(1986), is used in a later section when we attempt to assess the
biogeographical significance of our field work in Namibia.

LAWSON, SIMONS & ISAAC

Undoubtedly the most significant feature of the Namib
coast, as far as seaweeds is concerned, is that the Benguela
Current flows northwards along its length with the denser
waters of lower temperature closer inshore. The prevailing
southerly and south- easterly winds carry these surface waters
away from the coast and upwelling of water from 100-300 m
depths takes place at most times of the year (Anon., 1964).
This causes nutrient-rich waters of low temperature and
relatively low salinity to influence the coastal biota. Surface
temperatures as low as 10-15C in winter and 12-20C in
summer for this coast are on record (Anon., 1964), and Isaac
(1937a) points out that the average annual temperature at
Walvis Bay, just within the tropics, is as low as 16.1C. Bolton
(1986) includes 10 year averages of annual maximum and
minimum monthly temperatures for both Walvis Bay and
Liideritz. Any botanist visiting this area must be struck by the
contrast between the extreme paucity of the terrestrial vege-
tation on the hot and arid coastal plains and the relative
luxuriance of the marine vegetation.

According to Sverdrup, Fleming & Johnson (1942), the
effect of the Benguela Current is particularly marked as far
north as latitude 17; thus virtually the whole of the Namib
coast is included since the northern border at Kunene River
mouth is at 17 16' S. Somewhere near Cape Frio (18 30' S)
the Benguela Current begins to turn out westwards into the
Atlantic. More detailed accounts of the hydrography of the
region are given in, for example, Anon. (1963), Stephenson
& Stephenson (1972), and Richards et al. (1981), while
Shannon et al. (1987) describe a mobile front interfacing
Angolan waters and the Benguela System. Sand storms on
the coastal strip are frequent enough events, but far more
common and of greater significance to its desert life are the
'mist-rain' and chilling sea mists that blanket the coast during
darkness and early parts of the day (Anon., 1964: 77, 83).
These mists are more frequent in the north than in the south.
Thus, emersed shore life is perhaps very often shielded from
the worst effects of desiccation and overheating during the
early part of the mornings. One of the causes of this
condensation is the relatively low surface temperature of the
adjacent sea (Anon., 1964). Coastal humidity is relatively
high even in summer (certainly higher than at Cape Town)
and this, too, must have a protective role (Anon., 1964).

THE NAMIBIAN COAST AND ITS
ENVIRONMENTAL CONDITIONS

The long (approximately 1500 km) but relatively straight
coastline of Namibia (Fig. 1) lying between 17 16' S and 28
30' S is inhospitable, the Namib desert reaching directly to
the sea along much of its length. Considerable stretches
consist merely of wave-beaten beaches and are therefore
practically devoid of intertidal seaweed vegetation, though
broken rocky outcrops with more extensive areas of cliffs
occur at intervals; our observations have been limited to such
of these areas as we were able to reach. In general the coast is
exposed to the full force of the Atlantic rollers, but some
shelter from wave-action is found in places such as the natural
harbour at Liideritz and the artificial one at Swakopmund.

Tides are semi-diurnal and the amplitude is not great, vary-
ing most often from under 1.5 m (springs) to little more than
one metre (neaps). Exceptionally the range may be over 1.8 m.

PATTERNS OF LITTORAL ZONATION

In the following account we have used the littoral zonation
scheme of Lewis (1961). As faunal zonation is important in
Lewis' scheme we have, where necessary, added our own
limited observations on the vertical distribution of animals.

(a) The Liideritz area

At Liideritz we examined littoral rocks at three sites: firstly,
the recreational area adjacent to the town itself; secondly, at
Elizabeth Bay (visited through the courtesy of the regional
representative of Consolidated Diamond Mines) a 'ghost'
town about 30 km to the south of Liideritz, and at one time a
centre for diamond operations; lastly, Halifax Bay, on the
open coast about 8 km west of Liideritz.

i) Liideritz

This site is in the vicinity of the resort area. Since it is a bay
within a bay it is well protected from surf action, and waves
were never higher than about 0.5 m during our stay.

ape Frio ~
<J Rocky Point
Mo we Bay

NAMIBIA
Terrace Bay (SOUTH WEST

35 -

15

Fig. 1 Map of Namibia to show the localities mentioned in the text.

156

The Littoral Fringe

The most striking feature of this zone was the abundance of

the small snail Littorina knysnaensis (Philippi)* everywhere

present. Algae were few, mostly scattered plants of Porphyra

capensis and, in places, a sparse blackish crust of blue-green

algae. Species of Cladophora were often present in small rock

pools.

The Eulittoral

Barnacles were not abundant and there was much bare rock
in the upper levels, even on the outer relatively less sheltered
rocks. On the innermost sheltered rocks barnacles seemed
virtually absent. Penrith & Kensley (1970a) do not mention
them in their account of the 'rocks below the strand cafe' at
Luderitz and do not include them in any of their figs. 1-4,
showing the principal organisms of the localities visited. They
do, however, list two barnacles (Chthamalus dentatus Krauss
and Balanus algicola Pilsbry), noting that a few specimens
were present in sheltered localities. In our opinion, however,
there were sufficient numbers in 1957 to give an indication of
the upper limit of the Eulittoral, which was approximately the
same as the upper limit of a small brown limpet, Patella
granularis. The blackish crust of Cyanophyceae present in the
Littoral Fringe was more obvious here, interspersed among
the barnacles, and it is worth noting that the barnacles
themselves often had a greenish tinge, presumably due to
epizoic or boring algae. A few Littorina also extended down-
wards among the upper barnacles. It was in this upper
Eulittoral zone that the Porphyra capensis belt was found
below Littorina and mixed with the barnacles and brown
limpets. Hildenbrandia rubra was also found at this level, but
only in damp depressions. On some sheltered rocks, Ulva
formed an ill-defined belt below Porphyra.

The next lower algal belt was that of Nothogenia erinacea,
especially prominent in sheltered places followed by scattered
clumps of Aeodes orbitosa at the lower limits of Chthamalus
and of the upper Eulittoral algae.

Below Aeodes, no single algal species dominated and the
lower part of the Eulittoral was occupied by a mixed, mainly
red algal, community, chiefly Champia lumbricalis, Gigartina
stiriata, Gigartina radula, and Iridaea capensis, with some
Cladophora, Polysiphonia, and articulated and crustose
corallines.

Of the animals at lower Eulitttoral levels, limpets were
most evident, Patella granularis L. penetrated from higher
up, to be joined by the larger Patella granatina L. at about the
same level as Chthamalus dropped out, and next by P.
argenvillei Krauss. This last species extended into the Sub-
littoral Fringe where Patella barbara L. was also present.
Forming a striking feature on more sheltered rocks was a
thick encrustation of the gritty tubes of the polychaete worm,
Gunnarea capensis (Schmarda), which continued down to
low- water mark. Mussels were present but not common.

The Sublittoral Fringe

A well-marked Sublittoral Fringe was dominated by Laminaria
pallida and the hollow-stiped L. schinzii (but see note below
under L. pallida in the annotated list of species). Many of the

* The small snail of the littoral fringe was originally identified for us under this
name. We note that Penrith & Kensley (1970a,1970/>) and Kensley & Penrith
(1980) refer to what is almost certainly the same entity as Littorina punctata
Philippi. In our opinion, however, these animals and those also referred to as
Littorina punctata that commonly inhabit the littoral fringe in the Gulf of
Guinea are not identical and, to avoid confusion, we have tentatively retained
the original name provided.

LAWSON, SIMONS & ISAAC

species of the mixed red algal belt of the lower Eulittoral,
such as Champia lumbricalis, Gigartina stiriata, G. radula,
and geniculate and encrusting Corallinaceae, also extended
into the Sublittoral Fringe where they were joined by species
apparently restricted to that zone, such as Hypnea spp.
Hypnea spicifera was found only locally in the protected
shallows of the harbour. Gracilaria verrucosa occurred in the
drift, and its presence in quantity is reported by Rotmann
(1987).

ii) Elizabeth Bay

Very gently sloping rocks in the centre of the rather shallow
open bay were studied. Although the outermost rocks were
exposed to strong wave-action, those inshore were appreci-
ably sheltered but never as much as the rocks at Luderitz.

The Littoral Fringe

This level was occupied by rather large individuals of Littorina
knysnaensis, very common in some places and less frequent in
others. Porphyra capensis was present to some extent.

The Eulittoral

The upper part of this zone was occupied mostly by limpets
(Patella granularis), and barnacles were not observed at this
level. Porphyra capensis occurred sporadically.

Below the Porphyra belt but still within the limpet belt,
Aeodes orbitosa, associated with Splachnidium rugosum, was
commonly found. At this level Chordariopsis capensis was
also very frequent but mostly in small pools and often
accompanied by Scytosiphon lomentaria.

Algae overlapping with this Aeodes-Splachnidium belt
were Centroceras clavulatum, Caulacanthus ustulatus, and
occasional plants of Corallina, Ulva, and Codium. Below
these was a belt of Gunnarea capensis tubes, sometimes as
much as 0.30 m in thickness above the rock surface. Growing
on this were several algal species, notably Ceramium sp.,
Leathesia difformis, and Cladophora sp., the last-mentioned
sometimes carrying the Leathesia. It should be noted that a
second type of worm tube, probably a Pomatoleios, was
sometimes present. Much harder and smoother than Gunnarea,
it often supported coralline crusts or tufts of Gelidium
micropterum and occurred frequently mixed with mussels up
to the lower level of Aeodes.

Near the lower limit of the Gunnarea belt began a belt of
Champia lumbricalis, often epiphytized by Aristothamnion
collabens, which extended into the Sublittoral Fringe to form
an undergrowth to the kelps. Large limpets, often bearing
small barnacles, formed fairly distinct areas interspersed with
Gunnarea in the lower part of its belt.

The Sublittoral Fringe

This was dominated by Laminaria pallida and Laminaria
schinzii but Ecklonia maxima was present especially in pools
(Plate 1). Other plants commonly found were Pachymenia
carnosa, Gigartina stiriata, G. radula, and Champia lumbri-
calis. In addition there were patches of Petalonia debilis,
Centroceras clavulatum, Cladophora spp., and crustose coral-
lines. Animals included Patella granatina, often bearing small
barnacles, the red sponge Hymeniacidon perlevis (Montagu),
and the starfish Asterina exigua (Lam.). Some patches of
mussels were present but by no means enough to account for
the vast quantities of shells thrown up on nearby beaches.
Patella compressa L. and Polysiphonia virgata were found on
the stipes of cast-up Laminaria.

ALGAL FLORA OF NAMIBIA

157

Plate 1 Kelp beds of the sublittoral fringe exposed at low spring tides at Elizabeth Bay. Laminaria pallida, Laminaria schinzii, and Ecklonia

maxima are present.

iii) Halifax Bay

A very brief visit was made to Halifax Bay where there was a
considerable extent of rocky coast and low cliff. Despite some
protection from the offshore penguin-populated islands, wave
exposure was strong in this area. The species present, and
their distribution on the shore, were much more in accord
with the typical pattern of zonation found on the west coast of
South Africa. Several species not found at Liideritz itself
were conspicuous here. Splachnidium rugosum was fairly
common in the upper Eulittoral, whilst Ecklonia maxima was
abundant in the sublittoral kelp beds. This counteracted the
impression we had gained from our examination of the rocks
near Liideritz itself that a number of species characteristic of
the west coast flora of South Africa had already disappeared
at about the latitude of Liideritz, and indicated that the
sheltered conditions of the harbour itself were more likely to
be responsible for these apparent absences.

(b) The Swakopmund area

The most northerly locality visited was Swakopmund (22
40' S) just within the tropics. Intertidal zonation was ex-
amined on the concrete blocks of the harbour wall and at two
points south of the town where there were extensive outcrops
of beach rock. The contrast between the rich algal flora of
the intertidal region and the extremely barren desert lying

directly behind the beach was even more marked than at
Liideritz.

i) The harbour wall

The Littoral Fringe

This was occupied by Littorina knysnaensis, accompanied by
some Siphonaria capensis, though more of the latter were
found in the Eulittoral. Plants of Porphyra capensis, though
not extending as high as the snails, were scattered throughout
the intertidal.

The Eulittoral

The more wave-exposed south-western faces of the harbour
wall could be divided into two subzones, an upper one in
which the dominant animals were Chthamalus dentatus and
Patella granularis, and a lower one in which occurred dense
carpets of small specimens of the mussel Perna perna and
large limpets.

In the upper subzone, the main algae found were Chaeto-
morpha, which extended more or less throughout, with
Iridaea capensis and some Petalonia debilis in its lower part.
In the upper reaches of the lower Eulittoral subzone, Poly-
siphonia, mixed with occasional plants of Ulva and of Chylo-
cladia capensis, formed a marked belt. Iridaea capensis bridged
the upper and lower subzones. The lower part of the lower

158

LAWSON, SIMONS & ISAAC

Plate 2 Laminaria schinzii dominating the sublittoral fringe near the concrete pier at Swakopmund.

Plate 3 Porphyra capensis on sheltered rocks behind the concrete pier at Swakopmund.

ALGAL FLORA OF NAMIBIA

Eulittoral subzone was occupied by Chondria capensis down
to the beginning of the Laminaria belt.

The Sublittoral Fringe

Due to heavy wave-action it was not possible to investigate
this level except to note that it appeared to be dominated by
the hollow-stiped Laminaria schinzii (Plate 2).

ii) Sheltered rocks behind the pier

The algal distribution was very different here, being less
distinct due to the broken nature of the rocks; good shelter
from strong surf also meant that the algae were telescoped
into the lower part of the Eulittoral. Enteromorpha was a
conspicuous feature, sometimes accompanied by Vrospora
sp. on the upper parts of the rocks. Porphyra capensis
was also very common (Plate 3), though not attaining quite
the same height up the shore as did the green algae. Cyano-
phyceae were also spread over the rocks, especially in the
form of a blackish powdery covering. The animals most in
evidence were Littorina knysnaensis and mussels (Modiolus
sp.), though our collections also included Balanus algicola,
Siphonaria capensis Quoy & Gaimard, and Patella granularis.

iii) Rocks south of Swakopmund

Our study of two fairly extensive and irregular outcrops of
gently-sloping rock, interrupting the sandy beaches, yielded
the following information.

The Littoral Fringe

This zone was present only where the rocks stood high

enough to accommodate it. Littorina knysnaensis was again

159

the dominant organism though it occurred also well down into
at least the middle of the Eulittoral zone. Porphyra capensis
was present in fairly large quantity, forming a narrow band
which did not appear to extend downwards as at some other
localities; not all samples taken could be attributed with
certainty to P. capensis since there were puzzling variants,
especially among the smaller forms; in particular we draw
attention to a small form with a distinctive rounded margin
(Plate 4), possibly an undescribed species. We also collected
Siphonaria sp. and a black and white limpet from the Porphyra
belt; at a somewhat lower level small plants of Viva were
present, mainly in depressions.

The Eulittoral

Barnacles and the small brownish Patella granularis with
some Littorina knysnaensis occupied the upper portion of this
zone; algae were few but included some Chaetomorpha,
mostly confined to shallow pools, and scattered small plants
of Viva.

Near the middle a subzone was discernable; mussels began
to replace barnacles, especially on the somewhat more
exposed parts where the former were generally rather larger;
Patella granularis, however, extended down into the mussel
subzone, where it was joined by some of the larger limpet
species. The upper limit of mussels and large limpets marked
the beginning of a much greater algal cover in the upper part
of the subzone (Plate 5), forming a distinct girdle of red algae
of which Caulacanthus ustulatus and Gymnogongrus glomera-
tus were the chief components with Nothogenia erinacea,

Plate 4 Plants of a small form of Porphyra found at Swakopmund.

160

LAWSON, SIMONS & ISAAC

-__ ~

Plate 5 The Eulittoral zone on rocks south of Swakopmund. Note that the upper part is very arid and practically devoid of algae, whereas the
lower part is covered by a dense mat of mixed algae including Caulacanthus ustulatus, Gymnogongrus glomeratus, and Centroceras davulatum.

Chylocladia capensis, Acrosorium maculatum, and much
Centroceras davulatum, which often occurred epiphytically
on the Caulacanthus.

Below this Caulacanthus-Gymnogongrus girdle was an-
other slightly less distinct belt dominated by Chondria capen-
sis, but including Pterosiphonia doiophylla, Ralfsia expansa,
Cladophora spp.,and, continuing into the Sublittoral, genicu-
late and crustose corallines.

The lowermost parts of the Eulittoral bore, in addition, a
number of larger red algae, perhaps representing elements of
the Sublittoral Fringe flora though rising above the Laminaria
level. They included species of Plocamium, such as P. cornu-
tum and P. rigidum, as well as the large fleshy fronds of
Pachymenia carnosa. (Plate 6) and Aeodes orbitosa (Plate 7).

The Sublittoral Fringe

This final zone was dominated by Laminaria schinzii but
included many of the algae found in the lower parts of the
Eulittoral corallines, Plocamium spp., and Pachymenia,
which together formed an undergrowth to the kelps. We did
not find Ecklonia maxima. The dominance of L. schinzii was
also indicated by the large quantity of this species thrown up
on nearby beaches.

c) General comments

The general features of algal zonation in the Liideritz area are

similar to those of the South African coast as described by

Isaac (1937b). Both areas have an upper Porphyra belt. This
is followed downwards by a Nothogenia belt, which at
Liideritz consists of N. ovalis, though this belt did not seem to
be represented at Elizabeth Bay. An Aeodes orbitosa belt was
also present at both Liideritz and at Elizabeth Bay. Aeodes
was accompanied by Splachnidium rugosum and Chordariop-
sis capensis. Bifurcaria brassicaeformis was apparently al-
together absent from Namibia. A Champia lumbricalis belt,
accompanied by species of Gigartina and Iridaea capensis, at
Liideritz was also present at Elizabeth Bay; Iridaea thus
appears to be somewhat lower on the shore here than further
south where it occurs in the Aeodes belt. Finally, a kelp belt is
present. at all localities, with Laminaria pallida and a hollow-
stiped form identified as L. schinzii found everywhere. One
of us (R.H.S.) has established that similar hollow forms
dominate the sublittoral as far south as Cape Columbine, the
western headland of St. Helena Bay about 120 km north of
Cape Town. Ecklonia maxima was also present at all locali-
ties, except for the sheltered shores at Liideritz, but usually as
a small plant. Where present, epiphytes seemed to prefer
Ecklonia maxima to Laminaria schinzii; Laminaria pallida
was usually without epiphytes. On the west coast of South
Africa, Suhria vittata occurs mostly associated with Ecklonia
maxima but in the Liideritz region it was often found on rocks
as well as on limpets in the intertidal.

Northwards, at Swakopmund, the situation was noticeably
different. The algal belts appeared to be narrower and lower

ALGAL FLORA OF NAMIBIA

161

Plate 6 Pachymenia carnosa growing on rocks just south of Swakopmund.

on the shore in this area than they were further south. Both
the Porphyra and the Nothogenia belts were present, at least
on the rocks south of Swakopmund. The Aeodes belt was not
represented here though Iridaea capensis was recorded from
the harbour wall at an appropriate level. Isaac's Champia-
Gigartina belt was much modified: Champia lumbricalis was
present in limited quantity, but the larger species of Gigartina
(G. stiriata, G. radula, and G. scutellata) were not found at
all. Finally, in the kelp bed Ecklonia maxima had, so far as
we could ascertain, completely disappeared and Laminaria
schinzii, rather than L. pallida, appeared to be dominant.

THE PHYTOGEOGRAPHICAL STATUS OF
THE NAMIBIAN MARINE ALGAL FLORA

The biogeographical status of the marine flora and fauna of
the western coast of southern Africa has been a somewhat
controversial topic since the biota have been regarded by
some authors as cold temperate and by others as warm
temperate. Stephenson (e.g. 1948) was the leading proponent
of the former view and in this he was followed by Hedgpeth
(1957) who, largely on the basis of a synthesis by Ekman
(1953), produced a map of the littoral zoogeographic prov-
inces of the world. In this map Hedgpeth placed a cold
temperate zone on the western side of southern Africa

directly abutting on to, and indeed overlapping with, the
tropical zone of western Africa. In all other regions of the
world he had indicated a warm temperate zone lying between
the cold temperate and tropical zones. Others who followed
this view were Knox (1960) and Michanek (1979). Ekman
(1953) regarded the region in question as warm temperate, as
did Briggs (1974), Bolton (1986), Hoek (e.g. 1984), and
Liining (1985). It should be noted that the question is seen
from different standpoints by different authors; thus Ekman,
Briggs, and Hedgpeth were concerned soley with animal
distributions, whereas Bolton, Hoek, and Liining dealt with
plants. Stephenson took both into account. Again, Stephenson
was concerned only with intertidal organisms and his bound-
aries are decidedly sharper than those produced by some
other authors, who included consideration of the deep sub-
littoral where distributions show fewer differences between
east and west. This is due to the fact, as pointed out by Brown
& Jarman (1978), that sea temperatures are much less vari-
able in deeper waters, ranging from only about 12-14C
throughout the year from Liideritz on the west coast to Port
Elizabeth in the east. The subject is well reviewed by Brown
& Jarman (1978) who adhered to the Stephenson viewpoint,
and by Bolton (1986) who takes the opposite line; it is also
mentioned by John & Lawson (1989) and Lawson (1988).

Bolton (1986) points out that Stephenson (1948) con-
sidered large stands of kelps to indicate cold temperate
conditions, as in the northern hemisphere. Had Stephenson
realized that Ecklonia maxima and Laminaria pallida under

162

LAWSON, SIMONS & ISAAC

Plate 7 Aeodes orbitosa growing over encrusting corallines on the pier at Swakopmund.

laboratory conditions actually grow best at higher tempera-
tures than most other kelps (17.5-20C for optimal gameto-
phyte growth for the former and 15C for the latter), he might
have concluded differently. Similarly, Macrocystis angusti-
folia has an optimum for gametophytic growth at 14.16C and
17.5C for reproduction (Branch, 1974). But it should per-
haps be borne in mind that temperatures which are optimal
for growth in the laboratory do not always necessarily reflect
what actually happens in nature. The figures given by Bolton
(1986), for instance, indicate that Ecklonia occurs on parts of
the west coast of South Africa where the mean temperature
for the warmest month does not rise much above 13, which is
well below the temperatures for optimum growth given
above. Further possible support for the warm temperate
view, however, is given by Lawson (1988), who has sug-

gested, on the basis of ordination, that the marine flora of
the western side of southern Africa has more in common
with those of Uruguayo-Bonaerense and Uruguay in South
America than with that of the more strictly cold temperate
one of Patagonia (see Kiihnemann, 1972). If this is indeed a
warm temperate marine climate continuous with that of the
south coast of South Africa how may the biotic discontinuity
that exists between the two be explained? A possible reason
to explain the somewhat anomalous situation where two
largely different warm temperate floras lie adjacent to one
another without completely mixing, may be that mean sea
temperatures along the south coast are generally appreciably
higher than those of the west coast and also show a wider
range between coldest and warmest months (see fig. 2 in
Bolton, 1986).

ALGAL FLORA OF NAMIBIA

163

Table 1 Distribution of the Namibian marine algal flora within and Table 1 cont.

to the north and south of Namibia. The species are arranged in a

geographical sequence.

Angola Namibia South Africa Species

North Centre South West South and

Species south-east

Angola Namibia South Africa

North Centre South West South and
south-east

Codium decorticatum + + +

Viva uncialis + + H

Enteromorpha linza + + H

E. compressa + +

Caulacanthus ustulatus + + + H

Centroceras clavulatum + + + H

Gracilaria verrucosa + + + H

Plocamium suhrii + +

Acrosorium maculatum + + -\

Bryopsis plumosa + + H

Hypnea musciformis + +

Viva fasciata + +

Grateloupia filicina + H

Colpomenia sinuosa + H

Enteromorpha tubulosa +

Gigartina teedii + +

Cladophora capensis + + H

C. contexta + + H

C. flagelliformis + + H

C. mirabilis + + H

Laminaria schinzii + + H

Aristothamnion collabens + + -

Ceramium atrorubescens + + -

Gymnogongrus complicatus + +

G. dilatatus + + -

G. vermicularis + +

Hildenbrandia rubra + +

Kallymenia schizophylla + +

Pachymenia camosa + +

Phyllymenia belangeri + +

Polysiphonia urbana + +

Suhria vittata + +

Nothogenia ovalis + +

Chaetomorpha robusta + +

Streblocladia camptocladia + +

S. fasciculifera +

Hypnea ceramioides +

Codium fragile + +

Ralfsia expansa + +

Ceramium arenarium + +

Chondria capensis + +

Gymnogongrus glomeratus + +

Iridaea capensis + +

Nothogenia erinacea + +

Plocamium rigidum + +

Porphyra capensis + +

Pterosiphonia cloiophylla + +

Aeodes orbitosa + +

Ceramium flaccidum +

Hypnea spicifera +

Platysiphonia miniata +

Acrosorium acrospermum + +

Ceramium diaphanum + +

Heringia mirabilis + +

Tayloriella tenebrosa + +

Bryopsis caespitosa +

Hincksia confervoides +

Plocamium corallorhiza +

P. vulgare +

Gelidium pristoides Unspecified

Codium duthieae + +

Enteromorpha prolifera +

Endarachne binghamiae +

Arthrocardia palmata +

Hypnea ? tenuis +

Viva nematoides +

Hincksia granulosa +

Acrosorium cincinnatum
Antithamnion leptocladum
Audouinella hypneae
Cryptonemia hibernica
Cryptopleura calophylloides
Epymenia capensis
Erythrotrichia welwitschii
Gelidium pusillum
Iridaea elongata
Lomentaria patens
Nothogenia magnifica
Ophidocladus simplidusculus
Rhodophyllis reptans
Schottera nicaeensis
Stylonema alsidii
Enteromorpha atroviridis
E. bulbosa

Chordariopsis capensis
Laminaria pallida
Carpoblepharis flacdda
Ceramium obsoletum
C. planum complex
Epymenia obtusa
Gigartina bracteata
G. radula

Polysiphonia virgata
Schizymenia obovata
Viva capensis
Papenfussiella gradlis
Callithamnion hookeri
Carpoblepharis minima
Gymnogongrus corymbosa
Haraldiophyllum bonnemaisonii
Rhodymenia linearis
Petalonia debilis
Chylocladia capensis
Plocamium cornutum
P. glomeratum
Streblocladia corymbifera
Chaetomorpha linum
Heterosiphonia dubia
Rhodymenia natalensis
Arthrocardia carinata
Derbesia lamourouxii
Actinococcus latior
Euhymenia schizophylla
Herposiphonia heringii
Bryopsis tenuis
Codium isaadi
Desmarestia firma
Ecklonia maxima
Scytosiphon lomentarius
Arthrocardia filicula
A. setchellii
Botryocarpa prolifera
Botryoglossum platycarpum
Champia lumbricalis
Delesseria papenfussii
Gelidium micropterum
Gigartina scutellata
Hymenena venosa
Pleonosporium harveyanum
Polyopes constrictus
Polysiphonia atrorubescens
Trematocarpus flabellatus
Leathesia difformis
Splachnidium rugosum
Gigartina stiriata

164

The list of species given below incorporates all known
records and contains 136 identified species whose known
distributions on southern African shores are displayed in
Table 1 , as well as at least 35 taxa assigned only to genera. It
is evident that most (97) Namibian seaweeds occur also on the
west coast of South Africa: even in the north of Namibia 44
species out of 57 identified plants are species that occur on the
west coast of South Africa with 21 endemics involved; the
other two segments, namely central (Swakopmund area) and
south (Liideritz area), contain respectively 89 and 83 identi-
fied species. An interesting point is that whereas about 74%
of all the taxa present in the south segment are common with
the west coast of South Africa, the comparative figure for the
central segment is only 62% in fact, Swakopmund is notable
for having 20 (22% of the total identified) species apparently
not occurring on the west coast of South Africa. By contrast
elements of the Angolan seaweed flora are remarkable for
their very poor representation 14 in all, with eight of these
of very wide distribution or even cosmopolitan.

The overall concept of the Namibian seaweed flora that
emerges is of an attenuating northward extension of the flora
of the west coast of South Africa that abruptly terminates
somewhere near the border with Angola. Less noticeable, but
nevertheless significant, is the change of species composition
at Swakopmund. Others have also observed biotic changes in
that vicinity, for instance, a reduction in frequency of the
Cape rock lobster, Jasus lalandii, north of 25S (Pollock &
Beyers, 1981). A possible explanation for these changes is
that the relative wind stress and concomitant water circula-
tion is reduced here and to the north (Agenbag & Shannon,
1988).

Table 1 shows discontinuities that seem somewhat anoma-
lous. Thus Grateloupia filicina, for example, is apparently
absent between the Kunene River mouth and Luderitz, and
Colpomenia sinuosa is recorded only in the Luderitz area of
Namibia while being present in Angola and the southern
Cape. Again Aeodes orbitosa, and some othe species such as
Ceramium flaccidum, have been found all the way from
northern Namibia to the southern coast of Africa except at
Swakopmund. Two possible reasons for these anomalies are
that there have been misidentifications and/or inadequate
collection since it is somewhat presumptuous to discuss
discontinuities of distribution on the rather meagre data
available. It is hoped, however, that this presentation will
serve as a useful check-list for further studies of the respective
floras.

ANNOTATED LIST OF SPECIES

The Western African seaweed checklists of Lawson & Price
(1969) and Price et al. (1978, 1986, 1988) cite unpublished
lists prepared by one of us (R.H.S.). These lists include the
specimens collected on the trip with which this paper is
concerned, together with identifications of algae collected by
Penrith & Kensley, and other miscellaneous records from
Namibia. We have included all these in the list that follows
and have added all earlier records of marine algae from this
coast as well as algae mentioned by Penrith & Kensley (1970 a
& b), Kensley & Penrith (1973, 1980) and the recent identifi-
cations by Wynne (1986). Where records of other authors are
cited an asterisk(*) indicates that we have been able to

LAWSON, SIMONS & ISAAC

confirm independently the presence of the species in question
at the given locality. Sites may be identified by the following
letters:

AB Agate Beach, AF Angra Fria, CF Cape Frio (and
False Cape Frio), D Diaz Point (Luderitz area), EB
Elizabeth Bay, FB Flesh Bay, G Grossebucht, HB
Halifax Bay, HL Honolulu (near Kunene River mouth),
K Kaokoveld, KR Kunene River, L Luderitz, MB
Mowe Bay, N Namibia (used when locality is not men-
tioned), RP Rocky Point, S Swakopmund, TB Torra
Bay, TC Terrace Bay, TS Toscanini, U Unjab, WB
Walvis Bay.

Specimens are deposited in BOL, the Herbarium of the
Department of Botany, University of Cape Town.

CHLOROPHYCEAE

Bryopsis caespitosa Suhr ex Kiitz. MB

Bryopsis plumosa (Huds.) Agardh EB, N (Lawson & Price,

1969), S.

Bryopsis tenuis Levr. L, ?N (Lawson & Price, 1969).
Bryopsis spp. EB, L, RP, S, TB, TC.

Chaetomorpha aerea (Dillwyn) Kiitz. see Chaetomorpha linum.
Chaetomorpha linum (O. F. Mull.) Kiitz. S (Wynne, 1986, as

C. aerea)*.
Chaetomorpha robusta (Aresch.) Papenf. MB, RP, S, TB,

TC.

Chaetomorpha spp. D, S, TB.
Cladophora capensis (Agardh) De Toni D, EB, HB, L

(Penrith & Kensley, 1970a)*, MB, RP (Penrith & Kensley,

1970ft)*, S, TB,TC.

Cladophora contexta Levr. EB, L, MB, S.
Cladophora flagelliformis (Suhr) Kiitz. EB, S (Wynne,

1986)*, TB,L.
Cladophora mirabilis (Agardh) Rabenh. CF (Kensley &

Penrith, 1980), KR (Kensley & Penrith, 1980), L, N (Lawson

& Price, 1969), RP, S (Wynne, 1986)*, TB, TC.
Cladophora virgata (Agardh) Kiitz. see Cladophora

flagelliformis.
Cladophora sp. EB.
Codium decorticatum (Woodw.) M. Howe MB, N (Lawson &

Price, 1969), RP, S (Silva, 1960; Wynne, 1986)*.
Codium duthieae P. C. Silva L, MB, ?N (Lawson & Price,

1969), RP (Penrith & Kensley, 1970&), S.
Codium fragile subsp. capense P. C. Silva D, EB, L (Penrith

& Kensley, 1970a, Silva, 1959), N (Lawson & Price, 1969;

Schmidt, 1923 as C.fragile), RP, S (Silva, 1959; Wynne,

1986)*, TB,U.
Codium isaacii P. C. Silva L (Silva, 1959), N (Lawson &

Price, 1969).
Codium sp. S.

Derbesia lamourouxii (J. Agardh) Solier L.
Enteromorpha atro-viridis (Levr.) Wynne G, L (Penrith &

Kensley, 1970a as Ulva atroviridis*) , S (Wynne, 1986).
Enteromorpha bulbosa (Suhr) Kiitz. EB, S.
Enteromorpha compressa (L.) Grev. S.
Enteromorpha linza (L.) J. Agardh L, S.
Enteromorpha prolif era (O. F. Mull.) J. Agardh MB, RP.
Enteromorpha tubulosa Kiitz. RP.
Enteromorpha spp. AF (Kensley & Penrith, 1980), CF

(Kensley & Penrith, 1980), HB, L, N (Lawson & Price,

1969), S.
Viva atroviridis Levr. see Enteromorpha atro-viridis.

ALGAL FLORA OF NAMIBIA

Viva capensis Aresch. MB, L, S (Wynne, 1986).

Ulvafasciata Delile S.

Ulva nematoidea Bory S (Wynne 1986)*.

Ulva uncialis (Kiitz.) De Toni L (Pilger, 1908 as U. uncialis

Suhr), MB, N (Dinter, 1928 as Ulva capensis; Lawson &

Price, 1969).
Ulva spp. EB, L, MB, N (Lawson & Price, 1969), RP, S, TB,

TC, U.
Urospora sp. S.

PHAEPHYCEAE

Chordaria flagelliformis (O. F. Mull.) Agardh N (Dinter,
1919) probably Chordariopsis capensis according to Price
et al. (1978).

Chordariopsis capensis (Agardh) Kylin CF (Kensley &
Penrith, 1980)*, EB, L (Penrith & Kensley, 1970a)*, N
(Price et al., 1978), RP, S (Wynne, 1986)*, TB.

Colpomenia sinuosa (Roth) Derb. & Sol. AB, D, EB, N
(Price etal., 1978).

Desmarestia firma (Agardh) Skottsb. L (Price et al., 1978).
See notes in Price et al. (1978) on this entity.

Ecklonia maxima (Osbeck) Papenf. EB, HB, L, N (Price et
al., 1978), S (Wynne, 1986 'in the drift'. Wynne also
mentions that his species occurs at RP and cites a letter
from Kensley on this, but we have not been able to find it
referred to in Penrith and Kensley [1970ft]).

Ectocarpus spp. N (Price et al., 1978), RP, S.

Endarachne binghamiae J. Agardh RP.

Hincksia granulosa (Smith) P. C. Silva S (Wynne, 1986 as
Giffordia granulosa).

Laminaria digitata forma ensifolia (Kiitz.) Foslie N (Dinter,
1922), WB (Foslie, 1893). See note in Price et al.
(1978).

Laminaria ochroleuca Bach. Pyl. N (Delf & Michell, 1921).
See note in Price et al. (1978).

Laminaria pallida Grev. EB, L, N (Delf & Michell, 1921;
Price et al., 1978; Schmidt & Gerloff, 1957). WB (Delf &
Michell, 1921). It should be noted that some doubt attaches
to the presence of L. pallida in the Liideritz area since
differences between L. pallida and L. schinzii are not clear
cut and juvenile and inshore plants of the latter often have
solid stipes.

Laminaria schinzii Foslie EB, L, MB, N (De Toni, 1895;
Dinter, 1922; Feldmann, 1946; John et al., 1981; Lawson et
al., 1975; Price et al., 1978; Pilger, 1908; Schmidt &
Gerloff, 1957), RP (Penrith & Kensley, 1907ft)*, S (Wynne,
1986)*, TC, WB (Barton, 1893; Foslie, 1893).

Laminaria sp. L (Penrith & Kensley, 1970a).

Leathesia difformis (L.) Aresch. D, EB, L, N (Price et al.,
1978).

Myriogloia sp. EB, N (Price et al., 1978), S.

Papenfussiella gracilis Kylin N (Price et al., 1978), S.

Petalonia debilis (O. F. Mull.) Kuntze EB, N (Price et al.,
1978), S.

Ralfsia expansa (J. Agardh) J. Agardh CF, EB, N (see note in
Price et al. [1978]), S.

Ralfsia sp. KR (Kensley & Penrith, 1980).

Scytosiphon lomentaria (Lyngbye) Link EB, L, N (Price et
al., 1978).

Splachnidium rugosum (L.) Grev. EB, G, HB, L (Penrith &
Kensley, 1970a)*, N (Price et al., 1978).

?Zonaria tempta HB, N (see note in Price et al. [1978]).

165

RHODOPHYCEAE

Acrosorium acrospermum (J. Agardh) Kylin RP, TB, TC,

TS, N (Price etal., 1986), S.
Acrosorium cincinnatum Wynne S (Wynne, 1986), N (Price et

al., 1986)*.
Acrosorium maculatum (Kiitz.) Papenf. L, S, N (Lawson et

al., 1975; Price etal., 1986).

Acrosorium sp. L, MB, S, TB, TC, N (Price et al., 1986).
Actinococcus latior Schmitz L (Pilger, 1908), N (Price et al.,

1978), WB (Dinter, 1917).
Aeodes orbitosa (Suhr) Schmitz AF (Kensley & Penrith,

1980), D, EB, CF, (Kensley & Penrith, 1980), G, HB, KR

(Kensley & Penrith, 1980), L (Penrith & Kensley, 1970a)*,

MB, (Penrith & Kensley, 1970ft), N (Price et al., 1986),

RP, U.

Aeodes sp. N (Price et al., 1986).
Antithamnion leptocladum (Mont.) Wynne S (Wynne, 1986),

N (Price etal., 1986).

Antithamnion sp. L, RP, N (Price et al., 1986).
Aristothamnion collabens (Rudolphi) Papenf. EB, G, HB, L,

N (Dinter, 1918 as A. purpuriferum; Price et al., 1986), S

(Wynne, 1986)*, TC.

Arthrocardia attenuata Manza see Arthrocardia palmata.
Arthrocardia carinata (Kiitz.) Johansen L (Pilger, 1908 as

Corallina carinata), N (Price et al., 1986), S.
Arthrocardia filicula (Lam.) Johansen in Seagrief L, N (Price

etal., 1986).
Arthrocardia palmata (Ellis & Sol.) Aresch. MB, N (Price et

al., 1986, as Arthrocardia attenuata).
Arthrocardia setchellii Manza L.
Arthrocardia sp. D, L (Penrith & Kensley, 1970a)*, RP

(Penrith & Kensley, 1970ft)*, TB, N (Price et al., 1986),

TC.
Audouinella hypneae (B0rgesen) G. W. Lawson & D. John S

(Wynne, 1986), N (Price et al., 1986).
Bostrychia moritziana (Sender ex Kiitz.) J. Agardh N (Post,

1963; Price etal., 1986).

Botryocarpa prolifera Grev. EB, L, N (Price et al., 1986).
Botryoglossum platycarpum (Turner) Kiitz. EB, L (Simons,

1974; Price etal., 1986).
Callithamnion hookeri (Dillwyn) Gray S (Wynne, 1986), N

(Price etal., 1986).
Carpoblepharis flaccida (Agardh) Kiitz. L (Pilger, 1908)*, N

(Price et al., 1986), S, WB (Delf & Michell, 1921).
Carpoblepharis minima Bart. N (Dinter, 1918; Price et al.,

1986), S (Wynne, 1986).
Carpoblepharis sp. N (Price et al., 1986), S.
Carradoria virgata (Agardh) Kylin see Polysiphonia virgata.
Caulacanthus ustulatus (Mert.) Kiitz. EB, L, MB, N (Price et

al., 1986; Searles, 1968), RP, S (Wynne, 1986)*, TB.
Caulacanthus sp. N (Price et al., 1986), TB.
Centroceras clavulatum (Agardh) Mont. D, EB, L (Pilger,

1908)*, MB, N (Price et al., 1986; Searles, 1968), RP, S

(Wynne, 1986)*, TB,TC.
Ceramium arenarium Simons L (Simons, 1966), N (Price et

al., 1986; Simons, 1966), S (Wynne, 1986), TB.
Ceramium atrorubescens Kylin L (Simons, 1966), S Simons,

1966, Wynne, 1986), TB, N (Price et al., 1986), U, WB

(Simons, 1966).

Ceramium capense Kiitz. see Ceramium obsoletum.
Ceramium diaphanum (Lightf.) Roth TB (Dinter, 1919; Price

et al., 1986), WB (Delf & Michell, 1921).
Ceramium flaccidum (Kiitz.) Ardiss. G (Simons, 1966, as

166

Ceramium papenfussianum) HB, HL, KR, L, N (Price et

al., 1986, as Ceramium papenfussianum) .
Ceramium obsoletum^ Agardh EB (Simons, 1966), L (Simons,

1966), N (Dinter, 1917; Price et al., 1986 as C.capense),

WB (Delf & Michell, 1921 as C.obsolatum) (= Microdadia

capense Papenf.

Ceramium papenfussianum Simons See Ceramium flaccidum.
Ceramium planum complex sensu Simons 1968 L, N (Price et

al., 1986), S (Wynne, 1986).
Ceramium spp. S (Wynne, 1986), TC, TS, N (Price et al.,

1986).
Chaetangium erinaceum (Turner) Papenf. see Nothogenia

erinacea.

Chaetangium magnificum Pilger see Nothogenia magnifica.
Chaetangium ornatum (L.) Kiitz. see Nothogenia erinacea.
Chaetangium ovale (Suhr) Papenf. see Nothogenia ovalis.
Champia lumbricalis (L.) Desv. AB, D, EB, G, L (Penrith &

Kensley, 1970a; Simons, 1974), N (Price et al., 1986).
Chondria capensis (Harvey) Falkenb. CF, L, MB, N (Delf &

Michell, 1921; Lawson et al., 1975; Price et al., 1986;

Round, 1981; Schmidt & Gerloff, 1967; Simons, 1974), RP,

S (Wynne, 1986)*, WB (Dinter, 1919) TB, TC.
Chondria sp. CF (Kensley & Penrith, 1980), KR (Kensley &

Penrith, 1980), N (Price et al., 1986).
Chylocladia capensis Harvey L, N (Price et al., 1986), S.
Chylocladia sp. KR, N (Price et al., 1986), RP, TC.
Corallina carinata Kiitz. see Arthrocardia carinata.
Corallina sp. (Corniculariae) MB.
Corallina spp. S (Wynne, 1986), RP, N (Price et al., 1986), S,

TB, TS.
Cryptonemia hibernica Guiry & L. Irvine N (Price et al.,

1986), S (Wynne, 1986).
Cryptopleura calophylloides (J. Agardh) Wynne N (Price et

al., 1986), S (Wynne, 1986).

Delesseria papenfussii Wynne EB, N (Price et al., 1986).
Epymenia capensis (J. Agardh) Papenf. N (Dinter, 1926;

Schmidt & Gerloff, 1957, as Rhodymenia capensis Price et

al., 1986), WB (Delf & Michell, 1921, as Epymenia steno-

loba).
Epymenia obtusa (Grev.) Kutz. EB, L (Pilger, 1908)*, N

(Delf & Michell, 1921, Price et al., 1986, Simons, 1974), S

(Wynne, 1986)*, TC.

Erythrotrichia welwitschii (Rupr.) Batters S.
Euhymenia schizophylla Kiitz. L (Pilger, 1908), N (Price et

al., 1986).

Gelidium micropterum Kiitz. EB, L, N (Price et al., 1986).
Gelidium pristoides (Turner) Kiitz. N (Price et al., 1986).
Gelidium pusillum (Stackh.) Le Jolis N (Price et al., 1986), S.
Gelidium spp. MB, N (Price et al., 1986), RP.
Gigartina bracteata (Gmelin) Setch. & N. Gardner HB, S

(Wynne, 1986).
Gigartina radula (Esper) J. Agardh AB, D, EB, HB, L

(Penrith & Kensley, 1970a)*, S, TC.
Gigartina scabiosa (Kiitz.) Papenf. see G.scutellata.
Gigartina scutellata (Hering) Simons EB, L, N (Price et al.,

1988).
Gigartina stiriata (Turner) J. Agardh D, EB, G, L (Penrith &

Kensley, 1970a)*, N (Price et al., 1988).
Gigartina teedii (Roth) Lamouroux N (Dinter, 1921; Price et

al., 1988), RP, S (Wynne, 1986).

t The plant to which the name Ceramium obsoletum Agardh has been applied
is apparently unnamed since the Ceramium capense cited by Simons (1966: 164)
is not congeneric with Ceramium.

LAWSON, SIMONS & ISAAC

Gigartina spp. N (Price et al., 1988), RP (Penrith & Kensley,

19706)*.
Gracilaria verrucosa (Huds.) Papenf. L (Rotmann, 1987), N

(Isaac, 1956; Michanek, 1971, 1975; Price et al., 1988) S

(Wynne, 1986)*, TC, WB.
Grateloupiafilicina (Wulf.) J. Agardh EB, HB, L, N (Price et

al., 1988).
Griffithsia confervoides Suhr (including G. cymosa Simons)

MB, N (Price et al., 1988), TB, TC, TS.
Gymnogongrus complicatus (Kiitz.) Papenf. L, MB, N

(Price et al., 1988), S (Wynne, 1986)*, TB, TC, TS.
Gymnogongrus corymbosus J. Agardh N (Price et al., 1988),

5 (Wynne 1986).

Gymnogongrus dilatatus (Turner) J. Agardh L (Pilger,
1909)*, N (Price et al., 1988), S (Wynne, 1986)*, TC.

Gymnogongrus glomeratus J. Agardh AB, AF (Kensley &
Penrith, 1980), EB, D, L (Penrith & Kensley, 1970a)*, N
(Kensley & Penrith, 1980; Price et al., 1988), RP (Penrith

6 Kensley, 19706), S (Wynne, 1986)*, TC, TS.
Gymnogongrus vermicularis (Agardh) J. Agardh CF, HB,

KR, L, MB, N (Price et al.,1988), S, TC, TS.
Gymnogongrus spp. G, KR, MB, N (Price et al., 1988), S.
Haraldiophyllum bonnemaisonii (Grev.) Zinova prox. S

(Wynne, 1986).

Heringia mirabilis (Agardh) J. Agardh S (Wynne, 1986), TC.
Heringia sp. MB.

Herposiphonia heringii (Harvey) Falkenb. L.
Heterosiphonia dubia (Suhr) Falkenb. S (Wynne, 1986).
Heterosiphonia sp. TB.
Hildenbrandia rubra (Sommerfelt) Menegh. EB, L, RP, S,

TB.

Hymenena venosa (L.) Kylin EB, L.

Hypnea ceramioides Kiitz. RP (Penrith & Kensley, 19706).
Hypnea eckloni Suhr see Hypnea musciformis note in Delf &

Michell (1921).
Hypnea musciformis (Wulf.) Lamouroux S (Wynne, 1986),

WB (Delf & Michell, 1921, as Hypnea eckloni).
Hypnea spicifera (Suhr) Harvey AB, KR (Kensley & Penrith,

1980), L, RP (Penrith & Kensley, 19706)*.
Hypnea (?)tenuis Kylin KR.
Hypnea spp. KR, L, MB, RP, S, TC.
Hypoglossum sp. TC.
Iridaea capensis J. Agardh CF, D, EB, G, HL, L (Penrith

& Kensley, 1970a)*; MB, S (Wynne, 1986, as Iridaea

capensis)* , TC.

Iridaea elongata Suhr WB (Delf & Michell, 1921).
Kallymenia schizophylla J. Agardh L, S (Wynne, 1986)*, TC.
? 'Kallymenia sp. L.
Lithophyllum sp. RP.
Lithothamnion sp. D, L (Penrith & Kensley, 1970a), RP

(Penrith & Kensley, 19706).

Lomentaria patens Kiitz. WB (Delf & Michell, 1921).
Microdadia sp. TC.

Neuroglossum binderianum Kiitz. EB, L.
Nitophyllum fissum (Grev.) J. Agardh L (Pilger, 1908). See

Hymenena venosa.
Nothogenia erinacea (Turner) Parkinson CF, L, MB, RP

(Penrith & Kensley, 19706 as Chaetangium erinaceum), L

(Pilger, 1908 as Chaetangium ornatum), S (Wynne, 1986)*,

TB, WB (Delf & Michell, 1921, as Chaetangium ornatum).
Nothogenia magnifica (Pilger) J. Price S (Pilger, 1908; Price et

al.,1986).
Nothogenia ovalis (Suhr) Parkinson G, L (Penrith & Kensley,

1970a as Chaetangium ovale)* , MB.

ALGAL FLORA OF NAMIBIA

Ophidodadus simpliciusculus (P. Crouan & H. Crouan ex J.

Agardh) Falkenb. S (Wynne, 1986).
Orcasia pulla Simons see Streblodadia camptodada.
Pachymenia carnosa (J. Agardh) J. Agardh AF (Kensley &

Penrith, 1980), CF (Kensley & Penrith, 1980)*, EB, KR

(Kensley & Penrith, 1980), L, MB, S (Wynne, 1986)*, TC.
Phyllymenia belangeri (Bory) Setch.& N.Gardner CF, L, S

(Wynne, 1986)*, TB, TC.
Platysiphonia miniata (J. Agardh) B0rgesen EB, L, MB, RP,

TB.

Platysiphonia sp. MB.

Pleonosporium harveyanum (J. Agardh) De Toni L.
Plocamium corallorhiza (Turner) Harvey K.
Plocamium cornutum (Turner) Harvey D, EB, L (Pilger,

1 908) *,S( Wynne, 1986)*.

Plocamium glomeratum J. Agardh S (Wynne, 1986)*.
Plocamium rigidum Bory L, RP, S (Wynne, 1986)*, TB, TC,

MB.

Plocamium suhrii Kiitz. RP, TC.
Plocamium vulgare Lamouroux CF, TC, MB.
Poly opes constrictus (Turner) J. Agardh L.
Polysiphonia atrorubescens (Dillwyn) Grev. L.
Polysiphonia urbana Harvey CF, EB, HL, RP, TC, U, WB

(Delf & Michell, 1921 as Polysiphonia corymbiferd).
Polysiphonia virgata (Agardh) Sprengel EB, G, S (Wynne,

1986)*.

Polysiphonia sp. S (Wynne, 1986). Polysiphonia spp. L, S, U.
Porphyra capensis Kutz. AF (Kensley & Penrith, 1980), CF

(Kensley & Penrith, 1980), D, EB, G, HL, L (Penrith &

Kensley, 1970a)*, N (Dinter, 1926; John et al., 1979;

Piccone, 1884; Pilger, 1908; Schmidt & Gerloff, 1957), RP,

S (Wynne, 1986), TC, U. See remarks on p. 159.
Porphyra sp. S.
Pterosiphonia doiophylla (Agardh) Falkenb. AB, L, RP, S

(Wynne, 1986)*, TC.

Rhodophyllis reptans (Suhr) Papenf. S (Wynne, 1986).
Rhodymenia linearis J. Agardh S (Wynne, 1986).
Rhodymenia natalensis Kylin S (Wynne, 1986).
Rhodymenia sp. CF (Kensley & Penrith, 1980), KR.
Schizymenia obovata (J. Agardh) J. Agardh L, S (Wynne,

1986)*.
Schottera nicaeensis (Lamouroux ex Duby) Guiry & Hollen-

berg S (Wynne, 1986).
Streblodadia camptodada (Mont.) Falkenb. AF (Kensley &

Penrith, 1980 as Orcasia pulla), CF (Kensley & Penrith,

1980 as Orcasia pulla), KR (Kensley & Penrith, 1980 as

Orcasia pulla), MB, RP, S (Wynne, 1986), TC, TS, U.
Streblodadia corymbifera (Agardh) Kylin RP, S (Wynne,

1986).

Streblodadia fasciculi/era (Kutz.) Falkenb. RP, TC.
Stylonema alsidii (Zanard.) Drew S (Wynne, 1986).
Suhria vittata (L.) J. Agardh EB, L (Pilger, 1908)*, MB, S

(Wynne, 1986)*.

Tayloriella tenebrosa (Harvey) Kylin MB, RP, S, TB, TS.
Tayloriella virgata (Agardh) Papenf. see Polysiphonia virgata.
Tayloriella sp. RP, TC.
Trematocarpus flabellatus (J. Agardh) De Toni EB, L, N

(Price et al., 1986).

CYANOPHYCEAE

Lyngbya sp. L (Penrith & Kensley, 1970a).

ACKNOWLEDGMENTS. The authors are grateful to the State Depart-
ment of Agriculture of the Union of South Africa (now the Republic

167

of South Africa) for providing a four-wheel-drive vehicle and all its
fuel and maintenance needs. They also thank the Administration of
Consolidated Diamond Mines for permission to visit Elizabeth Bay.
The University of Cape Town and the Council for Scientific and
Industrial Research helped by funding some of the travelling ex-
penses of the party.

We are indebted to Miss Judith Graves for her participation in the
original field excursion of 1957, and to Dr D. M. John and Mr J. H.
Price of The Natural History Museum, London, who read the paper
critically and provided additional information and advice, and to the
Photographic Unit of the museum for providing black and white
prints from the colour slides made by one of us (G.W.L.).

REFERENCES

Anon. 1963. Africa pilot 2. London.

Anon. 1964. Handbuch der Westkiiste Afrikas II. Teil von Calabar River bis

Cape Hangklip. Hamburg.
Barton, E. S. 1893. A provisional list of the marine algae of the Cape of Good

Hope. /. Bot., Land. 31. 53-56, 81-84, 110-114, 138-144, 171-177, 202-210.

1896. Cape algae. /. Bot. Land. 34: 193-198.

Bolton, J. J. 1986. Marine phytogeography of the Benguela upwelling region
on the west coast of southern Africa: a temperature dependent approach.
Botanica mar. 29: 251-256.

& Anderson, R. J. 1988. Gradient analysis of seaweed communities

around the southern tip of Africa. Third Internal. Phycol. Congr., Monash
Univ., 79S8Abstr.:5.

Branch, M. 1974. Limiting factors for the gametes of three South African

laminariales. Investl. Rep. Sea Fish. Brch. S. Afr. 104: 1-38.
Briggs, J. C., 1974. Marine zoogeography. New York.
Bright, K. M. F. 1938. The South African intertidal zone and its relation to

ocean currents. III. An area on the northern part of the west coast. Trans.

R. Soc. S. Afr. 26: 67-88.
Brown, A. C. & Jarman, N.. 1978. Coastal marine habitats. In M.J.A. Werger

(Ed.), Biogeography and ecology of southern Africa 2: 1239-1277. [Mono-

graphiae Biologicae 31]. The Hague.
Delf, E. M. & Michell, M. R. 1921. The Tyson collection of marine algae. Ann.

Bolus Herb. 3: 89-119.
De Toni, G. B. 1895. Sylloge algarum omnium hucusque cognitarum III.

Sylloge Fucoidearum. Padova.
Dinter, K. 1919-1928. Index, der aus Deutsch-Sudwestafrika bis zum Jahre,

1917 bekannt gewordenen Pflanzenarten. Reprium Spec. nov. Regni veg. 16:

239-244; 17: 303-311; 18: 13-16, 248-256, 423-444; 19: 315-320; 22:

108-112, 375-383; 23: 130-137, 363-371; 24: 302-304.
Ekman, S. 1953. Zoogeography of the sea. London.
Feldmann, J. 1946. La flore marine des lies Atlantides. Mem. Soc. Biogeogr. 8:

395-435.
Koslie, M. 1893. Ueber eine neue Laminaria (Laminaria schinzii) aus Westafrika.

Bull. Herb. Boissier 1: 91-94.
Hedgpeth, J. W. 1957. Marine biogeography. Geol. Soc. Am. Memoir. 67:

359-382.
Hoek, C van den 1984. World- wide latitudinal and longitudinal distribution

patterns and their possible causes as illustrated by the distribution of

Rhodophycean genera. Helgolander wiss. Meeresunters. 38: 227-257.
Hommersand, M. H., 1986. The biogeography of the South African marine red

algae: a model. Botanica mar. 29: 257-270.
Isaac, W. E. 1937a. South African coastal waters in relation to coastal currents.

Geographical Rev. 27: 651-664.
19376. Studies of South African seaweed vegetation. 1 . West coast from

Lamberts Bay to the Cape of Good Hope. Trans. R. Soc. S. Afr. 25:

115-151.

1949. Studies of South African seaweed vegetation. II. South coast:

Rooi Els to Gansbaai, with special reference to Gansbaai. Trans. R. Soc. S.
Afr. 32: 125-160.

1956. The ecology of Gracilaria confervoides (L.) Grev. in South Africa
with special reference to its ecology in the Saldanha-Langebaan lagoon.
Proc.lnt. Seaweed Symp. (Trondheim, 1955) 2: 173-185. London.

1957. The distribution, ecology and taxonomy of Porphyra on South

African coasts. Proc. Linn. Soc. Land. 168: 61-65.

& Hewitt, F. 1953. The morphology, geographical distribution and
ecology of Hypnea spicifera (Suhr) Harv. // S. Afr. Bot. 19: 73-84.

John, D. M. 1986. Littoral and sublittoral vegetation. In G. W. Lawson (Ed.)
Plant ecology in West Africa: systems and processes: 215-246. Chichester.
& Lawson, G. W. 1990. Littoral ecosystems of tropical West Africa. In A.
C. Mathieson & P. H. Nienhuis (Eds), Intertidal and littoral ecosystems.
Elsevier Series Ecosystems of the World 24 (in press).

168

LAWSON, SIMONS & ISAAC

& Price, J. H. 1981. Preliminary results from a recent survey of the
marine algal flora of Angola (southwestern Africa). Proc. Int. Seaweed
Symp. (Bangor, 1974) 8: 367-371. Menai Bridge.

Price, J. H., Maggs, C. A. & Lawson, G. W. 1979. Seaweeds of the
western coast of tropical Africa and adjacent islands: a critical assessment.
III. Rhodophyta (Bangiophyceae).Bw//. Br. Mus. not. Hist. (Bot.) 7: 69-82.

Kensley, B. & Penrith, M.-L. 1973. The constitution of the intertidal fauna of
rocky shores of Moc.amedes, southern Angola. Cimbebasia A, 2: 113-123.

1980. The constitution of the fauna of rocky intertidal shores of
South West Africa. Part III. The north coast from False Cape Frio to the
Kunene River. Cimbebasia A, 5: 201-214.

Knox, G. A. 1960. Littoral ecology and biogeography of the southern oceans.

Proc. R. Soc. B. 152: 577-624.
Kiihnemann, O., 1972. Bosquejo fitogeografico de la vegetation marina del

litoral argentine. Physis 31: 117-142.
Lawson, G. W. 1978. The distribution of seaweed floras in the tropical and

subtropical Atlantic Ocean: a quantitative approach. Bot. J. Linn. Soc. 76:

177-193.
- 1988. Seaweed distribution patterns as revealed by ordination with

reference to the Atlantic and southern oceans. Helgolander wiss.

Meeresunters. 42: 187-197.

John, D. M. & Price, J. H. 1975. The marine algal flora of Angola: its
distribution and affinities. Bot. J. Linn. Soc. 70: 307-324.

& Price, J. H. 1969. Seaweeds of the western coast of tropical Africa and
adjacent islands: a critical assessment. I. Chlorophyta and Xanthophyta.
Bot. J. Linn. Soc. 62: 279-346.

Lewis, J. R. 1961. The littoral zone on rocky shores a biological or physical

entity? Oikos 12: 280-301.
Liming, K. 1985. Meeresbotanik. Stuttgart.
McLachlan, A. 1985. The ecology of two sandy beaches near Walvis Bay.

Madoqua 14: 155-163.
Michanek, G. 1971. A preliminary appraisal of world seaweed resources. Fish

Circ. 128. Roma.

1975. Seaweed resources of the ocean. Fish. Tech. Pap. 138. Roma.

1979. Phytogeographical provinces and seaweed distribution. Botanica
mar. 22: 375-391 .

Papenfuss, G. F. 1968. Notes on South African marine algae V. Jl S. Afr. Bot.

34: 267-287.
Penrith, M.-L. & Kensley, B. F. 1970a. The constitution of the intertidal fauna

of rocky shores of South West Africa. Part 1. Liideritzbucht. Cimbebasia A,

1: 191-239.

19706. The constitution of the fauna of rocky intertidal shores of
South West Africa. Part II Rocky Point. Cimbebasia A, 1: 243-268.

Piccone, A. 1884. Crociera del Corsaro alle isole Modern e Canarie del Capitano

Enrico d'Albertis. Alghe. Genova.
Pilger, R. 1908. Kleinere Beitrage zur Kenntnis der Meeresalgen 1. Hedwigia

48: 178-183.
Pollock, D. E. & Beyers, C. J. de B. 1981. Environment, distribution and

growth rates of west coast rock-lobster Jasus lalandii (H. Milne Edwards).

Trans. R. Soc. S. Afr. 44: 379-400.
Post, E. 1963. Zur Verbreitung und Okologie der Bostrychia-Caloglossa

Assoziation. Int. Rev. ges. Hydrobiol. 48: 47-152.
Price, J. H., John. D. M. & Lawson, G. W. 1978. Seaweeds of the western

coast of tropical Africa and adjacent islands: a critical assessment. II.

Phaeophyta. Bull. Br. Mus. not. Hist. (Bot.) 6: 87-182.

1986. Seaweeds of the western coast of tropical Africa and

adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 1.

Genera A-F. Bull. Br. Mus. not. Hist. (Bot.) 15: 1-122.

1988. Seaweeds of the western coast of tropical Africa and

adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 2.

Genera G. Bull. Br. Mus. nat. Hist. (Bot.) 18: 195-273.
Rotmann, K. W. G., 1987. The collection, utilization and potential farming of

red seaweeds in Namibia. Hydrobiologia 151/152: 301-305.
Round, F. E. 1981. The ecology of algae. Cambridge.
Searles, R. B. 1968. Morphological studies of red algae of the order Gigartinales.

Univ. Calif. Pubs. Bot. 43.
Schmidt, O. C. 1923. Beitrage zur Kenntnis der Gattung Codium Stackh.

Bibliotheca Bot. 91: 1-67.

& Gerloff, J. 1957. Die marine vegetation Afrikas in ihren Grundziigen
darstellt. Willdenowia 1: 709-756.

Seagrief, S. C. 1984. A catalogue of South African green, brown and red algae.
Mem. Bot. Surv. S. Afr. 47.

& Troughton, S. C. 1973. A guide to the literature on South African
marine algae. // S. Afr. Bot. 39: 95-109.

Shannon, L. V., Agenbag, J. J. & Buys, M. E. L. 1987. Large and mesoscale
features of the Angola Benguela front. In A. I. L. Payne, J. A. Gulland &
K. H. Brink (Eds) The Benguela and Comparable Ecosystems. S. Afr. J.
mar. Sci. 5: 11-34.

Nelson, G. & Jury, M. R. 1981. Hydrological and meteorological aspects
of upwelling in the south Benguela Current. In F. A. Richards (Ed.) Coastal
and estuarine sciences 1: 146-159.

Silva, P. C. 1959. The genus Codium (Chlorophyta) in South Africa. Jl S. Afr.
Bot. 25: 103-165.

1960. Codium (Chlorophyta) in the tropical western Atlantic. Nova

Hedwigia 1: 497-536.

Simons, R. H. 1964. Species of Plocamium on the South African coast. Bothalia
9: 153-168.

1966. The genus Ceramium in South Africa. Bothalia 9: 153-168.

1968. Notes on Ceramium planum Kiitzing, a red seaweed from South

Africa. // S. Afr. Bot. 34: 127-133.

1970. Marine algae from South Africa 1. Six new species from the inter-

and infra-tidal zones. Investl. Rep. Div. Sea Fish. Rep. S. Afr. 88: 1-13.
1974. Algae (including diatoms and seaweeds) In J. H. Day, N. A. H.

Millard & M. L. Penrith (Eds), A guide to the marine life on South African

shores. 2nd ed.: 239-261. Cape Town.
1983. The genus Trematocarpus (Sarcodiaceae, Rhodophyta) in southern

Africa and the exclusion of Sphaerococcus (Chondrus) scutellatus. Bothalia

14: 803-808.
& Hewitt, F. E. 1977. Marine algae from southern Africa 2. Morphology

and taxonomy of five foliaceous Floridiophyceae (Rhodophyta). Investl.

Rep. Div. Sea Fish. S. Afr. 110: 1-16.
Stephenson, T. A. 1939. The constitution of the intertidal fauna and flora of

South Africa. Part 1. J. Linn. Soc. (Zool.) 40: 478-536.

1944. The constitution of the intertidal fauna and flora of South Africa.
Part II. Ann. Natal Mus. 10: 261-358.

1948. The constitutiobn of the intertidal fauna and flora of South Africa.

Part III. Ann. Natal Mus. 11: 207-324.

& Stephenson, A. 1972. Life between tidemarks on rocky shores. San

Francisco.

& Day, J. H. 1940. The South African intertidal zone and its relation

to ocean currents. VIII Lambert's Bay and the west coast. Ann. Natal Mus.

9: 345-377.
Sverdrup, H. U., Johnson, M. W. & Hemming, R. H. 1942. The oceans. New

York.
Wynne, M. J. 1986. Report on a collection of benthic marine algae from the

Namibian coast (southwestern Africa). Nova Hedwigia 43: 311-355.

Bull. Br. Mus. nat. Hist.(Eot.) 20(2) 169-192 Issued 20 December 1990

The infrageneric classification of Gentiana
(Gentianaceae)

TING-NUNG HO ^nd SHANG-WU LIU

Northwest Plateau Institute of Biology, Academia Sinica, Xining, Qinghai, China

CONTENTS

Generic circumscription and status 170

Key to the genera in subtribe Gentianinae 170

Infrageneric classification 171

Taxonomic characters 172

Systematic treatment 178

Gentiana L 178

Key to the sections 178

Keys to the series 179

Classification 179

I. Sect. Otophora Kusn 179

1 . Ser. Otophorae Marquand 180

2. Ser. Decoratae Marquand 180

II. Sect. Cruciata Gaudin 180

III. Sect. Monopodiae (H. Smith) T.N.Ho 180

1. Ser. Monanthae (H. Smith) T.N.Ho 180

2. Ser. Verticillatae (Marquand) T.N.Ho 180

3. Ser. Ornatae (Marquand) T.N.Ho 180

4. Ser. Apteroideae (H. Smith) T.N.Ho 180

IV. Sect. Frigida Kusn 181

V. Sect. Gentiana 181

VI. Sect. Pneumonanthe (Gled.) Gaudin 181

VII. Sect. Phyllocalyx T.N.Ho 181

VIII. Sect. Calathianae Froelich 181

IX. Sect. Ciminalis (Adans.) Dumort 181

X. Sect. Isomeria Kusn 181

1 . Ser. Sikkimenses (Marquand) T.N.Ho 182

2. Ser. Depressae (H. Smith) T.N.Ho 182

3. Ser. Stragulatae T.N.Ho 182

4. Ser. Uniflorae (Marquand) T.N.Ho 182

XI. Sect. Microsperma T.N.Ho 182

1 . Ser. Suborbisepalae (Marquand) T.N.Ho 182

2. Ser. Tetramerae (Marquand) T.N.Ho 182

3. Ser. Annuae (Marquand) T.N.Ho 182

XII. Sect. Stenogyne Franchet 182

XIII. Sect. Dolichocarpa T.N.Ho 18 2

XIV. Sect. Chondrophyllae Bunge 182

1 . Ser. Fimbriatae Marquand 183

2. Ser. Rubicundae Marquand 18

3. Ser. Linearifoliae Marquand 183

4. Ser. Orbiculatae Marquand 18

5. Ser. Humiles Marquand 18

6. Ser. Capitatae T.N.Ho 18

7. Ser. Fastigiatae T.N.Ho 18

8. Ser. Napuliferae T.N.Ho 18

9. Ser. Conacre T.N.Ho I 83

10. Ser. Piasezkianae T.N.Ho 18

11. Ser. Grandiflorae Grossh. exT.N.Ho l g

XV. Sect. Fimbricorona T.N.Ho t8

Geographical distribution

Study materials

References . . 191

170

T.-N. HO&S.-W. LIU

SYNOPSIS. An infrageneric classification of the genus Gentiana L. is presented. The genus is divided into 15 sections,
of which five are subdivided into series, including 361 species. Two series, Coriaceae T.N. Ho and Grandiflorae
Grossh. ex T.N. Ho, are described as new, and the new combination, ser. Monanthae (H. Smith) T.N. Ho, is made.
Fimbricorona T.N. Ho is proposed as a new name for sect. Eurythalia (Borkh.) Griseb. Keys are given to the genera
in subtribe Gentianinae and the sections and series of Gentiana. The history of the infrageneric classification of
Gentiana, the characters used in infrageneric classification, and the geographical distribution of the genus are
discussed.

GENERIC CIRCUMSCRIPTION AND STATUS

The name Gentiana in its modern sense is generally accredited
to Tournefort (1700). His name was taken up by Linnaeus in
Genera plantarum (1737) and Species plantarum (1753).

Gentiana sensu lato, as traditionally circumscribed, is a
very heterogeneous assemblage of morphologically diverse
groups, including Tripterospermum, Crawfurdia, Megacodon,
and Gentianella sensu lato. Gentianella sensu lato further
consists of Gentianella sensu stricto, Comastoma, Gentianopsis,
and Pterygocalyx. Gentiana is circumscribed in the present
paper in the strict sense, as equivalent to subgenus Eugentiana
Kusn. This interpretation has been accepted widely by most
20th century authors on Gentianaceae, e.g. Smith (1936,
1965), Tutin (1972), Czerepanov (1973), Pringle (1977, 1978,
1979). It is narrower than Gentiana sensu lato mentioned
above, but much broader than Gentiana as defined by Holub
(1973) and Love & Love (1975) who restricted Gentiana to
the five European species treated by Tutin (1972) as sect.
Gentiana.

According to Gilg's comprehensive survey of the family
(1895), Gentiana is included in subfamily Gentianoideae,
tribe Gentianeae, subtribe Gentianinae. In subtribe
Gentianinae, it is more closely related to Crawfurdia,
Tripterospermum, and Megacodon than to Gentianella,
Comastoma, Gentianopsis, Lomatogonium, Lomatogoniopsis,
Swertia, and Halenia. It seems that these genera can be split
into two evolutionary groups the Gentiana group and the
Gentianella group. The two groups can be clearly distinguished
on the basis of the position of the floral glands: the Gentiana
group (Gentiana, Crawfurdia, Tripterospermum, and
Megacodon) has glands at the base of the ovary, while the
Gentianella group (Gentianella, Comastoma, Gentianopsis,
Lomatogonium, Lomatogoniopsis, Swertia, and Halenia} has
glands in foveae or spurs borne on the surface of the corolla
(epipetalous glands). In addition, the corolla bears plicae or
folds between the lobes in the Gentiana group (except in
Gentiana lutea L. and Megacodon), each lobe has 3 primary
vascular bundles, the calyx has a continuous intracalycular
membrane (except in Gentiana sect. Chondrophyllae and
sect. Calathianae which have obsolete membranes), and
calcium oxalate crystals have been reported in the leaf
mesophyll (except in sect. Calathianae}. In the Gentianella
group, there are no plicae or folds between the corolla-lobes
and there is no continuous intracalycular membrane, the
corolla-lobes have 5-9 vascular bundles, and calcium oxalate
crystals have not been found in the leaf mesophyll.
Crawfurdia and Tripterospermum are further characterized
by their climbing habit and by the calyx having 10 vascular
bundles in the former and 5 in the latter, and differ distinctly
from Gentiana in these characters.

Chromosome data have contributed little towards resolving
the problem of generic delimitation in the Gentianinae. From

previously reported chromosome data, we know that the
majority of perennial species of Gentiana are x = 6 or 13, 2n
= 12 or 26, less often x = 5, 7, 9, or 11, 2n = 10, 14, 18, 20,
22, 24, 30, 36, 40, 44, 48 or 52. The majority of annuals are x
= 5, 2n = 10, less often x = 13 or 19, 2n = 26 or 38. However,
in Gentianella, the chromosome number x = 9, 2n = 36 is
the commonest and most widespread one. The base number
is x = 9 or 13 in Swertia. Halenia has a base number of 11 , 2n
= 22. The diploid Comastoma has a base number of 5 in
common with Lomatogonium. Gentianopsis has two series, x
11 or 13, 2n = 44 or 78. However, in these genera,
sometimes more than one series of numbers may exist in the
same genus, even in the same section or within a group with
very similar morphological features. For example, Gentiana
frigida Haenke and G. froelichii Jan ex Reichenb., both
endemic to Europe and belonging to sect. Frigida, have
different chromosome base numbers, x 6 and x 7
respectively. Only about half the genera of Gentianaceae
have been investigated cytologically, and only selected
species rather than whole genera. Further study is clearly
necessary.

Most pollen grains in Gentianinae are 3-colporate,
less frequently 3-colpate (-colporoidate), or occasionally 4-
colporate (-colporoidate). The size of the pollen grains
(length of polar axis) ranges from 15 um to 65 urn. The shape
varies from peroblate to perprolate. The sexine patterns can
be classified into striate, striato-reticulate, reticulate, spin-
uliferous and smooth, and the diversity seems to follow the
generic or sectional classification. Most species of Gentiana
have striate sexine, rarely striato-reticulate (sect. Chondro-
phyllae and sect. Calathianae). The ora usually have
lateral extensions. Gentianopsis has typically reticulate,
heterobrochate pollen grains. The ora are distinctly delimited
or are visible only in lateral view. Swertia has heterogeneous
sexine features, i.e. striate, striato-reticulate, reticulate,
spinuliferous or smooth. Halenia has pollen grains which
are finely reticulate. Lomatogonium is characterized by the
presence of 'islands', and by the sexine distinctly overlapping
the ora. Its sexine patterns are striate to striato-reticulate or the
tectum is provided with spinules or is punctate. Comastoma
has pollen grains resembling those of Lomatogonium p. p.
and Halenia. The ora are lolongate to lalongate and relatively
large with lateral extensions. The sexine patterns are reticulate
with the bacula regularly arranged or with the bacula arranged
in no particular order, sometimes interconnected by thin con-
nections. It seems likely that further study of the group, using
the scanning electron microscope, will provide more taxon-
omically valuable pollen characters. Nilsson's (1967, 1976)
work is an example of the sort of studies that are needed.

Key to the genera in subtribe Gentianinae

la. Glands in a whorl attached to the base of the ovary:
2a. Corolla with plicae between the lobes (except in Gentiana

CLASSIFICATION OF GENTIAN A

171

luted), divided to less than halfway, the tube much longer than
the lobes, rarely divided to more than halfway and the tube
shorter than, or equalling, the lobes:
3a. Plants not climbing, with quadrangular, erect or ascending

stems Gentiana

3b. Plants climbing, with terete, twisted stems:
4a. Vascular bundles in calyx-tube 5; glands conspicuously
developed, forming a collar-like disc covering the base
of the ovary; stamens asymmetrical, unequal in length,
unilaterally and uniformly curved downwards at the apex,
the filaments linear or filiform; fruit a capsule or berry . . .

Tripterospermum

4b. Vascular bundles in calyx-tube 10; glands small, not
discoid; stamens symmetrical, equal in length, straight,

the filaments thickened below; fruit a capsule

Crawfurdia

2b. Corolla without plicae between the lobes, divided to more
than halfway, the tube shorter than, or equalling, the lobes:
5a. Ovary sub-bilocular, with a lamellate intrusion of the

placenta into the locular cavity Ixanthus

5b. Ovary unilocular, without a lamellate intrusion of the

placenta into the locular cavity:
6a. Stamens inserted on the sinus of the corolla-lobes;

capsule twisted above; flowers small Latouchea

6b. Stamens inserted on the upper part of the corolla-
tube; capsule straight; flowers large Megacodon

Ib. Glands in foveae, lobes or spurs attached to corolla-tube and
alternating with the stamens:
la. Corolla with spurs at the base of the corolla-tube, rarely

spurless; glands prolonged into the spurs Halenia

7b. Corolla without spurs; foveae on corolla-tube or on lobes:
8a. Corolla usually tubular, the tube longer than the lobes,

less often the tube shorter than, or as long as, the lobes:
9a. Buds large, slightly flattened, 4-angled; pairs of calyx-
lobes dissimilar, one longer and narrower than the
other; corolla frequently fringed or toothed; seeds

angular-papillose Gentianopsis

9b. Buds small, not flattened; calyx-lobes usually all

similar; corolla not fringed; seeds almost smooth:
lOa. Corolla bearing two evascular fimbriate scales at

the base of each lobe Comastoma

lOb. Corolla naked at the throat or bearing vascular

fimbriae, the fimbriae united into a single scale

extending across the base of each lobe (Gentianella):

lla. Stems twining; calyx conspicuously 4-winged;

seeds discoid-winged Pterygocalyx

lib. Stems erect; calyx unwinged; seeds unwinged:

12a. Corolla imbricate in bud; stamens inserted at,

or below, the middle of the corolla-tube; seeds

generally numerous per capsule .... Gentianella

12b. Corolla valvate in bud; stamens inserted on the

sinus between the lobes; seeds few per capsule

Jaeschkea

8b. Corolla rotate, the tube much shorter than the lobes:
13a. Dioecious, with unisexual flowers; stamens inserted on

the sinus between the corolla-lobes Veratrilla

13b. Flowers hermaphrodite; stamens inserted on the corolla-
tube:

14a. Style present, the stigmas never decurrent; corolla-
lobes concolorous Swertia

14b. Style absent, the stigmas decurrent along the ventral

suture of the ovary; corolla-lobes conspicuously bi-

colorous, half pale and half dark:

15a. Corolla- lobes bearing conspicuous foveae which

are sometimes connate below into a tube, but

lobed or lamellate above Lomatogonium

15b. Corolla-lobes efoveolate, bearing appendages
which are lamellate or galeate, and entire or erose
Lomatogoniopsis

INFRAGENERIC CLASSIFICATION

Some 40 botanists have studied Gentiana intensively since
Linnaeus formally established the genus. They not only
discovered large numbers of new species, but they have also
treated the genus in a wide variety of ways. The taxonomic
history has been well documented by Kusnezow (1894) and
Gillet (1957), and will therefore not be repeated here in
detail. The discussion that follows will just review some of the
more important works.

In 1737, when Linnaeus took up Tournefort's Gentiana he
divided the genus into seven groups. In 1753, he redivided it
less successfully into three unnamed sections, describing 23
species, of which the first, G. lutea L., has been taken as the
type species of Gentiana by later authors.

Froelich (1796) was the first to monograph Gentiana, and
split the genus into four groups of uncertain rank: Coelanthe,
Calathianae, Endotrichae, and Crossopetalae, with 47 species
altogether.

Grisebach's monographic study of Gentianaceae is one of
the landmarks of work on the family. In Genera et species
Gentianearum (1838) he divided the genus into seven sections
with 125 species. In his treatment of Gentianaceae in De
Candolle's Prodromus (1845) he redivided it into 15 sections, of
which seven (Asterias, Coelanthe, Pneumonanthe, Tretorrhiza,
Chondrophyllae, Thylacites, Cyclostigma, and Eurythalia}
belong to Gentiana sensu stricto, and increased the number of
species to 153.

Kusnezow's monograph of Gentiana subgenus Eugentiana
(1894) and his treatment of Gentiana in Engler & Prantl's Die
naturlichen Pflanzenfamilien (1895) provided the generally
accepted basis for the division of Gentiana into sections. His
subgenus Eugentiana corresponds to Gentiana sensu stricto as
a genus, and was divided into 10 sections (Coelanthe,
Pneumonanthe, Stenogyne, Frigida, Aptera, Chondrophyllae,
Otophora, Isomeria, Thylacites, and Cyclostigma}, with 162
species.

Marquand published several papers from 1928 to 1937,
notably his revision of the Chinese species of Gentiana
(1937a). He usefully subdivided some sections into series,
e.g. Otophora into ser. Otophorae and Decoratae; Frigida
into ser. Verticillatae, Omatae, Suborbisepalae, Confertifoliae,
Uniflorae, Sikkimenses, Multiflorae, and Annuae; and
Chondrophyllae into ser. Pubigerae, Fimbriatae, Orbiculatae,
Linearifoliae, Rubicundae, and Humiles; and included 163
species from China (excepting sect. Dipterospermum and
sect. Tripterospermum), of which 34 were new. Grisebach's
and Kusnezow's revisions contained only 71 and 162 species
respectively of Gentiana sensu stricto for the whole world.
However, Marquand combined the genera Crawfurdia and
Tripterospermum with Gentiana as two additional sections
and relegated Pterygocalyx to Gentianella, which had been
excluded from Gentiana by him in an earlier paper (1931).

In 1936, Smith raised Hemsley's sect. Megacodon to
generic rank and almost thirty years later (in 1965) he
reinstated Crawfurdia and Tripterospermum as two indepen-
dent genera. In 1961, when Smith studied the Gentiana

172

cachemirica group he subdivided sect. Frigida into two sub-
sections: Monopodiae and Sympodiae, based on differences
of branching. This is a very important character in perennial
Gentiana, which attracted the attention of Serebryakova
(1979), who demonstrated the basic models of shoot forma-
tion and indicated various evolutionary trends in the genus.
This character is further discussed in the present paper.

Ma (1951) segregated sect. Crossopetalum from Gentiana
as a separate genus, Gentianopsis.

In a series of papers published between 1967 and 1979,
Pringle not only monographed sect. Pneumonanthe and sect.
Chondrophyllae of eastern North America, central America
and Mexico, but also made a careful and clear revision of the
sectional and subsectional names.

In the present paper, Gentiana is split into 15 sections. Five
of these are further divided into a number of series. This
classification essentially agrees with that of Kusnezow's
monograph, but includes several significant changes. It is
based on Ho (1985), in which three new sections (Phyllocalyx,
Microsperma, and Dolichocarpd) and five new series (Stragu-
latae, Capitatae, Fastigiatae, Napuliferae, and Piasezkianae)
were proposed, and subsect. Monopodiae was raised to
sectional rank. These treatments are supported chiefly by
different branching patterns of the stem, various structures of
the testa, the presence or absence of a wing on the capsule,
features of the calyx, etc. Here we reinstate Grisebach's sect.
Eurythalia, which has been forgotten for about a century and
a half, as a section, but under a new name, Fimbricorona.
This section is very unusual in Gentiana and its fimbriate
corona borne at the throat of the corolla reminds one of the
genus Comastoma. In addition we describe two new series
(ser. Coriaceae T.N. Ho and ser. Grandiflorae Grossh. ex
T.N. Ho) and make one new combination, ser. Monanthae
(H. Smith) T.N. Ho, in sect. Monopodiae T.N. Ho. This is a
revision of the arrangement in Ho et al. (1988).

TAXONOMIC CHARACTERS

This section deals with those characters used in the infra-
generic classification that need explanation.

1. GROWTH HABIT. All species of Gentiana are herbs, varying
from dwarf plants, only one or a few centimetres high, to tall,
robust plants up to 120 cm high (G. luted). About half of the
species are annuals belonging to five sections; the other half
are perennials and belong to 10 sections.

2. ROOTS. The roots of Gentiana are of four types related to
the growth habit and the branching of the stems: fibrous
roots, fleshy stout tap-roots, fleshy linear-cylindrical roots,
and rhizomes or stolons with adventitious roots. The first type
is the commonest, the roots being fibrous or even woody. The
primary root is rather small but slightly thickened, forming a
slender tap-root with few secondary rootlets. This type
is found in the annual sections, i.e. Chondrophyllae,
Dolichocarpa, Microsperma, Stenogyne, and Fimbricorona.
The second type is confined to sect. Otophora, in which the
plant has a fleshy, stout cylindrical or fusiform, persistent tap-
root and a few smaller secondary rootlets. The third type, in
which the plant has several to many fleshy, linear-cylindrical
roots arising from the collar, is confined to sect. Monopodiae
and sect. Cruciata. The last type, as the second and third, is

T.-N. HO&S.-W. LIU

always associated with branching of the stem, the plants
having rhizomes or stolons with adventitious roots from the
nodes. This type is found in the majority of perennial
sections, i.e. Pneumonanthe, Gentiana, Phyllocalyx, Isomeria,
Frigida, Ciminalis, and Calathianae.

3. STEM AND BRANCHING. The stem is always erect or ascending,
terete, with striae or angles, sometimes having a conspicuous
caudex at the base. This is most conspicuous in sect. Cruciata,
where the plant has a stout caudex sheathed by fibrous old
petioles enabling this section to be recognized at a glance.
Sect. Monopodiae ser. Monanthae also has a stout caudex but
differs from sect. Cruciata in the caudex being sheathed by
membranous old petioles. In all the other groups the caudex
is indistinct. In the perennial groups, the branching of the
stem is a very constant and significant feature, and provides a
reliable diagnostic character for the definition and grouping
of sections and series. There are two main types as follows
(Fig. 1):

A. Monopodial branching. The plant consists of a basal
rosette with a terminal bud (growing point of the plant axis)
sunk in it. This terminal bud has great vitality and can grow
continuously, lasting throughout the life of the plant. The
flowering stems are produced each year from the axillary buds
of the basal leaves of the rosette. This monopodial branching
type falls into two subtypes:

a. Perennial flowering stems. The basal rosette is poorly
defined, with a few small, linear-triangular leaves. The
flowering stems grow continually and can last for several
years. This subtype is confined to sect. Otophora ser.
Decoratae.

b. Annual flowering stems. The basal rosette is well-
defined, with abundant large leaves. The flowering stems only
last for one year. This subtype is confined to sect. Monopodiae,
sect. Cruciata, and sect. Otophora ser. Otophorae.

B. Sympodial branching. In contrast to monopodial branch-
ing, in which the growing point is apical and everlasting, the
growing point in sympodial branching is lateral and constantly
renewed. The plant has rhizomes or stolons, which have
nodes, internodes, buds and adventitious roots, but no basal
rosette. After a time, growth of the terminal bud slows down
and stops, at which the buds on the nodes grow out instead.
The flowering shoots are produced annually from buds on the
nodes of the rhizomes or stolons, as well as from the apex of
the stolons. Sympodial branching is undoubtedly, as has often
been explained, more advanced evolutionarily than mono-
podial branching. Because the terminal bud dies and the
lateral buds renew the growth, the plant can produce progres-
sively more and more lateral buds to prodice a more
luxuriant form. The sympodial branching of Gentiana exhibits
two subtypes as follows:

a. Rhizome subtype. The rhizome is usually stout, fleshy
and horizontal or ascending, or more rarely vertical under the
ground. Adventitious roots and a lateral rosette arise first
from a node and then buds in the axils of the outer leaves of
the lateral rosette give rise to the flowering stems. This
subtype is found in sect. Frigida, sect. Gentiana, and some
species of sect. Pneumonanthe. However, most other species
of sect. Pneumonanthe are somewhat different from this
model in that the plants have no rosette and the buds of the
nodes give rise to the flowering stems directly.

b. Stolon subtype. This subtype is rather similar to the

Fig. 1 Types of stem branching (diagrammatic). 1-2 monopodial branching, 1. perennial flowering stem: G. decorate; 2. annual flowering
stem: G. veitchiorum. 3-5 sympodial branching: 3-4 rhizome subtype, 3. G. scabra; 4. G. algida; 5. stolon subtype: G. depressa var. stenophylla.

174

rhizome subtype but differs in having overground stolons and
runners. The stolons produce runners from the nodes and
these runners are ultimately terminated by a lateral rosette
with a few small leaves. The central bud of this rosette in time
produces the flowering stems. This subtype is the commoner,
being found in sect. Isomeria, sect. Phyllocalyx, sect.
Ciminalis, and sect. Calathianae.

4. LEAVES. The leaves are usually opposite, but in sect.
Monopodiae ser. Verticillatae they are in whorls of 3-8,
surrounding the 5-8-merous flowers. The leaves are generally
very variable in shape and size so that their taxonomic use is
limited, but in sect. Chondrophyllae leaf shape may be used
for the definition of series and species. Ser. Linear if oliae, for
example, is notable for its linear leaves.

5. INFLORESCENCE AND FLOWERS. The inflorescence is always
a cyme. The cyme may be simple, with 1-3 flowers, and this
may be reduced to a single terminal flower, sometimes also
with a few flowers in the axils of the uppermost leaves. This is
the commonest condition, occurring in almost all the annual
sections (except ser. Capitatae and some species of sect.
Chondrophyllae) and in some of the perennial groups. The
inflorescence may also be a complex cyme with few to many
flowers. The flowers are crowded and either pedicellate in
terminal and axillary cymes or sessile in terminal clusters, and
sometimes also in few-flowered axillary whorls. This condi-
tion is found in sect. Gentiana, sect. Pneumonanthe, sect.
Cruciata, etc.

6. CALYX. The calyx-tube is usually entirely tubular, but in
sect. Gentiana and some species of sect. Cruciata and sect.
Frigida it is split down one side to the middle or to near the
base, forming a spathe with small irregular teeth (Fig. 2). The
calyx-lobes are mostly triangular, ovate or lanceolate, but in
some groups other shapes occur: spathulate or suborbicular
and much narrowed at the base in sect. Isomeria ser. Stragulatae
and sect. Microsperma ser. Suborbisepalae; ovate and con-
tracted at the base in sect. Chondrophyllae ser. Orbiculatae;
filiform in sect. Chondrophyllae ser. Rubicundae; and subulate
with wide bases and acute apices in sect. Chondrophyllae ser.
Fimbriatae. The midribs are distinct, but the degree of
development varies considerably. The most prominent mid-
ribs are those forming keeled ribs on the outer face which are
decurrent into the angles or wings of the calyx-tube. This
condition is found in sect. Stenogyne, sect. Dolichocarpa,
sect. Calathianae, and sect. Microsperma ser. Tetramerae.

1. COROLLA. The corolla is important diagnostically and
provides a number of characters for the definition of species.
It varies in size from 3 mm to 70 mm, although the range is
usually relatively restricted and constant within each group.
Large corollas are 4-7 cm long, medium-sized ones 2.5-4 cm,
and small ones 0.3-2.5 cm. In general, all perennial sections
and the annual sect. Stenogyne have large or medium-sized
corollas, whereas the others (all the annual sections except
Stenogyne) have small corollas. The corolla is usually 5-lobed,
but 6-8-lobed corollas are found in sect. Monopodiae ser.
Verticillatae and in various species of the other sections. In
general, the corolla-tube is longer than the lobes, but the
reverse condition occurs in sect. Gentiana and sect.
Otophora, in which the corolla is divided to the middle or
near the base so that the corolla-tube is as long as, or much
shorter than, the lobes. The shape of the corolla varies
considerably and the terms used to describe it in this paper
are as follows (Fig. 2):

T.-N. HO &S. -W.LIU

Tubular. The corolla is tubular throughout, with slightly
divergent lobes.

Obconical. The corolla-tube is narrowed at the base but
gradually widened upwards with patent lobes.
Hypocrateriform. The corolla has a long, narrow, parallel-
sided tube and broad flat lobes spreading at right angles to the
tube.

Infundibuliform. The corolla-tube is narrowed in the lower
part but is gradually widened upwards, with erect-patent
lobes.

Campanulate. The corolla has a short, broad, parallel-sided
tube and patent lobes.

Urniform. The corolla-tube is very inflated in the middle but
contracted below and in the throat, with patent lobes.

These corolla shapes are not entirely clear-cut, and transitional
forms, which need not be discussed here, do occur.

The plicae of the corolla apparently exist in all species of
Gentiana except G. lutea, in which their absence is a particu-
larly diagnostic character. There are various degrees of
development of the plicae in the different groups (Fig. 2). In
sect. Gentiana and sect. Otophora the poorly developed
plicae are very small, asymmetrical and auriculate, whereas in
sect. Chondrophyllae and sect. Dolichocarpa the well-
developed plicae are large and symmetrical. These two types
represent the two extremes, the plicae of the other sections
being more or less intermediate. The margins of the plicae
vary from entire, erose, serrate, lacinulate or fimbriate to
setose-fimbriate. Sect. Chondrophyllae ser. Fimbriatae is
notable in that the corolla has fimbriate plicae.

8. STAMENS. The stamens are equal in number to the corolla-
lobes and alternate with them. They are generally 5 in
number, but in some species of sect. Monopodiae ser.
Verticillatae (where the flowers are 6-8-merous and the leaves
are in whorls of 6-8) they are 6-8. The great majority of
species have uniform stamens with more or less winged
filaments and free anthers, but in some species of sect.
Pneumonanthe and sect. Gentiana the anthers are contiguous.

9. PISTIL. The ovary, which is markedly uniform in general
structure, varies mainly in size. It is usually oblong or
elliptical and narrowed into the style, or the style may be
absent. The style is shortly cylindrical to narrowly linear,
much shorter than the ovary in most species, but in sect.
Stenogyne and a few other species (e.g. G. filistyla Balf. f. &
Forrest ex Marquand, G. tubiflora (G. Don) Wallich ex
Griseb., and G. vernayi Marquand) the style and stigmas are
very long and filiform, equalling, or only slightly shorter than,
the elongate ovary. The stigma-lobes in most sections are
uniformly oblong or linear and are free and recurved. How-
ever, in a few sections they depart from this norm and the
stigma-lobes are expanded and rounded, and either fimbriate
on the surface and contiguous into a small disc or infundibuli-
form structure but free after anthesis, or almost smooth and
either connate into a small disc or infundibuliform structure
and not free after anthesis, or contiguous into a small disc or
infundibuliform structure and free after anthesis. The first
condition is found in sect. Ciminalis, the second in sect.
Calathianae, and the third in sect. Phyllocalyx. These ex-
panded stigmas are diagnostic in Gentiana and form the most
important character separating the three sections. The gyno-
phore may elongate and become more apparent with age. In
some species the pistil is sessile or subsessile, but in others the
gynophore may be short in the young flower and elongate

Fig. 2 Calyx, corolla and plica shapes. 1-6 calyx types, 1. calyx-tube split down one side to the lower part to form a spathe with small teeth: G.
straminea; 2-6 calyx-lobe types, 2. filiform: G. rubicunda; 3. linear: G. dolichocalyx; 4. suborbicular: G. tongolensis; 5. ovate: G. squarrosa; 6.
triangular: G. pseudoaquatica. 7-12 corolla types, 7. tubular: G. prolata; 8. hypocrateriform: G. brachyphylla; 9. infundibuliform: G. farreri;
10. campanulate: G. alpina; 11. urniform: G. urnula; 12. rotate: G. lutea. 13-18 plica types, 13. plica absent: G. lutea; 14. truncate: G.
hexaphylla; 15. oblique: G. striata; 16. auriculate: G. doxiongshanensis; 17. symmetrical: G. pseudoaquatica; 18. fimbriate: G. panthaica.

176

T.-N. HO&S.-W. LIU

Fig. 3 SEM micrographs of seed types. 1-4 seeds with reticulately thickened testa, winged: 1-2 seeds surrounded by a discoid wing, 1 . G. lutea;
2. G. asclepiadea; 3. seed with wing on one side: G. pudica; 4. seed with wing at one end: G. bavarica. 5. seed with minutely reticulate testa,
triquetrous with the three edges winged: G. striata. 6. seed covered with membranous lamellae which form spongy complex pits: G. algida. (nos

1, 2, 5, 6, scale = 250 urn; nos 3, 4, scale = 50 urn)

considerably as the capsule matures. It seems that the gyno-
phore is of less taxonomic significance than the style or
stigma.

10. CAPSULE. The capsule is usually long and narrow, oblong
or cylindrical, and unwinged, but in sect. Chondrophyllae and
sect. Fimbricorona it is short and wide, obovoid or ellipsoid-

oblong, strongly compressed, and rounded at the apex with a
conspicuous broad wing which is narrowed towards the base.

11. SEEDS. There are six different seed types in the genus
(Figs 3-4):

a. The seed has a reticulately thickened testa, with either a
discoid wing or with a wing on one side or at one end of the

CLASSIFICATION OF GENTIANA

seed. This type is found in sect. Gentiana and sect.
Pneumonanthe, although some species of sect. Stenogyne
scarcely differ, their seeds having a minutely reticulate testa
and being triquetrous with the three edges winged.

b. The seed varies from almost smooth to distinctly reticulate
and is unwinged. This type is the commonest, being found
mainly in sect. Chondrophyllae, sect. Dolichocarpa, sect.
Cruciata, sect. Otophora, and sect. Calathianae.

c. The seed is covered with membranous lamellae which

177

form spongy, complex hexagonal pits. This type is confined to
sect. Frigida and some species of sect. Isomeria.

d. The seed is covered with membranous lamellae which
form honeycomb-like, simple, shallow, hexagonal pits. This
type occurs in sect. Monopodiae, sect. Microsperma, and
some species of sect. Isomeria.

e. Like d but the body of the seed is surrounded by a broad
wing. This type is found only in sect. Phyllocalyx.

f. The seed has a minutely and densely reticulate testa and

Fig. 4 SEM micrographs of seed types. 1-2 seeds covered with membranous lamellae which form honeycomb-like simple pits, 1 . G. farreri; 2.
G. 'tubiflora. 3-4 seeds minutely reticulate, 3. G. leucomelaena; 4. G. straminea. 5-6 seeds with minutely and densely reticulate testa, rugose, 5.

G. alpina; 6. G. clusii. (nos 1, 2, 4, 5, 6, scale = 250 urn; no. 3, scale = 50 urn)

178

is rugose and unwinged. This type is confined to sect.
Ciminalis.

These seed characters are certainly diagnostically important.
However, the different types do sometimes occur in the same
section, for instance, a, b, and d all occur in sect. Stenogyne, a
very natural group in general appearance; c and d both occur
in sect. Isomeria, a stoloniferous group. Seed characters must
therefore not be relied on too heavily, as Kusnezow and
Marquand did when delimiting sections. Seed structure is,
however, constant within species and may therefore be used
for defining species, as well as for the partial delimitation of
some sections (e.g. Cruciata, Chondrophyllae, Microsperma,
etc.).

SYSTEMATIC TREATMENT

Gentiana (Tourn., Inst. rei herb.: 80, t. 40 (1700) )L., Gen.
pi. : 69 (1737); Sp. pi. : 227 (1753). Type species: G. lutea L.

Subgen. Eugentiana Kusn. in Trudy imp. S.-Peterb. Obshch.
Estest. 24(2): 3 (1894).

Annual or perennial herbs. Roots fibrous with a primary root
and secondary rootlets, or slightly fleshy and forming a stout
tap-root with secondary rootlets, or several linear-cylindrical
roots from the collar. Stem erect or ascending, striate and
angled, simple or branched, in perennial species sometimes
both flowering stem and vegetative stems present; rhizomes
or stolons present or absent. Leaves opposite, rarely verticil-
late, sometimes forming a rosette at the base of the stem.
Cyme simple with 1-3 flowers, or complex with few many
flowers, the flowers pedicellate in terminal and axillary cymes
or sessile in terminal clusters or sometimes also in few-
flowered axillary whorls; flowers usually 5-merous, rarely 4-
or 6-8-merous; calyx-tube with a continuous or undeveloped
intracalycular membrane, the lobes triangular, ovate or filiform,
with prominent midribs; corolla tubular, infundibuliform,
obconical, hypocrateriform, or urniform, very rarely rotate,
usually lobed, the tube much shorter than the lobes, or rarely
the tube as long as or longer than the lobes, plicate between
the lobes (except in G. lutea L.); stamens alternate with the
corolla-lobes, the filaments more or less winged, the anthers
basifixed, free or rarely contiguous; ovary oblong or elliptic;
style often short, cylindrical or linear, less often long
and filiform, longer or slightly shorter than the ovary, the
stigma-lobes usually oblong to linear, free, recurved, rarely
expanded and rounded, connate or contiguous into a small
disc or infundibuliform structure; glands 5-10, at the base of
the ovary, the pistil sessile or with an elongate gynophore.
Capsule oblong or cylindrical and unwinged, or obovoid and
winged; seeds numerous, small, minutely reticulate, rugose,
simply areolate or with complex spongy areolation, winged or
unwinged.

DISTRIBUTION. Europe, Asia, America, NW. Africa (Morocco),
E. Australia.

Fifteen sections, 361 species.
Key to the sections

la. Perennials, with thick, slightly fleshy roots, rarely annuals with
fibrous roots (G. nivalis and G. utriculosa); flowers large (4-7 cm
long) or medium-sized (2.5^4 cm long); capsule unwinged:

T.-N. HO&S.-W. LIU

2a. Branching monopodial, the plant with a basal rosette with a

terminal bud sunk in it, the flowering stems arising annually

from the axillary buds of the rosette leaves:

3a. Corolla usually divided to the middle or below, the tube

shorter than, or equalling, the lobes; plicae small, auricu-

late; plant with a stout, cylindrical or fusiform tap-root

I. Sect. Otophora

3b. Corolla always divided to less than halfway, the tube much
longer than the lobes; plicae large, triangular to truncate;
plant with thickened, linear-cylindrical roots from the collar:
4a. Caudex with a fibrous sheath of brown remains of petioles;
roots usually contiguous and twisted into a stout, cylindri-
cal mass; seeds minutely but distinctly reticulate

II. Sect. Cruciata

4b. Caudex without a fibrous sheath of remains of petioles;
roots laxly divergent; seeds covered with honeycomb-like,
hexagonal, simple, shallow pits .... III. Sect. Monopodiae

2b. Branching sympodial, the plant with rhizomes or stolons with
nodes, internodes, membranous scales, buds, and adventitious
roots, but without a basal rosette with a terminal bud, the
flowering stems arising annually from the buds at the nodes or
apices of the rhizomes or stolons:

5a. Plants with a stout, slightly fleshy, horizontal or ascending,
rarely vertical, underground rhizome, or the rhizome some-
times abbreviated but conspicuous:
6a. Seeds covered with membranous lamellae which form

spongy, hexagonal complex pits, unwinged

IV. Sect. Frigida

6b. Seeds without membranous lamellae, but with a thick-
ened, reticulate testa, winged with either a discoid wing or
with a wing on one side or at one end of the seed, or rarely
unwinged:

7a. Plants with lateral rosettes which arise from the nodes
of the rhizome; flowering stems arising from the axils of
the lateral rosette-leaves, without scale-like leaves at
the base; corolla lobed to below the middle or to near
the base; 5-9-lobed, the plicae very small, auriculate

(absent in G. lutea) V. Sect. Gentiana

7b. Plants without lateral rosettes; flowering stems arising
directly from the nodes of the rhizome, usually with
scale-like leaves below; corolla lobed to less than half-
way, 5-lobed, the plicae large, obliquely truncate or
triangular VI. Sect. Pneumonanthe

5b. Plants with stolons and runners creeping on the surface:
8a. Stigma-lobes not expanded, linear or oblong, free

and recurved X. Sect. Isomeria

8b. Stigma-lobes expanded, connate or contiguous into a

small disc or infundibuliform structure:
9a. Calyx very small, completely enveloped by the upper-
most pair of broad, obovate, obtuse leaves; seeds
broadly winged, covered with honeycomb-like, hex-
agonal, simple, shallow pits VII. Sect. Phyllocalyx

9b. Calyx large, not enveloped by the uppermost pair of

leaves; seeds unwinged:

lOa. Corolla hypocrateriform; calyx-tube prominently
angled or winged; stigma-lobes almost smooth, con-
nate into a disc or infundibuliform structure, not

free after anthesis; seeds not rugose

VIII. Sect. Calathianae

lOb. Corolla obconical; calyx-tube unwinged; stigma-
lobes fimbriate on the surface, contiguous into a disc
or infundibuliform structure, but free after anthesis;
seeds rugose IX. Sect. Ciminalis

Ib. Annuals, with fibrous roots; flowers small (0.3-2.5 cm long),
or rarely large or medium-sized; rarely perennials with small
flowers; capsule winged or unwinged:

CLASSIFICATION OF GENTIAN A

179

lla. Capsule obovoid or ellispoid-oblong, rounded at the apex,

conspicuously winged, the wing strong and broad at the apex

but gradually narrowed towards the base; basal leaves longer

than the cauline; cauline leaves all more or less equal in size:

12a. Corolla naked in the throat . . . XIV. Sect. Chondrophyllae

12b. Corolla with a corona of multicellular hairs in the throat . . .

XV. Sect. Fimbricorona

lib. Capsule cylindrical to narrowly oblong, obtuse or attenuate

at the apex, unwinged; basal leaves small; cauline leaves

usually decreasing in size towards the base of the stem:

13a. Seeds covered with honeycomb-like, hexagonal, simple,

shallow pits; calyx usually not keeled or winged (if strongly

winged, then ser. Tetramerae flowers 4-merous)

XI. Sect. Microsperma

13b. Seeds with minute or thickened reticulation; calyx keeled

or winged:

14a. Flowers large or medium-sized; plicae obliquely
asymmetrical; style filiform, longer or slightly shorter
than the ovary; capsule enclosed in the corolla; seeds

usually winged XII. Sect. Stenogyne

14b. Flowers small; plicae symmetrical; style much shorter
than the ovary; capsule exserted from the corolla; seeds
usually unwinged XIII. Sect. Dolichocarpa

Keys to the series

Sect. Otophora Kusn.

la. Basal rosette well-developed, with abundant large leaves; flower-
ing stems annual; cauline leaves few, distant . . 1 . Ser. Otophorae

Ib. Basal rosette poorly developed, with few small leaves; flowering

stems perennial; cauline leaves numerous, crowded

2. Ser. Decoratae

Sect. Monopodiae (H. Smith) T.N. Ho

la. Vegetative axis sometimes prolonged and branched, with several
basal rosettes decumbent on the ground; flowers several in a

terminal inflorescence 4. Ser. Apteroideae

Ib. Vegetative axis not prolonged, with one basal rosette decumbent

on the ground; flowers sometimes solitary and terminal:
2a. Caudex with a brown membranous sheath of old petioles;
leaves broad; leaves and calyx-lobes with conspicuous carti-
laginous margins 1 . Ser. Monanthae

2b. Caudex without a sheath of old petioles; leaves narrow; leaves
and calyx-lobes without cartilaginous margins:

3a. Leaves in whorls of 3-8; flowers 5-8-merous

2. Ser. Verticillatae

3b. Leaves opposite; flowers 5-merous 3. Ser. Ornatae

Sect. Isomeria Kusn.

la. Flowers several together in an inflorescence; petioles of upper

cauline leaves of flowering stems conspicuously broadened

1 . Ser. Sikkimenses

Ib. Flowers typically solitary and terminal; petioles not broadened:
2a. Leaves and calyx-lobes with conspicuous, wide, membranous

or cartilaginous margins 2. Ser. Depressae

2b. Leaves and calyx-lobes without membranous or cartilaginous

margins:
3a. Calyx-lobes orbicular-spathulate, narrowed at the base into

a tongue 3. Ser. Stragulatae

3b. Calyx-lobes narrowly elliptical or triangular, not narrowed
at the base 4. Ser. Uniflorae

Sect. Microsperma T.N. Ho

la. Leaves and calyx-lobes spathulate; flowers small; gynoecium and

stamens usually exserted from the corolla

1 . Ser. Suborbisepalae

Ib. Leaves and calyx-lobes not spathulate; flowers medium-sized;
gynoecium and stamens included in the corolla:

2a. Flowers 4-merous; calyx-lobes with strongly keeled midribs

which are decurrent below into conspicuous broad wings

2. Ser. Tetramerae

2b. Flowers 5-merous; calyx unwinged 3. Ser. Annuae

Sect. Chondrophyllae Bunge

la. Capsule narrowly elliptical, attenuate or obtuse at the apex:
2a. Annuals; flowers 25-36 mm long; calyx-lobes with strongly
keeled midribs which are decurrent below into conspicuous

wings; corolla hypocrateriform 10. Ser. Piasezkianae

2b. Perennials; flowers 40-55 mm long; calyx-lobes without keeled
midribs; corolla infundibuliform 11. Ser. Grandiflorae

Ib. Capsule obovoid, spathulate or oblong, rounded or trun-
cate at the apex; calyx-lobes not keeled; corolla tubular or
infundibuliform:
3a. Annuals:

4a. Calyx-lobes usually very narrow, filiform or filiform-
subulate:
5a. Plicae usually fimbriate, rarely dentate; leaves and calyx

densely and minutely papillose 1 . Ser. Fimbriatae

5b. Plicae entire or denticulate; leaves and calyx glabrous,

smooth 2. Ser. Rubicundae

4b. Calyx-lobes broad, triangular or lanceolate:
6a. Stems ascending or decumbent, usually branched from
the base, without a conspicuous main stem, rarely
unbranched:
7a. Cauline leaves mostly long and very narrow, more or

less linear or subulate 3. Ser. Linearifoliae

7b. Cauline leaves short and broad, not linear or subulate:

8a. Calyx-lobes recurved or patent, rarely erect, ovate or

orbicular, rarely lanceolate, contracted at the base . .

4. Ser. Orbiculatae

8b. Calyx-lobes erect, triangular or lanceolate, not

contracted at the base 5. Ser. Humiles

6b. Stems erect, with a conspicuous main stem, branched or

rarely simple:

9a. Stems simple, densely papillose, winged or angled,
leafless in the lower half; much branched in the upper
half, with abbreviated internodes, numerous leaves and
flowering branches, all congested at the apex of the
stem into a subcapitate inflorescence . 6. Ser. Capitatae
9b. Stems almost glabrous, unwinged, fastigiately branched
from the base or the middle, with elongated internodes,
the branches lax and more or less corymbosely disposed

7. Ser. Fastigiatae

3b. Perennials:

lOa. Roots rather fleshy; leaves herbaceous

8. Ser. Napuliferae

lOb. Roots fibrous; leaves coriaceous 9. Ser. Coriaceae

CLASSIFICATION

I. Sect. Otophora Kusn. in Engl. & Prantl, Nat. Pflanzenfam.
4(2): 82 (1895). Type species: G. Otophora Franchet ex F.
B. Forbes & Hemsley.

Perennials, with a stout, cylindrical or fusiform, slightly fleshy
tap-root; branching monopodial; corolla usually lobed to the
middle or below, the tube shorter than, or equalling, the
lobes (except in G. doxiongshanensis T.N. Ho); plicae very
small, auriculate, attached on the right side of each lobe as
seen from within; seeds usually minutely reticulate, rarely
covered with honeycomb-like, hexagonal, simple, shallow
pits.

180

DISTRIBUTION. C. and E. Himalaya*, E. India (Assam), NE.
Burma, SW. Chinat-

Two series, 12 species.

1. Ser. Otophorae Marquand in Bull. misc. Inf. R. hot. Gdns,
Kew for 1937: 152 (1937). Type species: G. otophora
Franchet ex F. B. Forbes & Hemsley.

Plants with well-developed basal rosette; flowering stems
annual.

DISTRIBUTION. As for the section.
Eight species.

2. Ser. Decoratae Marquand in Bull. misc. Inf. R. hot. Gdns,
Kew for 1937: 152 (1937). Type species: G. decor ata Diels.

Plants with poorly developed basal rosettes; flowering stems
perennial.

DISTRIBUTION. As for the section.
Four species.

II. Sect. Cruciate Gaudin, Fl. helv. 2: 269 (1828). Type
species: G. cruciata L.

Sect. Erythaliae Bunge in Nouv. Mem. Soc. Imp. Nat. Mosc.

1(7): 207 (1829), non (Borkh.) Griseb. (1840). Type species:

G. cruciata L.
Sect. Tretorrhlza Griseb. in Hook., Fl. bor.-amer. 2: 60

(1837). Lectotype species: G. cruciata L.
Sect. Kurroo C. B. Clarke in J. Linn. Soc. (Bot). 14: 433

(1875). Type species: G. kurroo Royle.
Sect. Aptera Kusn. in Trudy imp. S.-Peterb. hot. Sada 13: 62

(1893). Lectotype species: G. biflora Regel.

Perennials; roots usually contiguous and twisted into a stout,
cylindrical structure; branching monopodial; caudex with a
brown, fibrous sheath of old petioles; basal rosette well-
developed, with large leaves; seeds minutely but distinctly
reticulate.

DISTRIBUTION. E., C., and S. Europe, U.S.S.R. European
part; W. Asia (Turkey, Iraq, Iran, Syria, Afghanistan), N.
and central part of Asia (Kashmir-Himalayan region, SW.
and N. China, U.S.S.R. (Caucasus, C. Asia, W. and E.
Siberia, Far East), Mongolia).

Twenty-one species.

III. Sect. Monopodiae (H. Smith) T.N.Ho in Bull. hot. Res.
Harbin 5(4): 10 (1985).

Subsect. Monopodiae H. Smith in Kew Bull. 15(1): 43 (1961).
Type species: G. cephalantha Franchet ex F. B. Forbes &
Hemsley.

Perennials; branching monopodial; seeds covered with
honeycomb-like, hexagonal, simple, shallow pits.

DISTRIBUTION. Kashmir-Himalayan region, Burma, NW.,
SW., C., S., and E. China, Japan, Thailand, Vietnam,
Malaya, Indonesia.

* Includes Nepal, Sikkim, Bhutan, and S. Tibet.

t The regions of China are NW.: Qinghai, Gansu, Shaanxi, Ningxia-Hui,
Xinjiang Uygur Zizhiqu; N.: Shanxi, Hebei, Nei Mongol Zizhiqu; NE.:
Heilongjiang, Jilin, Liaoning; C.: Henan, Hubei, Hunan; E.: Shandong,
Jiangsu, Anhui, Zhejiang, Jiangxi, Fujian, (and Taiwan); SW.: Tibet, Yunnan,
Guizhou, Sichuan; S.: Guangdong, Guangxi-Zhuang.

T.-N. HO&S.-W. LIU

Four series, 37 species.

1. Ser. Monanthae (H. Smith) T.N. Ho, comb. nov.

Sect. Frigida ser. Rosulatae subser. Monanthae H. Smith in
Kew Bull. 15(1): 54 (1961). Type species: G. szechenyii
Kanitz.

Caudex with a brown, membranous sheath of old petioles;
leaves broad; leaves and calyx-lobes both with conspicuous
cartilaginous margins.

DISTRIBUTION. Kashmir-Himalayan region, SW. and NW.
China.

Four species.

2. Ser. Vertidllatae (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 11(1985).

Kudoa Masam. in /. trop. Agric. 2: 29 (1930). Type species:

G. yakushimensis Makino.
Sect. Frigida ser. Vertidllatae Marquand in Bull. misc. Inf. R.

hot. Gdns, Kew for 1937: 158 (1937). Type species: G.

hexaphylla Maxim, ex Kusn.
Sect. Kudoa (Masam.) Satake & Toyok. ex Toyok. in /. Jap.

Bot. 35: 202 (1960).
Sect. Frigida ser. Ornatae subser. Vertidllatae H. Smith in

Kew Bull. 15(1): 54 (1961). Type species: G. hexaphylla

Maxim, ex Kusn.

Leaves in whorls of 3-8, narrow; flowers solitary, terminal, 5-
8-merous.

DISTRIBUTION. China (SW. and NW. regions, except Xinjiang
Uygur Zizhiqu, Taiwan), Japan.

Eight species.

3. Ser. Ornatae (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 11 (1985).

Sect. Frigida ser. Ornatae Marquand in Bull. misc. Inf. R.

hot. Gdns, Kew for 1937: 159 (1937). Type species: G. sino-

ornata Balf. f.
Sect. Frigida ser. Ornatae subser. Oppositifoliae H. Smith in

Kew Bull. 15(1): 54 (1961). Type species: G. sino-ornata

Balf. f.

Leaves opposite, narrow; flowers solitary, terminal, 5-merous.

DISTRIBUTION. C. & E. Himalayan region, Burma, SW. and
NW. China.

Sixteen species.

4. Ser. Apteroideae (H. Smith) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 11(1985).

Sect. Frigida ser. Rosulatae subser. Apteroideae H. Smith in
Kew Bull. 15(1): 54 (1961). Type species: G. cephalantha
Franchet ex F. B. Forbes & Hemsley.

Vegetative stems sometimes prolonged and branched, with
several basal rosettes decumbent on the ground; flowers
several together in infloresences at the apex of the flowering
stems.

DISTRIBUTION. Burma, SW., C., E., and S. China, Thailand,
Vietnam, Malaya, Indonesia, Sabah.

Nine species.

CLASSIFICATION OF GENTIAN A

IV. Sect. Frigida Kusn. in Trudy imp. S.-Peterb. hot. Sada 13:
61 (1893). Type species: G. frigida Haenke.

Sect. Frigida subsect. Sympodiae ser. Algidae H. Smith in
Kew Bull. 15(1): 55 (1961). Type species: G. algida Pallas.

Favargera A. & D. Love in Bot. Notiser 125: 256 (1972). Type
species: F. froelichii (Kusn.) A. & D. Love.

Gentianodes A. & D. Love in Bot. Notiser 125: 256 (1972).
Type species: G. frigida Haenke.

Perennials; branching sympodial, the plants with a short
rhizome, 1-few non-flowering rosettes and 1-few flowering
stems; seeds covered with membranous lamellae which form
spongy, hexagonal, complex pits.

DISTRIBUTION. Europe (Alps, Carpathians, and SW. Bulgaria);
Asia (Kashmir-Himalayan region, SW., NW., and ME.
China, U.S.S.R. (Caucasus, C. Asia, W. and E. Siberia, Far
East), Mongolia, Korea, Japan); N. America (western part of
Canada and the United States).

Eighteen species.

V. Sect. Gentiana. Type species: G. lutea L.

Sect. Coelanthae Froelich, Gentiana: 15 (1796). Type species:

G. lutea L.
Sect. Gentianotypus Dumort., Fl. belg.: 51 (1827). Type

species: G. lutea L.
Sect. Asteria Gaudin, Fl. helv. 2: 269 (1828). Lectotype

species: G. lutea L.

Perennials; branching sympodial, the plants with a short,
stout rhizome, a non-flowering basal rosette, and a flowering
stem; corolla lobed to the middle or below, plicae very small,
auriculate, attached to the right side of each lobe as seen from
within, or absent (G. luted); seeds with thickened reticulate
testa, and a discoid wing.

DISTRIBUTION. C. and S. Europe and S. Norway; Turkey.
Five species.

VI. Sect. Pneumonanthe (Gled.) Gaudin, Fl. helv. 2: 269
(1828).

Pneumonanthe Gled., Syst. pi. stamin. situ: 238 (1764). Type
species: G. pneumonanthe L.

Gentiana Pneumonanthe Link, Enum. hort. berol. alt. 1:

258 (1821). Lectotype species: G. pneumonanthe L.

Subgen. Pneumonanthe Raf., Med. fl. 1: 208 (1828). Lecto-
type species: G. pneumonanthe L.

Sect. Cyane Griseb. in Hook., Fl. bor.-amer. 2: 54 (1837).
Type species: G. pneumonanthe L.

Perennials; branching sympodial, the plants with a rhizome
and 1-few flowering stems, but non-flowering basal rosettes
usually absent; flowering stems bearing scale-like leaves
below; seed with a reticulately thickened testa, usually
winged.

DISTRIBUTION. Much of Europe; N. and E. Asia; N. and C.
America.

Thirty-eight species.

VII. Sect. Phyllocalyx T.N. Ho in Bull. hot. Res. Harbin 5(4):
14 (1985). Type species: G. phyllocalyx C. B. Clarke.

Sect. Frigida p. Phyllocalyx Kusn. in Trudy imp. S.-Peterb.
hot. Sada 15: 287 (1898).

181

Perennials; branching sympodial, the plants with short
stolons; calyx very small, completely enveloped by the upper-
most pair of leaves; stigma-lobes expanded, contiguous into a
small disc or infundibuliform structure, but free after anthe-
sis; seeds with honeycomb-like, hexagonal, simple, shallow
pits, with a discoid wing.

DISTRIBUTION. C. and E. Himalayan region, mountains of
SW. China.

A monotypic section.

VIII. Sect. Calathianae Froelich, Gentiana: 57 (1796). Lecto-
type species: G. nivalis L.

Hippion F. W. Schmidt, Fl. boem. 2: 18 (1793). Type species:
H. aestivum F. W. Schmidt.

Sect. Ericoila Dumort., Fl. belg.: 51 (1827). Type species: G.
nivalis L.

Sect. Gentianella Gaudin, Fl. helv. 2: 270 (1828), non
Dumort. (1827). Lectotype species: G. nivalis L.

Sect. Hippion (F. W. Schmidt) Gaudin, Fl. helv. 2: 270
(1828).

Sect. Cydostigma Griseb., Gen. sp. Gent.: 259 (1838). Lecto-
type species: G. nivalis L.

Perennials, less often annuals; branching sympodial, the
plants with stolons and runners; calyx-tube prominently
angled or winged; corolla hypocrateriform; stigma-lobes
expanded, almost smooth on the surface, contiguous into a
disc or infundibuliform structure, but not free after anthesis;
seeds usually minutely reticulate, unwinged, rarely reticulately
thickened and with a wing at one end.

DISTRIBUTION. W., C., and S. Europe, U.S.S.R. (Arctic
European part); W. Asia (Turkey, Iran, Iraq), N. Asia
(U.S.S.R. (Caucasus, Altai), Mongolia); N. America
(Labrador, Greenland); Africa (Morocco).

Eight species.

IX. Sect. Ciminalis (Adans.) Dumort., Fl. belg.: 51 (1827).

Ciminalis Adans., Fam. pi. 2: 504 (1763). Lectotype species:
G. acaulis L.

Sect. Megalanthe Gaudin, Fl. helv. 2: 270 (1828). Lectotype
species: G. acaulis L.

Sect. Thylacites Griseb. in DC., Prodr. 9: 115 (1845). Lecto-
type species: G. acaulis L.

Perennials; branching sympodial, the plants with stolons and
runners; corolla obconical; stigma-lobes expanded, fimbriate
on the surface, contiguous into a small disc or infundibuliform
structure, free after anthesis; seeds rugose, with a minutely
reticulate testa.

DISTRIBUTION. C. and S. Europe.
Five species.

X. Sect. Isomeria Kusn. in Trudy imp. S.-Peterb. Obshch.
Estest. 24(2): 196 (1894). Lectotype species: G. depressa D.
Don.

Perennials; branching sympodial, the plants with stolons and
runners; stigma-lobes free, recurved, linear or oblong; seeds
with honeycomb-like, hexagonal, simple, shallow pits, rarely
with spongy, hexagonal, complex pits.

DISTRIBUTION. Asia (Kashmir-Himalayan region, NE. Burma,

182

SW., NW., and C. China, U.S.S.R. (E. Siberia, Far East),
Japan); N. America (western part of Canada and the United
States).

Four series, 18 species.

1. Ser. Sikkimenses (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 13 (1985).

Sect. Frigida ser. Sikkimenses Marquand in Bull. misc. Inf.
R. hot. Gdns, Kew for 1937: 162 (1937). Type species: G.
sikkimensis C. B. Clarke.

Leaves obovate, spathulate or oblong, rounded or obtuse at
the apex, petioles of cauline leaves gradually broadened
towards the top of the stem; flowers in fascicles of 3-8 in a
capitate infloresence.

DISTRIBUTION. As for the section, except NW. China.
Six species.

2. Ser. Depressae (H. Smith) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 13 (1985).

Sect. Frigida ser. Confertifoliae Marquand in Bull. misc. Inf.

R. hot. Gdns, Kew for 1937: 161 (1937). Type species: G.

confertifolia Marquand.
Sect. Frigida subsect. Sympodiae ser. Depressae H. Smith in

Kew Bull. 15(1): 55 (1961). Type species: G. depressa D.

Don.

Leaves and calyx-lobes with conspicuous cartilaginous mar-
gins; flowers usually solitary, terminal.

DISTRIBUTION. C. and E. Himalayan region, SW. and NW.
China.

Six species.

3. Ser. Stragulatae T.N. Ho in Bull. hot. Res. Harbin 5(4): 13
(1985). Type species: G. stragulata Balf. f. & Forrest ex
Marquand.

Calyx-lobes orbicular, spathulate, abruptly contracted at the
base into a tongue.

DISTRIBUTION. E. Himalayan region, SW. China.
Two species.

4. Ser. Uniflorae (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 14 (1985).

Sect. Frigida ser. Uniflorae Marquand in Bull. misc. Inf. R.
hot. Gdns, Kew for 1937: 161 (1937). Type species: G.
filistyla Balf. f. & Forrest ex Marquand.

Flowers solitary, terminal; calyx-lobes narrowly elliptical or
triangular.

DISTRIBUTION. Himalayan region, SW. China.
Four species.

XI. Sect. Microsperma T.N. Ho in Bull. hot. Res. Harbin
5(4): 14 (1985). Type species: G. delavayi Franchet.

Annuals; capsule oblong; seed with honeycomb-like, hex-
agonal, simple, shallow pits.

DISTRIBUTION. Nepal, Bhutan, SW. China.
Three series, 10 species.

T.-N. HO&S.-W. LIU

1. Ser. Suborbisepalae (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 15 (1985).

Sect. Frigida ser. Suborbisepalae Marquand in Bull. misc. Inf.
R. hot. Gdns, Kew for 1937: 160 (1937). Type species: G.
suborbisepala Marquand.

Leaves and calyx-lobes spathulate; flowers small; gynoecium
and stamens usually exserted from the corolla.

DISTRIBUTION. Nepal, Bhutan, SW. China.
Six species.

2. Ser. Tetramerae (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 15 (1985).

Sect. Frigida ser. Annuae subser. Tetramerae Marquand in
Bull. misc. Inf. R. hot. Gdns, Kew for 1937: 165 (1937).
Type species: G. lineolata Franchet.

Flowers 4-merous, medium-sized; calyx-lobes with strongly
keeled midribs which are decurrent below into strong wings.

DISTRIBUTION. China (Yunnan, Sichuan).
Two species.

3. Ser. Annuae (Marquand) T.N. Ho in Bull. hot. Res.
Harbin 5(4): 15 (1985).

Sect. Frigida ser. Annuae Marquand in Bull. misc. Inf. R.
hot. Gdns, Kew for 1937: 165 (1937). Type species: G. picta
Franchet ex F. B. Forbes & Hemsley.

Flowers 5-merous, small; calyx-lobes linear or oblanceolate.
DISTRIBUTION. China (Yunnan, Sichuan).
Two species.

XII. Sect. Stenogyne Franchet in Bull. Soc. hot. Fr. 31: 375
(1884). Type species: G. primulaefolia Franchet.

Annuals, rarely perennials; flowers large or medium-sized;
calyx keeled or winged; plicae asymmetrical, extremely oblique;
style filiform, longer or slightly shorter than the ovary; seeds
usually with a minutely reticulate testa, triquetrous with three
winged edges, rarely unwinged or areolate.

DISTRIBUTION. E. Burma, SW., NW., N., C., and S. China,
Thailand.

Fourteen species.

XIII. Sect. Dolichocarpa T.N. Ho in Bull. hot. Res. Harbin 5
(4): 16 (1985). Type species: G. prostrata Haenke.

Varasia Philippi, Fl. atacam.: 35 (1860). Type species: V.
podocarpa Philippi.

Annuals; flowers small; plicae symmetrical, large; capsule
oblong, unwinged; seeds minutely reticulate, unwinged at
maturity, rarely winged when young.

DISTRIBUTION. Europe (E. Alps); Asia (W. Asia (Iran,
Afghanistan), Pamir-Himalayan region, SW. and NW. China,
U.S.S.R. (Caucasus, Arctic region, C. Asia, W. and E.
Siberia, Far East)); N., C., and S. America.

Twelve species.

XIV. Sect. Chondrophyllae Bunge in Nouv. Mem. Soc. imp.
Nat. Mosc. 1(7): 207, 231 (1829) (Changed to Chondrophylla
by most later authors). Lectotype species: G. aquatica L.

CLASSIFICATION OF GENTIAN A

Chondrophylla (Bunge) Nelson in Bull. Torrey hot. Club 31:

245 (1904).
Holubia A. & D. Love in An. Inst. hot. A. J. Cavanilles 32:

226 (1975), nom. illeg. (Art. 64.1), non Holubia Oliver

(1884). Type species: H. pyrenaica (L.) A. & D. Love.
Holubogentia A. & D. Love in Bot. Notiser. 131: 385 (1978),

based on Holubia A. & D. Love.

Annuals, rarely perennials; flowers small; plicae symmetrical,
large; capsule obovoid or ellipsoid-oblong, conspicuously
winged, the wing strong and broad at the apex but gradually
narrowed towards the base; seeds minutely reticulate.

DISTRIBUTION. Europe, Asia, America, Morocco, Australia.
Eleven series, 158 species.

1. Ser. Fimbriatae Marquand in Bull. misc. Inf. R. hot. Gdns,
Kew for 1937: 169 (1937). Lectotype species: G. panthaica
Burk. The type designation of 'G. recurvata C. B . Clarke' by
Ho (1985) is incorrect (Art. 8) because Marquand's original
series circumscription (1937) did not include G. recurvata.

Leaves and calyx densely and minutely papillose; calyx-lobes
filiform-subulate; plicae usually fimbriate, rarely dentate.

DISTRIBUTION. C. and E. Himalayan region, N. Burma, SW.,
C., and S. China, Indonesia (Sumatra).

Twenty species.

2. Ser. Rubicundae Marquand in Bull. misc. Inf. R. hot.
Gdns, Kew for 1937: 171 (1937). Type species: G. rubicunda
Franchet.

Calyx-lobes filiform; plicae entire or denticulate.
DISTRIBUTION. China (except N. and NE.), Philippines.
Ten species.

3. Ser. Linearifoliae Marquand in Bull. misc. Inf. R. hot.
Gdns, Kew for 1937: 170 (1937). Type species: G. aristata
Maxim.

Cauline leaves long and very narrow, at least the uppermost
linear or subulate.

DISTRIBUTION. Europe (E. Pyrenees, Carpathians, SW.
Bulgaria); Asia (W. Asia (Turkey, Iran, Afghanistan), C.
and E. Himalayan region, SW., S., and NW. China,
U.S.S.R. (Caucasus)).

Fifteen species.

4. Ser. Orbiculatae Marquand in Bull. misc. Inf. R. hot.
Gdns, Kew for 1937: 170 (1937). Type species: G. squarrosa
Ledeb.

Calyx-lobes recurved or patent, rarely erect, ovate or orbicular,
rarely lanceolate, contracted at the base.

DISTRIBUTION. Kashmir-Himalayan region, China, U.S.S.R.
(C. Asia, W. and E. Siberia, Far East), Mongolia, Korea,
Japan, Indian Peninsula, Sri Lanka, Indonesia; E. Australia.

Twenty species.

5. Ser. Humiles Marquand in Bull. misc. Inf. R. hot. Gdns,
Kew for 1937: 172 (1937). Lectotype species: G. leucomel-
aena Maxim, ex Kusn. The type designation of 'G. aquanca
L.' by Ho (1985) is incorrect (Art. 8) because Marquand's
original circumscription (1937) did not include this species.

183

Ser. Pubigerae Marquand in Bull. misc. Inf. R. hot. Gdns,
Kew for 1937: 168 (1937).

Stems ascending or decumbent, usually branched from the
base, without a conspicuous main stem, or rarely
unbranched; calyx-lobes erect, triangular or lanceolate.

DISTRIBUTION. Europe (Spain); Asia (W. Asia, Kashmir-
Himalayan region, Burma, China, U.S.S.R. (Caucasus, C.
Asia, W. and E. Siberia), N. Mongolia, Korea, Japan); N.
America (western part of Canada and the United States).

Thirty-three species.

6. Ser. Capitatae T.N. Ho in Bull. hot. Res. Harbin 5(4): 18
(1985). Type species: G. capitata Buch.-Ham. ex D. Don.

Stems simple, densely papillose, winged or angled and leaf-
less in the lower half; in the upper half much branched with
abbreviated internodes, numerous leaves and flowering
branches, all congested at the apex of the stem into a
subcapitate inflorescence.

DISTRIBUTION. Asia (Kashmir-Himalayan region, N. Burma,
SW. China, Indo-China); C. America (Mexico and
Guatemala); Africa (Morocco).

Sixteen species.

7. Ser. Fastigiatae T.N. Ho in Bull. hot. Res. Harbin 5(4): 18
(1985). Type species: G. intricata Marquand.

Stems fastigiately branched from the base or the middle, the
main stem very conspicuous, with lax, more or less corym-
bosely disposed branches.

DISTRIBUTION. Asia (W. Asia, Pamir-Himalayan region,
China, U.S.S.R. (C. Asia, W. and E. Siberia, Far East),
Mongolia, Korea, Japan, Indonesia (Sumatra)); western part
of N. America.

Twenty-four species.

8. Ser. Napuliferae T.N. Ho in Bull. hot. Res. Harbin 5(4): 18
(1985). Type species: G. napulifera Franchet.

Perennials, with slightly fleshy, fusiform roots; leaves
herbaceous.

DISTRIBUTION. Bhutan, E. India (Assam), SW., C., and S.
China, Thailand, Vietnam.

Three species.

9. Ser. Coriaceae T.N. Ho, ser. nov. Type species: G.
cruttwellii H. Smith.

Herba perennis, radicibus fibrosis; folia coriacea.

DISTRIBUTION. S. China (Taiwan), Philippines, Sabah, New
Guinea.

Thirteen species.

10. Ser. Piasezkianae T.N. Ho in Bull. bot. Res. Harbin 5(4):
18 (1985). Type species: G. piasezkii Maxim.

Calyx-lobes with keeled midribs which are decurrent below
into conspicuous wings; corolla hypocrateriform; capsule
narrowly elliptical, attenuate or obtuse at the apex.

DISTRIBUTION. China (N. Sichuan, Gansu, Shaanxi).
Three species.

184

T.-N. HO&S.-W. LIU

11. Ser. Grandiflorae Grossh. ex T.N. Ho, ser. nov. Type
species: G. grandiflora Laxm.

Ser. Grandiflorae Grossh. in V. Komarov, Fl. URSS 18: 572
(1952), nom. inval. (Art. 36.1).

Herba perennis; flores grandi, 40-55 mm longi; capsula
anguste elliptica, apice attenuata vel obtusa.

DISTRIBUTION. U.S.S.R. (W. and E. Siberia, C. Asia),
Mongolia.

One species.

XV. Sect. Fimbricorona T.N. Ho, nom. nov.

Sect. Eurythalia (Borkh.) Griseb. in Getting, gel. Anz. for
1840 (2): 815 (1840), nom. illeg. (Art. 64.1), non Bunge
(1829). Lectotype species: G. coronata Royle.

Annuals; corolla bearing a corona of multicellular hairs in the
throat; plicae well-developed; capsule obovate, rounded at
the apex, winged, the wing strong and broad at the apex but
gradually narrowed towards the base of the capsule; seeds
minutely reticulate.

DISTRIBUTION. Kashmir-Himalayan region, mountains of
SW. China.

Four species.

GEOGRAPHICAL DISTRIBUTION

Gentiana is a subcosmopolitan genus. It is distributed widely
in Europe and the temperate zone of Asia and N. America,
but a few species extend northwards to the Arctic region
(arctic Siberia, Iceland and to 73 N in Greenland), south-

wards to C. and S. America (the southernmost to Santa Cruz
(c. 49 58'S, 69W), Argentina), through S. Asia to the South
Tableland (c. 35 58'S, 149 25'E) of Australia, and from SW.
Europe to Djebel Toubkal in C. Morocco.

There are 14 sections in Asia, eight in Europe, and six in
America, comprising 93.3%, 53.3%, and 40% respectively of
the world total, while there are 312 species in Asia, 27 in
Europe, and 35 in America, comprising 86.4%, 7.5%, and
9.7% respectively of the world total,

In Asia (Table 1) sect. Ciminalis is entirely absent; sect.
Gentiana and sect. Calathianae contain only one or two
species in this area and are restricted to W. Asia, with a few
subspecies scattered as far as Lake Baikal; sect. Pneumonanthe
also has few (only 12) here. Conversely, the other 11 sections
contain their highest numbers of species in Asia, i.e. Otophora
12 species in Asia/ 12 species in the world, Cruciata 21/21,
Monopodiae 37/37, Frigida 16/18, Phyllocalyx 1/1, Isomeria
18/18, Microsperma 10/10, Stenogyne 14/14, Dolichocarpa 10/
12, Chondrophyllae 154/158, and Fimbricorona 4/4. More-
over, of these 11 sections, six (Otophora, Monopodiae,
Phyllocalyx, Microsperma, Stenogyne, and Fimbricorona) are
endemic to Asia. In respect of species, of the 312 species 299
are endemic to Asia, 95.8% of the total for Asia, and only 13
are common to Asia and Europe or America. It is clear that
Asia is the major centre for Gentiana with the greatest
number of species as well as endemics. The highest con-
centration of these occurs in the mountains of SW. China
(NW. Yunnan, W. Sichuan, and SE. Tibet) and in the NE.
Burma area, the square between 25-34 N and 91-105 E. In
this area there are 1 1 sections and 190 species, of which 98 are
endemic to the mountains of SW. China and the NE. Burma
area; thus three quarters of all the sections and half the
species of the world occur in this small area, indicating that it
is a major centre of diversity and endemism of Gentiana. In
addition, 10 sections and 61 species (13 of which are local

Table 1 The numbers of species of Gentiana in different regions.

Sections

<a

u

S *

<u

i

|

1

1

2

efl

'B
o

03

c

-3

o g

CO

C

a

a

_CS

&

u
c o
>> o

n.
o

8

I
o

6

Crucial

G.

O

03
T3

Gentia

o

13 r"!

(U >>

'

5
"
U

cfl

C

E
ij

(U

o

89

Micros

eo .c
o u
c
<u o

$5 Q

o

c
o

U

Fimbri

1

The world

12

21

37

18

5

38 1

8

5

18

10

14 12

158

4

361

Total

12

21

37

16

1

12 1

2

-

18

10

14 10

154

4

312

1 Mts of SW. China

10

10

25

15

-

1

-

-

15

9

9 6

88

2

190

C. & E. Himalaya

5

3

8

3

-

1

-

-

9

2

5

23

2

61

Total

_

2

_

2

5

2

8

5

_

_

1

2

_

27

<u

o Pyrenees-Alps

-

1

-

2

5

2 -

8

5

-

-

1

2

-

26

W Carpathians

-

1

-

1

2

2

4

2

-

-

-

1

-

13

Total

-

-

-

1

-

26

1

-

1

-

3

3

-

35

g N. America

-

-

-

1

-

20

1

-

1

-

1

2

-

26

w C. America

-

-

-

-

-

6

-

-

-

-

2

1

-

9

^v, c A,rticric3.

-i

i

Africa (Morocco)

-

-

-

-

-

- -

1

-

-

-

1

-

j

2

A.ustr3.1i2

i

.

i

CLASSIFICATION OF GENTIAN A

185

Fig. 5 Isoflor map of species of Gentiana. \ . 88-100 species; 2. 60-64 species; 3. 27-42 species; 4. 1 1-18 species; 5.4-10 species; 6. 1-3 species.

endemics) are found in the Himalayan region (Bhutan,
Sikkim, Nepal, and S. Tibet), which forms a second centre of
diversity and endemism. However, the scores drop more
sharply in moving from the main centre to the adjacent
regions. The isoflor map (Fig. 5) shows the maximum con-
centration of species and this trend in distribution.

Of the eight sections in Europe, sect. Cruciata occurs also
in Asia, while three others (Pneumonanthe, Frigida, and
Dolichocarpa) also occur in both Asia and America, and sect.
Chondrophyllae also occurs in Asia, America, Australia, and
Morocco (these sections, however, contain only one or two
species each). Sect. Ciminalis is a strictly European endemic
and sect. Gentiana is almost endemic. There are 27 species in
Europe, of which 17 are endemic and 10 more widespread. Of
the 10 widespread species, all also occur in Asia, but only two
reach America and one NW. Africa (Morocco). The maxi-
mum concentration of sections and species in Europe is in the
Alps and Pyrenees, with 26 species belonging to eight sec-
tions; the second concentration is in the Carpathians, with 13
species belonging to seven sections. It is suggested that the
floristics of European Gentiana is closely related to, and
derived from, that of Asia, and has reached its highest
development in the most favourable environment of the Alps
and Pyrenees.

In America, in contrast to Asia and Europe, there are no
endemic sections, all six being widespread; one also occurs in
Asia, and the others in Asia and Europe. Five of the six
sections (Frigida, Calathianae, Isomeria, Dolichocarpa, and
Chondrophyllae) contain only 1-3 species each. Sect.
Pneumonanthe contains 26 species in America, more than
elsewhere, and all are endemic. There are 35 species in all in
America, of which 30 are endemic and five widespread. All
five widespread species also occur in Asia, while two occur in
Europe as well. The floristics of American Gentiana can thus
be said to be dominated by sect. Pneumonanthe and to be

derived from Asia, although there is a lesser development
than in Asia, or even than in Europe, without any endemic
sections but with more endemic species. Most species occur in
N. Ameria (26 species), but some in C. America (9 species)
and S. America (3 species). However, there is no obvious
centre of diversity, although there are relatively many species
(10) in Pennsylvania, Virginia, N. and S. Carolina, Georgia,
and Florida.

According to the literature, there is no Gentiana in Australia,
but we have seen one in SE. Australia, G. quadrifaria Blume, a
rather common species of Indo-China, Malaya, and Indonesia.
The Australian occurrence is clearly an extension from S.
Asia. We have also seen specimens of two species from
Morocco: G. brachyphylla Vill. and G. atlantica Litard. &
Maire. The former presumably spread from Europe, while
the latter is endemic to Morocco.

The geographical patterns within Gentiana can be further
indicated by the floristic analysis of sections and species. The
15 sections and 361 species may be arranged in 14 geo-
graphical elements and variant patterns.

No section is tropical but an overwhelming majority, 14
(about 93.3% of the world total), have a temperate distribu-
tion pattern. 326 species (about 90.3% of the world total)
have a temperate distribution pattern, and only 35 species
(about 9.7%) have a tropical distribution pattern. Gentiana is
thus a predominantly temperate genus.

Of those sections with a temperate distribution, most (6
sections, 40% of the total) belong to the E. Asia element and
to its variant, the Sino-Himalaya element, then to the North
temperate element (4 sections), then to the European en-
demic element (2 sections), Old world temperate element (1
section), E. Asia and N. America disjunct element (1 section)
and cosmopolitan element (1 section). The distribution pat-
tern of the species is essentially similar to that of the sections.
It is dominated by the E. Asia element, containing 259

186

species, about 71.8% of the world's total, but the percentage
of endemic and stenochorous species is very high, whereas
that of the widespread species is quite low. It is clear that
these widespread distribution patterns (North temperate, Old
world temperate, and E. Asia and N. America disjunct)
indicate the relationship between the floras of Asia, Europe,
and America. The mountains of SW. China and NE. Burma
comprise not only a strong modern centre of distribution, but
may also be the main area of origin for the genus, as the
greatest number of taxa occur there, as well as probably the
primitive type of the genus, sect. Stenogyne. Through this
primitive section, Gentiana shows a certain affinity with
Crawfurdia and Tripterospermum. Sect. Stenogyne,
Crawfurdia, and Tripterospermum share asymmetrical plicae
between the corolla-lobes, glands at the base of the ovary,
and winged seeds. Their geographical distributions all belong
to the E. Asia pattern and most species are centred on the
mountains of SW. China and NE. Burma.

The geographical distribution of the sections may be sum-
marized as follows:

A. Wide distribution

1. Cosmopolitan. There is only one section with this distribu-
tion pattern, Chondrophyllae . This section contains some 158
species and has the widest distribution of any section of
Gentiana. It occurs in. Europe, Asia, America, NW. Africa,
and Australia. The great majority (154) of species are centred
in Asia (especially in the mountains of SW. China and NE.
Burma, where there are 88 species) and occur all over Asia,
with one species (G. quadrifaria Blume) extending to SE.
Australia, and another (G. aquatica L.) to western North
America. There are two species in Europe, three in America,
and one in Africa (Morocco). One European species, G.
boryi Boiss., is endemic to Spain, and another, G. pyrenaica
L., occurs in the E. Pyrenees, the Carpathians and the

T.-N. HO&S.-W. LIU

mountains of SW. Bulgaria, and extends to W. Asia. Of
the three American species, G. douglasiana Bong, is en-
demic to western N. America, G. pumilio Standley &
Steyerm. is endemic to Guatemala and Mexico, and G.
aquatica L. occurs in the western part of N. America as well
as in N. and W. Asia (Fig. 6). The Moroccan species, G.
atlantica Litard. & Maire, is endemic to the western coast of
that country.

B. Tropical distribution

No section can be described as having a tropical distribution,
although a few species do.

C. Temperate distribution

The overwhelming majority of sections and species have
various types of temperate distribution pattern.

1 . North temperate pattern. Four sections have this distribu-
tion pattern, which covers Europe and the temperate zone of
Asia and N. America.

Sect. Frigida. Although this section with 18 species occurs
in Europe, Asia, and N. America, there is a disjunction
across the eastern European plain of the U.S.S.R. The two
western species are endemic to the Alps, Carpathians, and
SW. Bulgaria, whereas 16 species occur in the East, Asia, and
America. The distribution of these species is centred on the
mountains of SW. China and NE. Burma, where of 15
species, ten are extremely local, the others extending both
east and west, i.e. one species to the Himalayan region, two
just to Qinghai, and one to both Qinghai and the Himalayan
region. G. algida Pallas, which is more widespread, links the
continents of Asia and America, extending from the moun-
tains of SW. China westwards to Sikkim, northwards across
NW. China to C. Asia, and through W. and E. Siberia,

Fig. 6 Distribution area of sect. Chondrophyllae Bunge, the cross-hatched area showing the maximum concentration of species.

CLASSIFICATION OF GENTIANA

187

Fig. 7 Distribution area of sect. Frigida Kusn., the cross-hatched area showing the maximum concentration of species.

Fig. 8 Distribution area of sect. Pneumonanthe (Gled.) Gaudin, the cross-hatched area showing the maximum concentration of species.

Korea, Japan, and the Far East into Alaska, and south along
the Rocky Mts to Colorado. G. apiata N. E. Br. is restricted
to Tai-Pei-Shan of Shaanxi, China (Fig. 7).

Sect. Pneumonanthe. This section, consisting of 38 species,
is distributed widely through much of Europe, N. and E.
Asia, and N. and C. America, and disjunctly in Kamchatka

and the Kolyma-Anadyr region of the U.S.S.R. It is centred
in America where the majority of species (26) occur, 20 being
endemic to N. America and six to C. America. Fewer species
(\2) occur in Asia and Europe, six being restricted to E. Asia,
occurring in NE., N., and E. China, U.S.S.R. (E. Siberia,
Far East), Korea, and Japan, four occurring in W. and C.
Asia, and two being common to both Asia and Europe,

188

T.-N. HO&S.-W. LIU

Fig. 9 Distribution area of sect. Dolichocarpa T.N. Ho, the cross-hatched area showing the maximum concentration of species.

Fig. 10 Distribution area of sect. Cruciata Gaudin, the cross-hatched area showing the two subcentres of distribution of the species.

CLASSIFICATION OF GENTIANA

distributed over much of Europe, W. and C. Asia, and W.
and E. Siberia (Fig. 8).

Sect. Calathianae. This rather small section, comprising
eight species and five subspecies, occurs in C., S., and N.
Europe, although G. brachyphylla Vill. extends south-east to
Turkey, Iraq, and N. Iran, and south-west to Morocco, with
two subspecies extending further to C. Asia and the Altai; G.
nivalis L. also extends south-east to Turkey and the Caucasus,
as well as north to Labrador in Canada and Greenland to
about 73 N, where it reaches the northernmost distribution
limit of the genus (Fig. 12-1).

Sect. Dolichocarpa. Although this section, with 12 species,
belongs to the North temperate pattern, it extends more into
South America than the sections mentioned above. It is
centred in NW. and SW. China and the Himalayan region,
where nine species occur. The three other species occur
exclusively outside this area, G. podocarpa (Philippi) Griseb.
being endemic to South America, G. sedifolia Kunth occur-
ring in S. and C. America, and G. prostrata Haenke having a
remarkably widespread distribution, being scattered inter-
ruptedly in the E. Alps, W. Asia, the Caucasus, C. Asia, W.
and E. Siberia, the Far East, the Kolyma-Anadyr region,
Alaska, and along the mountains of the western part of N.,
C., and S. America south to Santa Cruz, at about 49 58' S, in
Argentina (Fig. 9).

2. Old world temperate pattern. Only one section has this
distribution pattern, which covers Europe and the temperate
and cold-temperate zones of Asia.

Sect. Cruciata. This section comprises 21 species and has
two sub-centres of distribution: the first is in the Himalayan
region and the Qinghai-Tibet plateau (including the mountains
of SW. China) with 13 species, of which 11 are restricted to
this area and two extend eastwards and link the Himalayan
region, the Qinghai-Tibet plateau and the north part of

189

China, C. Asia, and E. Siberia; the second is in C. Asia,
where three species are endemic and six more widespread,
extending east and west and linking Europe, W. Asia, C.
Asia, the northern part of China, Siberia, and the Far East.
The most widespread of these, G. cruciata L. , extends west to
E., C., and S. Europe, east to W. Siberia, and south-west to
W. Asia and the Caucasus. Apart from these species, one
other, G. tenuicaulis Ling, occurs away from the two sub-
centres, in Hebei, China (Fig. 10).

3. E. Asia and N. America disjunct pattern. Only one
section has this distribution pattern, which covers the tem-
perate and subtropical zones of E. Asia and North America.

Sect. Isomeria. This section consists of 18 species. It is
centred in the mountains of SW. China and the Himalayan
region, where there are 17 species, three of which extend
slightly east and south, i.e. G. chinensis Kusn., occurring
from Sichuan to W. Hubei in China, G. urnula H. Smith from
E. Tibet to SW. Qinghai, and G. depressa D. Don from the
Himalayan region, with a disjunct southern area in the Indian
Peninsula. The remaining species, G. glauca Pallas, is some-
what separated from the main area of distribution and occurs
in E. Asia and N. America, extending from Japan, E. Siberia,
the Far East, and the Kolyma-Anadyr region into Alaska, the
Yukon, and along the Rocky Mts to Montana in the United
States (Fig. 11).

4. E. Asia pattern. Two sections have this distribution
pattern, which covers Kashmir, the Himalaya, China, Korea,
Japan, E. Siberia, and the Far East of Russia, extending
south to Indonesia.

Sect. Monopodiae. This section consists of 37 species and
occurs from the Himalayan region and China to Japan, and
extends south to Indonesia. The greatest concentration of
species occurs in the SW. provinces of China and the Himalayan
region (to Kashmir) with 32 species, with two of them

Fig. 11 Distribution area of sect. Isomeria Kusn., the cross-hatched area showing the maximum concentration of species.

190

T.-N. HO&S.-W. LIU

Wig. 12 Distribution areas of sections: 1. Calathianae Froelich; 2. Microsperma T.N. Ho; 3. Stenogyne Franchet; 4. Monopodiae (H. Smith)

T.N. Ho.

Fig. 13 Distribution areas of sections: 1. Gentiana; 2. Ciminalis (Adans.) Dumort.; 3. Phyllocalyx T.N. Ho and Otophora Kusn.

CLASSIFICATION OF GENTIAN A

191

extending from Yunnan in China through Burma to Thailand,
and from Guangxi-Zhuang in China to Vietnam. The remain-
ing five species occur exclusively outside the main area and
are all endemic, one to Japan and Taiwan, three to Indonesia,
and one restricted to S. and E. China, with two varieties in
Guizhou, Fujian, and Taiwan (Fig. 12-4).

Sect. Stenogyne. This section consists of 14 species, and
occurs in SW., NW., N., C., and S. China, E. Burma, and
Thailand. It is centred on SW. China where there are 13
species, of which 10 are endemic; three species are more
widespread, extending north to SE. Qinghai, S. Gansu,
Ningxia-Hui (G. striata Maxim.), to S. Shanxi, Henan,
Hubei, Guangxi-Zhuang (G. rhodantha Franchet ex F. B.
Forbes & Hemsley), and south to E. Burma (G. melvillei S.
Moore). The final species, G. australis Craib, is the southern-
most of this section, being endemic to Thailand (Fig. 12-3).

Sino-Himalaya variant of the E. Asia distribution pattern.
Four sections have this distribution pattern, which covers the
area from the mountains of SW. China to the Himalayan
region.

Sect. Otophora. This section of 12 species has a typical
Sino-Himalayan pattern, occurring in the mountains of SW.
China and NE. Burma (10 species) and the E. and C.
Himalayan region (5 species). It has a marked western
distribution (Fig. 13-3).

Sect. Phyllocalyx. This monotypic section is restricted to
the mountains of SW. China, NE. Burma, and the E. and C.
Himalayan region (Fig. 13-3).

Sect. Microsperma. This section of 10 species is almost
restricted to the SW. provinces of China, being found in
Yunnan, Sichuan, SE! Tibet, and Guizhou, but with one
species, G. vernayi Marquand, extending from SE. Tibet to
Nepal (Fig. 12-2), and another, G. rotundisepala T.N. Ho,
endemic to Nepal and Bhutan.

Sect. Fimbricorona. This small section of four species also
has a typical Sino-Himalaya pattern. Two species occur from
NW. Yunnan and SE. Tibet to Bhutan, and the other two
from NW. India (Garhwal, Punjab) to the NW. Himalaya,
Kashmir, and Lahul.

5. Endemic to Europe. Two sections.

Sect. Gentiana. This section with five species is almost
endemic to Europe, being centred in the mountains of C. and
S. Europe, with G. purpurea L. extending to S. Norway, and
G. lutea to Turkey (Fig. 13-1).

Sect. Ciminalis. This section of five species is strictly
endemic to Europe, occurring in the mountains of C. and S.
Europe, from SW. Germany and the Carpathians to the Alps,
Pyrenees, Spain, C. Italy, and SW. Yugoslavia, but is absent
from Greece (Fig. 13-2).

STUDY MATERIALS

This study has been based primarily on standard herbarium
techniques and on field observations in natural habitats in
Yunnan, Sichuan, Tibet, and Qinghai provinces, China.
Nearly 20,000 specimens have been examined from the

following herbaria in Europe, China, America, Japan, and
Australia, either by visiting these herbaria or by the loan of
specimens: A, AU, BM, CANB, CCTC, CIB, COLO, E,
FMPI, FUS, FUTU, GMI, HBDB, HGBG, HHBG, HHU,
HIB, HMPI, HNDB, HNTC, HNWP, HSNU, IBSC, K,
KUN, L, LE, LINN, LZU, MAK, MEL, MO, N, NAS, NK,
NY, P, PE, S, SDB, SHM, SMMI, SZ, TI, TIM, UC, UPS,
US, W, WH, WU, WUK, YNDB, YUKU, ZHDB, and
ZHM.

ACKNOWLEDGEMENTS. We acknowledge with gratitude the directors
of the herbaria listed above for facilities and for the loan of specimens
during the preparation of this paper and 'Flora of China, volume 62,
Gentianaceae'.

In particular, we are most grateful for the opportunity to make a
visit to Great Britain in connection with the Gentiana project and
extend here our thanks to the Academia Sinica with whose financial
support this visit was made by T.N. Ho from March 1986 to May
1987. We sincerely thank the directors and staff of the British
Museum (Natural History), the Royal Botanic Gardens, Kew, the
Royal Botanic Garden, Edinburgh, and the Linnean Society of
London for permission to consult their collections, for facilities
provided, and for the loan of specimens. We are much indebted to
Mr A. O. Chater and Mr J. F. M. Cannon for many kindnesses,
useful discussions, correcting the English manuscript and preparation
for publication, and to Mr A. R. Vickery, Dr C. E. Jarvis, Miss C.
Whitefoord, Mr F. Nigel Hepper, Miss Sally Bidgood, Dr I. C.
Hedge, and Miss Kit Tan for hospitality, much advice and assistance.

We warmly thank the Swedish Institute for enabling T.N. Ho to
pursue our research on Gentiana in Sweden and we are grateful to Dr
R. Moberg and Dr B. Jonsell for arranging this visit.

Many people and organizations, too numerous to name individually
here, have helped during the time when we were struggling with the
Gentiana project. However, special thanks go to the American Rock
Garden Society, the Rhododendron Species Foundation of America,
the Herbarium of Arnold Arboretum of Harvard University, the
United States Herbarium, Department of Botany, Smithsonian In-
stitution, the Herbarium of the Missouri Botanical Garden, and
Denver Botanic Gardens for financial assistance or facilities provided,
and to Prof. Howard Mason, Mrs Judy Young, Dr Peter S. Ashton,
Dr Peter F. Stevens, Dr Peter H. Raven, and Dr Bruce Bartholomew
for hospitality and assistance.

Finally, we would like to thank our colleagues Mrs Cui-lan Van,
Mr Jin-jun Liu and Mr Xie-fen Lu for cooperation, illustrations, help
with the scanning electron microscopy of seeds, and assistance with
field work.

REFERENCES

Czerepanov, S. K. 1973. Additamenta et corrigenda ad 'Floram URSS' (tomi I-

XXX). Leningrad. [In Russian; Gentianaceae on pp. 288-292]
Froelich, J. A. 1796. De Gentiana libellus sistens specierum cognitarum

descriptionibus cum observationibus. Erlangen.
Gilg, E. 1895. Gentianaceae. In A. Engler & K. Prantl (Eds), Die naturlichen

Pflanzenfamilien 4(2): 50-62. Leipzig.
Gillett, J. M. 1957. A revision of the North American species of Gentianella

Moench. Ann. Mo. hot. Gdn 44(3): 195-269.
Grisebach, A. H. R. 1838 ['1839']. Genera et species Gentianearum ...

Stuttgart.
1845. Gentianaceae. In A. de Candolle (Ed.), Prodromus systematis

naturalis regni vegetabilis ... 9: 38-141. Paris.
Ho, T.-N. 1985. A study on the genus Gentiana of China IV. Bull. hot. Res.

Harbin 5(4): 1-22.
Ho, T.-N., Liu, S.-W. & Wu, C. J. 1988. Gentianaceae. Flora reipublicae

popularis sinicae 62. Beijing.
Holub, J. 1973. New names in Phanerogamae 2. Folia geobot. phytotax. 8:

155-179.

192

T.-N. HO&S.-W. LIU

Kusnezow, N. I. 1894. Subgenus Eugentiana of genus Gentiana Tourn. Trudy
imp. S.-Peterb. Obshch. Estest. 24(2): 1-531. [Latin translation in Trudy
imp. S. -Peterb. hot. Sada 15: 1-507 (1896-1904)]

1895. Gentiana Tourn. In A. Engler & K. Prantl (Eds), Die naturlichen
Pflanzenfamilien 4(2) : 80-86. Leipzig.

Linnaeus, C. 1737. Genera plantarum, ed. 1. Leiden.

1753. Species plantarum, ed. 1. Stockholm.

Love, A. & Love, D. 1975. The Spanish gentians. An. Inst. hot. A. J. Cavanilles

32(2): 221-232.
Ma, Y. C. 1951. Gentianopsis a new genus of Chinese Gentianaceae. Acta

phytotax. 1: 5-19, pis I-V.
Marquand, C. V. B. 1928. New Asiatic gentians. Bull. misc. Inf. R. hot. Gdns,

Kew for 1928: 49-62.

1931. New Asiatic gentians: II. Bull. misc. Inf. R. hot. Gdns, Kew for
1931: 68-88.

1932. The cultivated gentians of China and the Himalaya. // R. hort. Soc.
57: 188-211.

1937a. The gentians of China. Bull. misc. Inf. R. hot. Gdns, Kew for 1937:
134-180.

\937b. Contributions to the gentian flora of southern Tibet, N. E. Burma
and Bhutan. Bull. misc. Inf. R. hot. Gdns, Kew for 1937: 180-191.

Nilsson, S. 1967. Pollen morphological studies in the Gentianaceae-Gentianinae.
Grana paly not. 7: 46-143.

1976. Notes on pollen morphological variation in Gentianaceae-Gentianinae.

Pollen Spores 9: 49-58.
Pringle, J. S. 1967. Taxonomy of Gentiana, Section Pneumonanthae, in

eastern North America. Brittonia 19: 1-32.
1977. Taxonomy and distribution of Gentiana (Gentianaceae) in Mexico

and Central America. I. Sect. Pneumonanthe. Sida Contr. Bot. 7(2):

174-217.

1978. Sectional and subgeneric names in Gentiana (Gentianaceae). Sida
Contr. Bot. 7(3): 232-247.

1979. Taxonomy and distribution of Gentiana (Gentianaceae) in Mexico
and Central America. II. Sect. Chondrophyllae. Sida Contr. Bot. 8(1):
14-33.

Serebryakova, T. I. 1979. Models of shoot formation and some evolutionary

trends in the genus Gentiana L. Byull. Mask. Obshch. Ispyt. Prir., Otdel

Biol. 84(6): 97-109. [In Russian]
Smith, H. 1936. Gentianaceae. In H. Handel-Mazzetti (Ed.), Symbolae sinicae

7: 948-988. Wien.
1961. Problems relating to the Gentiana cachemirica of the 'Flora of

British India'. Kew Bull. 15(1): 43-55.

1965. Notes on Gentianaceae. Notes R. bot. Gdn Edinb. 26(2): 237-258.
Tournefort, J. P. 1700. Institutions rei herbariae. Editio altera, . . . Paris.
Tutin, T. G. 1972. Gentiana. In T. G. Tutin, V. H. Heywood et al. (Eds),

Flora Europaea 3: 59-63. Cambridge.

Bull. Br. Mas. nat. Hist. (Bot.) 20(2): 193-236 Issued 20 December 1990

Revision of Piper (Piperaceae) in the New

World

2. The taxonomy of Piper section Churumayu

MARGARET CECILIA TEBBS ,

* tv
Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD

CONTENTS

Introduction ...................................................................................... 194

Distribution and habitat ............................................................................ 194

Morphology ...................................................................................... 194

Habit .......................................................................................... 194

Leaves ......................................................................................... 195

Venation ....................................................................................... 195

Indumentum and pellucid glands ................................................................... 195

Inflorescences ................................................................................... 195

Bracts ......................................................................................... 195

Fruit ........................................................................................... 195

Section Churumayu (Miq.) C. DC .................................................................... 195

Key to the species in section Churumayu ............................................................ 196

Species descriptions ................................................................................ 197

21 . Piper guanacostense C. DC .................................................................... 197

22. P. trigonum C. DC ........................................................................... 197

23. P. rupununianum Trel. & Yuncker ............................................................. 199

24. P. atroglandulosum Tebbs ..................................................................... 200

25. P. divaricatum G. Meyer ...................................................................... 200

26. P. dumosum Rudge .......................................................................... 202

27. P. imberbeTre}. & Standley ................................................................... 202

28. P. xylosteoides (Kunth) Steudel ................................................................ 203

29. P. urophyllum C. DC ......................................................................... 204

30. P. subrepens Trel ............................................................................. 204

31 . P. mikanianum (Kunth) Steudel ............................................................... 206

32. P. carpunya Ruiz Lopez & Pavon .............................................................. 208

33. P. grande M. Vahl ........................................................................... 208

34. P. lepturum (Kunth) C. DC .................................................................... 209

35. P. aequale M. Vahl ........................................................................... 211

36. P. verruculosum C. DC ....................................................................... 212

37. P. corrugatum Kuntze ........................................................................ 213

38. P. schiedeanum Steudel ....................................................................... 213

39. P. cadenaensis Tebbs ......................................................................... 218

40. P. regnellii (Miq.) C. DC ...................................................................... 219

41. P. veraguense C. DC .......................................................................... 221

42. P. perbrevicaule Yuncker ..................................................................... 223

43. P. reptabundum C. DC ........................................................................ 223

44. P. fortunaensis Tebbs ......................................................................... 223

45. P. dolichotrichum Yuncker .................................................................... 225

46. P. subsessilifolium C. DC ...................................................................... 225

47. P. brachypodon (Benth.) C. DC ................................................................ 227

48. P. xanthostachyum C. DC ..................................................................... 227

49. P. ottoniifolium C. DC ........................................................................ 229

50. P. septuplinervium (Miq.) C. DC ............................................................... 231

51 . P. coruscans Kunth .......................................................................... 232

52. P. scutilimbum C. DC ......................................................................... 232

53. P. auritum Kunth .......................................................................... 234

References

.236

Index . , 237

194

M. C. TEBBS

SYNOPSIS. The paper presents a taxonomic treatment of Piper section Churumayu, and 33 species are recognized,
keyed, and described. There are two new species, P. cadenaensis Tebbs and P. fortunaensis Tebbs, and the new
name P. atroglandulosum Tebbs is proposed for P. obovatum Ruiz Lopez & Pavon .

INTRODUCTION

This second part of a revision of New World species of Piper
deals with section Churumayu (Miq.) C. DC.; see Tebbs
(1989a) for part 1 which deals with section Macrostachys.
Section Churumayu, as accepted here, is essentially Miquel's
(1844) Artanthe section Churumayu which he describes as
'Petioli nudi. Folia rotundato-vel ovato-oblonga, inf. aequila-
tera, saepe cordata, sup. basi inaequali rotundata, costis
versus basin approximatis, supra glabra, raro puberula'.
Although it is regrettable that he defines his section by leaf
characters alone, the types of inflorescences, bracts, and
fruits in at least half of his species conform with those of the
taxa covered in the present treatment. The species accepted
in section Churumayu have slender, erect, arching or pendu-
lous inflorescences, fleshy orbicular to triangular bracts, and
small, trigonous, obovoid fruits, sometimes with a minute
style. Prophylls are usually prominent and often several
centimetres long. In this treatment, 33 species have been
accepted in section Churumayu, and 128 names have been
reduced to synonymy. The species are differentiated using
inflorescence characters supported by vegetative characters,
habitat, and general distribution. To this end many speci-
mens, including type material, have been examined, and the
full extent of the variation in the section has become
apparent. Many of the problems encountered in attempts to
define the species limits in Piper are due to the wide and
frequently continuous range of variation in characters such as
leaf size and pubescence density. Narrow taxonomic concepts
that give undue weight to these characters, such as those
adopted by Trelease (1927, 1929), and to a lesser extent by de
Candolle (1869), only serve to create many new and unneces-
sary taxa, adding to the general confusion within the genus.
As in the first part of this treatment of Piper (Tebbs, 1989a), I
have attempted to minimise and clarify this confusion by
adopting a broad species concept. This is reflected in the
extensive synonymies included under the species accepted
here.

I question the taxonomic value of varieties based on
vegetative characters in a genus that is phenotypically plastic.
Trelease & Yuncker (1950) described many new varieties in
their treatment of Piper. For example, P. ottoniifolium C.
DC. was separated by them into var. frutescens Trel. &
Yuncker, a shrub with leaves 12-15 cm long and 5.5-6 cm
wide, and var. latifolium Trel. & Yuncker, a scandent shrub
with leaves 15-22 cm long and 6-9.5 cm wide. So many
intermediate forms can be found in the characters used to
distinguish these varieties that it becomes pointless to attempt
to separate them. Clearly, the characters popularly employed
in the past to segregate varieties, such as slight differences in
habit, leaf shape, or the presence or absence of indumentum,
overlap too much to be used with any certainty, and no
varieties are recognized in this revision.

DISTRIBUTION AND HABITAT

Piper species are usually plants of moist shady ground in
disturbed places, and are often rapid colonisers of road- and
trail-sides, edges of pasture, and forest clearings. Most
species in section Churumayu follow this general pattern, and
can be found at altitudes ranging from sea-level to more than
2000 m. For example, Piper guanacostense C. DC. is confined
to the drier Pacific slopes of Costa Rica, growing from
0-1000 m, in fairly close proximity to the coast in shady moist
soil at forest edges, while the widespread P. schiedeanum
Steudel ranges from Mexico to Brazil, growing in a variety of
disturbed places from 0-1600 m. Piper aequale M. Vahl, P.
dolichotrichum Yuncker, P. grande M. Vahl, P. ottoniifolium,
P. subsessilifolium C. DC., P. veraguense C. DC., and P.
xanthostachyum C. DC. typically occur in moist forest or
damp shady places of remnant forest in areas cleared for
pasture, from sea-level to over 2000 m. Piper brachypodon
(Benth.) C. DC. is confined to lowland areas of Narino
Province in Colombia (including Isla Gorgona), where it
grows in dense moist forest. Piper septuplinervium (Miq.) C.
DC. is a plant of brackish places, often growing in mangrove
swamps and on riverbanks near the coasts of Colombia and
Ecuador, while P. coruscans Kunth occurs on high, wet, steep
slopes and cliff bases, from Colombia to Brazil. Piper verru-
culosum C. DC. is only found in Costa Rica, growing in the
shade of cloud forests between 800-1800 m. Piper scutilim-
bum C. DC. also occurs in cloud forest, but is more widely
distributed, from Panama to Ecuador. Two species, P.
mikanianum (Kunth) Steudel and P. regnellii (Miq.) C. DC.,
have a more southerly distribution than the others in this
section and are found at 400-1300 m altitude, from Brazil to
Argentina. Piper mikanianum grows in gallery forest, while
P. regnellii can be found on steep wooded slopes, boggy
ground, and streamsides. Piper auritum Kunth is a successful
coloniser of tracksides, clearings, riverbanks, and ground
disturbed by landslides, and ranges from Mexico to the West
Indies and Colombia, while P. subrepens Trel. is a rare
species only found on mountain slopes between 700 and 800
m, in Code Province, Panama.

MORPHOLOGY

Habit

The majority of species in section Churumayu are shrubs or
slender trees, with erect to spreading branches. Piper auritum
forms a thin-trunked tree some 6 m in height, while P.
verruculosum is a densely branching shrub 1-3 m high. Piper
mikanianum is a spreading or decumbent shrub with branches

PIPER IN THE NEW WORLD

that root at the lower nodes, as do those of P. urophyllum C.
DC. Five species, P. dolichotrichum, P. ottoniifolium, P.
reptabundum C. DC., P. subsessilifolium and P. xantho-
stachyum, are true climbers or scramblers, with copious
adventitious roots at the nodes that attach the stems to a
support (Tebbs, \9S9b). One species, P. perbrevicaule
Yuncker, is a terrestrial, sometimes epiphytic, trailing herb
often rooting at the nodes.

Leaves

As in the rest of the genus, leaves of the species belonging to
section Churumayu are very variable in shape and size. Piper
guanacostense and P. trigonum C. DC. have elliptic-oblong to
lanceolate leaves, P. urophyllum has elliptic leaves that
narrow abruptly to a long slender apex, while those of P.
auritum are asymmetrically oblong-cordate and may reach 55
cm in length. Piper schiedeanum is an example of a species in
this section displaying a confusing variety of leaf shapes, from
ovate-elliptic or lanceolate to deeply cordiform. The leaves
often vary on the same plant, and older leaves may be wider
and have more prominent basal lobing than juvenile foliage.
This variation in leaf shape has resulted in many superfluous
taxa being described. Gomez Pompa (1966) has examined the
variable shapes of the leaves and the number of secondary
veins in P. schiedeanum, and on this basis has sunk several
taxa into synonymy. Two species, P. scutilimbum and P.
veraguense, have distinctive peltate leaves.

There is a certain amount of variation between species in
thickness and texture of the leaves. Piper auritum has mem-
branous leaves, while those of P. carpunya Ruiz Lopez &
Pavon and P. ottoniifolium are coriaceous. Piper corrugatum
Kuntze and P. perbrevicaule often have bullate or rugose leaf
surfaces.

The petioles range from a few millimetres to 5 cm long, and
may be vaginate at the base, e.g. P. corrugatum, or partially
or wholly sheathing, e.g. P. auritum, P. regnellii.

Venation

Most species in this section have 1-5 pairs of secondary veins
arising from, or near, the leaf base, and 1-3 pairs of veins
arising from the middle to upper part of the midrib. The
number of basal veins is related to the shape and extent of
lobing of the leaf base. Several species, e.g. P. guanacostense
and P. trigonum, have leaves with secondary veins arising
along the length of the midrib, indicating a phylogenetic
relationship with P. arboreum Aublet, of section
Macrostachys. The venation of P. ottoniifolium is distinctly
camptodromous, with secondary veins curving strongly from,
or near, the base to the apex.

Indumentum and pellucid glands

The hairs found in Piper are simple, either unicellular or
multicellular. Several species, e.g. P. dolichotrichum and P.
perbrevicaule, have stems and leaves that are densely covered
with multicellular hairs 1-3 mm long. Piper regnellii and P.
subsessilifolium are often minutely pubescent on young
growth and undersides of leaves. The leaf margins of P.
auritum are fringed with minute pale hairs, a character that is
also found in P. marginatum Jacq.

Prominent pellucid glands are quite common in Piper, and
may be pale yellow, orange, red, brown, or black. The leaves

195

of P. trigonum have both hairs and conspicuous pellucid
glands. Piper atroglandulosum Tebbs has copious black
glands on the stems and leaves. Piper ottoniifolium has pale
glands on the floral bracts and the fruits.

Inflorescences

The inflorescences are mostly erect in section Churumayu,
e.g. Piper aequale and P. schiedeanum. P. auritum has erect
or slightly arching spikes, another character that indicates its
relationship with P. marginatum. There are several species in
which the inflorescences become pendulous with maturity.
This is mainly because they are borne on long, thin peduncles,
which bend under the weight of the developing fruit (e.g.
those of P. dolichotrichum and P. reptabundum). The floral
parts are closely congested in this section, and difficult to
segregate. Burger (1972) indicates that the development of
slender congested inflorescences can be associated with the
activities of small bees, which collect pollen from the surface
of the spike.

There is a limited amount of colour variation between
species in section Churumayu. P. subsessilifolium has pink to
maroon inflorescences, while those of P. xanthostachyum
are yellow-white. Generally, however, most of the species
treated here have greenish-yellow to dull white spikes.

Bracts

The floral bracts found in this section are triangular to round,
often fleshy and glabrous, glandular or pubescent. Several
species have bracts that partially cover the fruit, at least in
early stages of development, e.g. P. schiedeanum and P.
veraguense. The distinctive bracts of P. grande are glabrous,
umbonate, and completely cover the developing fruits.

Fruit

The fruits in this section are trigonous and obovoid with
angular sides, on average about 1 mm across, with occasion-
ally a domed apex with a minute, scarcely noticeable style.
The exception is P. coruscans with fruits that become orbicu-
lar as they mature. There are 2-4 slender stigmas. Fruits may
be glabrous, e.g. P. aequale, minutely pubescent, e.g. P.
verruculosum, or with prominent pellucid glands, e.g. P.
brachypodon. A few species have a mealy or granular sur-
face, e.g. P. trigonum. Species with small trigonous fruits can
also be found in section Radula, e.g. P. dilatatum C. DC. and
P. lanceifolium Kunth. However, these species also display
other characters that are diagnostic of that section, such as
dark, often scabrous leaves, dense indumentum, and stipule-
like developments on the bases of the petioles, and on that
basis are excluded from section Churumayu.

Section CHURUMAYU (Miq.) C. DC.
Type: Piper churumayu Poeppig

Herbs, shrubs, and trees, scramblers or climbers, rarely
epiphytic, glabrous, minutely to densely pubescent or glandu-
lar. Leaves ovate, ovate-elliptic to lanceolate, or deeply
asymmetrically oblong-cordate to regularly cordiform, occa-
sionally peltate, glabrous, or minutely to densely pubescent
or glandular with yellow, orange, brown, or black glands,
glossy above, sometimes glaucous or coriaceous. Venation

196

M. C. TEBBS

with 1-8 pairs of secondary veins arising from or near base,
and 1-3 pairs from middle to upper part of midrib. Petioles
vaginate at the base, or partially to completely sheathing the
stem. Prophylls prominent, narrow to wide, acute or blunt-
tipped, glabrous, glandular, or pubescent. Infloresences
erect, sometimes arching, or pendulous on long, slender
peduncles; flowers densely crowded on rachis. Floral bracts
orbicular, crescent-shaped, triangular or T-shaped, some-
times umbonate, fleshy, glabrous, minutely ciliate or with
dense margins of hairs, sometimes partially or completely
covering developing fruits. Anthers minute, dehiscing
laterally or upward. Fruits obovoid, trigonous, rarely subor-
bicular, glabrous, glandular, granular, or minutely puberu-
lent, occasionally with domed apex and minute style; stigmas
2-4.

Key to the species in section Churumayu

la. Climbers or scramblers:

2a. Inflorescences on long slender peduncles, pendulous in fruit:
3a. Leaves asymmetrically lanceolate to oblong-elliptic, glabrous

43. P. reptabundum

3b. Leaves symmetrical, broadly ovate to elliptic, densely long-
pubescent 45. P. dolichotrichum

2b. Inflorescences not on long slender peduncles, erect:

4a. Leaves coriaceous, with 2-3 pairs of prominent secondary

veins arising from base and curving to apex

49. P. ottoniifolium

4b. Leaves membranous, with 3-5 pairs of secondary veins
arising from near the base to middle part of midrib, ascend-
ing rather sharply to apex:
5a. Inflorescences pink or maroon; bracts 0.6-1 mm wide,

pink-to purple-ciliate 46. P. subsessilifolium

5b. Inflorescences yellow-white; bracts 0.3-0.5 mm wide,
yellow-white-ciliate 48. P. xanthostachyum

Ib. Herbs, shrubs or trees:
6a. Herbs, erect or prostrate:

7a. Stems erect, 1-2 m high, glabrous; leaves peltate; inflore-
scences 7-16 cm long, on short peduncles

41 . P. veraguense

7b. Stems prostrate or trailing, 15-50 cm long, pubescent; leaves
not peltate; inflorescences 2-4 cm long, on long slender
peduncles 42. P. perbrevicaule

6b. Shrubs or trees:

8a. Petioles sheathing the stems:

9a. Leaves peltate, base cuneate or caudate

52. P. scutilimbum

9b. Leaves not peltate, base shallowly to deeply cordate:
lOa. Leaves deeply unevenly cordate with one basal lobe
conspicuously lower than other, margins densely cili-
ate; inflorescences arching 53. P. auritum

lOb. Leaves widely shallowly cordate, margins glabrous;

inflorescences erect 40. P. regnellii

8b. Petioles not sheathing the stems:

1 la. Floral bracts partially or completely concealing fruits:
12a. Floral bracts umbonate, completely concealing fruits

33. P. grande

12b. Floral bracts not umbonate, only partially concealing

fruits:

13a. Leaves symmetrically ovate to broadly, evenly
cordiform, glabrous; floral bracts 0.4-0.7 mm,
orbicular-triangular, fleshy, glabrous or minutely

ciliate 38. P. schiedeanum

13b. Leaves asymmetrically oblong-cordate to sub-
cordate, glabrous or with long multicellular hairs,

sometimes bullate; floral bracts 0.2-0.5 mm, oblong-
triangular, upper part slightly protruding, minutely
pubescent below 37. P. corrugatum

lib. Floral bracts neither completely nor partially concealing

fruits:

14a. Fruits shortly pubescent or minutely granular:
15a. Leaves asymmetrically narrowly to broadly elliptic;
floral bracts oblong-triangular with gibbous bases . .

44. P. fortunaensis

15b. Leaves symmetrically ovate- or oblong-elliptic;

floral bracts triangular, glabrous to densely ciliate:
16a. Stems with tubercles; leaves glabrous on both

surfaces, eglandular 32. P. carpunya

16b. Stems smooth, without tubercles; leaves puberu-
lent on undersurfaces, with prominent orange or
brown glands:
17a. Leaves with 3-4 pairs of secondary veins;

peduncles 15-30 mm long

23. P. rupununianum

17b. Leaves with 4-7 pairs of secondary veins;
peduncles 6-15 mm long 22. P. trigonum

14b. Fruits glabrous:

18a. Fruits with pellucid glands visible on upper surface:
19a. Stems and leaves red-streaked, eglandular; veins

prominent 47. P. brachypodon

19b. Stems and leaves with copious glands; veins faint:
20a. Leaves oblanceolate . . .24. P. atroglandulosum
20b. Leaves ovate-lanceolate to oblong-elliptic:
21 a. Leaves glabrous:
22a. Leaves membranous, apex long-acuminate

34. P. lepturum

22b. Leaves subcoriaceous, apex acute, blunt

25. P. divaricatum

21b. Leaves sparsely to moderately pubescent
26. P. dumosum

18b. Fruits without pellucid glands visible on upper
surface:

23a. Leaves lanceolate to narrowly elliptic:

24a. Fruits glabrous 35. P. aequale

24b. Fruits pubescent:
25a. Leaves with orange or brown glands

36. P. verruculosum

25b. Leaves eglandular . 39. P. cadenaensis

23b. Leaves ovate, sub-orbiculate, ovate-
elliptic, base obtuse to cordate or
subcordate:

26a. Leaves sub-orbiculate; fruits sub-
orbicular 51 . P. coruscans

26b. Leaves not sub-orbiculate; fruits

trigonous:

27a. Subshrubs, spreading or
decumbent, 0.5-1 m high:

28a. Leaves peltate

27. P. imberbe

28b. Leaves not peltate

31. P. mikanianum

27b. Shrubs, erect, at least 1 m high:

29a. Floral bracts T-shaped ...

50. P. septuplinervium

29b. Floral bracts not T-shaped:

30a. Leaf bases thickened,

forming dark 'lips'; fruits

1-2 mm wide

29. P. urophyllum

PIPER IN THE NEW WORLD

197

30b. Leaf bases not thick-
ened; fruits 0.5-1 mm
wide:

3 la. Leaves with scattered
glands; floral bracts

glabrous

... 28. P. xylosteoides

31b. Leaves eglandular;

floral bracts minutely

ciliate:

32a. Inflorescences 2-3

cm long

. . 30. P. subrepens
32b. Inflorescences 3-9

cm long

21 . P. guanacostense

SPECIES DESCRIPTIONS

21. Piper guanacostense C. DC. in Linnaea 37: 356 (1872).

Type: Costa Rica, Oersted 862 (C!-holotype).
Fig. 2A, a, b.

Shrubs or slender trees, 1-3 m high, glabrous or minutely
puberulent. Leaves 10-14 (-18) cm long, 3-5 cm wide,
lanceolate, glossy, apex acuminate, base acute to obtuse,
glabrous. Venation with 6-12 pairs of secondary veins arising
along the length of the midrib, loop-connecting to the apex.
Petioles 3-10 mm long. Prophylls 4-12 mm long, glabrous.
Inflorescences 3-9 cm long; peduncles 6-12 mm long. Anthers
0.2-0.3 mm long. Floral bracts 0.5-0.7 mm wide, triangular,
ciliate. Fruits obovoid, 1 mm wide, trigonous to orbicular,
glabrous; stigmas sessile.

Growing in shade in moist places or along streams; 0-1000 m.
DISTRIBUTION. Costa Rica. Fig. 1.

Costa Rica, Guanacaste: Liesner 5013 (MO); Puntarenas:
Burger & Ramirez 4044 (F, BM).

Piper guanacostense is restricted to the Pacific side of Costa
Rica and grows in the shade of moist forest on hills fairly close

Fig. 1 Distribution of P. guanacostense.

to the coast. It has similar leaf shape and venation to that
of Piper arboreum Aublet, but does not have that species'
sheathing petioles, and the inflorescence is erect rather
than pendulous. Unlike P. arboreum it has a well-developed
prophyll.

Piper guanacostense, along with P. atroglandulosum and
P. trigonum, can be distinguished from other species in
this section by its distinctly pinnate venation loop-connecting
to the apex, lacking the prominent pairs of veins arising
from the base of the leaf. The glabrous fruits distinguish it
from P. trigonum, which has glandular, pubescent, or granular
fruits.

22. Piper trigonum C. DC. in /. Bot., Lond. 4: 212 (1866).

Type: Colombia, Prov. Barbacoas, Triana 44 (G-holotype;

BM!-isotype).
Fig. 2B, c, d.

P. schlimii C. DC., in DC., Prodr. 16(1): 274 (1869). Type:

Venezuela, Funcke & Schlim 591 (G-holotype;

BM!-isotype).
P. arieianum C. DC. in An. Inst. fis.-geogr. C. Rica 9: 166

(1897). Type: Costa Rica, Bords de 1'Ariei, Talamanca,

Pittier9381 (G-holotype; MO!-photograph).
P. saltuum C. DC. in Urban Symb. antill. 7: 185 (1912).

Type: Trinidad, Cuparo, Broadway 3207 (G-holotype).
P. acutissimum Trel. in Contr. U.S. natn. Herb. 26: 25 (1927).

Type: Panama, Chagres, prov. Colon, Fendler 266 (US-

holotype; K!, MO!-isotypes).
P. trichopus Trel. in Contr. U.S. natn. Herb. 26: 25 (1927).

Type: Panama, Sutton Hayes 797 (NY!-holotype).
P. altevaginans Trel. in Contr. U.S. natn. Herb. 26: 140

(1929). Type: Costa Rica, Guapiles, Limon, Standley 37316

(US-holotype).
P. generalense Trel. in Publs Field Mus. not. Hist. (Bot.) 18:

344 (1937). Type: Costa Rica, vicinity of El General, prov.

San Jose, Skutch 2846 (US-holotype; NYl-isotype).
P. marginatibaccum Trel. ex Standley in Publs Field Mus.

not. Hist. (Bot.) 18: 349 (1937). Type: Costa Rica, vie. of

El General, San Jose, Skutch 2183 (US-holotype; MO!,

NY!-isotypes).
P. seducentifolium Trel. in Ann. Mo. hot. Gdn 27: 297 (1940).

Type: Panama, Woodson et al. 781 (MO!-holotype).
P. acutissimum var. trichopus (Trel.) Yuncker in Ann. Mo.

hot. Gdn 37: 66 (1950). Type as for P. trichopus.
P. gorgonae-insulae Trel. & Yuncker, Piperac. N. South

Amer. 1: 331 (1950). Type: Colombia, south end of Gorgona

Island, Narina, Killip & Garcia 33095 (US-holotype;

BM!-isotype).
P. portachueloense Trel. & Yuncker, Piperac. N. South

Amer. 1: 327 (1950). Type: Venezuela, El Portachuelo,

Aragua, Pittier 11815 (US- holotype; NY!-isotype).
P. riitosenseTrel. & Yuncker, Piperac. N. South Amer. 1: 396

(1950). Type: Venezuela, Aragua, Killip & Lasser 37755

(US-holotype, photograph!).

Shrub 0.5-1 (-2) m high, stems minutely pubescent, especi-
ally on younger growth. Leaves 10-20 cm long, 3-6.5 cm
wide, elliptic-oblong to lanceolate, apex acuminate, base
acute to obtuse, glabrous above, puberulent on veins below,
with conspicuous brown or black pellucid glands. Venation
with 4-7 pairs of secondary veins arising along the length of
the midrib, loop-connecting to apex. Petioles 3-15 mm
long, puberulent. Prophylls 6-15 (-20) mm long, glabrous
to minutely pubescent. Inflorescences 4-8 (-11) cm long,

198

M. C. TEBBS

Fig. 2 A: P. guanacostense, habit; a: prophyll; b: bract and fruit. B: P. trigonum, habit; c: prophyll; d: bract and fruit. C: P. rupununianum,

habit; e: prophyll; f: bract and fruit; g: leaf section.

PIPER IN THE NEW WORLD

199

r

Fig. 3 Distribution of P. trigonum.

becoming pendulous in fruit; peduncles 6-15 mm long,
puberulent. Anthers 0.2-0.3 mm long. Floral bracts 0.5-
0.8 mm wide, triangular, sparsely pubescent. Fruit obovoid,
1-1.5 (-2) mm wide, trigonous, shortly pubescent or
granular, glandular on upper surface; stigmas 3, in small
depression.

Growing in shade at forest edges, trail sides, and along banks
of streams; 0-1200 m.

DISTRIBUTION Mexico-Brazil. Fig. 3.

Mexico, Oaxaca: Chavelas & Perez 294 (MEXU); Chiapas:
Gilly & Xolocotzi 189 (MEXU). Guatemala, Quetzaltenango:
Croat & Hannon 63423 (BM, MO). Honduras, Morazan:
Martinez 1 (BM). Nicaragua, Zelaya: Vincelli 157 (MO).
Costa Rica, Alajuela: Liesner 14115 (MO); Heredia: Opler
92 (F); Limon: Burger & Antonio 11228 (F); Puntarenas:
Liesner 1901 (MO). Panama, Bocas del Toro: Kirkbride &
Duke 773 (NY); Code: Hammel4191 (MO); Colon: Kennedy
2787 (MO); Darien: Croat 37710 (MO); Panama: Croat 17308
(MO); Perlas, San Jose Is.: Johnston 1215 (BM); San Bias:
Duke 14875 (MO); Veraguas: Nee 11269 (MO). West Indies,
Trinidad: Broadway 6206 (BM). Colombia, Santa Marta:
Smith 2736 (K, NY); Santander: Killip & Smith 14875 (BM,
US). Venezuela, Aragua: Croat 54482 (BM,MO). French
Guiana, Cayenne: de Granville 6561 (K). Peru, Loreto: Jones
7175 (ILL). Brazil, Amapa, Rio Oiapoque: Irwin et al. 47222
(K).

Piper trigonum occurs in the deep shade of swamp forest,
lowland rain-forest, and secondary vegetation. Its leaf shape,
venation, and the presence of pellucid glands on the stems,
leaves, and fruits show an affinity with P. atroglandulosum.
Piper trigonum can be distinguished by its pubescent young
stems, sparsely pubescent floral bracts, and widely triangular,
glandular, pubescent, or granular fruits. Specimens from

Trinidad are generally less pubescent than those on the
mainland, and the fruits have a less pronounced granular
surface.

23. Piper rupununianum Trel. & Yuncker, Piper ac. N. South
Amer. 1: 303 (1950). Type: Guyana, Tumatumari, Gleason
107 (NY-holotype; K!-isotype).

Fig. 2C, e, f, g.

Spreading shrub or subshrub, 1-2 m high, stems glandular.
Leaves 14-22 cm long, 4-8 cm wide, oblong-elliptic to lanceo-
late, glabrous, glossy above, puberulent on veins beneath,
with orange or brown glands on both surfaces, apex acute-
acuminate, base acute to obtuse. Venation with 3-4 pairs
of secondary veins arising from the lower to middle part
of midrib, loop-connecting to apex. Petioles 5-10 mm long.
Prophylls 5-15 mm long, glabrous, glandular. Inflorescences
8-11 cm long, erect, sometimes pendulous in fruit;
peduncles 15-30 mm long, slender. Anthers 0.2-0.3 mm long.
Floral bracts 0.8-1 mm wide, triangular, densely yellow-
ciliate, with dark, glabrous centre. Fruit obovoid, 1-1.5 mm
wide, trigonous, glandular, slightly granular; stigmas 3,
sessile.

Clearings, damp places, secondary mixed forest, and shady
rocky areas; 0-400 m.

DISTRIBUTION. Guyana. Fig. 4.

Guyana, Aremoo Creek: Bartlett 8408 (K); Oko Creek: Tutin
318 (BM); Mabaruma: Forest Dept. F2509 (K); Potars district:
Whitton 102 (K).

This small, spreading shrub is apparently confined to Guyana.
It can be recognized by its large glossy leaves with orange or
brown pellucid glands, and inflorescences with long peduncles.
The floral bracts are similar to those of P. atroglandulosum or

200

Fig. 4 Distribution of P. rupununianum.

P. divaricatum, with yellow-ciliate margins and dark centres.
The glandular, granular fruits are similar to those of P.
trigonum.

24. Piper atroglandulosum Tebbs, nom. nov.
Fig. 5A, a, b, c.

P. obovatwn Ruiz Lopez & Pavon, Fl. peruv. 1: 36, tab. 53
(1798), non M. Vahl, Eclog. amer. 1: 5 (1796) (Art. 64.1).
Type: Peru, Huanuco, Cochero & Chinchao, Ruiz Lopez
& Pavon s.n. (B-holotype; Mi-photograph).

Schilleria obovata Kunth in Linnaea 13: 686 (1839).

Artanthe obovata (Kunth) Miq., Syst. piperac.: 508 (1844).

Shrub 1-2 m high, stems glabrous, rarely sparsely pubescent,
copiously black- or brown-glandular. Leaves 9-14 cm long, 3-
6 cm wide, oblanceolate to elliptic, both surfaces with dark
glands, upper surface glossy, apex narrowly long- acuminate,
base cuneate. Venation with 5-9 pairs of veins arising along
the length of the midrib and curving towards apex. Petioles 5-
12 (-15) mm long. Prophylls 1-2.5 cm long, glandular.
Inflorescences 3-8 cm long, 4-5 mm wide, curving in fruit;
peduncle 6-10 mm long. Anthers 0.2-0.4 mm long. Floral
bracts 1 mm wide, triangular, densely pale yellowish-ciliate.
Fruit obovoid, 1-2 mm wide, trigonous, glabrous, glandular;
stigmas 3.

Edges of forest; 0-800 m.
DISTRIBUTION. Colombia-Bolivia. Fig. 6.

Colombia, El Meta: Haught 2759 (BM, US). Peru, Casapi:
Mathews 1692 (K). Ecuador, Napo-Pastaza: Mexia 7230 (BM,
US). Bolivia, Tumupasa: Williams 465 (K).

This species is closely related to P. divaricatum, with similar
leaf shape, type of venation, and prominent glands. It differs
from that species mainly by its obovate leaves with long
narrow apices, dark glands, much smaller prophyll and west-
ern Andean distribution. It is also similar to P. heterophyllum
Ruiz Lopez & Pavon, but the latter has a short style and

M. C. TEBBS

round fruit. However, these species can be difficult to segre-
gate in dried herbarium specimens without mature
inflorescences.

Piper obovatum Ruiz Lopez & Pavon is a later homonym of
Piper obovatum M. Vahl, published two years earlier in 1796,
and is hence illegitimate. Piper obovatum M. Vahl, however,
was placed in the genus Peperomia by A. Dietrich (1831) in
the synonymy of Peperomia obversa (M. Vahl) A. Dietr.
(basionym Piper obversum M. Vahl, Enum. pi. 1: 354
(1804)). Casimir de Candolle (1902) correctly placed
Peperomia obversa into synonymy under Peperomia obovata
(M. Vahl) C. DC. as the earliest published name, leaving the
Ruiz Lopez & Pavon species without a legitimate name.
Unusually for this genus, no later name has been found that
could be adopted as a name for Piper obovatum Ruiz Lopez
& Pavon. It was thought at first that it should be placed in P.
divaricatum, but its distinctive leaf shape, western distribu-
tion, and different habitat have prevented this; therefore, it
has been necessary to provide a new name for this species.

25. Piper divaricatum G. Meyer, Prim. fl. esseq.: 15 (1818).
Type: Surinam, Weigelt s.n. (W-holotype; SP!-photograph).
Fig. 5B, d, e, f.

P. nitidum M. Vahl, Enum. pi. 1: 314 (1804), non Swartz

(1788) (Art. 64.1). Type: Vahl s.n. (C-holotype).
P. colubrinum Link, Jb. Gewachsk. 1 (3): 61 (1820), nom.

nud. (Art. 32.1).
P. impunctatum Link, Jb. Gewachsk. 1 (3): 62 (1820). Type:

Brazil, Willdenow 645 (B-holotype).
P. crassum Veil. Cone., Fl. flumin.: 25, t.59 (1825). Type:

Brazil, Vellozo s.n. (R-holotype).
Schilleria colubrina Link ex Kunth in Linnaea 13: 685 (1839).

Type: Brazil meridionali, Gaudichaud 86 (G-holotype).
Piper densispicatum Steudel, Nomencl. hot. ed. 2, 2: 340

(1841), nom. nud. (Art. 32.1).
Peltobryon attenuatum Klotzsch ex Miq., Syst. piperac.: 371

(1844). Type: Brazil, Martins 1209 (K!-isosyntype).
Artanthe colubrina (Link ex Kunth) Miq., Syst. piperac.: 512

(1844).
A. glabella Miq., Syst. piperac.: 371 (1844). Type: Cayenne,

Poiteau s.n. (G-holotype).
A. tejucensis Miq., Syst. piperac.: 496 (1844). Type: Brazil ad

Tejuco, Vauthier4Sl (G-holotype).
A. adenophylla Miq. in Linnaea 20: 170 (1847). Type: Brazil,

Coralfalga, Pohl 5026 (W-holotype).
A. colubrina var. crassifolia Miq. in Linnaea 20: 170 (1847).

Type: Brazil, Pohl 53 (W-holotype).
A. subaequalis C. Presl, Epimel. hot.: 226 (1849). Type:

Brazil, Rio de Janeiro, Mikan s.n. (W-holotype).
Piper apiculatum C. DC. in DC., Prodr. 16(1): 273 (1869).

Type: Brazil, Riedelll (K!-isotype).
P. crassifolium (Miq.) C. DC. in DC., Prodr. 16(1): 301

(1869).
P. glabellum (Miq.) C. DC. in DC., Prodr. 16(1): 312 (1869),

non Swartz (1788) (Art. 64.1).
P. leiophyllum C. DC. in DC., Prodr. 16(1): 297 (1869).

Type: Brazil, Bahia, Blanchet3\52 (W-holotype).
P. subaequale (C. Presl) C. DC. in DC., Prodr. 16(1): 371

(1869).

P. tejucense (Miq.) C. DC. in DC., Prodr. 16(1): 287 (1869).
P. glabrilimbum C. DC. in Verh. hot. Ver. Brandenb. 47: 107

(1905). Type: Brazil, Amazonas, Ule 47b (B-holotype).
P. belemense C. DC. in Notizbl. hot. Gart. Mus. Berl. 6: 458

(1917). Type: Brazil, Belem, Huber 145 (G-holotype).

PIPER IN THE NEW WORLD

201

Fig. 5 A: P. atroglandulosum, habit; a: prophyll; b: bract and fruit; c: leaf section. B: P. divaricatum, habit; d: prophyll; e: bract and fruit; f:

leaf section. C: P. dumosum, habit; g: prophyll; h: bract and fruit.

202

Fig. 6

vX

Distribution of P. atroglandulosum.

P. guedesii C. DC. in Notizbl. hot. Gart. Mus. Berl. 6: 436

(1917). Type: Brazil, Ilha mexicana, Guedes s.n. (G-

holotype).
P. oblongilimbum C. DC. in Notizbl. hot. Gart. Mus. Berl. 6:

448 (1917). Type: Brazil, Alto Purus, Huber 4418 (G-

holotype).
P. insipiens Trel. & Yuncker, Piperac. N. South Amer. 1: 224

(1950). Type: Cayenne, Poiteau s.n. (G-holotype).

Shrub 1-3 m high, sometimes straggling, stems glabrous, with
yellow, red, or brown glands. Leaves 13-18 cm long, 4-7 cm
wide, ovate-lanceolate to ovate-elliptic, subcoriaceous, glossy
above, glabrous, both surfaces covered with yellow or reddish
glands, apex acute, blunt, base ovate to sub-obtuse. Venation
with 4-6 pairs of secondary veins arising mostly from the
lower to middle part of midrib, loop-connecting to apex.
Petioles 10-15 mm long. Prophylls 1-3 cm long, glabrous,
glandular. Inflorescences 5-15 cm long, 5-8 mm wide when
mature, curving or pendulous in fruit; peduncles 6-12 mm
long. Anthers 0.3-0.4 mm long. Floral bracts 0.8-1 mm wide,
triangular, fleshy, broadly and densely yellow-white-ciliate.
Fruit obovoid, 1-2 mm wide, trigonous, glabrous, glandular;
stigmas 3, sessile.

Forest, wet soil along river banks, and in riverine woodlands;
0-300 m.

DISTRIBUTION. Guyana-Brazil. Fig. 7.

Surinam, Hostmann 709 (BM). Guyana, Demerara River:
Jenman 4206 (NY); Parika: Davis 1967 (K). Brazil, Amapa:

M. C. TEBBS

Egler & Irwin 46024A (K); Amazonas: Prance et al. 16325
(K); Bahia: Glocker 1 (BM); Goias: Prance & Silva 59663
(K); Para: Baker 413 (E); Rondonia: Cid et al. 4554 (K);
Roraima: Prance et al. 11103 (K).

Trelease & Yuncker (1950) placed Piper nitidum M. Vahl
into synonymy under P. divaricatum, without citing type
material, although they may have seen it. Yuncker (1972)
reduced 16 species of Piper to synonymy under P. divari-
catum, and noted the 'considerable variation in the size,
shape and texture of the leaves, and, to some extent, also in
the nervation.' Piper divaricatum can be recognized by its
ovate-lanceolate to ovate-elliptic, glabrous leaves, both sur-
faces of which are copiously glandular. It is also found in
wetter conditions, most frequently along river banks, and its
easterly distribution separates it further from P. atroglandu-
losum, which is mostly confined to the western side of the
Andes.

26. Piper dumosum Rudge, PI. Guian. 1: 13 (1805). Type:
Guiana, Martin s.n. (BM!-holotype).

Fig. 5C, g, h.

Shrub or treelet, 1-3 (^4)m high, stems glabrous or with
scattered pale hairs 1-1.5 mm long, glandular. Leaves 15-23
cm long, (4.5-) 7-11 cm wide, narrowly to broadly elliptic or
oblong-elliptic, upper surface sparsely pubescent, lower sur-
face sparsely to moderately pubescent, copiously red- or
black-glandular, apex acute-acuminate, base obtuse or sub-
obtuse. Venation with 5-9 pairs of secondary veins arising
from along most of the length of the midrib, curving towards
apex. Petioles 0.5-1.5 (-2) cm long, glabrous or sparsely
pubescent, vaginate at or near base. Prophylls 1-2 cm long,
pale, leaf-like, covered in brown or black glands. Inflores-
cences 5-10 cm long, 3-5 mm wide when mature, yellowish
white or brown, usually pendulous; peduncles 5-12 mm long,
with black glands. Anthers 0.3-0.4 mm long. Floral bracts
0.5-0.8 mm wide, triangular, shortly yellowish-ciliate. Fruit
obovoid, 1-2 mm wide, trigonous, glabrous, glandular; stig-
mas 3, sessile.

Shady moist places in primary and secondary forest and along
track sides; 0-600 m.

DISTRIBUTION. French Guiana to Brazil. Fig. 8.

French Guiana, Poiteau s.n. (K). Guyana, Pakaraima mts:
Maas et al. 5603 (K). Brazil, Acre: Prance et al. 7630 (K);
Amapa: Irwin et al. 48433 (K); Amazonas: Lleras et al.
P16615 (K); Para: Prance & Pennington 2020 (K); Rondonia:
Prance et al. 9006 (K).

Piper dumosum can be identified by its moderately pubescent
leaves with red or black glands, and the large, rather leaf-like,
copiously glandular prophylls.

27. Piper imberbe Trel. & Standley in Fieldiana Bot. 24 (3):
303 (1952). Type: Guatemala, Izabal, between Escobas &
waterfall, Steyermark 39857 (F!-holotype).

Fig. 15A, a.

Subshrub, sometimes epiphytic, 0.5-1 m high, stems spread-
ing or decumbent, strongly ridged when dry, glabrous or
sometimes minutely puberulent on young growth. Leaves
17-23 cm long, 4.5-9.5 cm wide, lanceolate, peltate, glab-
rous, sparsely glandular, subcoriaceous, dull green above,
paler beneath, narrowing to long acuminate apex; peltate

PIPER IN THE NEW WORLD

203

Fig. 7 Distribution of P. divarication.

Fig. 8 Distribution of P. dumosum.

base round. Venation with 2-3 pairs of secondary veins
radiating from petiole attachment, and 4-6 pairs arising along
the length of the midrib. Petioles 3-5 cm long. Prophylls 1-3
cm long, narrow. Inflorescences erect, 7-9 cm long;
peduncles 10-12 mm long. Anthers 0.2-0.3 mm long. Floral
bracts 0.4-0.6 mm wide, triangular to orbicular, fleshy,
glabrous. Fruit obovoid, 0.7-0.9 mm wide, trigonous, glab-
rous, apex domed; stigmas 3, linear.

Moist forest; 0-100 m.

DISTRIBUTION. Guatemala, Honduras. Fig. 9.

Guatemala, Izabal: Steyermark 39857 (F, NY). Honduras,
Olancho: Blackmore & Heath 2034 (BM).

This distinctive plant has only been collected a few times. It
can be distinguished mainly by its diminutive habit and
peltate lanceolate leaves.

28. Piper xylosteoides (Kunth) Steudel, Nomencl. hot. ed. 2,

2: 344 (1841).
Fig. 11A, a, b, c.

Schilleria xylosteoides Kunth in Linnaea 13: 677 (1839). Type:

Brazil, Sellow s.n. (K!-isotype).
Artanthe xylosteoides (Kunth) Miq., Syst. piperac.: 422

(1844).

204

Fig. 9 Distribution of P. imberbe.

A. concinna Miq. in Linnaea 20: 151 (1847). Type: Brazil,

Po/z/5032 (W-holotype; SP!-photograph).
Piper concinnum (Miq.) C. DC. in DC., Prodr. 16(1): 328

(1869), non Haworth (1821) (Art. 64.1).
P. pabstii C. DC. in Notizbl. hot. Gart. Mus. Berl. 6: 448

(1917). Type: Brazil, Santa Catarina, PabstAQS (B-holotype).
P. concinnatoris Yuncker in Bolm Inst. hot., S. Paulo 3: 33

(1966). Type as for Artanthe concinna Miq.

Slender shrub 2-3 m high, stems glabrous. Leaves ovate-
elliptic to oblong-elliptic, 7-11 cm long, 2-4 (-5) cm wide,
glossy above, glabrous, with scattered pellucid glands, apex
acute to acuminate, base acute. Venation with 2 pairs of
secondary veins arising from or near the base and curving to
apex, and 1-2 pairs from further up the midrib. Petioles 6-10
mm long. Prophylls 6-8 mm long. Inflorescences 3-6.5 cm
long, erect; peduncles 5-10 mm long. Anthers 0.2-0.3 mm
long. Floral bracts 0.5-0.8 mm wide, triangular-conchiform,
fleshy, glabrous. Fruit obovoid, 0.8-1 mm wide, trigonous,
apex slightly domed, sometimes with minute style 0.5 mm,
glabrous; stigmas 3.

M. C. TEBBS

Cool mountain areas; 700-1600 m.
DISTRIBUTION. South-eastern Brazil. Fig. 10.
Brazil, Rio de Janeiro: Burchell 6686 (BM).

This species is very similar in leaf-form to P. urophyllum, but
lacks the distinctive thickening of the leaf-base. The fruits are
smaller and less prominent in P. xylosteoides.

29. Piper urophyllum C. DC. in An. Inst. fis.-geogr. C. Rica
9: 170 (1897). Type: Costa Rica, Bois dans la vallee du Rio
Tuis (bassin du Reventazon), 600 m, Tonduz 8159 (G-
holotype; M!-isotype).

Fig. 11B, d, e, f, g.

P. sarapiquinum C. DC. in An. Inst. fis.-geogr. C. Rica. 9:
166 (1897). Type: Costa Rica, Hacienda Vieja, bord du
Sarapiqui, Biolley 7437 (?P-holotype).

Fig. 10 Distribution of P. xylosteoides.

Slender, sometimes lax shrub, 1.5^1 m, stems glabrous, often
producing roots at nodes. Leaves 7-14 cm long, 2.5-6 cm
wide, elliptic, glabrous, greyish green, narrowing abruptly to
slender apex 1-2 cm long, base thickened at petiole. Venation
with 4-7 pairs of secondary veins arising from along the
length of the midrib, curving towards apex. Petioles 5-10 mm
long. Prophylls 4-8 (-15) mm long, pale green, apex blunt.
Inflorescences 4-8 cm long, pale green; peduncles 1-2 cm
long. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.8 mm
wide, triangular, fleshy, glabrous. Fruit obovoid, 1-2 mm
wide, broadly trigonous, with minute style; stigmas 3, linear.

Moist forests, remnant woods, and stream sides; 0-1000 m.
DISTRIBUTION. Nicaragua-Panama. Fig. 12.

Nicaragua, Zelaya: Pipoly 3754 (MO, BM). Costa Rica,
Alajuela: Lent 2654 (NY); Cartago: Liesner et al. 15287
(US); Guanacaste: Standley & Valeria 45913 (US); Heredia:
Opler 1597 (NY); Limon: Davidson & Donahue 8684 (CAS);
Puntarenas: Burger & Stolze 5525 (MO). Panama, San Bias:
Sugden 513 (K).

The habit of this slender, lax shrub, often rooting at the lower
nodes, is similar to that of P. mikanianum, although the latter
is generally smaller and more spreading. P. urophyllum has
distinctive leaves that narrow abruptly at the top to form long
narrow apices, and bases that are thickened at their attach-
ment to the petiole. This thickening gives the appearance of
small 'lips', that dry to a darker colour than the rest of the
leaf. When ripe, the fruit is prominent on the inflorescence,
and, with the distinctive leaves, makes this species easily
recognizable. Piper subrepens has similar leaves, but lacks the
thickening at the base, and has smaller, less prominent
mature fruits. Burger (1971) united P. sarapiquinum with P.
urophyllum and therefore the latter name should be used
(Art. 57.2).

30. Piper subrepens Trel. in Ann. Mo. hot. Gdn. 24: 187
(1937). Type: Panama, C. Valle Chiquito, Code, Seibert
504 (MO!-holotype; K!, NY!-isotypes).

Fig. 11C, h, i, j.

Glabrous, nodose shrub, stems glossy. Leaves 4-8 cm long,
2-3.5 cm wide, ovate-lanceolate, upper surface glossy, apex
acuminate, base obtuse, symmetrical or slightly asym-
metrical. Venation with 3-4 pairs of secondary veins, loop-
connecting towards apex. Petioles 7-12 mm long, vaginate to
middle, glabrous. Prophylls 3-6 mm long, slender, acute.

PIPER IN THE NEW WORLD

205

Fig. 11 A: P. xylosteoides, habit; a: prophyll; b: part of inflorescence; c: bract and fruit. B: P. urophyllum, habit; d: prophyll; e: leaf-base; f:
part of inflorescence; g: bract and fruit. C: P. subrepens, habit; h: prophyll; i: part of inflorescence; j: bract and fruit. D: P. mikanianum, habit;

k: prophyll; 1: part of inflorescence; m: bract and fruit.

206

M. C. TEBBS

Fig. 12 Distribution of P. urophyllum.

Inflorescences 2-3 cm long; peduncles 6-10 mm long, glab-
rous. Anthers 0.2 mm long. Floral bracts 0.5 mm wide,
triangular, fleshy, minutely ciliate. Fruit obovoid, 0.5-0.8 mm
wide, trigonous, with domed apex; stigmas 3, recurved.

Mountain slopes; 700-800 m.
DISTRIBUTION. Panama. Fig. 13.

There is no record of the height of P. subrepens, either in the
original description or in collection data. Piper subrepens
resembles P. urophyllum in inflorescence, floral bracts, and
fruit structure, although the mature fruits are smaller and less
prominent in the former. The leaves are ovate-lanceolate as
opposed to elliptic, and do not have the abruptly narrowed
apex and thickened base of P. urophyllum. This species is not
well represented in herbaria, and only type material has been
examined. It is an endemic of Panama, occurring only on the
western slope and summit of Cerro Valle Chiquito in Code.

31. Piper mikanianum (Kunth) Steudel, N amend, bot. ed. 2,

2: 342 (1841).
Fig. HD,k,l,m.

Fig. 13 Distribution of P. subrepens.

Fig. 14 Distribution of P. mikanianum.

Schilleria mikaniana Kunth in Linnaea 13: 719 (1839). Type:

Brazil, Sellow s.n. (B-holotype; K!-isotype).
Artanthe mikaniana (Kunth) Miq., Syst. piperac.: 383 (1844).
Piper parthenium C. DC. in DC., Prodr. 16(1): 330 (1869).

Type: Brazil meridionali, Sellow s.n. (B-holotype).
P. debile C. DC. in Bull. Herb. Boissier 2 (2): 400 (1903).

Type: Paraguay, in sylva Pacoba, Hassler 4517 (BAF-

holotype, photograph!).
P. parthenium var. pilosius C. DC. in Bull. Soc. bot. Geneve

II, 6: 108 (1914). Type: Paraguay, Cordillera de Villa Rica,

Hassler 8706 (G-holotype).
P. misionesanum C. DC. in Reprium Spec. nov. Regni veg.

14: 395 (1916). Type: Argentina, Santa Ana Missiones,

Hassler s.n. (BAF-holotype).
P. mikanianum forma clausum Yuncker in Bolm Inst. bot. , S.

Paulo 3: 55 (1966). Type: Brazil, Rio Branco do Sul,

Parana, Hatschbach 7261 (Herbario Hatschbach, Curitiba-

holotype, photograph!).
P. mikanianum var. peltatum Yuncker in Bolm Inst. bot., S.

Paulo 3: 54 (1966). Type: Brazil, Minas Gerais, Regnell II

256 (S-holotype).

Spreading or decumbent shrub, 0.5-1 m high, stems glabrous
to minutely pubescent, rooting at lower nodes. Leaves 6-14
cm long, 3-8 (-11) cm wide, ovate-lanceolate, apex acute-
acuminate, glabrous to sparsely pubescent, base obtuse to
shallowly or deeply cordate. Venation with 2-3 pairs of
secondary veins arising from the base, and 1-2 pairs from
further up the midrib. Petioles 0.5-3 (-5) cm long, glabrous to
sparsely pubescent. Prophylls 5-15 mm long. Inflorescences
3-6 cm long, fragrant; peduncles 8-20 mm long, glabrous to
minutely pubescent. Anthers 0.2-0.3 mm long. Floral bracts
0.5-0.6 mm wide, triangular, fleshy, glabrous or minutely
ciliate. Fruit obovoid, 0.81 mm wide, trigonous, apex con-
vex, glabrous; stigmas 3, linear.

Margins of woods and gallery forest; 400-1000 m.
DISTRIBUTION. Brazil, Paraguay, Argentina. Fig. 14.

Brazil, Montenegro: Sehnem 5055 (B); Gravatai: Rambo
42741 (BM); Minas Gerais: Irwin et al. 26862 (E); Parana:
Kummrow 1305 (MU); Rio Grand do Sul: Saint-Hilaire 2796
(K); Santa Catarina: Smith & Klein 15519 (B). Paraguay, Del

PIPER IN THE NEW WORLD

207

Fig. 15 A: P. imberbe, habit; a: bract and fruit. B: P. carpunya, habit; b: bract and fruit. C: P. grande, habit; c: part of inflorescence; d: bract

and fruit.

208

Guaira: Monies 15879 (BM); Balansa 2313 (K). Argentina,
Candelaria: Monies 2363 (LIL); Misiones: Schwindt 5006
(LIL); Posadas: Ekman 1233 (US).

Piper mikanianum is one of the few species of Piper to grow
in the far south of Latin America, extending well into
Argentina. It shares similarities, such as inflorescence, bract,
and fruit shape, with P. urophyllum. P. mikanianum is a
sprawling or decumbent shrub, rooting at the lower nodes, a
character also shared by P. urophyllum. This may also
indicate a scandent habit, but there is insufficient evidence
available to confirm this.

Piper mikanianum is another species in this section with
leaves that range in shape from ovate to cordiform, often on
the same plant. The taxa cited above were originally placed
into synonymy under P. mikanianum by Yuncker (1972).

32. Piper carpunya Ruiz Lopez & Pavon, Fl.peruv. 1: 37, t.
63b (1798). Type: Peru, Huanuco, Chinchao et Cuchero,
Pavon s.n. (MA-holotype, photograph!; F!, BM!-? isotype).

Fig. 15B, b.

Ottonia carpunya (Ruiz Lopez & Pavon) Miq., Syst. piperac. :

547 (1844).
O. lessertiana Miq., Syst. piperac.: 546 (1844). Type: 'Habit

in America meridionali', Matthews 1696 (G-holotype;

BM!,E!,K!-isotypes).

Carpunya lessertiana (Miq.) C. Presl, Epimel. hot.: 229 (1851).
C. peruviana C. Presl, Epimel. hot.: 229 (1851). Type as for

P. carpunya.
Piper lenticellosum C. DC. in J. Bot., Lond. 4: 166 (1866).

Type: Colombia, prov. Barbacoas, Triana 3 (G-holotype;

BM!-isotype).
P. lessertianum (Miq.) C. DC. in DC., Prodr. 16(1): 258

(1869), non C. DC. in /. Bot., Lond. 4: 164 (1866) (Art.

64. 1).
P. cocherense Trel. in Publs Field Mus. nat. Hist. (Bot.) 13:

151 (1936). Type: Peru, Huanuco, Poeppig D1354 (W-

holotype) .
P. subdurum Trel. ex Standley in Publs Field Mus. nat. Hist.

(Bot.) 18: 362 (1937). Type: Costa Rica, vicinity of El

General, Prov. San Jose, Skutch 2993 (US-holotype; NY!-

isotype).

Shrub or small tree 2-3 (-5) m high, stems glabrous, with
tubercles. Leaves 9-17 cm long, 5-11 cm wide, ovate-elliptic
to oblong-elliptic, coriaceous, glabrous, slightly glossy above,
apex acute-acuminate, base round to acute. Venation with 2-
5 pairs of secondary veins, arising from lower to middle part
of midrib and curving towards the apex, with smaller con-
necting veins between. Petioles 5-10 mm long, glabrous.
Prophylls 1-2.5 cm long, glabrous. Inflorescences erect, 5-10
(-12) cm long; peduncles 5-8 mm long. Anthers 0.3 mm long.
Floral bracts 0.7-0.9 mm wide, triangular-cupulate, fleshy,
glabrous, or occasionally minutely ciliate. Fruit obovoid, 1
mm wide, upper surface slightly domed, granular or minutely
pubescent; stigmas 3, prominent.

Moist woods, remnant rain-forest and scrub, slopes and steep
hillsides; 1000-2000 m.

DISTRIBUTION. Costa Rica, Colombia, Ecuador, Peru. Fig.
16.

Costa Rica, Cartage: Webster & Miller 12294 (F); Heredia:
Poveda 955 (F). Colombia, Popayan: Hartweg 1400 (BM).
Ecuador, Chimborazo: Camp 3525 (K); mentis Pululagua:

M. C. TEBBS

Fig. 16 Distribution of P. carpunya.

Andre 3313 (K). Peru, Amazonas: Young & Eisenberg 348
(MO, NY); Chachapoyas: Mathews 3226 (BM); Chili: Poeppig
1354 (BM); Huacafistana: Sandeman 4412 (K).

Piper carpunya can be recognized by its prominently nodose,
tuberculate stems, glabrous, coriaceous, pungent-aromatic
leaves, short, erect inflorescences, and cupulate bracts. The
species is confined to forested areas at high altitudes; only
three specimens have been seen from Central America, all
collected from mountainous areas in Costa Rica. The first
collection is from remnant rain-forest north-east of Paraiso,
Cartago Province, at an altitute of 1600 m; the second,
originally misidentified as P. verruculosum C. DC., is from
between San Miguel and Cariblanco, Heredia Province, and
does not have altitude or habitat details; the third, the type of
P. subdurum Trel., was collected from a clearing in the
vicinity of El General, San Jose Province. These collections
indicate that P. carpunya is more widespread than was
previously thought, and not restricted to northern South
America.

33. Piper grande M. Vahl, Eclog. amer., fasc. 2: 3, t. 11
(1798). Type: New Granada, Santa Martha, Von Rohr
46 labelled as Piper coriaceum (C!-holotype;
BM!-?isotype).

Fig. 15C, c, d.

P. grandifolium Kunth in Humb., Bonpl. & Kunth, Nov. gen.

sp. 1: 46 (1816). Type: Colombia, montis Quindiu,

Humboldt & Bonpland s.n. (P-holotype).
P. riparium Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp.

1: 48 (1816). Type: Colombia, fluminis Magdalenae,

Humboldt & Bonpland s.n. (P-holotype).

PIPER IN THE NEW WORLD

Schilleria riparia (Kunth) Kunth in Linnaea 13: 707 (1839).
5. grandifolia (Kunth) Kunth in Linnaea 13: 709 (1839).
Artanthe grandifolia (Kunth) Miq., Syst. piperac. : 408 (1844).
A. riparia (Kunth) Miq., Syst. piperac.: 409 (1844).
Piper borucanum C. DC. in Bull. Soc. r. Bot. Belg. 30(1): 219

(1892). Type: Costa Rica, forets pres de Boruca, Pittier

3609 (G!-holotype).
P. pseudo-variabile Trel. in Contr. U.S. natn. Herb. 26: 23

(1927). Type: Panama, Loma de la Gloria, Pittier 3845

(US-holotype, photograph!).
P. pseudo-variabile var. collium Trel. in Contr. U.S. natn.

Herb. 26: 23 (1927). Type: Panama, Rio Tapla, Standley

26130 (US-holotype, photograph!).
P. pseudo-variabile var. pachypus Trel. in Contr. U.S. natn.

Herb. 26: 24 (1927). Type: Panama, between Frijoles &

Monte Lirio, Killip 12145 (US-holotype, photograph!).
P. cercidiphyllum Trel. in Contr. U.S. natn. Herb. 26: 146

(1929). Type: Costa Rica, near Tilaran, Guanacaste,

Standley & Valerio 46110 (US-holotype).
P. subvariabile Trel. in Contr. U.S. natn. Herb. 26: 145

(1929). Type: Costa Rica, near Tilaran, Guanacaste,

Standley & Valerio 44970 (US-holotype).
P. marmoreum Trel. in Ann. Mo. hot. Gdn 24: 186 (1937).

Type: Panama, vicinity of Arenoso, Seibert 622 (MO!-

holotype; K!, Y!-isotypes).
P. spissinervium Trel. fide Bro. Tomas Alberto in Colegio de

San Jose, Medellfn, Colombia Publ. 53-70, p. 18 (1942),

nom. nud. (Art. 32.1).

Shrub or small tree 1-3 m high, stems glabrous. Leaves 12-28
cm long, 8-17 cm wide, broadly elliptic to ovate-elliptic,
slightly asymmetric, glabrous, apex acuminate, base some-
times slightly cordate. Venation with 4-9 pairs of secondary
veins arising along the length of the midrib, those from or
near the base ascending more sharply than the rest. Petioles
1-5 cm long. Prophylls 1-3 cm long, glabrous. Inflorescences
8-15 cm long; peduncles 6-15 mm long, glabrous. Anthers
0.2-0.3 mm long. Floral bracts 0.5-0.8 mm wide, oblong-
triangular, fleshy, umbonate, minutely ciliate. Fruit obovoid,
0.8-1 mm wide, trigonous to orbicular, glabrous, covered by
bracts; stigmas 3, sessile.

Moist forest, edges of roads, trails and stream sides;
0-1 000m.

DISTRIBUTION. Nicaragua to Colombia. Fig. 17.

Nicaragua, Bunting & Licht 381 (NY). Costa Rica, Puntarenas:
Williams et al. 24213 (MO). Panama, Canal Zone: Croat 4380
(MO); Code: Allen 2001 (MO); Colon: Dodge et al. 16924
(MO); Darien: Duke 14168 (BM); Panama: Garwood &
Tebbs 1932A (BM); San Bias: Duke 8578 (MO); Veraguas:
Antonio 2315 (MO, BM).

This species is characterised by its distinctive pale inflores-
cence with fleshy, umbonate bracts. These conceal the
developing fruit, a feature that occurs to a lesser extent in
other species in this section, such as Piper veraguense. How-
ever, even in early stages of development, the floral bracts of
P. grande are distinctive in their structure. P.grande is a
common species, often found growing in open areas of
secondary forest.

Several species have been synonymised here by previous
authors, notably Trelease & Yuncker (1950), the types of
which have not been seen by the present author. There seems
no reason to dispute this synonymy, however, as the original
descriptions compare well with that of P. grande.

209

Fig. 17 Distribution of P. grande.

34. Piper lepturum (Kunth) C. DC. in DC., Prodr. 16(1): 320

(1869).
Fig. ISA, a.

Schilleria leptura Kunth in Linnaea 13: 679 (1839). Type:

Brazil, Rio de Janeiro, Gaudichaud s.n. (?B-holotype).
S. jussiaeana Kunth in Linnaea 13: 680 (1839). Type: Brazil,

Serra d'Estrella. Luschnath 292 (B-holotype; SP!-

photograph).
Piper jussiaeanum Kunth in Linnaea 13: 680 (1839) as

synonym (Art. 34.1).

Artanthe jussiaeana (Kunth) Miq., Syst. piperac.: 420 (1844).
A. leptura (Kunth) Miq., Syst. piperac.: 523 (1844).
A. meyeniana Klotzsch ex Miq., Syst. piperac.: 524 (1844).

Type: Brazil, Rio de Janeiro, Gaudichaud s.n. (B-holotype).
A. denudans Miq. in Linnaea 20: 172 (1847). Type: Brazil,

monte Corcovado, Pohl 4778, 5028 (W-syntypes).
Piper fuscum C. DC. in DC., Prodr. 16 (1): 319 (1869). Type:

Brazil, Riedel 291 (LE).

P. denudans (Miq.) C. DC. in DC., Prodr. 16 (1): 324 (1869).
P. denudans var. angustifolium C. DC. in DC., Prodr. 16 (1):

324 (1869). Type: Brazil, Corcovado, Pohl 5028 (W-

holotype).
P. dupasquieri C. DC. in Linnaea 37: 361 (1872). Type:

Brazil, Dupasquier s.n. (G-holotype).
P. glaziovii C. DC. in Annu. Conserv. Jard. hot. Geneve 2:

265 (1898). Type: Brazil, Rio de Janeiro. Glaziou 7848 (G-

holotype).
P. sievekingii C. DC. in Notizbl. hot. Gart. Mus. Berl. 6: 435

(1917). Type: ?
P. lepturum var. angustifolium (C. DC.) Yuncker in Bolm

Inst. hot., S. Paulo 3: 103 (1966).

Shrub 2-3 m high, stems glabrous, glandular. Leaves 11-20
cm long, 3-5 cm wide, lanceolate to oblong-lanceolate,
glabrous, with brown or black glands on both surfaces; apex
long-acuminate, base obtuse or round. Venation with 1-2
pairs of secondary veins arising steeply from or near the base,
and 4-6 pairs arising along the length of the midrib, ascending
steeply to apex. Petioles 10-25 mm long, glabrous. Prophylls
8-14 mm long, narrow, apex blunt. Inflorescences 6-10 cm
long; peduncles 5-15 mm long, glabrous. Anthers 0.2-0.3 mm
^^rq] bracts 0.6-0.8 mm wide, triangular-cucullate,

210

M. C. TEBBS

Fig. 18 A: P. lepturum, habit; a: bract and fruit. B: P. aequale, habit; b: bract and fruit; c: fruit. C: P. verruculosum, habit; d: bract and fruit.

PIPER IN THE NEW WORLD

Fig. 19 Distribution of P. lepturum.

fleshy, glabrous or minutely ciliate. Fruit obovoid, 1 mm
wide, glabrous, glandular, with rounded apex; stigmas 3,
linear.

Forest; 0-500 m.
DISTRIBUTION. Brazil. Fig. 19.

Brazil, unlocalized: Bowie & Cunningham s.n. 1814-17 (BM);
Sellow 8 (BM); Sellow 137 (BM).

Piper lepturum has similarly shaped leaves to P. aequale, but
can be distinguished by its glandular stems and leaves, and
steeply ascending secondary veins. The floral bracts are
smaller, and the fruits have a glandular surface. The names
cited above were first placed into synonymy under P. lepturum
by Yuncker (1972). Although not all of the types of these taxa
have been examined for this study, the original descriptions
agree with that of P. lepturum.

35. Piper aequale M. Vahl, Eclog. amer. 1: 4, pi. 3 (1796).
Type: Montserrat, Ryan s.n. (C!-holotype; BM!-isotype).
Fig. 18B, b, c.

P. caripense Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp.

1: 48 (1816). Type: Caripense Prov. Cumanae, Humboldt

& Bonplands.n. (P-holotype).
P. costaricense C. DC. in DC., Prodr. 16(1): 328 (1869).

Type: Costa Rica, Hoffmann 678 (B-holotype).
P. micranthera C. DC. in Linnaea 37: 354 (1872). Type:

Costa Rica, Oersted 877 (C!-holotype).
P. donnell-smithii C. DC. ex J.D. Smith, Enum. pi. guatem.

2: 95 (1891). Type: Guatemala, Pansamala, Tuerkheim 975

(US?-holotype; NY!-isotype).
P. cabagranum C. DC. in An. Inst. fis.-geogr. C. Rica. 9: 173

(1897). Type: Costa Rica, forets de Cabagra, Pittier 6528

(G-holotype; MO!- photograph).
P. johnstoni C. DC. in Proc. Am. Acad. Arts Sci. 40: 685

(1905). Type: Venezuela: Isla Margarita, Juan Griego

Trail, Johnston 19 (MO!- holotype; KMsotype).
P. aequale var. elliptico-lanceolatum C. DC. in Bot. Gaz. 70:

171 (1920). Type: Costa Rica, San Ramon, Pittier 14185,

14186 (US-syntypes, photograph!).
P. chiriquinum C. DC. in Smithson. misc. Collns 71: 2 (1920).

Type: Panama, El Bocquete, Pittier 2930 (US-holotype).

211

P. tenuispicum C. DC. in Bot. Gaz. 70: 170 (1920). Type:

Costa Rica, Tonduz 13187 (G-holotype; MO!-photograph).
P. pablense Trel. in Contr. U.S. natn. Herb. 26: 23 (1927).

Type: Costa Rica, San Pablo de Tarrazu, Jimenez 1094

(US-holotype; NY!-isotype).
P. caeruleifolium Trel. in Contr. U.S. natn. Herb. 26: 145

(1929). Type: Costa Rica, Quebrada Serena near Tilaran,

Standley & Valeria 46134 (US-holotype).
P. concinnifolium Trel. in Contr. U.S. natn. Herb. 26: 167

(1929). Type: Costa Rica, Colinas de Piedades, Brenes

14194 (US-holotype).
P. oppressumTrel. in Contr. U.S. natn. Herb. 26: 161 (1929).

Type: Costa Rica, Nicoya Peninsula, Cook & Doyle 697

(US-holotype, photograph!).
P. seductum Trel. in Contr. U.S. natn. Herb. 26: 135 (1929).

Type: Costa Rica, Madre de Dios (Atlantique), Pittier

10308 (US-holotype, photograph!).
P. atrichopus Trel. in /. Wash. Acad. Sci. 19: 330 (1929).

Type: Honduras, Atlantida, Standley 54594 (F-holotype;

ILL!-isotype).
P. colemanense Trel. ex Standley in Publs Field Mus. nat.

Hist. (Bot.) 18: 337 (1937). Type: Costa Rica, Dodge &

Goerger 9580 (MO! -holotype).
P. crispans Trel. ex Standley in Publs Field Mus. nat. Hist.

(Bot.) 18: 339 (1937). Type: Costa Rica, vicinity of San

Ramon, Alajuela, Brenes 5236 (F-holotype; NY!-isotype).
P. rubripes Trel. ex Standley in Publs Field Mus. nat. Hist.

(Bot.) 18: 358 (1937). Type: Costa Rica, Cerro San Jose de

Libano, Dodge, Hanckel & Thomas 6382 (GH-holotype;

F!-isotype).
P. epilosipes Trel. in Publs Field Mus. nat. Hist. (Bot.) 17:

344 (1938). Type: Honduras, Yuncker et al. 6296 (ILL-

holotype; MO!-isotype).
P. paso-anchoenseTrel. in Ann. Mo. hot. Gdn 27: 295 (1940).

Type: Panama, Allen 1579 (ILL-holotype; MO!, NY!-

isotypes).
P. jericoense Trel. & Yuncker, Piperac. N. South Amer. 1:

397 (1950). Type: Colombia, Alejandro 2730 (US! -holotype).
P. perlasense Yuncker in Ann. Mo. hot. Gdn 37: 67 (1950).

Type: Panama, San Jose Island, Johnston 1215 (ILL-
holotype; MO!-isotype).
P. cheyennense Trel. & Standley in Fieldiana Bot. 24: 292

(1952). Type: Guatemala, Steyermark 39154 (F!-holotype).
P. frioense Standley & Steyerm. in Fieldiana Bot. 24: 299

(1952). Type: Guatemala, Standley 90328 (F!-holotype).

Shrub 1-3 (-5) m high, stems glabrous or sometimes minutely
puberulent. Leaves 6-20 cm long, 2-9 cm wide, narrowly
elliptic, apex acuminate, base acute, equally attached to
petiole. Venation with 2-8 pairs of veins arising from along
the length of the midrib, 2-3 prominent pairs arising from or
near the base. Petioles 6-15 cm long. Prophylls 10-15 mm
long, slender, glabrous. Inflorescences erect, pale, 5-10 cm
long; peduncles 5-12 mm long, glabrous. Anthers 0.1-0.3 mm
long. Floral bracts 0.8-1 mm wide, triangular, fleshy, glab-
rous or minutely ciliate. Fruit obovoid, 0.8-1 mm wide,
trigonous, rounded at apex, glabrous; stigmas 3.

Moist forests, wet ravines, and shaded sides of roads and
tracks; 0-2000m.

DISTRIBUTION. Southern Mexico-South America. Fig. 20.

Mexico, Chiapas: Breedlove & McClintock 34184 (MO);
Tabasco: Cowan et al. 2850 (MEXU). Belize, Rio Grande:
Schipp S-452 (BM). Guatemala, Alta Verapaz: Molina 12163

212

M. C. TEBBS

Fig. 20 Distribution of P. aequale.

(F, NY); Izabal: Jones & Facey 3249 (F); Peten: Ortiz 2100
(MO). Honduras, Atlantida: Standley 56757 (F); Puerto
Sierra: Wilson 601 (NY). Nicaragua, Jinotega: Standley 10709
(F); Matagalpa: Williams et al. 24813 (F). Costa Rica,
Alajuela: Smith 1848 (NY); Cartago: Skutch 4598 (NY);
Heredia: Opler 729 (MO); Puntarenas: Croat 26682 (MO);
San Jose: Skutch 4774 (NY). Panama, Bocas del Toro:
Kirkbride & Duke 768 (NY); Canal Zone: Mori 7939 (MO);
Chiriqui: Correa & Lazor 1443 (MO); Code: Nee & Dwyer
9145 (MO); Darien: Duke 14172 (MO); Panama: Gentry 6126
(MO); San Jose Island: Johnston 134 (MO); Veraguas: Croat
27547 (MO). Colombia, Antioquia: Ewan 15812 (BM);
Magdalena: Jaramillo et al. 5214 (B); Santa Marta: Smith
1231 (BM). Venezuela, Apure: Davidse & Gonzalez 21879
(MO); Carabobo: Steyermark 94315 (K). Surinam, Juliana
Top: Irwin et al. 54610 (K); French Guiana, Cayenne:
Granville 6547 (K). Guyana, Marudi Hills: Staffers et al. 224
(K). Brazil, Amazonas: Pipoly et al. 6930 (K); Minaes
Geraes: Mexia 4666 (B). West Indies, Dominica: Whitefoord
5625 (BM); Guadeloupe: Proctor 20020 (BM); Montserrat:
Howard 15165 (BM); St Kitts: Proctor 19243 (BM); St Lucia:
Proctor 18162 (BM); Tobago: Broadway 4272 (BM); Trinidad:
Broadway 5804 (BM).

It is possible for the leaves of Piper aequale to be confused
with those of P. schiedeanum, which may also be elliptic-
lanceolate in shape. However, the leaf -bases of P. schiedeanum
are round rather than acute or cuneate as in P. aequale. While
P. schiedeanum often has elliptic, ovate or cordiform leaves
on the same plant, the leaves of P. aequale are generally

uniform in shape. Furthermore, P. aequale has a short,
slender, acute prophyll, which often dries to a dark brown,
while that of P. schiedeanum is usually oblong with a blunt
tip, and remains pale green on drying. The floral bracts of P.
aequale do not obscure the young fruits.

36. Piper verruculosum C. DC. in J. Bot., Lond. 4: 215
(1866). Type: Costa Rica, Candelaria, Hoffmann 8 (B-
holotype).

Fig. 18C, d.

P. nudicaule C. DC. in An. Inst. fis.-geogr. C. Rica 9: 162

(1897). Type: Costa Rica, forets de Tablazo, Pittier 7922

(G-holotype; F!-photograph).
P. carnosicaule Trel. in Contr. U.S. natn. Herb. 26: 168

(1929). Type: Costa Rica, El Muneco, Cartago, Standley &

Valeria 51050 (US-holotype).
P. zingiberinum Trel. ex Standley in Publs Field Mus. nat.

Hist. (Bot.) 18: 370 (1937). Type: Costa Rica, Santa Clara

Hills, Stork 2594 (ILL-holotype).

Densely branching shrub 1-3 m high, stems glabrous, glandu-
lar. Leaves 7-16 (-19) cm long, 2-7 cm wide, obliquely
narrowly elliptic, dark green and slightly glossy above, paler
beneath, glabrous, with prominent orange or brown pellucid
glands, apex acuminate, base unequally obtuse, Venation
with 2-3 pairs of secondary veins arising fairly sharply to
apex, with cross-veins. Petioles 3-10 mm long. Prophylls 8-16
mm long, narrow, blunt-tipped. Inflorescences erect, 4-11 cm
long, narrow; peduncles 5-10mm long. Anthers 0.2-0.3 mm

PIPER IN THE NEW WORLD

Fig. 21 Distribution of P. verruculosum.

long. Floral bracts 0.3-0.5 mm wide, triangular, fleshy,
minutely ciliate. Fruit obovoid, 0.5-0.8 mm wide, trigonous,
upper surface granular-pubescent, slightly domed; stigmas 3,
slender, recurved.

Shade in cloud forest; 800-2000 m.
DISTRIBUTION. Costa Rica. Fig. 21.

Costa Rica, Guadeloupe de Zarcero, Alajuela: Smith HI 127
(F); slopes of Volcan Arenal, Alajuela: Lent 2888 (MO).

Only found in mountainous regions of Costa Rica, growing in
cloud forest. Piper verruculosum is similar to P. aequale,
sharing characters such as fruit and bract shape, and a nar-
row prophyll. It differs in its somewhat asymmetric leaves
with unequally obtuse bases, distinctly glandular stems and
leaves, blunt-tipped rather than acute prophyll, and granular-
pubescent fruits.

37. Piper corrugatum Kuntze, Revis. gen. pi. 2: 565 (1891).
Type: Costa Rica, Port Limon, Kuntze s.n. (G-holotype).
Fig. 22.

P. nemorense C. DC. in Bull. Soc. r. Bot. Belg. 30(1): 222

(1892). Type: Forets de Juan Vinas, Pittier 1849 (G-

holotype; US-isotype, photograph!)
P. riparense C. DC. in An. Inst. fis.-geogr. C. Rica 9: 173

(1897). Type: Costa Rica, forets des bords du R. Hacum

pres de Buenas Aires, Tonduz 4893 (G-holotype; BM!-

isotype).
P. tsakianum C. DC. in An. Inst. fis.-geogr. C. Rica 9: 174

(1897). Type: Costa Rica, forets de Tsaki, Talamanca,

Tonduz 9532 (G-holotype; F!-isotype).
P. partiticuspe Trel. in Ann. Mo. hot. Gdn 27: 294 (1940).

Type: Panama, Code, Allen 1195 (ILL-holotype; MO!-

isotype).

Shrub 1.5-3 m, stems glabrous to densely hirsute, with
yellowish multicellular hairs, 0.5-1.5 mm long. Leaves 16-34
(40) cm long, (7-) 10-28 cm wide, asymmetrically narrowly
to broadly ovate-lanceolate to cordiform, coriaceous or sub-
coriaceous, sometimes bullate, upper surface glabrous to
sparsely pubescent, lower surface densely pubescent, apex
acuminate, base shallowly to deeply, unevenly lobed, lower
leaves usually more deeply lobed than upper. Venation with

213

3-6 secondary veins arising from or near leaf-base and
curving towards apex, 1-2 pairs arising from further
up midrib, curving upwards. Petioles 3-5 cm long, glabrous
to pubescent, vaginate at base. Prophylls 2-5 cm long,
apex blunt. Inflorescences erect, 13-22 cm long, 4-5 mm
wide; peduncles 1-2.5 cm long. Anthers 0.2-0.4 mm long.
Floral bracts 0.2-0.5 mm wide, oblong-triangular, fleshy,
upper part slightly protruding, minutely pubescent on
lower margins, sometimes covering fruits in early stages.
Fruit obovoid, 0.8-1 mm wide, trigonous, glabrous;
stigmas 3.

Woods and forest, shaded areas along roads and trails, and
steep banks; 0-1200m.

DISTRIBUTION. Nicaragua to Colombia. Fig. 23.

Nicaragua, Bluefields: Proctor et al. 27222 (NY). Costa Rica,
Alajuela: Burger et al. 11842 (NY); Cartago: Lent 3042 (F);
Limon: Jimenez 2794 (F); Puntarenas: Allen 6038 (F); San
Jose: Burger & Stolze 5307 (F); Panama, Chiriqui: Nee 10662
(MO); Code: Gentry 7419 (MO); Panama: Croat 12156
(MO); Veraguas: Mori & Kallunki 5320 (MO). Colombia,
Cauca: Alston 8153 (BM).

Piper corrugatum is closely related to P. schiedeanum, with
more or less identical inflorescence characters, and glabrous
forms can be very difficult to separate from that species. It has
been maintained here as a separate species only because the
leaf shape is consistently obliquely oblong-cordate, while P.
schiedeanum has generally symmetrically elliptic, ovate, or
cordiform leaves, often together on the same plant. Further-
more, the leaves of P. corrugatum are coriaceous or sub-
coriaceous, occasionally bullate, and sparsely to copiously
pubescent. P. corrugatum also has similar inflorescences to
the peltate-leaved P. veraguense.

Piper riparense appears to be a pilose form of P. corrugatum,
being otherwise identical morphologically. De Candolle men-
tions this at the end of his original description of P. riparense:
'Species P. nemorense C. DC. proxima forsan ejus forma
longius pilosior et foliis majoribus.' A wide range of speci-
mens from Nicaragua to Panama has been examined, and no
other characters can be found to separate these two species
satisfactorily. The leaves of the Panamanian specimens tend
to be rougher and more bullate, but this again is variable. The
more densely pubescent forms of P. corrugatum are
commonly found between and 1200 m altitude, while
glabrous or sparsely pubescent forms occur between 500 and
1700m.

38. Piper schiedeanum Steudel, Nomencl. hot. ed. 2, 2: 343
(1841). Type: Mexico, in sylvis Misantlae, Schiede 1102
(BM!-holotype).

Fig. 24.

P. tiliifolia Schldl. & Cham, in Linnaea 6: 352 (1831), non

Desv. ex Ham., Prodr. pi. Ind. occid.:4 (1825) (Art. 64.1).
Artanthe benthamiana Miq. in Lond. J. Bot. 4: 447 (1845).

Type: Venezuela, Caracas, Linden 216 (K!-holotype).
Piper glanduligerum C. DC. in /. Bot., Lond. 4: 215 (1866).

Type: Venezuela, Tovar, Fendler 1137, 2400 (G-syntypes;

K!-isosyn types).
P. benthamianum (Miq.) C. DC. in DC., Prodr. 16(1): 264

(1869).
P. tenuifolium C. DC. in DC., Prodr. 16(1): 321 (1869).

Type: Mexico, Pavon s.n. (G-holotype; MO!-photograph).

214

M. C. TEBBS

Fig. 22 P. corrugatum A: habit; a: pubescent stem; b: rugose leaf
surface; c: prophyll; d: part of inflorescence; e: bract and fruit.

PIPER IN THE NEW WORLD

Fig. 23 Distribution of P. corrugatum.

P. zacuapanum C. DC. in DC., Prodr. 16(1): 330 (1869).

Type: Mexico, Zacuapan, Schiede 1102 (B-holotype).
P. megalophyllum C. DC. in Linnaea 37: 357 (1872). Type:

Mexico, Mirador in sylvis humidis, Liebmann 69, 70 (C!-

syntypes).
P. megalophyllum var. connivens C. DC. in Linnaea 37: 357

(1872). Type: Mexico, Mirador, Liebmann 71 (C!-

holotype).
P. casimirianum Hemsley, Biol. cent.-amer., Bot. 3: 46

(1882). Type: Nicaragua, Chontales, Tate 383 (K!-holotype;

BM!-isotype).
P. carrilloanum C. DC. in Bull. Soc. r. Bot. Belg. 30 (1): 209

(1892). Type: Costa Rica, Bord de la route a Carrilo, Pittier

2525, 2526 (G-syntypes; BM!, K!-isosyntypes of both

specimens).
P. vallicolum C. DC. in Bull. Soc. r. Bot. Belg. 30 (1): 222

(1891). Type: Costa Rica, bord de la vallee du Rio Jaris

pres de Pacaca, Pittier 3298 (G!-holotype).
P. variabile C. DC. ex J.D. Smith in Bot. Gaz. 19: 259 (1894).

Type: Guatemala, Coban, Tuerckheim 434 (US-holotype;

F!-photograph).
P. paulownifolium C. DC. in An. Inst. fis.-geogr. C. Rica 9:

173 (1897). Type: Costa Rica, forets de la Palma, Pittier

6739 (G!- holotype).
P. levilimbum Trel. in J. Wash. Acad. Sci. 19: 333 (1929).

Type: Honduras, Lancetilla Valley, Standley 53483 (F-

holotype) .
P. omega Trel. in Contr. U.S. natn. Herb. 26: 146 (1929).

Type: Costa Rica, La Hondura, Standley 37800 (US-
holotype, photograph!).
P. middlesexense Trel. in Publs Field Mus. nat. Hist. (Bot.)

12: 103 (1936). Type: Belize, Middlesex, Schipp 286 (F-

holotype; NY!- isotype).
P. atrichopus var. arborescens Trel. in Publs Field Mus. nat.

Hist. (Bot.) 17: 347 (1938). Type: Honduras, Yunckeret al.

8578 (MO!-holotype).
P. collinum var. flagellicuspe Trel. in Publs Field Mus. nat.

Hist. (Bot.) 17: 347 (1938). Type: Panama, Miller 1773

(MO! -holotype).
P. exigens Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 345

(1938). Type: Honduras, Yunckeret al. 6124 (ILL-holotype;

C!, MO!-isotypes).

215

P. exigens var. subovatum Trel. in Publs Field Mus. nat. Hist.

(Bot.) 17: 345 (1938). Type: Honduras, Yunckeret al. 5997

(ILL-holotype).
P. zarceroense Trel. ex Standley in Publs Field Mus. nat. Hist.

(Bot.) 18: 1548, f.9 (1938). Type: Costa Rica, Zarcero,

Smith H49 (F!-holotype).
P. variabile var. obtentum Trel. in Publs Field Mus. nat. Hist.

(Bot.) 9: 281 (1940). Type: Honduras, Atlantida, Yuncker,

Koepper & Wagner 8495 (ILL!-holotype).
P. pergamentifolium Trel. & Standley in Fieldiana Bot. 24(3):

314 (1952). Type: Guatemala, Volcan de Zunil, Skutch 876

(F!-holotype).

Shrub or small tree, 2-3 (-5)m high, stems glabrous. Leaves
12-32 cm long, 7-27 cm wide, narrowly to broadly ovate-
elliptic to ovate-lanceolate, glabrous or minutely puberulent
on veins beneath, apex acuminate, base ovate to deeply
cordate. Venation with 4-7 pairs of secondary veins mostly
arising from or near the base, curving towards apex, with
prominent cross veins. Petioles 2-12 (-20) cm long, glabrous.
Prophylls 2-5 cm long, obtuse, glabrous, pale when dry.
Inflorescences erect, 10-22 cm long; peduncles 6-20 mm long,
glabrous. Anthers 0.2 mm long. Floral bracts 0.4-0.7 mm
wide, orbicular to triangular, fleshy, glabrous or minutely
ciliate, sometimes covering fruit in early stages. Fruit obovoid,
0.8-1 mm wide, trigonous, glabrous or minutely granular;
stigmas 3, sessile.

Disturbed places in moist forest, wooded slopes, roadsides,
and trails; 0-1500 m.

DISTRIBUTION. Mexico-Brazil. Fig. 25.

Mexico, Chiapas: Matuda 4273 (MO); Hidalgo: King 4231
(NY); Oaxaca: Calderon 1381 (MEXU); San Luis Potosi:
King 4295 (NY); Tabasco: Cowan & Zamudio 3356 (MO);
Veracruz: Purpus 7767 (NY). Belize, Cayo: Gentle 9020
(NY); Stann Creek: Gentle 8238 (F); Toledo: Gentle 5105
(F). Guatemala, Alta Verapaz: Standley 89745 (NY); Baja
Verapaz: Standley 90970 (F); Chimaltenango: Standley 62133
(F); Escuintla: Standley 64566 (F); Izabal: Steyermark 41807
(F); Peten: Steyermark 45527 (F); Quezaltenango: Steyermark
33532 (F); Retalhuleu: Standley 88482 (F); San Marcos:
Standley 68701 (F); Santa Rosa: Standley 78352 (F). Honduras,
Atlantida: Molina 25654 (MO); Comayagua: Standley 13691
(MO); Cortez: Molina 6708 (NY); Olancho: Standley 18682
(F); Puerto Sierra: Wilson 395 (NY). Nicaragua, El Cabo:
Molina 15068 (NY); Zelaya: Stevens 6826 (F). Costa Rica,
Alajuela: Lent et al. 3384 (MO); Cartago: Lent 2355 (NY);
Guanacaste: Standley & Valeria 45050 (ILL); Heredia: Opler
745 (MO); Puntarenas: Haber 1127 (MO, BM). Panama,
Bocas del Toro: Croat & Porter 16244 (MO); Canal Zone:
Croat 6319 (MO); Chiriquf: Mori & Kallunki 5918 (MO);
Darien: Gentry & Mori 13893 (MO); Panama: Croat 14520
(MO). Colombia, Antioquia: Haught 4729 (ILL); Boyaca:
Lawrence 681 (K); Caldas: Penned 9075 (ILL); El Meta:
Philipson et al. 2136 (BM); Valle: Cuatrecasas 21639 (ILL).
Venezuela, Bolivar: Steyermark 88978 (K, NY); Caracas:
Buschel s.n. (K); Falcon: Steyermark 99339 (K). Surinam,
Lucie River: Irwin et al. 54650 (K, NY). Bolivia, without
location: Bang 2640 (K). Ecuador, Los Rios: Mexia 6616
(ILL); Pichincha: Holmgren 835 (B). Peru, Hualgayoc:
Hutchison & Bismarck 6386 (K); Huanuco: Mexia 8130 (K);
Pachitea: Schunke 1619 (F, K). Brazil, Rio de Janeiro:
Glaziou 4922 (K).

216

M. C. TEBBS

Fig. 24 P. schiedeanum A: habit; a-d: leaf variation; e: prophyll; f:
part of inflorescence; g: bract and fruit.

PIPER IN THE NEW WORLD

217

Fig. 25 Distribution of P. schiedeanum.

This species has an extensive range that stretches from
Mexico to Brazil, and is one of the most phenotypically
plastic species in this section. It grows in shaded places of
moist lowland forest, lower montane forest, wet ravines, and
on the banks of streams and rivers. It displays great variation
in leaf size and shape, ranging from elliptic-lanceolate or
ovate to cordiform, frequently on the same plant; this plastic-
ity can make correct identification difficult, something that is
reflected in the large number of taxa that have been described
in the past. Gomez Pompa (1966) has described the extreme
vegetative variations to be found in this species. He reduced
four species to synonymy (P. megalophyllum C. DC., P.
variabile C. DC., P. zacuapanum C. DC., and P. middle-
sexense Trel.), and suggested that P. carrilloanum C. DC.
and P. casimirianum Hemsley might also belong to P.
schiedeanum.

For this study, large numbers of specimens of P. carrillo-

anum (including type material), and the type of P. casimiri-
anum, were examined. It has proved impossible to separate
these species satisfactorily from P. schiedeanum for, while
there is a continuous variation in leaf shape and size, the
inflorescence structure remains constant. The bracts partially
cover the fruit in early stages of development, and the fruit is
trigonous and glabrous. Piper schiedeanum has frequently
been confused with P. aequale, and the inflorescence struc-
tures are very similar. Piper aequale can be distinguished by
its dark, acute prophylls, bracts that do not appear to cover
the fruits at any stage, and leaves that are acute to cuneate at
the base. Piper schiedeanum has pale, blunt-tipped prophylls,
bracts that cover the fruits at early stages, and leaves
with ovate to deeply cordate bases. Piper corrugatum and
P. veraguense are similar in inflorescence structure to P.
schiedeanum, with floral bracts that often cover the fruits in
early stages, but can be separated by their distinctive leaves.

218

M. C. TEBBS

cm

mm

Fig. 26 P. cadenaensis A: habit; a: prophyll; b: part of
inflorescence; c: bract and fruit.

39. Piper cadenaensis Tebbs, sp. nov. Type: Guatemala,
Poptun Road, Cadenas, Peten, Lundell & Contreras 19010
(MO!-holotype).

Fig. 26.

Frutex 2-3 m altus. Folia anguste lanceolata glabra subcori-
acea, apice acuminata basin rotundata interdum inaequalia.

Spica erecta, bracteis ciliatis triangularibus. Bacca trigona
apicem versus granulosa vel pilis minutis obtecta.

Shrub 2-3 m high, stems glabrous. Leaves 13-22 cm long,
3-6 (-7.5) cm wide, lanceolate, slightly curving, glabrous,
subcoriaceous, pale green, apex narrowly acuminate, base
rounded or slightly unequally lobed. Venation with 1-3 pairs

PIPER IN THE NEW WORLD

Fig. 27 Distribution of P. cadenaensis.

of secondary veins arising from the lower half of the promi-
nent midrib and ascending to apex, and numerous cross-veins
arising along the length of the midrib. Petioles 48 mm long,
glabrous. Prophylls 4-12 mm long, narrow, glabrous, apex
blunt. Inflorescences 3.5-7 cm long, erect; peduncles 4-8 mm
long. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.8 mm
wide, triangular, fleshy, densely ciliate. Fruit obovoid, 0.7-1
mm wide, trigonous, minutely pubescent or granular; stigmas
3, linear.

Forest-covered hillsides; 0-200 m.
DISTRIBUTION. Guatemala. Fig. 27.

Guatemala, Izabal: Contreras 9051 (MO); Peten: Lundell &
Contreras 19010 (MO).

This species superficially resembles P. schiedeanum, but can
be distinguished by its distinctive lanceolate leaves with
numerous cross-veins, shorter inflorescences, and pubescent
or granular fruits. It is confined to the drier lowland areas of
north-eastern Guatemala.

40. Piper regnellii (Miq.) C. DC. in DC., Prodr. 16(1): 307

(1869).
Fig. 29A, a.

Artanthe regnellii Miq. in Linnaea 22: 574 (1849). Type:

Brazil, prov. Minas Gerais, Caldas, Regnell II 256 (S-

holotype; C!, K!, US!-isotypes).
Piper epunctulatum C. DC. in DC., Prodr. 16(1): 338 (1869).

Type: Brazil, Minas Gerais, Lindberg 486 (M-holotype;

S-isotype).
P. pallescens C. DC. in Linnaea 37: 362 (1872). Type: Brazil,

Lagoa Santa. Warming s.n. (C!-holotype).
P. fulvescens C. DC. in Ann. N.Y. Acad. Sci. 7: 214 (1892).

Type: Central Paraguay, Morong 760 (NY-holotype;

K!-isotype).
P. fulvescens var. subglabrum C. DC. in Mem. Soc. Phys.

Hist. nat. Geneve 32: 8 (1893). Type: Paraguay, Villa Rica,

Balansa 2321 (G-holotype).
P. flabellinerve C. DC. in Bot. Jb. 65: 26 (1901). Type: Brazil,

Goias, Ule 461 (P-isotype).
P. fulvescens var. geraense C. DC. in Notizbl. hot. Gart. Mus.

219

Bed. 6: 436 (1917). Type: Brazil, Serra de Caldas, Mosen

1027 (S-holotype).
P. regnellii var. glabrius C. DC. in Candollea 1: 273 (1923),

nom. nud. (Art. 32.1).
P. regnellii var. parvifolium C. DC. in Candollea 1: 273

(1923), nom. nud. (Art. 32.1).
P. regnellii var. pallescens (C. DC.) Yuncker in Hoehnea 2:

172 (1972).

Shrub (1-) 2-4 m high, stems glabrous to shortly pubescent.
Leaves 9-25 cm long, 8-22 cm wide, pale green, glabrous to
minutely pubescent or velutinous, apex short-acuminate,
base shallowly to deeply cordate. Venation with 3-6 pairs of
secondary veins, mostly arising from the base, ascending at an
acute angle to the apex. Petioles 49 cm long, vaginate to
middle or above, sheathing the stems, glabrous to minutely
pubescent. Prophylls 2-3 cm long. Inflorescences 6-15 cm
long; peduncles 8-25 mm long. Anthers 0.5 mm long. Floral
bracts 0.7-0.8 mm wide, orbicular-triangular, fleshy, white-
ciliate. Fruit obovoid, 0.8-1 mm wide, glabrous; stigmas 3,
linear.

Moist places in forest shade, steep wooded slopes, boggy
ground, and stream sides; 400-1100 m.

DISTRIBUTION. Brazil to Argentina. Fig. 28.

Fig. 28 Distribution of P. regnellii.

Brazil, Distrito Federal: Prance & Silva 59025 (K); Goias:
Heringer 8467 (NY); Matto Grosso: Lankester s.n. (K);
Minas Gerais: Bareto 1816 (NY); Parana: Dusen 9620 (K,
NY); Rio de Janeiro: Glaziou 11577 (K); Santa Catarina:
Kummrow & Oliveira 2318 (MU); Sao Paulo: Hoehne &
Gehrt s.n. (NY). Paraguay, Central: Pederson 4238 (C);
Cordillera: Schwarz 11083 (BM); Misiones: Pedersen 6551

220

M. C. TEBBS

Fig. 29 A: P. regnellii, habit; a: bract and fruit. B: P. veraguense,
habit; b: part of inflorescence; c: bract and fruit.

PIPER IN THE NEW WORLD

(B,K); Paraguari: Sparre & Vervoorst 641 (K); San Pedro:
Woolston 1336 (K); Sierra de Maracayu: Hassler 4788 (BM).
Argentina, Itusaingo: Meyer 6263 (F); Santo Tome: Pedersen
2909 (C).

Piper regnellii grows in moist, partially shaded places in
gallery forest, mostly at about 1000 m altitude. It is one of the
few species of Piper with a distribution stretching as far south
as Argentina. The wide, usually shallowly cordiform leaves,
together with the 2-3 cm long prophylls, and partly sheathing
petioles, distinguish this species from other members of the
section. There is considerable variation in the depth of the
basal lobing of the leaves, often on the same individual.

41. Piper veraguense C. DC. in DC., Prodr. 16: 294 (1869).

Type: Costa Rica et Veragua, Warscevicz 1 (G-holotype;

MO !-photograph) .
Fig. 29B, b, c.

P. peltaphyllum C. DC. in Bull. Soc. r. Bot. Belg. 30 (1): 220

(1892). Type: Costa Rica, Route de Carillo, Pittier 2522

(G-holotype).
P. grandilimbum C. DC. in Bot. Gaz. 70: 172 (1920). Type:

Guatemala, Turckheim 8743 (US-holotype; F! -photograph).
P. albert-smithii Trel. & Yuncker, Piperac. N. South. Amer.

1: 413 (1950). Type: Colombia, vicinity of El Roble, Dept.

Santander, Killip & Smith 19360. (US-holotype; NY!-

isotype).
P. mutisii Trel. & Yuncker, Piperac. N. South Amer. 1: 414

(1950). Type: Colombia, Mutis 3463 (MA-holotype; ILL!-

photograph).
P. veraguense var. mutisii (Trel. & Yuncker) Steyerm. in Fl.

Venezuela 2: 590 (1984).
P. veraguense var. venezuelense Steyerm. in Fl. Venezuela 2:

221

590 (1984). Type: Venezuela, Trujillo, Breteler 4082 (VEN-
holotype).

Herb 1-2 m high, stems few, slender, erect, glabrous,
sometimes rooting at nodes. Leaves 16-34 cm long, 10-22
(-28) cm wide, peltate or subpeltate, hanging vertically from
petiole, ovate-lanceolate, coriaceous, glabrous or sparsely
pubescent on the undersides, tapering to long-acuminate
apex. Venation with 4-6 pairs of secondary veins arising from
or near petiole attachment, radiating outwards and curving
towards apex, and 2-3 pairs of secondary veins arising from
further up the midrib, curving towards apex. Petioles 9-22 cm
long. Prophylls 1.5-4 cm long, glabrous, apex blunt. Inflores-
cences 7-16 cm long, erect; peduncles 1-2 cm long. Anthers
0.2-0.3 mm long. Floral bracts 0.6-1 mm wide, triangular-
rhomboid, minutely ciliate, concealing fruit in early stages.
Fruit obovoid, trigonous, 1-1.2 mm wide, glabrous; stigmas
3.

Moist forest, shaded disturbed ground, and stream sides;
500-1900 m.

DISTRIBUTION. Mexico to Peru. Fig. 30.

Mexico, Chiapas: Breedlove 26096 (LAM, MO). Guatemala,
Alta Verapaz: Turckheim II 1063 (MO); Huehuetenango:
Steyermark 49216 (F); Peten: Steyermark 45372 (F). Honduras,
Atlantida: Standley 53357 (F). Nicaragua, Jinotega: Stevens
et al. 16531 (MO, BM). Costa Rica, Cartago: Lent 2745
(NY). Panama, Darien: Folsom 4299 (MO). Venezuela,
Merida: Nees & Whalen 16942 (VEN); Trujillo: Liesner et al.
12840 (MO, VEN). Peru, San Martin: Schunke 6258 (CAS).

Although the inflorescence of Piper veraguense is very simi-
lar to that of P. schiedeanum and P. corrugatum, the distinc-
tive peltate leaves, hanging vertically from the petioles, make
this species easily recognizable. Piper scutilimbum also has
peltate leaves, but these are oblong-elliptic, with abruptly
narrowed bases and long sheathing petioles; it also has much
smaller (0.4 mm) floral bracts than P. veraguense.

Steyermark (1984) divided Venezuelan P. veraguense into
three varieties. In his treatment, varieties mutisii and vene-
zuelense were separated from the type variety by the differing
degrees of pubescence on the veins of their leaves. This
pubescence can be found to a greater or lesser extent in many
specimens of P. veraguense outside Venezuela, and in this

Fig. 30 Distribution of P. veraguense.

Fig. 31 Distribution of P. perbrevicaule.

222

M. C. TEBBS

Fig. 32 A: P. perbre vicaule, habit; a: part of stem; b: part of inflorescence; c: bract and fruit. B: P. reptabundum, habit; d: part of

inflorescence; e: bract and fruit; f: fruit.

PIPER IN THE NEW WORLD

223

treatment the variation is considered insufficiently distinct to
maintain any division within the species.

42. Piper perbrevicaule Yuncker in Ann. Mo. hot. Gdn 37:
51, fig. 9 (1950). Type: Panama, vicinity of Chiriquf
Lagoon, Water Valley, Bocas del Toro, von Wedel 971
(MO!-holotype).

Fig. 32A, a, b, c.

P. perbrevicaule var. subglabrilimbum Yuncker in Ann. Mo.
hot. Gdn 37: 51 (1950). Type, Panama, Bocas del Toro,
von Wedel 1074 (MO!-holotype).

Terrestrial or epiphytic herbs, stems 15-50 cm long, prostrate
or trailing, often rooting at nodes, densely pubescent with
long multicellular hairs. Leaves 6-12 cm long, 2.5-5 cm wide,
asymmetrically oblong-elliptic to ovate-elliptic, often long-
villous on both sides, sometimes bullate, apex acute-acuminate,
base with one side lobed and overlapping petiole. Venation
with 34 pairs of secondary veins loop-connecting to apex.
Petioles 5-10 mm long, densely pubescent. Prophylls 8-12
mm long, sparsely to densely pubescent. Inflorescences 2-4
cm long, erect in early stages, greenish, white, pink or
reddish-purple; peduncles (1-) 2-5 cm long, slender, curving.
Anthers 0.3-0.4 mm long. Floral bracts 0.3-0.4 mm wide,
triangular, glabrous to minutely pubescent. Fruit obovoid,
0.8-1 mm wide, trigonous, glabrous or granular, with domed
apex; stigmas 3.

Moist places on floor of lower montane rain-forest, edges of
forest trails, and banks of streams; 0-700 m.

DISTRIBUTION. Costa Rica, Panama. Fig. 31.

Costa Rica, Alajuela: Grayum et al. 6305 (BM, MO); Heredia:
Grayum et al. 6155 (BM, MO). Panama, Bocas del Toro:
Wedel 1469 (MO); San Bias: Nevers et al. 5048 (MO, BM);
Veraguas: Croat 25648 (MO).

Piper perbrevicaule is a decumbent, succulent plant with
stems barely reaching 50 cm in length, creeping along the
banks of streams and growing on fallen logs, or occasionally
found as an epiphyte. It is easily recognized by its densely
pubescent stems, often bullate leaves, and inflorescences on
long, slender peduncles. The scandent P. reptabundum also
has long, slender peduncles and similarly shaped leaves, but
does not share the copious pubescence and the sometimes
bullate leaves of P. perbrevicaule. Long, slender peduncles
are also found in P. dolichotrichum, a shrubby climber, and in
P. urostachyum Hemsley of section Callianira.

A form of P. perbrevicaule from Costa Rica has peduncles
of only 1 cm or less, rather than the 2-5 cm normally found.
Although the specimens that have been seen are not fully
mature, they are herbs of the forest understorey, and display
the other characters found in this species, such as asymmetri-
cal leaves, long, multicellular hairs, minute, triangular bracts,
and glabrous or granular fruits.

43. Piper reptabundum C. DC. in Bot. Gaz. 70: 169 (1920).
Type: Costa Rica, Forets de Shirores, Talamanca, Limon,
Tonduz 9277 (G-holotype; F!, USMsotypes).

Fig. 32B, d, e, f.

Scandent, with glabrous stems. Leaves 10-18 cm long, 3-6 cm
wide, narrowly asymmetrically oblong-elliptic to oblong-
lanceolate, rough above, glabrous below, apex narrowly
long-acuminate, base sometimes slightly lobed on one side.

Fig. 33 Distribution of P. reptabundum.

Venation with 3-5 pairs of secondary veins arising from the
length of the midrib, loop-connecting to apex. Petioles 1-5
mm long, glabrous. Prophylls 5-10 mm long, slender. Inflor-
escences 5-10 cm long, greenish pink, curving and becoming
pendulous in fruit, on slender, glabrous peduncles 4-6 (-7)
cm long. Anthers 0.3-0.5 mm long. Floral bracts 0.5-0.8 mm
wide, broadly triangular, fleshy, glabrous or minutely ciliate.
Fruit obovoid, 0.8-1 mm wide, trigonous, minutely granular
above, with domed apex; stigmas 3, recurved.

Growing in moist places in forest shade, often near streams;
0-600 m.

DISTRIBUTION. Costa Rica. Fig. 33.

Costa Rica, Limon: Burger et al. 10400 (F, MO); San Jose:
Croat 35274 (MO).

Piper reptabundum is closely related to P. perbrevicaule, with
asymmetric leaves, curved or arching inflorescences on long
slender peduncles, and fruits with minutely granular surfaces.
It can be distinguished by its much longer, narrow leaves with
scarcely any basal lobing, absence of multicellular hairs, and
more robust habit. Piper dolichotrichum also has long,
slender peduncles, but can be distinguished from P. repta-
bundum by its vigorous climbing habit up to 7 m high, elliptic-
ovate leaves, long multicellular hairs, and bracts thickly
fringed with hairs. Unfortunately P. reptabundum has not
been widely collected, and only a few specimens from scat-
tered locations in Costa Rica have been seen. Although there
is some variation in leaf width between these populations,
they all share the distinctive leaf asymmetry, narrow, sharply
pointed apices, and inflorescences with long filiform peduncles.

44. Piper fortunaensis Tebbs, sp. nov. Type: Panama, vicinity
of Fortuna Dam, Chiriquf, McPherson 9832 (MO!-holotype).
Fig. 34.

Frutex 1-1.5 m altus. Folia elliptica asymmetrica glabra
glandulosa apice acuminata basin acuta vel cuneiformia.
Spica erecta, bracteis gibbosis triangularibus. Bacca trigona
apicem versus granulosa.

Spindly shrub 1-1.5 m high, stems glabrous. Leaves 14-21 cm
long, 3-8.5 cm wide, asymmetrically narrowly to broadly

224

M. C. TEBBS

Fig. 34 P. fortunaensis A: habit; a: prophyll; b: part of inflorescence; c: bract and fruit.

PIPER IN THE NEW WORLD

elliptic, glabrous, glandular, apex long-acuminate, base acute
to cuneate. Venation with 2-4 pairs of secondary veins arising
from the lower half of the midrib, and 1-2 pairs from the
upper half, curving to apex. Petioles 4-8 mm long, glabrous.
Prophylls \-b cm long, narrow, apex acute. Inflorescences 4-
8 cm long, erect, green or white; peduncles 5-10 mm long.
Anthers 0.2-0.3 mm long. Floral bracts 0.7-0.8 mm wide,
fleshy, oblong-triangular, gibbous, glabrous. Fruit obovoid,
1 mm wide, trigonous, upper surface granular, glandular;
stigmas 3.

Damp places in high forest; 600-1500 m.
DISTRIBUTION. Costa Rica, Panama. Fig. 35.

Costa Rica, Heredia: Grayum et al. 5067 (MO). Panama,
Bocas del Toro: Gordon 53D; McPherson 9667 (MO);
Chiriquf: D'Arcy 16358B; (MO); Veraguas: Antonio 3536
(MO).

Piper fortunaensis has similarly shaped leaves, bracts, and
fruits to P. reptabundum. It can be identified by its shrubby
habit, long narrow prophylls, and short peduncles. Its distinc-
tive, fleshy, glabrous bracts also serve to distinguish it from
those of P. aequale, which are ciliate. Most of the specimens
examined were collected from the Fortuna Dam area, which
has produced a number of interesting plants. It has been
recorded on one collection (Gordon 53D) that this plant is a
'fish-poisoning shrub.'

225

Fig. 35 Distribution of P. fortunaensis.

45. Piper dolichotrichum Yuncker in Amer. J. Bot. 39: 635
(1952). Type: Colombia. Narino, Gorgona Island, Collenette
524 (K!-holotype; US!-isotype).

Fig. 37A, a.

Shrubby liana 1-5 (-7) m high, occasionally epiphytic, stems
usually covered with multicellular hairs 1-3 mm long. Leaves
12-24 cm long, 6-12 cm wide, narrowly to widely elliptic-
ovate, upper surface with scattered hairs, undersurface
sparsely to densely pubescent, glandular, apex long-acuminate,
base unequally shallowly lobed. Venation with 5-7 pairs of
prominent secondary veins ascending sharply to apex. Petioles
3-12 mm long, densely long-pubescent. Prophylls 7 mm long,
narrow, with long multicellular hairs. Inflorescences 3.5-7 cm
long, pendulous, sometimes with a short sterile tip, greenish
white; peduncles 2-3 cm long, slender, densely covered in
multicellular hairs. Anthers 0.1-0.3 mm long. Floral bracts

Fig. 36 Distribution of P. dolichotrichum.

0.8 mm wide, lunulate to widely triangular, ciliate. Fruit
obovoid, 1-1.2 mm wide, trigonous, often glandular; stigmas
3, sessile.

In marshy or damp areas of moist forest; 0-800 m.
DISTRIBUTION. Costa Rica-Ecuador. Fig. 36.

Costa Rica, Heredia: Burger 8074 (F); Limon: Soils 23435
(BM); Puntarenas: Burger & Liesner 7310 (F). Panama,
Chiriquf: Him & Gordon 500 (MO); Veraguas: Maas &
Dressier 1643 (F). Colombia, Choco: Hugh-Jones 306 (K).
Ecuador, Esmeraldas: Mexia 8411 (BM).

Piper dolichotrichum is one of several climbing species in
Piper, and is capable of reaching 7 m in height. It is similar to
the shrubby P. urostachyum of section Callianira, which also
has long multicellular hairs, pendulous inflorescences on
slender peduncles, and long prophylls. However, the fruits of
P. dolichotrichum are obovoid, trigonous and lack a style,
whereas those of P. urostachyum are globose or hemispheri-
cal, and have a distinct elongated style. Sterile specimens of
P. dolichotrichum appear similar to the scandent P. brachy-
podon, which also has ovate-elliptic leaves with prominent
veins. However, P. brachypodon has glabrous, often macu-
late stems, and the leaves are larger and thicker than those of
P. dolichotrichum.

46. Piper subsessilifolium C. DC. in Bull. Soc. r. Bot. Belg.

30(1): 216 (1892). Type: Costa Rica, Rio Navarrito, Pittier

2435 (G-holotype, photograph!).
Fig. 39A, a, b, c.

P. flavirameum C. DC. in Bot. Gaz. 70: 181. (1920). Type:

Costa Rica, forets de la Palma, Pittier 12510 (G!-holotype).
P. arundinetorum Trel. in Contr. U.S. natn. Herb. 26: 159

(1929). Type: Costa Rica, near Laguna de la Escuadra, San

Jose, Standley 41967 (US-holotype).
P. concepcionis Trel. in Contr. U.S. natn. Herb. 26: 159

(1929). Type: Costa Rica, La Conception, Donnell Smith

6749 (US!-holotype).

226

M. C. TEBBS

a Vf

/ p

fk.'

.o *

Fig. 37 A: P. dolichotrichum, habit; a: bract and fruit. B: P. brachypodon, habit; b: bract and fruit.

PIPER IN THE NEW WORLD

P. pendens Trel. in Contr. U.S. natn. Herb. 26: 160 (1929).

Type: Costa Rica, Yerba Buena, San Isidro, Heredia,

Standley & Valeria 49822 (US-holotype, photograph!).
P. pendens var. infaustum Trel. in Contr. U.S. natn. Herb. 26:

160 (1929). Type: Costa Rica, El Muneco, S. of Navarro,

Cartago, Standley 33651 (US-holotype, photograph!).
P. sulcinervosum Trel. in Contr. U.S. natn. Herb. 26: 159

(1929). Type: Costa Rica, Cerro de la Carpintera, Cartago,

Standley 35569 (US-holotype).
P. flavirameum var. obscurum Trel. in Publs Field Mus. nat.

Hist. (Bot.) 18: 343 (1937). Type: Costa Rica, Brenes 20629

(F!-holotype).
P. debilicaule Trel. & Yuncker, Piperac. N. South. Amer. 1:

333 (1950). Type: Ecuador, Parroquia Conception, Playa

Rica, Esmeraldas, Mexia 8411 (US-holotype; BM!, K!-

isotypes).

Climbing or scrambling liana, 2-4 (-5) m high, stems glabrous
to minutely puberulent, zig-zagging. Leaves 13-28 cm long,
6-17 cm wide, dark green and lustrous above, membranous,
paler on underside, narrowly to widely elliptic-ovate, glab-
rous to minutely puberulent, apex acute-acuminate, base
narrowing to small, rounded, unequal lobes. Venation with
3-5 pairs of secondary veins arising from lower half of midrib,
ascending sharply to apex, prominent beneath. Petioles 3-10
mm long, pubescent. Prophylls 1-4 cm long, minutely pubes-
cent, sometimes glandular. Inflorescences 3.5-8 (-10) cm
long, erect, pink to purple, sometimes pendulous in fruit;
peduncles 5-12 mm long. Anthers 0.2-0.3 mm long. Floral
bracts 0.6-1 mm wide, triangular-lunulate, pink- to purple-
ciliate. Fruit obovoid, 0.8-1.5 mm wide, trigonous, with
minute style; stigmas 3.

Tropical wet forest, cloud and elfin forest, and damp shady
spots in land cleared for agriculture or by logging; 0-2200 m.

DISTRIBUTION. Costa Rica to Ecuador. Fig. 38.

Costa Rica, Alajuela: Lent 2815 (F); Cartago: Gomez 18872
(CR); Heredia: Garwood et al. 939 (BM); Limon: Davidson

Fig. 38 Distribution of P. subsessilifolium.

227

& Donahue 8827 (NY); Puntarenas: Haber 5726 (MO); San
Jose: Lent 3233 (F). Panama, Bocas del Toro: Croat 60165
(MO); Chiriqui: Hampshire & Whitefoord 18 (BM); Colon:
Dressier 4656 (MO); Darien: Kirkbride & Duke 1160 (MO);
Panama: Croat 22748 (F); Veraguas: Mori & Kallunki 2593
(MO). Colombia, El Valle: Killip 11814 (NY). Venezuela,
Yaracuy: Davidse et al. 20689 (MO). Ecuador, Esmeraldas:
Asplund 16304 (K); Imabura: Acosta Soils 12213 (F).

A vigorous liana climbing trees, shrubs, and tree-ferns, with
lateral branches often hanging downwards. The new leaves
and flowering spikes are often wine-red or maroon to violet,
and blend into the deep shade of the forest. The stem has a
spicy or pungent aroma when crushed. It is related to P.
xanthostachyum, but differs from this species by having a pink
or purple, sometimes pendulous inflorescence, as opposed to
a yellow- white and erect one, and larger, less angular floral
bracts fringed with pink to purple hairs. Piper concepcionis
has been reduced to synonymy here, as it is considered to be
merely a broad-leaved form of P. subsessilifolium. It has the
same type of leaf venation, and identical inflorescence charac-
ters, notably purple or pink colouring, bracts fringed with
pink hairs, and trigonous fruits.

47. Piper brachypodon (Benth.) C. DC. in DC., Prodr. 16(1):
327(1869).

Fig. 37B, b.

Artanthe brachypoda Benth., Bot. voy. Sulphur. 167, pi. 54
(1844). Type: Colombia, Gorgona Island, Hinds s.n. (K!-
holotype).

Straggling shrub, 1-5 m high, stems glabrous, red-streaked.
Leaves (13-) 17-29 (-32) cm long, 8-16 (-20) cm wide, widely
ovate-elliptic, glabrous, subcoriaceous, apex acute-acuminate,
base slightly unequal. Venation with 4-7 pairs of secondary
veins arising from lower to middle part of midrib, curving
towards apex, prominent on underside. Petioles 7-15 mm
long, glabrous. Prophylls 1-2 cm long, 8-12 mm wide.
Inflorescences 3-5 (-7) cm long, erect; peduncles 2-5 mm
long. Anthers 0.2-0.3 mm long. Floral bracts 0.8-1 mm wide,
widely triangular, sometimes with pellucid glands, minutely
ciliate. Fruit obovoid, 1 mm wide, apex domed, glabrous,
with prominent glands; stigmas 3, recurved.

Dense moist forest; 0-300 m.
DISTRIBUTION. Colombia. Fig. 40.

Colombia, Barbacoas: Triana 17; Gorgona Island: Collenette
714 (K).

Piper brachypodon is a glabrous plant, with maculate stems,
sub-coriaceous leaves with prominent veins, and glandular
fruits. It can be compared with both P. subsessilifolium and
P. dolichotrichum in leaf shape and venation, although it is a
straggling shrub, rather than a climber. Piper dolichotrichum
is copiously pubescent and has a slender inflorescence on a
long, thin peduncle, making it easily separable from P.
brachypodon when spikes are present. P. subsessilifolium can
be distinguished by its deep red to purple, often pendulous
inflorescences, while those of P. brachypodon are white or
green, and erect. Both species have broadly triangular bracts,
fringed with hairs.

48. Piper xanthostachyum C. DC. in An. Inst. fis.-geogr. C.
Rica 9: 169 (1897). Type: Costa Rica, Bois de la vallee du
Rio Tuis, Pittier 8158 (G-holotype; Pi-photograph).

228

M. C. TEBBS

Fig. 39 A: P. subsessilifolium, habit; a: variation in leaf shape; b: part of inflorescence; c: bract and fruit. B: P. xanthostachyum, habit; d: part

of inflorescence; e: bract and fruit.

PIPER IN THE NEW WORLD

Fig. 40 Distribution of P. brachypodon.

Fig. 39B, d, e.

P. matinanum C. DC. in An. Inst. fis.-geogr. C. Rica. 9: 170

(1897). Type: Costa Rica, Pittier 9759 (G-holotype; US-

isotype ; MO ! -photograph) .
P. bryogetum C. DC. in Bot. Gaz. 70: 175 (1920). Type:

Costa Rica, Tonduz 12939 (G-holotype; MO!-photograph).
P. opacibracteum Trel. in Contr. U.S. natn. Herb. 26: 148

(1929). Type: Costa Rica, N. of Tilaran, Standley & Valeria

45899 (US-holotype, photograph!).

Climbing or scrambling over low shrubs, or epiphytic, 1-2 m
high, stems glabrous or minutely pubescent. Leaves 10-18 cm
long, 3-8 cm wide, oblong to elliptic-lanceolate, glabrous on
both surfaces, apex acute-acuminate, base narrowly obtuse.

229

Venation with 3-4 pairs of secondary veins arising mainly
from the lower half of midrib, arcuate-ascending. Petioles 4-
10 mm long, glabrous. Prophylls 8-12 mm long, glabrous.
Inflorescences erect, yellow-white, 5-12 cm long; peduncles
5-12 mm long, glabrous. Anthers 0.2-0.3 mm long. Floral
bracts 0.3-0.5 mm wide, narrowly triangular to crescent-
shaped, densely yellow-white-ciliate. Fruit obovoid, 0.8-1
mm wide, trigonous, with pellucid glands; stigmas 3, linear.

Pre-montane rain-forest, wet evergreen forest, or moist
secondary forest; 0-2400 m.

DISTRIBUTION. Guatemala-Peru. Fig. 41.

Guatemala, Alta Verapaz: Standley 91383 (F); San Marcos:
Standley 68570 (F). El Salvador, Santa Ana: Molina 12525
(F). Honduras, Intibuca: Molina 13990 (F); Francisco
Morazan: Morton 7287 (F). Costa Rica, Alajuela: Lent 2653;
Cartago: Liesner 14429 (MO); Puntarenas: Liesner 2018
(MO); San Jose: Molina et al. 18074 (MO). Panama, Darien:
Kirkbride & Duke 1197 (MO). Ecuador, Pichincha: Cazalet
& Pennington 5086 (K). Peru, San Martin: Knapp & Mallet
8385 (K, MO).

The leaves of this species are similar in shape and venation to
those of Piper subsessilifolium, P. dolichotrichum, and P.
brachypodon, and it can be difficult to distinguish sterile
specimens of these taxa. However, the inflorescence charac-
ters of narrowly triangular to crescent-shaped bracts fringed
with yellow to white hairs, and glandular fruits, easily separ-
ate P. xanthostachyum from these species. Comments on
collectors' labels say that the leaves have a very spicy odour
like Lindera benzoin Meissner (Spice Bush) when first
crushed, or a lemon or lime fragrance. Burger (1971) united
P. matinanum and P. xanthostachyum under the latter name
and this choice should be followed (Art. 57.2).

49. Piper ottoniifolium C. DC. in /. Bot., Lond. 4: 213 (1866)
Type: Colombia, prov. del Choco, Triana 32 (G-holotype;
BM!-isotype).

Fig. 43 A, a.

Fig. 41 Distribution of P. xanthostachyum.

Fig. 42 Distribution of P. ottoniifolium.

230

M. C. TEBBS

Fig. 43 A: P. ottoniifolium, habit; a: bract and fruit. B: P. septuplinervium, habit; b: bract and fruit.

PIPER IN THE NEW WORLD

P. scleromyelum C. DC. in An. Inst. fis.-geogr. C. Rica 9: 167
(1897). Type: Costa Rica, Bois de Tsuritkub, Talamanca,
100 m, Tonduz 8675 (G-holotype; F!, MO!-photographs).

P. ledezmae Trel. & Yuncker, Piperac. N. South Amer. 1: 82
(1950). Type: Colombia, near Quibdo, Choco, Klllip &
Ledezma 35314 (US-holotype, photograph!).

P. ottoniifolium var. frutescens Trel. & Yuncker, Piperac. N.
South Amer. 1: 87 (1950). Type: Colombia, Klllip &
Cuatrecasas 39164 (US-holotype; F!-isotype).

P. ottoniifolium var. latifolium Trel. & Yuncker, Piperac. N.
South Amer. 1: 87 (1950). Type: Ecuador, Balao, Eggers
14434 (US-holotype).

Shrubby, scandent or epiphytic, stems glabrous or minutely
puberulent, aromatic. Leaves 9-21 cm long, (3-) 5-9 (-11) cm
wide, narrowly to broadly oblong-elliptic, coriaceous or
subcoriaceous, glossy above, pale green or brown beneath,
glabrous, glandular, apex acute-acuminate, occasionally
mucronate, base unequally obtuse to cuneate, sometimes
with very small lobes overlapping petiole. Venation with 2-3
(-4) pairs of prominent secondary veins arising from base, the
upper pair curving to apex, crossed with reticulate veins.
Petioles 2-10 mm long, glabrous. Prophylls 8-20 mm long,
glabrous. Inflorescences 5-11 cm long, erect, pale green,
often becoming pendulous in fruit; peduncles 6-10 mm long,
glabrous. Anthers 0.2-0.5 mm long. Floral bracts 0.5-0.8 mm
wide, sharply triangular to crescent-shaped, glandular, densely
yellowish- white-ciliate. Fruit obovoid, 1 mm wide, trigonous,
with prominent glands, apex conical; stigmas 3, linear.

Moist forest, and thickets by roads or tracks; 0-1300 m.
DISTRIBUTION. Costa Rica to Ecuador. Fig. 42.

Costa Rica. Limon: Gomez et al. 20390 (BM); Puntarenas:
Grayum & Hammel 5637 (BM, MO). Panama, Colon:
Dressier 3862 (MO); Darien: Mori & Kallunki 5468 (MO);
Panama: Busey 908 (NY); Veraguas: Gentry 6219 (F).
Colombia, Antioquia: Araque & Barkley 18c751 (BM);
Cauca: Cuatrecasas 14168 (F); Choco: Triana 32 (BM); Valle:
Cuatrecasas 15728 (F). Ecuador, near village of Bucay: Camp
3767 (BM); Carchi: Madison et al. 4578 (F).

The distinctive pale, coriaceous leaves with camptodromous
venation and the usually climbing or scandent habit readily
distinguish P. ottoniifolium from other species of Piper. It is
most similar to the shrubby P. septuplinervium, which also
has glossy, coriaceous leaves with similar venation. However,
P. septuplinervium can easily be identified by its inflorescence
characters of distinctive T-shaped bracts fringed with white
hairs, and eglandular fruits. Burger (1971) suggested that P.
scleromyelum might be synonymous with P. ottoniifolium,
differing only in having much broader and thicker leaves.
Only two specimens have been seen labelled as P. scleromye-
lum, both from Puntarenas in Costa Rica and both described
as shrubs or small trees 2-3 m high. In characters such as leaf
shape and venation, inflorescence, shape of floral bracts and
glandularity of fruits, these specimens are certainly identical
with P. ottoniifolium. As the original description and the
photograph of the type compare favourably with those of P.
ottoniifolium, P. scleromyelum has been placed here into
synonymy.

50. Piper septuplinervium (Miq.) C. DC. in DC., Prodr.

16(1): 313 (1869).
Fig. 43B, b.

231

Artanthe septuplinervia Miq. in Seemann, Bot. voy. Herald:

199, pi. 40 (1854). Type: Panama, without locality, Seemann

1066 (K-holotype; BM!-isotype).
Piper novo-granatense C. DC. in DC., Prodr. 16(1): 313

(1869). Type: Colombia, Barbacoas, Triana 30 (G-holotype;

BM!-isotype).
P. septuplinervium var. subpubinervium Yuncker in Ann.

Mo. bot. Gdn 37: 46 (1950). Type: Panama, Sutton Hayes

783 (NY!-holotype).

Slender shrub or small tree, 1-3 m high, stems glandular,
glabrous or occasionally minutely pubescent on younger
growth. Leaves 10-16 cm long, 5-8 cm wide, glabrous,
glandular, lustrous above, paler beneath, coriaceous or sub-
coriaceous, elliptic-lanceolate, apex acuminate, base slightly
unequally rounded. Venation with 3-5 pairs of secondary
veins arising from lower to middle part of midrib, curving
towards apex, with prominent cross-veins. Petioles 3-8 mm
long, glabrous. Prophylls 10-15 mm long. Inflorescences 3-6
(-8) cm long; peduncles 4-10 mm long. Anthers 0.2 mm long.
Floral bracts 0.6-0.8 mm wide, narrowly T-shaped, densely
white-ciliate. Fruit obovoid, 0.8-1 mm wide, trigonous, glab-
rous; stigmas 3, sessile.

Mangrove swamps, clearings along river banks, and forest
near coast; 0-400 m.

DISTRIBUTION. Panama-Ecuador. Fig. 44.

Fig. 44 Distribution of P. septuplinervium.

Panama, Colon: Pittier 2477 (G). Colombia, Altaquier,
Cordillera Central: Andre 3355 (K); Choco: Killip & Cuatre-
casas 39156 (US, BM); El Valle: Killip 34965 (BM). Ecuador,
Esmeraldas: Jativa & Epling 2212 (S).

Piper septuplinervium can be distinguished by its shrubby
habit, short inflorescence, and distinctive T-shaped bracts
with a dense fringe of white hairs. It has similar leaves to the
climbing P. ottoniifolium, with prominent secondary and
cross-venation and lustrous upper surfaces, but the inflores-
cences of P. ottoniifolium are longer, and the bracts are
crescent-shaped and fringed with yellowish white hairs. There
can also be some confusion with the leaves of the shrubby P.
carpunya, which are also glossy and coriaceous, but the bracts
of this species are fleshy, cupulate, and glabrous.

232

51. Piper coruscans Kunth in Humb., Bonpl. & Kunth, Nov.
gen. sp. 1: 53 (1816). Type: Crescit in ripa fluminis
Magdalenae, inter S. Bartholeme et Garapatas, Humboldt
1629 (B-?holotype; Fl-photograph).

Fig. 46A, a.

Schilleria coruscans (Kunth) Kunth in Linnaea 13: 708 (1839).
Steffensia pseudochurumayu Kunth in Linnaea 13: 662

(1839). Type: In Brazilia ad flumen Amazonum, Poeppig

3059 (B-holotype; G!-isotype).

Artanthe coruscans (Kunth) Miq., Syst. piperac.: 411 (1844).
A. pseudochurumayu (Kunth) Miq., Syst. piperac.: 407

(1844).
A. amazonica Miq. in Linnaea 20: 146 (1847). Type: Brazil,

Costa de Mattari, prov. Rio Negro, Martins s.n. (M-

holotype, ILL!-photograph).
Piper amazonicum (Miq.) C. DC. in DC., Prodr. 16(1): 272

(1869).
P. baryanum C. DC. in DC., Prodr. 16(1): 272 (1869). Type:

Brazil, Riedel 49 (L-holotype; K!-isotype).
P. pseudochurumayu (Kunth) C. DC. in DC., Prodr. 16(1):

288 (1869).
P. pseudochurumayu var. membranaceum C. DC. in DC.,

Prodr. 16(1): 288 (1869). Type: Ad flumina Casiquari,

Vasiva et Pamoni, Spruce 3433 (G!-holotype; K!-isotype).
P. orenocanum C. DC. in Bull. Herb. Boissier 6: 564 (1898).

Type: Venezuela, ad flumen Orenoco, Rusby & Squires

267 (NY-holotype; ILL!-photograph).
P. wurdackii Yuncker in Mem. N.Y. hot. Gdn 10(5): 41

(1964). Type: Venezuela, Cerro de la Neblina, Maguire et

al. 42007 (NY-holotype; VEN-isotype) .
P. coruscans var. membranaceum (C. DC.) Steyerm., Fl.

Venezuela 2: 383 (1984).

Shrub 1-4 m high, stems glabrous to minutely puberulent.
Leaves 10-24 (-30) cm long, 6-19 cm wide, ovate to ovate-
suborbiculate, membranous, lustrous and glabrous above
with pellucid glands often present, minutely puberulent on
veins below, sometimes rugose, apex acute-acuminate, base
round to subcordate. Venation with 5-8 pairs of secondary
veins mostly arising from the lower half of the midrib,
ascending steeply to apex, prominent on underside. Petioles
0.5-6 cm long, glabrous to minutely puberulent. Prophylls
2-3 cm long, glabrous. Inflorescences 7-15 cm long, erect,
pale green, yellow or white; peduncles 1-3 cm long. Anthers
0.3 mm long. Floral bracts 0.5 mm wide, triangular-orbicular,
densely ciliate. Fruits obovoid, 0.8-1 mm wide, suborbicular-
trigonous, with slight depression in centre, glabrous; stigmas
3.

Moist places in tropical evergreen forest, cliff bases and wet
steep slopes, stream and river banks; 0-1700 m.

DISTRIBUTION. Colombia to Brazil. Fig. 45.

Colombia, Choco: Molina & Barkley 19Ch048 (K); Popayan:
Hartweg 1398 (BM); Putumayo: Klug 1832 (BM); Santander:
Naught 1699 (BM); Valle: Cuatrecasas 16658 (F). Venezuela,
Amazonas: Nee 30828 (F); Apire: Davidse & Gonzalez 14070
(MO); Bolivar: Morillo & Liesner 8960 (MO); Casiquari:
Spruce 3433. Ecuador, Guayas: Camp 3586 (NY). Peru,
Loreto: Jones & Davidson 9534 (LAM); Madre de Dios:
Foster & Wachterl336 (F,MO). Bolivia, Morillo: Croat 51417
(MO); Pando: Nee 31730 (NY). Brazil, Acre: Steward et al.
P13209 (K); Mato Grosso: Berg et al. P19831 (K); Rondonia:
Prance et al. 5253 (K).

M. C. TEBBS

Fig. 45 Distribution of P. coruscans.

Piper coruscans can be distinguished by its distinctive ovate
to suborbiculate leaves with steeply ascending, prominent
secondary veins, and fruits that become suborbicular as they
mature. Steyermark (1984) was responsible for reducing
three species (P. amazonicum, P. pseudochurumayu, and P.
wurdackii) into the synonymy of P. coruscans, explaining that
the differences between them were not sufficient to maintain
them as separate taxa. There are some slight differences in
the shape of the leaf base, the degree of rugosity of the leaf
surface, and the quantity of pellucid glands present, but these
are within the range of variation for this species, and Steyer-
mark's view is followed here.

52. Piper scutilimbum C. DC. in Annu. Conserv. Jard. hot.
Geneve 21: 242 (1920). Type: Colombia, Santa Marta,
Smith 1235 (G-holotype; BM!, MO!, NY!, S!-isotypes).

Fig. 46B, b, c.

Thin-stemmed shrub or small tree, 1.5-4 m high, stems
glabrous. Leaves 16-27 (-31) cm long, 8-15 cm wide, oblong-
elliptic, glandular, apex acuminate, base abruptly narrowed,
cuneate or caudate, peltate. Venation with 6-10 pairs of
secondary veins arising along the length of the midrib,
arcuate-ascending, and 4-6 veins radiating around peltate
base. Petioles sheathing, 2-5 cm long. Prophylls not appar-
ent, if present hidden by sheathing petiole. Inflorescences
5-20 cm long, erect; peduncles 1.5-3.5 cm long. Anthers
0.2-0.3 mm long. Floral bracts 0.3-0.4 mm wide, triangular,
ciliate. Fruits obovoid, 0.8-1.2 mm wide, trigonous, apex
domed, glabrous or minutely granular; stigmas 3.

Tropical wet forest, cloud forest, and by sides of streams; 0-
2000m.

PIPER IN THE NEW WORLD

233

Fig. 46 A: P. coruscans, habit; a: bract and fruit. B: P. scutilimbum, habit; b: leaf with caudate base; c: bract and fruit.

234

Fig. 47 Distribution of P. scutilimbum.

DISTRIBUTION. Panama-Ecuador. Fig. 47.

Panama, Bocas del Toro: Kirkbride & Duke 620 (MO);
Chiriqui: Hamilton & Stockwell 3610 (MO); Colon: Knapp et
al. 4464 (MO, BM); Veraguas: Croat 27424 (MO). Ecuador,
Napo: Cerron 638 (MO); Jipa et al. DI959 (F).

This species has often been confused with Piper veraguense C.
DC., because both species have peltate leaves. P. scutilimbum
can be identified by its sheathing petioles and oblong-elliptic
leaves narrowing abruptly towards the peltate bases. Some of
the specimens from Ecuador (e.g. Cerron 638) have caudate
leaf bases 1-2 cm long and about 0.5 cm wide. P. veraguense
has ovate-lanceolate, peltate leaves without sheathing peti-
oles. The bracts of P. veraguense are 0.6-1 mm wide,
triangular-rhomboid and often cover the fruits in early stages,
while those of P. scutilimbum are minute, 0.3-0.4 mm wide,
and do not cover the fruits at any stage of development.

53. Piper auritum Kunth in Humb., Bonpl. & Kunth, Nov.

gen. sp. 1: 54 (1816). Type: Crescit in regni Novae Hispaniae

temperatis, Humboldt s.n. (P-holotype).
Fig. 48.

Schilleria aurita (Kunth) Kunth in Linnaea 13: 713 (1839).

Artanthe aurita (Kunth) Miq., Syst. piperac.: 400 (1844).

A. seemanniana Miq. in Seemann, Bot. voy. Herald: 199, t.39

(1854). Type: Panama, southern Darien, Seemann 1069

(BM!-holotype).
Piper perlongipes Trel. in Contr. U.S. natn. Herb. 26: 154

(1929). Type: Costa Rica, Pejivalle, Cartago, Standley &

Valeria 46834 (US-holotype, photograph!).
P. alstoni Trel. in Ann. Mo. hot. Gdn 27: 287 (1940). Type:

Panama, El Valle, Code, Alston & Allen 1842 (ILL-

holotype; MO!-isotype).
P. auritilimbum Trel. in Publs Field Mus. not. Hist. (Bot.) 9:

277 (1940). Type: Honduras, La Ceiba, Atlantida, Yuncker,

Koepper & Wagner 8763 (ILL-holotype; K!, MO!,

NY!-isotypes).
P. auritilaminum Trel. in Publs Field Mus. nat. Hist. (Bot.) 9:

277 (1940). Type: Honduras, La Ceiba, Atlantida, Yuncker,

Koepper & Wagner 8263 (ILL-holotype; NY!-isotype).
P. heraldi var. cocleanum Trel. in Ann. Mo. hot. Gdn 27: 292

M. C. TEBBS

(1940). Type: Panama, vicinity of El Valle, Code, Allen
1192 (ILL-holotype; MO!-isotype).

Shrub or small tree, 1-6 m high, stems glabrous. Leaves 18-
50 (-55) cm long, (11-)14-30 cm wide, oblong-elliptic, ciliate,
upper surface with soft scattered hairs, undersurface with
short hairs on veins, apex acute to obtuse, basal lobes deeply
cordate, with one lobe considerably lower than other, not
overlapping petiole. Venation with 4-6 pairs of secondary
veins mostly arising from the lower part of the midrib.
Petioles sheathing, 4-9 (-10) cm long, glabrous or with
minute hairs. Prophylls not apparent, if present hidden by
sheathing petiole. Inflorescences 15-30 (-37) cm long, erect,
often arching, white or cream; peduncles 4-11 cm long,
glabrous. Anthers 0.2-0.4 mm long. Floral bracts 0.5-0.8 mm
wide, orbicular or triangular, white-ciliate. Fruits 0.6-0.8
mm wide, obovoid, trigonous, glabrous; stigmas 3.

Moist soil in disturbed areas such as tracks and roadsides,
forest clearings, stream banks, landslides and edges of pas-
ture; 0-1200 (-2000) m.

DISTRIBUTION. Mexico-Colombia, West Indies. Fig. 49.

Mexico, Chiapas: Matuda 4743 (NY); Quintana Roo: Tellez
3074 (BM); San Luis Potosi: Pringle 3693 (MU); Tabasco:
Cowan 2334 (NY); Veracruz: Gentry & Lott 32620 (BM,
MO); Yucatan: Gaumer 23276 (MO). Belize, Cayo: Dwyer
12715 (MO); Corozal: Gentle 442 (LA); Stann Creek: Gentry
8004 (MO); Toledo: Whitefoord 1662 (BM). Guatemala, Alta
Verapaz: King 3314 (NY); Izabal: Jones & Facey 3219 (NY);
Peten: Ortiz 412 (NY); San Marcos: Croat 40796 (MO);
Retalhuleu: Cosminsky 114 (F). Honduras, Atlantida: Webster
& Miller 12685 (MO); Colon: Hagen 1363 (NY); Comayagua:
Yuncker, Dawson & Youse 6224 (MO); Cortes: Blackmore &
Charley 4004 (BM); Morazan: Trochez 45 (NY); Olancho:
Blackmore & Heath 1769 (BM); Yoro: MacDougal et al. 3216
(BM,MO). El Salvador, Sonsonate: Standley 19328 (NY).
Nicaragua, Bluefields: Proctor et al. 27282 (NY); Chinandega:
Baker 85 (NY); Chontales: Pipoly 1589 (CAS); Granada:
Guzman et al. 1705 (BM); Jinotega: Grijalva & Araquistain
229 (BM); Matagalpa: Stevens 11991 (M); Rio San Juan:
Araquistain 3208 (BM); Zelaya: Araquistain 3035 (BM).
Costa Rica, Alajuela: Webster & Miller 12210 (MO); Cartago:
Croat 567 (MO); Heredia: Bawa 613 (MO); Limon: Davidson
6867 (LA); Puntarenas: Raven 22032 (DS); San Jose: Tonduz
2386 (CM). Panama, Bocas del Toro: Blum 1341 (MO);
Canal Zone: Croat 5231 (NY); Chiriqui: Liesner 5 (MO);
Code: Correa 55 (MO); Colon: Nee 7273 (MO); Darien:
Garwood et al. 172 (BM); Panama: Carrasquilla et al. 224
(MO); San Bias: Sugden 535 (MO); Veraguas: Folsom &
Edwards 3403 (BM). West Indies, Jamaica: Houston s.n.
(BM). Colombia, Antioquia: Scolnik et al. 19An332 (BM);
Choco: Killip & Garcia 33550 (BM); El Valle: Pennell et al.
8578 (K, NY, US); Santander: Haught 1806 (BM).

Piper auritum is one of the most common New World species
of Piper, growing at the edges of woods or in clearings and
open sites. Its distinctive large, deeply-lobed leaves with
sheathing petioles, and pale arching inflorescences make it
easy to recognize. When crushed, the stems and leaves emit a
strong smell of aniseed. Sterile specimens have sometimes
been confused with species such as P. obliquum of section
Macrostachys, which also has sheathing petioles. However,
the inflorescence of P. auritum is slender and erect rather
than substantial and pendulous as in P. obliquum, and the

PIPER IN THE NEW WORLD

Fig. 48 P. auritum A: habit; a: part of leaf showing pubescent margin; b: bract and fruit.

236

M. C. TEBBS

Fig. 49 Distribution of P. auritum.

fruits are small and trigonous. The leaves of P. auritum have a
distinct margin of short white hairs, a character that is shared
with P. marginatum of section Radula. Piper auritum also has
other characters in common with this species, such as sheath-
ing petioles and arching, white or cream inflorescences. The
fruits of P. marginatum, dark in colour and round from above
with a raised rim, are considerably different from those found
in section Churumayu. Arching inflorescences are also found
elsewhere in section Radula, e.g. P. aduncum L. The vigor-
ous climber, P. multiplinervium C. DC., also in section
Radula, has sheathing petioles and similar fruits to those of P.
marginatum. P. auritum displays characters linking sections
Churumayu and Radula.

Excluded species

Piper abutiloides (Kunth) C. DC. in DC., Prodr. 16 (1):329

(1869).

This species compares well vegetatively with P. regnellii
(Miq.) C. DC. However, Kunth's original description of P.
abutiloides (as Schilleria abutiloides) emphasized that the
fruits were not available ('haud suppetunt'). The only speci-
men seen for this study (not the type), identified as P.
abutiloides, has an immature inflorescence. Without the pres-
ence of mature inflorescences, it is impossible to make a
definite statement regarding the combination of P. abutiloides
and P. regnellii, especially as P. abutiloides would take
precedence as the earliest published name.

ACKNOWLEDGEMENTS. My grateful thanks are due to Mr A. O.
Chater, Dr C. Jarvis, and Mr J. R. Press of the Department of

Botany, The Natural History Museum, and Dr A. J. Bornstein,
Assistant Professor of Biology, Southeast Missouri State University,
for advice and helpful comments. I would also like to thank the
following herbaria for the loan of their specimens-B, C, CAS, E, F,
ILL, K, MO, MU, S, and US.

REFERENCES

Burger, W. C. 1971. Piperaceae. Flora costaricensis. Fieldiana Bot. 35: 5-227.

1972. Evolutionary trends in the Central American species of Piper

(Piperaceae). Brittonia 24(4): 356-362.

Candolle, C. de. 1869. Piperaceae. In A. P. de Candolle, Prodromus systematis

natural is regni vegetabilis 16(1): 235471.

1902. Piperaceae. In I. Urban, Symbolae antillanae 3: 161-274.

Dietrich, A. 1831. Species plantarum 1: 173.

Gomez I'ompa, A. 1966. Estudios botanicas en la region de Misantla, Veracruz:

140-152. Mexico City.

Miquel, F. A. 1844. Sy sterna piperacearum. Rotterdam.
Steyermark, J. A. 1984. Flora de Venezuela. Fundacion para la Educacion

Ambiental.
Tebbs, M. C. 1989a. Revision of Piper (Piperaceae) in the New World 1.

Review of characters and taxonomy of Piper section Macrostachys. Bull. Br.

Mus. not. Hist. (Bot.) 19: 117-158.

19896. The climbing species of New World Piper (Piperaceae). Will-

denowia 19: 175-189.

Trelease, W. 1927. Piperaceae of Panama. Contr. U.S. natn. Herb. 26(2): 15-

50.

1929. Piperaceae of Costa Rica. Contr. U.S. natn. Herb. 26(4): 115-226.

Trelease, W. & Yuncker, T. G. 1950. Piperaceae of northern South America 1:

1-434. Urbana.
Yuncker, T. G. 1972. Piperaceae of Brazil. Piper Groups 1-4. Hoehnea 2:

23-366.
1973. Piperaceae of Brazil. Piper Group 5. Hoehnea 3: 29-284.

PIPER IN THE NEW WORLD

237

INDEX

Accepted names are in roman and synonyms in italic, new names and principal
references are in bold. An asterisk (*) denotes a figure.

Artanthe 194

Artanthe adenophylla Miq. 200

A. amazonica Miq. 232

A. aurita (Kunth) Miq. 234

A. benthamiana Miq. 213

A. brachypoda Benth. 227

A. colubrina (Link) Miq. 200

A. colubrina var. crassifolia Miq. 200

A. concinna Miq. 204

A. coruscans (Kunth) Miq. 232

A. denudans Miq. 209

A. glabella Miq. 200

A. grandifolia (Kunth) Miq. 209

A. jussiaeana (Kunth) Miq. 209

A. leptura (Kunth) Miq. 209

A. meyeniana Klotzsch ex Miq. 209

A. mikaniana (Kunth) Miq. 206

A. obovata (Kunth) Miq. 200

A. pseudochurumayu (Kunth) Miq. 232

A. regnellii Miq. 219

A. riparia (Kunth) Miq. 209

A. seemanniana Miq. 234

A. septuplinervia Miq. 231

A. subaequalis C. Presl 200

A. tejucensis Miq. 200

A. xylosteoides (Kunth) Miq. 203

section Callianira 223, 225
Carpunya lessertiana (Miq.) Presl 208
C. peruviana C. Presl 208
section Churumayu 194, 195, 236

Lindera benzoin Meissner 229
section Macrostachys 194, 195, 234

Ottonia carpunya (Ruiz Lopez & Pavon) Miq. 208
O. lessertiana Miq. 208

Peltobryon attenuatum Klotzsch ex Miq. 200

Peperomia 200

P. obversa (M. Vahl) A. Dietr. 200

P. obovata (M. Vahl) A. Dietr. 200

Piper abutiloides (Kunth) C. DC. 236

Piper acutissimum Trel . 197

P. acutissimum var. trichopus (Trel.) Yuncker 197

P. aequale M. Vahl 194, 195, 196, 210*, 211, 212, 212*, 213, 217, 225

P. aequale var. elliptico-lanceolatum C. DC. 211

P. albert-smithiilreA. & Yuncker 221

P. alstoni Trel. 234

P. altevaginansTrel. 197

P. amazonicum (Miq.) C. DC. 232

P. apiculatum C. DC. 200

P. arboreum Aublet 195, 197

P. arieianum C. DC. 197

P. arundinetorum Trel. 225

P. atrichopus Trel. 211

P. atrichopus var. arborescens Trel. 215

P. atroglandulosom Tebbs 194, 195, 196, 197, 199, 200, 201*, 202, 202*

P. auritilaminum Trel. 234

P. auritilimbum Trel . 234

P. auritum Kunth 194, 195, 196, 234, 235*, 236*

P. baryanum C. DC. 232

P. belemense C. DC. 200

P. benthamianum (Miq.) C. DC. 213

P. borucanum C. DC. 209

P. brachypodon (Benth) C. DC. 194, 195, 196, 225, 226*, 227, 229, 229*

P. bryogetum C. DC. 229

P. cabagranum C. DC. 211

P. cadenaensis Tebbs 194, 196, 218, 218*, 219*

P. caeruleifolium Trel. 211

P. caripense Kunth 21 1

P. carpunya Ruiz Lopez & Pavon 195, 1%, 207*, 208, 208*, 231

P. carnosicauleTre\. 212

P. carrilloanum C. DC. 215, 217

P. casimirianum Hemsley 215, 217

P. cercidiphyllum Trel. 209

P. cheyennense Trel. & Standley 211

P. chiriquinum C. DC. 211

P. churumayu Poeppig 195

P. cocherenseTre\. 208

P. colemanense Trel. 211

P. collinum var. flagellicuspe Trel. 215

P. colubrinum Link 200

P. concepcionis Trel. 225, 227

P. concinnatoris Yuncker 204

P. concinnifolium Trel. 211

P. concinnum (Miq.) C. DC. 204

P. corrugatum Kuntze 195, 196, 213, 214*, 215*, 217, 221

P. coruscans Kunth 194, 195, 196, 232, 232*, 233*

P. coruscans var. membranaceum (C. DC.) Steyerm. 232

P. costaricense C. DC. 211

P. crassifolium (Miq.) C. DC. 200

P. crassum Veil. Cone. 200

P. crispans Trel. ex Standley 21 1

P. debile C. DC. 206

P. debilicaule Trel. & Yuncker 227

P. densispicatum Steudel 200

P. denudans (Miq.) C. DC. 209

P. denudans var. angustifolium C. DC. 209

P. dilatatum C. DC. 195

P. divaricatum Meyer 196, 200, 201*, 203*

P. dolichotrichum Yuncker 194, 195, 196, 223, 225, 225*, 226*, 227, 229

P. donnell-smithii C. DC. ex J.D. Smith 211

P. dumosum Rudge 1%, 201*, 202, 203*

P. dupasquieri C. DC. 209

P. epilosipes Trel. 211

P. epunctulatum C. DC. 219

P. exigens Trel. 215

P. exigens var. subovatum Trel. 215

P. flabellinerve C. DC. 219

P. flavirameum C. DC. 225

P. flavirameum var. obscurum Trel. 227

P. fortunaensis Tebbs 194, 1%, 223, 224*, 225, 225*

P. frioense Standley & Steyerm. 21 1

P. fulvescens C. DC. 219

P. fulvescens var. geraense C. DC. 219

P. fulvescens var. subglabrum C. DC. 219

P. fuscum C. DC. 209

P. generalense Trel. 197

P. glabellum (Miq.) C. DC. 200

P. glabrilimbum C. DC. 200

P. glanduligerum C. DC. 213

P. gorgonae-insulae Trel. & Yuncker 197

P. glaziovii C. DC. 209

P. grande M. Vahl 194, 195, 196, 207*, 208, 209, 209*

P. grandifolium Kunth 208

P. grandilimbum C. DC. 221

P. guanacostense C. DC. 194, 195, 197, 197*, 198*

P. guedesii C. DC. 202

P. heraldi var. cocleanum Trel. 234

P. heterophyllum Ruiz Lopez & Pavon 200

P. imberbe Trel. & Standley 196, 202, 204*, 207*

P. impunctatum Link 200

P. insipiens Trel. & Yuncker 202

P. jericoense Trel. & Yuncker 21 1

P. johnstom C. DC. 211

P. jussiaeanum Kunth 209

P. lanceifolium Kunth 195

P. ledezmae Trel. & Yuncker 231

P. leiophyllum C. DC. 200

P. lenticellosum C. DC. 208

P. lepturum (Kunth) C. DC. 196, 209, 210*, 211, 211*

P. lepturum var. angustifolium (C. DC.) Yuncker 209

P. lessertianum (Miq.) C. DC. 208

P. levilimbum Trel. 215

P. marginatibaccum Trel. ex Standley 197

P. marginatum Jacq. 195, 236

P. marmoreum Trel. 209

P. matinanum C. DC. 229

P. megalophyllum C. DC. 215, 217

P. megalophyllum var. connivens C. DC. 215

P. micranthera C. DC. 211

P. middlesexense Trel. 215, 217

P. mikanianum (Kunth) Steudel 194, 196, 204, 205*, 206, 208

P. mikanianum f . clausum Yuncker 206

P. mikanianum var. peltatum Yuncker 206

P. misionesanum C. DC. 206

P. multiplinervium C. DC. 236

238

M. C. TEBBS

P. mutisii Trel. & Yuncker 221

P. nemorense C. DC. 213

P. nitidum M. Vahl 200, 202

P. novo-granatense C. DC. 231

P. nudicauleC. DC. 212

P. obliquum Ruiz Lopez & Pavon 234

P. oblongilimbum C. DC. 202

P. obovatum M. Vahl 200

P. obovatum Ruiz Lopez & Pavon 194, 200

P. obversum M. Vahl 200

P. omega Trel. 215

P. opacibracteum Trel. 229

P. oppressum Trel. 211

P. orenocanum C. DC. 232

P. ottoniifolium C. DC. 194, 195, 196, 229, 229*, 230*

P. ottoniifolium var. frutescens Trel. & Yuncker 194, 231

P. ottoniifolium var. latifolium Trel. & Yuncker 194, 231

P. pablense1re\.2\\

P. pabstii C. DC. 204

P. pallescens C. DC. 219

P. partiticuspe Trel. 213

P. parthenium C. DC. 206

P. parthenium var. pilosius C. DC. 206

P. paso-anchoenseTrel. 211

P. paulownifolium C. DC. 215

P. peltaphyllum C. DC. 221

P. pendens Trel. 227

P. pendens var. infaustum Trel. 227

P. perbrevicaule Yuncker 195, 196, 221*, 222*, 223

P. perbrevicaule var. subglabrilimbum Yuncker 223

P. pergamentifolium Trel. & Standley 215

P. perlasense Yuncker 21 1

P . perlongipes Trel. 234

P. portachueloense Trel. & Yuncker 197

P. pseudochurumayu (Kunth) C. DC. 232

P. pseudochurumayu var. membranaceum C. DC. 232

P. pseudo-variabile TTe\. 209

P. pseudo-variabile var. collium Trel. 209

P. pseudo-variabile var. pachypus Trel. 209

P. regnellii (Miq.) C. DC. 194, 195, 196, 219, 219*, 220*, 221, 236

P. regnellii var. glabrius C. DC. 219

P. regnellii var. pallescens (C. DC.) Yuncker 219

P. regnellii var. parvifolium C. DC. 219

P. reptabundum C. DC. 195, 196, 222*, 223, 223*, 225

P. riitoense Trel. & Yuncker 197

P. riparense C. DC. 213

P. riparium Kunth 208

P. rubripes Trel. ex Standley 211

P. rupununianum Trel. & Yuncker 196, 198*, 199, 200, 200*

P. saltuum C. DC. 197

P. sarapiquinum C. DC. 204

P. schiedeanum Steudel 194, 195, 196, 212, 213, 216*, 217, 217*, 219, 221

P. schlimii C. DC. 197

P. scleromyelum C. DC. 231

P. scutilimbum C. DC. 194, 195, 196, 221, 232, 233*, 234*

P. seducentifolium Trel. 197

P. seductum Trel. 21 1

P. septuplinervium (Miq.) C. DC. 194, 196, 230*, 231, 231*

P. septuplinervium var. subpubinervium Yuncker 231

P. sievekingii C. DC. 209

P. spissinervium Trel. 209

P. subaequale (C. Presl) C. DC. 200

P. subdurum Trel. ex Standley 208

P. subrepens Trel. 194, 197, 204, 205*, 206, 206*

P. subsessilifolium C. DC. 194, 195, 1%, 225, 227, 227*, 228*, 229

P. subvariabile Trel. 209

P. sulcinervosum Trel. 227

P. tejucense (Miq.) C. DC. 200

P. tenuifolium C. DC. 213

P. tenuispicum C. DC. 211

P. tiliifolia Schldl. & Cham. 213

P. trichopus Trel. 197

P. trigonum C. DC. 195, 196, 197, 198*, 199, 199*

P. tsakianum C. DC. 213

P. urophyllum C. DC. 195, 196, 204, 205*, 206, 206*, 208

P. urostachyum Hemsley 223, 225

P. vallicolum C. DC. 215

P. variabile C. DC. ex J. D. Smith 215, 217

P. variabile var. obtentum Trel. 215

P. veraguense C. DC. 194, 195, 196, 209, 213, 217, 220*, 221, 221*, 234

P. veraguense var. mutisii (Trel. & Yuncker) Steyerm. 221

P. veraguense var. venezuelense Steyerm. 221

P. verruculosum C. DC. 194, 195, 196, 210*, 212, 213, 213*

P. wurdackii Yuncker 232

P. xanthostachyum C. DC. 194, 195, 196, 227, 228*, 229, 229*

P. xylosteoides (Kunth) Steudel 197, 203, 204, 204*, 205

P. zacuapanum C. DC. 215, 217

P. zarceroense Trel. ex Standley 215

P. zingiberinum Trel. ex Standley 212

section Radula 195, 236

Schilleria abutiloides Kunth 236

5. aurita (Kunth) Kunth 234

5. colubrina Link ex Kunth 200

5. coruscans (Kunth) Kunth 232

5. grandifolia (Kunth) Kunth 209

S. jussiaeana Kunth 209

5. leptura Kunth 209

S. mikaniana Kunth 206

5. obovata Kunth 200

5. riparia (Kunth) Kunth 209

5. xylosteoides Kunth 203

Steffensia pseudochurumayu Kunth 232

OBITUARY
Ivan Mackenzie Lamb 1911-1990

239

Ivan Mackenzie Lamb was born in London on 10 September 1911 and died in
Massachusetts, U.S.A., on 28 January 1990 after a long illness. He was educated
at the University of Edinburgh, where he obtained a Ph.D. for research in
mycology. In the mid-1930s three additional posts of Assistant Keeper were
created in the Department of Botany at the British Museum (Natural History),
London. Lamb was appointed on 14 January 1935 to the first of these, which was
for work on lichens. The other two of these posts were for work on flowering
plants, filled later in the same year by W. R. Philipson, and on diatoms, filled by
myself in October 1936. 1 was, in consequence, a colleague of Lamb's from then
until mid- 1940 when I was transferred to other work for the duration of the war.
I cannot say, though, that I came to know him well. He was a quiet and reserved
person, always perfectly amiable and polite and never difficult to deal with, but
not inclined to chat. Although never abrupt or dismissive, he was obviously
anxious to get back to his work as soon as the matter that had caused one to
interrupt him had been dealt with. In this he was a great contrast to his nearest
neighbour in the Cryptogamic Herbarium, Geoffrey Tandy.

Lamb was a conscientious objector and continued to work in the museum
until 20 September 1943. During this period he completed a monograph on
Neuropogon, especially its Antarctic species, and followed this by work on
Placopsis, another genus with many Antarctic representatives. This brought him
in touch with J. W. S. Marr, a biologist with Discovery Investigations, who

became commander of Operation Tabarin, an expedition to the Antarctic mounted to safeguard British interests in that area.
Marr recruited Lamb as the botanist on this expedition and he spent two years in the Antarctic, one at Port Lockroy and one at
Hope Bay. It is clear from the obituaries in The Daily Telegraph (20 February 1990) and The Independent (5 March 1990), both
obviously written by other members of the expedition, that Lamb earned the respect and affection of his companions not only
for his personality but also for his physical toughness. For his service on this expedition, Lamb was awarded the Polar Medal in
silver with Antarctic clasp.

Only about a year after his return to the museum, Lamb resigned in 1946 and accepted the post of Professor of
Cryptogamic Botany at Tucuman University in Argentina in 1947. He had by then married and his wife suffered badly from
arthritis. He thought that the change in climate would help to alleviate this. In 1950, however, he moved to the Canadian
National Museum, Ottawa, as curator of cryptogams. This, too, was a post that he held for only three years, for in 1953 he
became Director of the Farlow Herbarium of Harvard University at Cambridge, Mass., U.S.A., a position he held until he
left in 1972. Afterwards he changed his name to Hike Mackenzie. During these years he continued his work and varied
correspondence on lichens and published his monumental Index Nominum Lichenum (1963), Antarctic lichens (1964,
1968), and monographs of Leprocaulon (1974, with A. Ward) and Stereocaulon (1977, 1978). Soon after his retirement he
also did some work on marine algae of the New England coast, and published short papers on these. All Lamb's
publications have a very high reputation. He will be long remembered for his dedicated, if isolated, research on Antarctic
matters.

R. Ross

British Museum (Natural History)
Publications

FLORA MESOAMERICANA

Flora Mesoamericana, a joint project of the Institute de Biologia, Universidad Nacional Autonoma de Mexico,
Missouri Botanical Garden, St. Louis, and The Natural History Museum, London, will start in 1991 with the
publication of Vol 6, covering half of the monocotyledonous families. The entire work will be in 7 volumes
published in Spanish at approximately 2-yearly intervals. It will comprise a concise identification manual for the
vascular plants of Mesoamerica. The work incorporates much new research and will form a data base for the
floristics and taxonomy of the region.

A Preliminary Bibliography of the Mesoamerican Flora

R. J. Hampshire, D. A. Sutton

1988, 245x1 75mm, 194pp.

565 01068 9 Paperback 15.00

THE CORALLINE RED ALGAE
An analysis of the genera and subfamilies of the nongeniculate Corallinaceae

W. J. Woelkerling

Nongeniculate coralline red algae (Corallinales, Rhodophyta) are both abundant in and vital to coral reefs,
cementing the coral animal skeletons into a durable reef framework. In colder waters they form pavements of
many square kilometres on the sea bed. It is one of the aims of this study to help resolve outstanding difficulties
relating to taxonomy of the group. Extremely well illustrated with over 60 pages of scanning electron
micrographs.

1988, 245 x 176mm, 268pp, 82 scanning electron micrographs, 259 figs, 19 tables.

Co-published with Oxford U.P.

1 98 54249 6 Hardback 40.00

Taxonomy and Biogeography of Macquarie Islands Seaweeds

R. W. Richer.

1987, 175 x 250mm, 344pp, 126 figs.

565 00998 2 Hardback 40.00

SEAWEEDS OF THE BRITISH ISLES

Vol. 1. Part 1. Rhodophyta. Introduction, Nemaliales, Gigartinales. P. S. Dixon & L. M. Irvine.

1977,xi + 252pp,90figs.

565 0078 1 5 Paperback 15.00

Vol. 1. Part 2A. Cryptonemiales (sensu stricto), Palmariales, Rhodymeniales. L. M. Irvine.

1983,xii + 115pp, 28 figs.

565 0087 1 4 Paperback 15.00

Vol. 3. Part 1. Fucophyceae (Phaeophyceae). R. L. Fletcher.

1987, 145 x 210mm, 359pp,90figs, 15 plates.

565 00992 3 Paperback 30.00

Vol. 4. Tribophyceae (Xanthophyceae). T. Christensen.

1987, 145 x 210mm, 36pp, 8 figs.

565 00980 X Paperback 7.50

Vol. 2. Chlorophyta. E. M. Burrows.

To be published in 1991 . Details to be announced.

CONTENTS

1 53 The marine algal flora of Namibia: its distribution and affinities

George W. Lawson, Richard H. Simons & William Edwin Isaac

169 The infragenic classification of Gent/ana (Gentianaceae)

Ting-Nung Ho & Shang-Wu Liu

193 Revision of P/per (Piperaceae) in the New World
2. The taxonomy of Piper section Churumayu

Margaret Cecilia Tebbs

lulletm british Museum
BOTANY SERIES

Vol.20, December 1 990