Bulletin of the

British Museum (Natural

BRITISH MUSEUM
(NATURAL HISTORY)

25FEBS988

PRESENTED
GENERAL LIBRARY

istory)

Botany series Vol 16 1987

British Museum (Natural History)
London 1987

Dates of publication of the parts

Nol 29 January 1987

No 2 25 June 1987

No 3 27 August 1987

No 4 26 November 1987

\

ISSN 0068-2292

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

Contents
Botany Volume 16

Page

No 1 Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)
N. K. B. Robson 1

No 2 The lichen genus Ramalina in Australia

G. N. Stevens 107

No 3 An annotated list of vascular plants collected in the valleys south of
Mt Everest
G. Miehe 225

No 4 Further genera of the Biddulphiaceae (diatoms) with interlocking
linking spines
R. Ross and P. A. Sims 269

> 11 4. f*1~

Bulletin of the

British Museum (Natural History)

Studies in the genus Hypericum L.

(Guttiferae)

7. Section 29. Brathys (part 1)

Norman K. B. Robson

Botany series Vol 16 No 1 29 January 1987

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

Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum's resources. Many of the papers are
works of reference that will remain indispensable for years to come.

Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW75BD,
England.

World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)

Trustees of the British Museum (Natural History), 1987

The Botany series is edited in the Museum's Department of Botany
Keeper of Botany: Mr J. F. M. Cannon

Editor of Bulletin: Mr J. R. Laundon

Assistant Editor: Dr A. J. Harrington

Editor's Assistant: Miss M. J. Short

ISBN 565 08014 8
ISSN 0068-2292

British Museum (Natural
Cromwell Road
London SW7 5BD

v
2 9 JAN

X

I Botany series
7 Vol 16 No 1 pp 1-106

Issued 29 January 1987

(NATURAL H1S T ORY)

Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)

Norman K. B. Robson

Department of Botany, British Museum (Natural History), Cromwell Road, London SW7
5BD

Contents

Synopsis 1

1 . Introduction 2

Morphology 3

(a) Leaves 3

(b) Inflorescences 5

(c) Flowers and fruits 5

Cytology and hybrids 7

Distribution and evolution 7

(a) Wide disjunctions 7

(b) Species with leaves persistent or deciduous near the base

(Spp. 3-23) 9

(c) Species with leaves with sheathing leaf-bases or marginal

secretions (Spp. 24-26,61) 10

(d) Species with incurved-cucullate leaves but no marginal

secretions (Spp. 27-30) 10

(e) Species with sessile leaves deciduous above the base but

not incurved-cucullate (Spp. 31p.p.-48) 11

(f) Species with pseudopetiolate leaves deciduous above the base

but not incurved-cucullate (Spp. 31p.p. , 49-60) 12

2. Systematic treatment 12

3 . Acknowledgements 103

4. References 104

5. Systematic index 104

Synopsis

A systematic account is given of the 61 shrubby species of Hypericum sect. 29 Brathys, which occur from
Belize and the Greater Antilles to Peru and Bolivia. A discussion of the morphology, chromosome number
(only one known), distribution, and evolution of the members of the section is included.

Sect. 30 Spachium sensu Robson (1977) is shown to comprise three distinct groups: (i) sect. Spachium
sensu stricto (Central America, Greater Antilles, eastern U.S.A.), which is included in sect. 29. Brathys;
(ii) sect. Knifa (Adanson) N. Robson, stat. nov. (S. American Cordillera, Central and North America,
Cuba, tropical and southern Africa, east and south-east Asia, Australia); (iii) sect. Trigynobrathys (Y.
Kimura) N. Robson, comb. nov. (S. America south of Amazonia and along the Cordillera, Australasia,
south-east Asia).

The following other new taxa (sp. nov. or subsp. nov.) are described: H. phellos subsp. oroqueanum N.
Robson (Colombia: Norte de Santander/Cesar), H. acostanum Steyerm. ex N. Robson (Ecuador: Loja,
Morona-Santiago), H. irazuense Kuntze ex N. Robson (Costa Rica), H. simonsii N. Robson (Colombia:
Magdalena), H. papillosum N. Robson (Colombia: Boyaca), H. martense N. Robson (Colombia:
Magdalena), H. maguireiN. Robson (Ecuador: Azuay, Loja, El Oro), H. gleasoniiN. Robson (Colombia:
Norte de Santander), H. prietoi N. Robson (Ecuador: Azuay), H. cassiopiforme N. Robson (Peru:
Amazonas), H. magdalenicum N. Robson (Colombia: Magdalena; Venezuela: Zulia, Merida), H.
valleanum N. Robson (Colombia: Valle de Cauca), H. sprucei N. Robson (Ecuador: widespread; Peru:
Piura), H. recurvum N. Robson (Peru: Amazonas, Junin, Pasco), H. wurdackii N. Robson (Peru:
Amazonas), H. costaricense N. Robson (Costa Rica; Panama; Colombia: Bolivar, Antioquia), H.
bolivaricum N. Robson (Colombia: Bolivar), H. parallelum N. Robson (Colombia: Norte de Santander/

Bull. Br. Mus. nat. Hist. (Bot.) 16 (1): 1-106 Issued 29 January 1987

2 NORMAN K. B. ROBSON

Cesar), H. marahuacanum N. Robson (three subspecies): subsp. marahuacanwn (Venezuela: Amazo-
nas), subsp. strictissimum N. Robson (Colombia: Boyaca; Venezuela: Merida), subsp. chimantaicum N.
Robson (Venezuela: Bolivar), H. horizontal N . Robson (Colombia: Norte de Santander, Santander), H.
jaramilloi N. Robson (Costa Rica; Colombia: Magdalena, Cesar/Norte de Santander), H. llanganaticum
N. Robson (Ecuador: Tungurahua, Napo), H. selaginella N. Robson (Colombia: Boyaca, Arauca,
Cundinamarca), H. cymobrathysN. Robson (Colombia: Boyaca).

The following changes of rank (stat. nov.) are also made: H. phellos subsp. platyphyllum (Gleason) N.
Robson, H. loxense subsp. aequatoriale (R. Keller) N. Robson, H. caracasanum subsp. turumiquirense
(Steyerm.) N. Robson. Two others are also new combinations (comb, et stat. nov.): H. strictum subsp.
compactum (Triana & Planchon) N. Robson, H. lancioides subsp. congestiflorum (Triana & Planchon) N.
Robson.

1. Introduction

It has proved to be impracticable to complete the systematic part of this monograph of
Hypericum L. in the order in which the sections were treated in part 1 (Robson, 1977). This
account of sect . 29 . Brathys , which is part 7 of an estimated nine parts * , has been completed next
after part 3 (sects l-6a) as a result of work on it for Flora Neotropica. The distributional area of
the shrubby members of sect. Brathys (Belize and the Greater Antilles to Bolivia) falls wholly
within the area of that Flora.

The publication of part 7 out of order has necessitated the abandoning of the double
enumeration started in part 3, where each species has a running generic number as well as
a sectional one. In parts 4-8, therefore, the species will have only sectional numbers; but a
continuous enumeration will be included in part 9, along with addenda, corrigenda, and a
revised sectional key.

The division of sect. Brathys sensu lato into two sections, 29 Brathys (trees, shrubs, and
shrublets, centred north of Amazonia) and 30 Spachium (subshrubs and herbs, centred south of
Amazonia) has proved to be incorrect. Sect. Brathys in fact includes a group of subshrubs and
wiry herbs that contains the type of sect. Spachium (H. gentianoid.es (L.) Britton, Sterns &
Poggenb.), thus extending the area of the section northward to eastern U.S.A. In addition, the
herbaceous H. piriai Arechav. (including H. hilaireanum L.B.Sm. = H. tenuifolium St. Hil. non
Pursh) represents a development from Sect. Brathys in which the flower has apparently become
adapted for specialized insect pollination in a manner partially analogous to those of sects Elodes
andAdenotrias. It has been treated as an extension of variation of Sect. Brathys, and its inclusion
extends the area of this section south of Amazonia to south-eastern Brazil and Uruguay (Fig. 1).

The remainder of sect. Brathys sensu lato divides into two sections. One of these is centred
(i.e. has the species with the most primitive characters) in the Andes of Venezuela and adjacent
Colombia, extending southwards along the Cordillera to Chile and northwards to Canada; and it
also occurs in SE Asia and Australasia (H. japonicum Thunb. ex Murray) and Africa and
Madagascar (H. lalandii group). Until now this section, which includes both shrubs and herbs,
has not been recognized as distinct and therefore has no name. It does, however, include the
type species of the genera Knifa Adanson (H. mutilum L.) and Tridia Korthals (H. japonicum
Thunb. ex Murray), both of which were treated as series of Sarothra sect. Spachium by Kimura
(1951). I have therefore chosen Knifa as the epithet for this new section:

Hypericum section Knifa (Adanson) N. Robson, stat. nov.

Knifa Adanson, Fam. PL 2: 44, 541 (1763).

Tridia Korth. inTijdschr. Natiirl. Gesch. Physiol. 3: 17(1836).

Sarothra sect. Spachium series Knifa (Adanson) Y. Kimura in Nakai & Honda, Nova Fl.
Japonica 10: 233 (1951) 'Kniffa'.

Sarothra sect. Spachium series Japonica Y. Kimura, torn. cit. : 233 (1951).
Type: H. mutilum L. (Y. Kimura, 1951, lectotype).

The other section, comprising shrubs, suffrutices, and herbs, is centred in south-eastern Brazil
and extends in area to Uruguay, northern Argentina, and Peru and thence northward along the

* For part 1 see Robson (1977); for part 2 see Robson (1981); for part 3 see Robson (1985).

THE GENUS HYPERICUM L. 3

Cordillera to Venezuela. It also occurs in Australasia and SE Asia (H. gramineum Forster f.).
The type of Kimura's Sarothra sect. Trigynobrathys (H. myrianthum Cham. & Schlecht.)
belongs to this section (although that epithet is not especially appropriate to the concept adopted
here), as well as the type of sect. Brathys subsect. Connatum R. Keller (H. connatum Lam.):

Hypericum section Trigynobrathys (Y. Kimura) N. Robson, comb. nov.

Sarothra section Trigynobrathys Y. Kimura in Nakai & Honda, Nova Fl. Japonica 10: 233

(1951).
Type: H. myrianthum Cham. & Schlecht. (Sarothra myriantha (Cham. & Schlecht.) Y.

Kimura).

The characters by which species in these three sections may be recognized can be summarized
as follows:

1 . Leaf lamina with short , sometimes thick basal insertion , pairs free or united by interfoliar ridge

or very rarely more broadly united; inflorescence-branching usually pseudo-dichotomous or

monochasial sect. 29. Brathys

Leaf lamina with long narrow basal insertion, pairs free or united by interfoliar ridge or by
lamina proper, sometimes with bases decurrent or leaves perfoliate; inflorescence-branching
pseudo-dichotomous or monochasial or dichasial 2

2. Leaves narrowed at the base or wholly narrow, united by interfoliar ridge, with basal vein

single, venation pinnate or 1-nerved, or, if (secondarily) amplexicaul and 3-5-nerved (H.

majus and H. lalandii groups p.p.), then midrib branches absent of weak .... sect. 30. Knifa

Leaves amplexicaul with 3-7 basal veins and midrib clearly branched, the leaf pairs usually

decurrent or sometimes perfoliate sect. 31 . Trigynobrathys

Morphology (Fig. 2)

The species in sect. 29 Brathys vary from small trees (up to 4 m in height) to prostrate shrublets
or wiry herbs. They all lack dark glands completely and have no resin glands on the lower surface
of the leaf (the latter are common in sect. 3 Ascyreia but not in sect. 1 Campylosporus), and there
are no fringing glands on leaves, sepals, and petals. It appears that all have marginal punctiform
leaf glands (although these are often very small), and the laminar glands in the leaves are also
always punctiform. In the sepals and petals, however, the laminar glands vary from wholly linear
to wholly punctiform.

(a) Leaves

The reduction of the laminar leaf glands from lines to dots is not here associated with the
development of reticulate venation, as it is in sect. 1 Campylosporus (Robson, 1981: 80, Fig. 11).
In the species of sect. Brathys with the most primitive characters (1 H. terrae-firmae) the
venation is parallel and open, even though the laminar glands are punctiform. The primary
venation remains open in nearly all the woody species, thus allowing reduction to 1-nerved
leaves to occur in several parts of the section. Only in some of the relatively primitive species,
e.g. 24 H. magniflorum, is there a clear tertiary reticulum.

In sect. Brathys the ultimate fate of the leaves has proved to be of major taxonomic
importance. In the most primitive state (found in 1 H. terrae-firmae and 2 H. styphelioides) the
leaves are free and completely deciduous, leaving a rhombic leaf scar. In all other species they
are wholly or partly persistent, the persistent part being eventually shed with the cortex of the
stem internode; and the leaf bases are usually united by an interfoliar ridge. The ultimate fate of
these leaves depends on their thickness and size. Where they are thick and long, they break off
without fading, leaving a perceptible part of the base (25 H. gleasonii to 61 H. cymobrathys
except 54 H. millefolium) . Where they are thin and long, they tend to droop and wither on the
stem (3a H. phellos subsp. phellos in part, 10 H. simonsii, 11 H. papillosum, and 24 H.
magniflorum). Where these thin leaves have (in evolution) become smaller or narrower and
thicker, they either fall at the apparent base (3 H. phellos and 4 H. garciae) or persist and wither
(Spp. 5-9, 12-23), tending to become secondarily appressed if the base is narrow (e.g. 8 H.
stenopetalum) and depressed if it is broad (e.g. 16 H. cuatrecasii) . Finally, in 54 H. millefolium

NORMAN K. B. ROBSON

THE GENUS HYPERICUM L. 5

the leaves are very small and remain attached to the stem, falling with it either when it
disarticulates at the nodes or with the cortex.

In those taxa in which the leaves fall at the apparent base, the true base has become united
with the stem, surviving only as a protuberance (e.g. in 4 H. garciae). Such leaves can be
distinguished from the wholly deciduous leaves of H. terrae-firmae and H. styphelioides by the
absence of a true leaf scar.

The order of these leaf characters in evolution would appear to have been: wholly deciduous -
wholly persistent - partly deciduous - wholly persistent again (Spp. 5-9 and 62-80), see Fig. 2.

The leaf base mostly remains parallel-sided or only slightly narrowed before widening to form
the interfoliar ridge; but in one group (31 H. pimeleoides in part and Spp. 49-53, 55, 59 in part) it
narrows perceptibly to form a pseudopetiole. Where the lamina has narrowed to the width of the
pseudopetiole, however, members of this group cease to appear petiolate and have to be
described as (secondarily) sessile (Spp. 54, 56-59 in part).

(b) Inflorescences

The inflorescence in woody members of sect. Brathys is nearly always 1-flowered (the primitive
state in the genus). Only in a few of the more primitive species (e.g. 3 H. phellos, 26 H.
mexicanum, and 61 H. cymobrathys) is there any development of cymose branching; and even
then it is not usually constant and never elaborate. Apart from this relatively weak tendency to
produce monochasia, the inflorescence in woody species is always strictly a synflorescence with
variation resulting from different types and aggregations of 1-flowered branches. The basic form
is the pseudo-dichotomy , i . e . the development of equal foliate branches from the axils of the leaf
pair immediately below the flower. This is the only type of acropetal inflorescence development
in the section; but the upper lateral branches are also often floriferous, and the relative
frequency of these two modes of branching can be of taxonomic value. Thus in 10 H. simonsii, for
example, the flowers are solely on lateral branches, whereas in 21 H. martense the majority of
the pulviniform plant consists of a pseudo-dichotomous synflorescence, with the branches of the
latest pseudo-dichotomies bearing lateral flowering branches (Fig. 8).

The development of a suffruticose or herbaceous habit (6 H. piriai and Spp. 62-80) is
accompanied by the elaboration of monochasial cymose inflorescences.

(c) Flowers and fruits

In section Brathys the perianth is normally pentamerous, with the sepals appressed to the petals.
These are longer than the sepals and usually spreading ('flowers stellate'), but in small flowers
they may remain ascending ('flowers obconic'). The sepals vary in width and hence in the
number of nerves, which may be of taxonomic significance, as may also be the aspect (plane to
incurved-cucullate). Variation in petal shape and size may help to separate related species; but
the degree of development of the apiculus, though striking, is not discontinuous enough to be of
taxonomic value.

The number and size of the stamens are correlated in general with flower size as a whole, but
the woody species with the fewest stamens (35 H. decandrum, which can have as few as 5) does
not have the smallest flowers. That distinction belongs to 42 H. prostratum, in which they can be
as small as 4 mm in diameter. Among the herbs H. gentianoides can have as few as 5 stamens and
also has the smallest flowers.

The styles vary in number from 5 (H. terrae-firmae, H. styphelioides) to 3, 4 as usual being an
apparently unstable number, not characteristic of any one species. The majority of species have
3 styles, but in three advanced woody species this trend has been reversed so that an increased
number (4-5) of styles is associated with very small flowers (41 H. juniperinum = H. brathys), 42
H. prostratum, and 60 H. selaginella) . The form of the styles (outcurving, incurving, etc.) may be
taxonomically useful. The stigma is primitively narrow but has frequently become broader,
usually in a capitate form; in one group (Spp. 51-53) the style apex is clavate. The ovary and
capsule remain enclosed in the perianth except when the flower is open, and so it is not always
easy to observe them in herbarium specimens. Nevertheless, their shape varies and can be
utilized taxonomically.

NORMAN K. B. ROBSON

THE GENUS HYPERICUM L. 1

Cytology and hybrids

Very little cytological work has been done on the major part of Sect. Brathys, the only recorded
chromosome count for a woody species being n = 12for7//. irazuense(Part2: table?). As this is
the primitive number for the whole genus (Part 2: 150), there is no available information about
possible chromosome variation within the woody members of the section. The numbers n = 6,
12, 24 have been recorded for herbaceous species (see part 8).

Hybrids in sect. Brathys are apparently very rare. The only specimen seen that might be a
hybrid was collected in Costa Rica (/. & C. Taylor 11757 (NY)) and looks like H. irazuense x
costaricense.

Distribution and evolution (Fig. 3)

In parts 2 and 3 it was shown that sect. 29 Brathys is directly related to the basic, African sect. 1
Campy losporus, the taxon in it that is most closely related to H. terrae-firmae being H.
revolutum subsp. keniense from the East African mountains (Figs 1-3). Sect. 30 Spachium, as
has been shown above, must be divided into three parts, the name-giving part being included in
sect. 29 Brathys. The other two parts, sects 30 Knifa and 31 Trigynobrathys, are more closely
related to each other than either is to sect. Brathys. All three sections, 29-31, would seem to
have stemmed from an early westward development from the original Hypericum stock in the
eastern part of west Gondwanaland, i.e. in what was to become Africa. This western group
appears to have divided into a northern (sect. Brathys) and a southern subgroup, the latter also
having divided into a northern group (sect. Knifa) and a southern group (sect. Trigynobrathys)
as a result of (climatic?) changes in the Amazon region. It clearly migrated into what is now
South America while contact between Africa and eastern South America was still possible, i.e.
until the Lower Cretaceous (Turonian-lOOm yrs BP) (Raven & Axelrod, 1974; Owen, 1976;
Howarth, 1981); but whether it had split into two before or after reaching that region is
uncertain. From the evolutionary divergence pattern of Hypericum as a whole (Robson, 1981:
Fig. 73; Robson, 1985; Fig. 1), it seems likely that the first split occurred in Africa.

For a temperate genus such as Hypericum that is confined in tropical latitudes to high
altitudes, the ranges of the Cordillera act as highways of dispersion to which the species are
restricted and along which they migrate and differentiate. It is therefore often possible to trace
presumed evolutionary lines (clades or, more accurately, morphoclines) in sequence along one
or more of the Andean ranges. If the variation is continuous, as it is for examples in 20 //.
laricifolium, no infraspecific taxa can be recognized, even though the forms at the extreme ends
of the range of this species ('//. laricoides' in Venezuela and '//. acerosum' in Peru) look very
different from the primitive form from northern Ecuador.

If the variation becomes discontinuous, on the other hand, one finds a geographical
replacement series of taxa each more advanced (apomorphic) than the one before. In such a
clade it is frequently difficult or impossible to discover any apomorphic character in the less
advanced taxon that would enable one to conclude that it had continued to evolve independently
subsequent to its isolation from the more advanced one. In such clades, therefore, where it is
difficult to believe in the Hennigian maxim that one taxon is automatically altered when it gives
rise to another (i.e. that at each speciation the cladogram must dichotomize), I have depicted the
relationship (Figs 1-3) by a line, not a dichotomy.

(a) Wide disjunctions

The majority of the primitive species in sect. Brathys occur in, or are confined to the mountains
near, the Colombian- Venezuelan border area or to adjacent areas in Boyaca (Colombia) and
Merida, Tachira or Trujillo (Venezuela); but the most primitive (plesiomorphic) characters are
found in the species from Belize (1 H. terrae-firmae) and Cuba (2 H. styphelioides), species that
are so closely related that they have sometimes been united under the latter name, e.g. by
Standley & Williams (1961). The question therefore arises as to whether (i) the original
members of sect. Brathys migrated and differentiated southward from Belize/Cuba to Col-
ombia/Venezuela before isolation of the two groups, or (ii) the differentiation took place

NORMAN K. B. ROBSON

co 3;

\

\

x\

CO

\

\

*

CN ^

OO
*

f

'x"

X

N ,

S

j

"~

- ' O

'

col

" 1>
^J5

?.s

8

_ o>

"O C/3

3

O ">

t; <u

T3

rt rt

1 "

i <U

<U T3

" G

iyj

S5 (fl

O D

s

ra tso

n

THE GENUS HYPERICUM L. 9

northwards from South America to the Greater Antilles before isolation, or (iii) a uniform
population was split and the two groups evolved in isolation (Fig. 3a/b). What is clear is that this
distribution gives strong support to the theory that the first land link between South America and
North American was by way of the Greater Antilles (Rosen, 1976, 1985; Robson, 1981: 213;
Melville, 1981:247).

The remaining evolution of the section has been wholly within South America, except for: (i)
four species that have reached Costa Rica/Panama after the establishment of the second land
link (Isthmus of Panama), (7 H. irazuense, 38 H. costaricense , 49 H. jaramilloi, and 50 H.
cardonae), (ii) three shrubby species in Hispaniola (52 H. ekmanii, 53 H. pycnophyllum, and 54
H. millefolium) , and (iii) the derivatives of 61 H. cymobrathys, which occur throughout Central
America and the Greater Antilles and extend their area of distribution into eastern U.S.A. The
three Hispaniolan shrubs terminate a clade that originates with two of the Costa Rican species,
which also occur in the basic area of the Colombian- Venezuelan border (49 H. jaramilloi, 50 H.
cardonae) (Fig. 3, clade xxviii). Occupying an intermediate position is 51 H. caracasanum of
northern Venezuela, of which the more advanced subspecies (51b subsp. turmiquirense) is found
at the north-eastern extreme of the Andean range and is also morphologically close to 52 H.
ekmanii. These data suggest that the ancestor of the Hispaniolan species reached that island
either directly by long-distance dispersal or via the Lesser Antilles, which, according to the
above mentioned theory, would by then have arisen as a result of volcanic action; but overland
access before the separation of the Greater Antilles from South America cannot be ruled out. A
three-fold origin of Hispaniola from parts previously associated respectively with Yucatan,
eastern Cuba, and the Bahamas, such as has been proposed by Rosen (1985), would make the
overland route less likely (see also p. 91).

The immediate derivatives of 61 H. cymobrathys are three species of the Colombian and
Venezuelan Andes (see Part 8). Of these the Colombian H. chamaemyrtus is the sister-species of
the H. gnidioides - H. gentianoides group, which radiates from the Honduras Republic: south to
Panama (H. gnidioides), north to north-eastern U.S.A. (H. gentianoides), and east to the
Greater Antilles (H. diosmoides group). On the other hand, there is a wide geographical
disjunction between the Venezuelan H. pseudocaracasanum and its sister-group in the south-
eastern U.S.A. (H. denticulatum et aff.).

(b) Species with leaves persistent or deciduous near the base (Spp. 3-23)
In this group (Fig. 3 clade (c)) the leaves are rarely broadest above the middle, and so the
leaf-venation is parallel to pinnate or 1-nerved, not flabellate. All species have leaves that are
either wholly persistent or deciduous at the apparent base. They form two subgroups: clade (i)
with leaves that are free (Spp. 3-9) ; clade (ii) with leaves that are united by an interfoliar ridge or
rarely more intimately (Spp. 10-23).

Subgroup (i) radiates from 3 H. phellos of north-eastern Colombia and adjacent Venezuela.
The variation in this species can be described; but it has not been possible to express this
variation in a formal classification, apart from the recognition of two derivative subspecies.
Subspecies H. phellos has therefore had to be differentiated plesiomorphically (i.e. by the
possession of primitive characters), viz. by having plane or recurved (not incurved or re volute)
leaves that are deciduous at the base of the free part (see p. 22).

Four separate derivative clades (iv, vi, vii, viii) end with taxa in which the leaves are
secondarily wholly persistent, and in three of these (all but vii), the leaves are plane or incurved.
The taxa in these three are all distributed in or near the Colombia- Venezuela border area near
H. phellos except 7 H. irazuense (Costa Rica and adjacent Panama), which is confined to the
nearest high ground in Central America to the Colombian Cordillera and is apparently derived
from H. phellos subsp. oroqueanum (Cesar/Norte de Santander border).

In clade (vii) the tendency for the margin of coriaceous leaves to become reflexed (3a iii H.
phellos subsp. phellos 'patens') is exaggerated, so that in 4 H. garciae it is revolute, hiding most
of the lower lamina. In this species, which has a disjunct distribution in the central Colombia/
western Venezuela area, the leaves tend to become narrow and sometimes longer, a tendency
that is more evident in 5 H. acostanum, a local species of southern Ecuador that is smaller and

10 NORMAN K. B. ROBSON

less woody than H. garciae. Although isolated from the nearest H. garciae population by a wide
disjunction (central Colombia to southern Ecuador), H. acostanum is separated from the
species that terminates clade (vii) by an even wider one. 6 H. piriai is a woody herb of
south-eastern Brazil and Uruguay in which the flowers have apparently become adapted to
specialized insect-pollination. Thus the stamen fascicles have the filaments united for more than
halfway (as in, say, H. elodes), and two pairs of fascicles are more or less completely united. The
petals are oblique to erect, sometimes making the flower pseudo-tubular, again as in H. elodes.
How the gap between southern Ecuador and south-eastern Brazil was traversed is not clear;
quite possibly it was by ancient long-distance dispersal. I have not yet discovered any similar
disjunctions, but the morphocline southward along the Andes to Ecuador suggests that a
vicariant interpretation is unlikely to be correct.

An early offshoot of clade (ii) comprises two isolated species (clade ix) in which the relatively
large herbaceous leaves fade and droop while on the stem. 10 H. simonsii is confined to the
northern part of the Sierra de Santa Marta (Magdalena), whilst 11 H. papillosum has a restricted
distribution in Boyaca. In the remainder of the clade (ii) the leaves are smaller and either
broad-based and deflexed (clade xi) or narrow-based and spreading only (clade xii). At the
extremes of clade (xi) the leaves become too narrow to droop and so remain spreading (15 H.
myricariifolium, 18 H. loxense in part). The species at the base of these clades (13 H. thuyoides)
has a vicariad that is scarcely more than subspecifically distinct (12 H. lycopodioides) , in which
the leaves have become coriaceous. H. thuyoides itself, from central Colombia (Cundinamarca),
is variable, with the broader-leaved (typical) form near clade (xi) and the narrower-leaved form
('genistoides') at the base of clade (xii). Clade (xii), in turn, divides into two distinct subclades:
in one (xiv), the areas of 22 H. hartwegii and 23 H. maguiret represent a major disjunction to
southern Ecuador, whereas in the other (xiii), H. thuyoides is near the origin of two trends
initially in opposite directions, respectively northwards to Boyaca (19 H. sabiniforme) and
southward to northern Ecuador (20 H. laricifolium). As has already been mentioned, H.
laricifolium shows continuous variation southwards to central Peru and northwards to western
Venezuela; but the morphologically reduced population further north, in the Sierra Nevada de
Santa Marta (Magdalena) is sufficiently distinct to warrant specific recognition (21 H. martense).

(c) Species with leaves with sheathing leaf-bases or marginal secretions (Spp. 24-26, 61)

In contrast to the early offshoot (f) in Boyaca in which the oblong to elliptic leaves of the basic
species have parallel venation (61 H. cymobrathys), nearly all early members of clade (d) (Fig.
3) have leaves broader above the middle with flabellate venation. A partial exception is 24 H.
magniflorum, in which the Boyaca population has parallel- veined leaves, but the more northern
one (straddling the Colombia- Venezuela border) has flabellate venation. H. magniflorum is also
exceptional in that the leaves are relatively thin and droop when fading, instead of breaking off
above the base like those of all the other species. It forms the base of one sub-branch of Branch
(e) (clade xv), in which the broad, flabellate-veined leaves have sticky marginal leaf secretions,
capitate stigmas, and usually a cymose inflorescence. 25 H. gleasonii occurs in the border area of
Colombia/ Venezuela, whereas 26 H. mexicanum has its primitive form in Boyaca and Cundina-
marca and differentiated back northwards into the area of//, gleasonii.

(d) Species with incurved-cucullate leaves but no marginal secretions (Spp. 27-30)

Apart from those species with marcescent or secreting leaves in clade (xv) there remains clade
(e), which splits into two subclades, (xvi) with leaves cucullate at the apex and (xvii) in which
they have a plane apex (apart from a few of the more specialized species where the apex is
somewhat cucullate).

The basic species of clade (xvi) , 27 H. stuebelii, is a northern Peruvian relict, the area of which
is thus separated from that of its nearest relatives in north-central Colombia by a considerable
disjunction. All the other members of this group are confined to the south Ecuador/north Peru
region except 30 //. decandrum, which has extended its area into north Ecuador. H. stuebelii has
leaves broader than those of H. magniflorum, with flabellate venation; but the leaves of the
other species (Spp. 28-30) are smaller and narrower with only a midrib or, at most, one pair of

THE GENUS HYPERICUM L. 11

vein branches. Like H. stuebelii, 28 H. prietoi and 29 H. cassiopiforme appear to be relicts, but
30 H. decandrum is more widespread.

(e) Species with sessile leaves deciduous above the base but not incurved-cucullate (Spp.
31p.p.-48).

The primitive form of 31 H. pimeleoides, which is found in the Cerro de Oroque (Cesar/Norte de
Santander), has large, broad, sessile, plane leaves; but within the range of this species are forms
where the leaf base has narrowed to a pseudopetiole. The remaining major woody clades in sect.
Brathys are respectively characterized by these two leaf forms, the species of clade (xviii) (Spp.
32-48) having sessile leaves, whereas in all but some extreme species of clade (xix) they are
pseudopetiolate.

In clade (xviii) the basic species, 32 H. magdalenicum, is almost confined to the isolated
northern Sierra de Santa Marta (Magdalena) and the adjacent Sierra de Peri j a on the border
with Venezuela. Apart from 43 H. parallelum, a close relative with smaller, densely imbricate
leaves isolated on the Cerro de Oroque, this variable species has given rise to two clades, the
members of which differ by a complex of characters. Clade (xx), which derives from the more
primitive, broader-leaved form of the species, includes species with leaves that twist as they
spread. This character is evident only where (a) the leaves are long enough to twist at all and (b)
the internodes are not too short to preclude individual twisting. In two extreme species, 35 H.
aciculare and 42 H. prostratum, the leaves are often too short to twist, while in 36 H. recurvum
and 37 H. wurdackii the internodes are too short to allow it.

The two subsidiary clades of clade (xx) each has a relict basic species, both of which occur at a
considerable distance to the south of the H. magdalenicum area. In clade (xxii), 33 H. valleanum
(Colombia: Valle de Cauca) forms a stepping-stone to the species of clade (xxiv), which are all
from Ecuador and Peru. Of these, 34 H. sprucei occurs from northern Ecuador to northern
Peru, with the distribution of the derivative 35 H. aciculare overlapping its area in south Ecuador
and just extending into Peru. The species with recurving leaves (36 H. recurvum, 37 H.
wurdackii) are in north and central Peru, H. wurdackii being a bizarre relict with densely
imbricate leaves forming four 'wings'.

Clade (xxv) proceeds along the Cordillera from H. valleanum in the opposite direction to that
of clade (xxiv). A small population of 38 H. costaricense is found on the Bolivar/Antioquia
border as well as the main one in Costa Rica and adjacent Panama, there being no suitably high
ground between them. 39 H. bryoides, clearly a reduced version of//, costaricense, is confined to
the South American Cordillera (Norte de Santander to Boyaca).

In the relict basic species of clade (xxiii), 40 //. bolivaricum (Colombia: Bolivar), twisting
leaves are combined with a condensed synflorescence and capitate stigmas, all characters that
are found in the variable 41 H. juniperinum (= H. brathys). In the latter species, which is
distributed from western Venezuela (Trujillo) to southern Colombia (Putumayo), there is a
trend towards an increase in the number of styles and placentae from 3 to 4 and then to 5. The
population from Cauca shows another trend, among 3-4-styled forms, from tall erect shrubs to
smaller shrubs with decumbent stems. This trend is continued back north-eastward in 42 H.
prostratum, a dwarf shrub or shrublet from the northern half of the Colombian Cordillera
Oriental with stems decumbent to prostrate.

The remaining derivatives of 32 H. magdalenicum are all related to more advanced forms of
that species (32' in Fig. 3). Apart from 43 H. parallelum they form two clades: (xxvi) of which
the members have polished, acicular leaves, rounded-incurved in section and twisting when
large enough, and (xxvii) in which the species have leaves that are dull to metallic and flat to
incurved but not usually rounded in section and become recurved not twisted. Clade (xxvi)
comprises three geographically separate taxa that I have treated as subspecies of 44 //.
marahuacanum. The subspecies with the largest flowers (44a subsp. marahuacanum) is confined
to the Cerro Marahuaca in southern Venezuela (Amazonas), an isolated massif from which no
other Hypericum species are known; and one of the derivative subspecies, 44c subsp. chiman-
taicum, is equally isolated further east, on the Chimanta Massif and Cima de Roraima (Bolivar)
near the Guyana border. The remaining derivative subspecies, 44b subsp. strictissimum, occurs

12 NORMAN K. B. ROBSON

in two quite separate areas (Venezuela: Merida and Colombia: Boyaca) and differs from subsp.
marahuacanum in size and condensation of parts. In comparison with H. marahuacanum, the
species of the H. strictum group (Spp. 45-48) are south-western in distribution, being confined
to the area from Cundinamarca and Meta (Colombia) to western Venezuela. The relatively
broad-leaved 45 H. lancifolium, from the Colombia- Venezuela border area, has a dwarf,
high-altitude derivative, 46 H. horizontale. on the Colombian side of the frontier and a southern
derivative complex (47 H. tetrastichum, 48 H. strictum) with H. tetrastichum extending from
Boyaca northward into Venezuela and H. strictum occurring farther south.

(f ) Species with pseudopetiolate leaves deciduous above the base but not incurved-cucullate (Spp .
31 p.p., 49-60)

The above heading is not strictly accurate in that a few of the advanced members of this group
(clade xix) have developed a somewhat cucullate apex. But their leaves are linear and would not
be confused with those of species in clade (xvi).

The species with pseudopetiolate leaves and their derivatives in clade (xix) (Spp. 49-60)
would appear to have evolved relatively early in the history of sect. Brathys, as they have spread
to almost the furthest limits of the shrubby part of the section (Hispaniola and Costa Rica to
Bolivia). The basic species of this clade, 49 H. jaramilloi, has a disjunct distribution in northern
Colombia (Cerro de Oroque, Sierra de Santa Marta) and Costa Rica; and an early species in the
eastern subclade (xxviii), 50 H. cardonae, has a similar but wider distribution, in Costa Rica and
the Colombia/Venezuela border region south to Boyaca. The rest of the species of this clade
show a geographical replacement pattern eastward: 51 H. caracasanum in the mountains of
northern Venezuela, with 51b subsp. turumiquirense at their extremity in Sucre, and the
remainder (Spp. 52-54) in Hispaniola (see p. 9).

The second (westward) derivative clade of 49 H. jaramillo, clade (xxix), is based on 55 H.
ruscoides, from south Colombia and north Ecuador, which gave rise southward to clade (xxx), a
series of sessile-leaved species (56 H. llanganaticum in central Ecuador, 57 H. struthiolifolium in
Peru, and 58 H. andinum in south Peru and Bolivia) with acute leaves and (at least initially) long
styles with relatively narrow stigmas. In another direction, evolution resulted in two species with
obtuse, cucullate leaves and short styles with relatively broad stigmas. Of these, 59 H. lancioides
comprises a southern subspecies with a pseudo-dichotomous synflorescence (59a subsp. lan-
cioides) and a northern one in which the flowering branches are lateral and congested (59b
subsp. congestiflorum) . A final southward development from the latter has resulted in the dwarf
60 H. selaginella, with prostrate or ascending stems and 3-5 styles and placentae.

2. Systematic treatment

Sect. 29. BRATHYS (Mutis ex L.F.) Choisy
Prodr. Monogr. Fam. Hyperic.:38, 58 (1821).*

Small trees, shrubs, shrublets or wiry herbs up to 6 m tall, evergreen, glabrous or very rarely with
simple hairs, without dark glands; branching below inflorescence lateral. Stems 4(6)-lined and
compressed (ancipitous) when young, sooner or later becoming terete, eglandular; cortex
exfoliating in strips or scales or irregularly; bark fissured, smooth. Leaves opposite, decussate,
sessile to shortly petiolate, free or deeply united to form interfoliar ridge or very rarely bases
of lamina proper united, base sometimes sheathing, deciduous at base or (more usually) above
base or persistent and (in woody species) either fading or shed with cortex; lamina entire, with
venation parallel or flabellate to pinnate or 1-veined, open or closed, with tertiary venation
laxly reticulate; laminar glands punctiform; marginal gland dots dense, often very small; ventral

* Including Sarothra L. (1753), Mania sensu Sprengel (1826) pro parte, Hypericum sect. Brathys subsect. Multista-
mineum R. Keller (1893). Hypericum sect. Brathys subsect. Spachium R. Keller (1893), Sanidophyllum Small (1924),
Sarothra sect. Spachium (R. Keller) Y. Kimura (1951), Sarothra sect. Spachium series Eusarothra Y. Kimura (1951),
see Part 1 (Robson, 1977: 338); also Hypericum sect. Sarothra (L.) Reichardt in Martius, Fl. Brasil. 12(1): 186 (1878).

THE GENUS HYPERICUM L. 13

resin glands absent. Inflorescence 1 -flowered, with branching pseudo-dichotomous (usually
from uppermost node) or 2-15-flowered and then with branching (within inflorescence)
dichasial/monochasial from uppermost node, often with subsidiary branches from lower nodes;
bracts and bracteoles foliar or reduced or sometimes transitional in form to sepals. Flowers
stellate or occasionally obconic, homostylous. Sepals 5, free, persistent, erect in fruit, with
margin entire; veins 3- 11 (-19); laminar glands linear to punctiform; marginal, submarginal and
inframarginal glands absent. Petals 5, persistent, with apiculus subterminal to obsolete, acute to
acuminate or obtuse; margin entire; marginal glands absent; laminar glands linear to punc-
tiform. Stamen fascicles basically 5, very rarely distinguishable individually, usually forming a
continuous narrow ring of 5-250 stamens (i.e. 1-50 per fascicle) or very rarely fascicles united
2 + 2+1, persistent; filaments very shortly united or very rarely united above midway; anthers
yellow to orange, gland amber; pollen type VIII. Ovary with 3-5 parietal placentae, -ovulate;
styles 3-5, free with bases contiguous; stigmas small to broadly capitate or clavate. Capsule
3-5-valved, coriaceous to chartaceous, with valves obscurely narrowly longitudinally vittate,
occasionally with 1-3 flattish vesicles. Seeds narrowly cylindric, shallowly carinate or not,
without apical expansion; testa scalariform-reticulate to ribbed-scalariform.

BASIC CHROMOSOME NUMBER (x): 12; ploidy 1,2,4.

HABITAT: open grassland or among stones or sometimes in scrub or moist montane woodland,
often along streams, in dry to locally wet areas of the Cordilleran paramo and subparamo
vegetation; 1600-4875 m. Also in open Pinus or Pinus-Quercus woodland or Pinus savannah or
on sterile white sand (Spp. 1 and 2, in Belize and Cuba, 0-800 m); or in open Pinus forest or in
cloud forest among limestone rocks (Spp. 52-54 in Hispaniola); 1300-3175 m. See part 8 for
herbaceous species.

DISTRIBUTION: Belize, Cuba, Hispaniola; Costa Rica, Panama; Guyana (Roraima), Venezuela
(Roraima, Chimanta, Marahuaca) and Andes of Venezuela, Colombia, Ecuador, Peru, and
Bolivia: also (subshrubs and herbaceous species) Porto Rico, Honduras, Guatemala, Mexico,
and south-eastern and eastern U.S.A.

80 species (+8 subspecies).

Key to sect. 29. Brathys

1 Leaves deciduous at base, leaving rhombic scar on stem; stem not articulate or with

interfoliar ridges 2

Leaves persistent (falling with cortex) or deciduous above base; stem articulate and/or with

interfoliar ridges 5

2(1) Leaves narrowly elliptic, acute, 16-30 mm long, chartaceous to subcoriaceous; sepals

14-22 mm long 1 . terrae-firmae(p. 18)

Leaves narrowly obovate or, if oblong to elliptic, then acuminate or shorter (10-16 mm
long) or thickly coriaceous ; sepals 5-15 mm long (2 . styphelioides) 3

3(2) Leaves narrowly elliptic to narrowly oblong, usually thinly coriaceous, spreading, often

outcurving 2a. styphelioides subsp. c/arense(p. 21)

Leaves oblanceolate to narrowly obovate, thickly coriaceous, densely imbricate,
straight 4

4(3) Leaves with 7-11 basal veins, glands not prominent

2b. styphelioides subsp. styphelioides(p. 21)
Leaves with 5-7 basal veins, glands prominent 2c. styphelioides subsp. moaense(p. 22)

5(1) Leaves persistent or deciduous immediately above base of free part; pseudo-dichotomous

branches often absent or sparse 6

Leaves deciduous above base, i.e. leaving base protruding from stem; pseudo-
dichotomous branches often predominant 38

6(5) Leaf bases either (i) sheathing or (ii) amplexicaul with stem fragile and articulated, the

internodes and leaves falling as one

Leaf bases free to perf oliate or shallowly cupped ; internodes and leaves not as above 8

14 NORMAN K. B. ROBSON

7(6) Leaf bases sheathing; leaves 10-18 mm long, withering; flowers 25-40 mm in diam.

24. magnifforum(p. 53)
Leaf bases amplexicaul; leaves up to 2 mm long, falling with internodes; flowers 5-7 mm in

diam 54. millefolium (p. 93)

8(6) Leaves free, the base sometimes indurated or fused with the stem 9

Leaves united by interfoliar ridge or lamina proper

9(8) Leaves becoming deflexed, marcescent; stem internodes without or almost without corky

emergences 3a. phellos subsp. phellos (i) 'marcescens' (p. 25)

Leaves becoming deflexed to appressed but not marcescent; stem internodes with or
without corky emergences 10

10(9) Leaves revolute, lower surface obscured, bases forming woody projections; stem

internodes smooth 11

Leaves recurved to incurved, lower surface always visible; stem internodes usually with

corky emergences 13

11(10) Leaves deciduous, 4-10 mm long, elliptic-oblong to linear; sepals ovate to lanceolate;

shrub to 3 m tall 4. garciae(p. 26)

Leaves persistent, (8)10-40 mm long, linear; sepals narrowly lanceolate to linear or

oblong; dwarf shrub to 0-8 m tall or suffrutex or herb 12

12(11) Flowers solitary, inflorescence-branching pseudo-dichotomous; flowers stellate; stamens

not fasciculate; dwarf shrub 5. acostanum(p. 27)

Flowers 1-25, inflorescence-branching monochasial; flowers obconic to pseudotubular
stamens 3(5)-fasciculate ; suffrutex or perennial herb 6. p/r/ai(p. 28)

13(10) Leaves with 3 basal veins, subcoriaceous

Leaves with 1 basal vein (venation wholly pinnate) , chartaceous 19

14(13) Leaves recurved before falling 3a. phellos subsp. phellos (ii) 'angustum' (p. 25)

Leaves spreading to appressed before falling 15

15(14) Leaves deciduous

Leaves persistent 18

16(15) Leaves spreading, not or scarcely becoming appressed, elliptic to oblanceolate

3a. phellos subsp. phellos (iii) 'patens' (p. 25)
Leaves becoming or remaining appressed, oblanceolate to narrowly oblong 17

17(16) Leaves spreading, then appressed, before falling

3a. phellos subsp. phellos (iv) 'phellos' (p. 25)
Leaves remaining appressed, tetrastichous 3b. phellos subsp. oroqueanum(p. 25)

18(15) Leaves with 3 unbranched basal veins; flowers 25-30 mm in diam.; styles 9-10 mm long,

4-6 x as long as ovary 7. irazuense(p. 30)

Leaves with at least midrib branched; flowers 15-20 mm in diam. ; styles 5-9 mm long, 2-4

x as long as ovary 8. stenopetalum(p. 31)

19(13) Leaves deciduous or subpersistent , midrib branched 20

Leaves persistent, 1-nerved 9. carinosum(p. 33)

20(19) Leaves with margins plane or recurved, spreading or becoming ascending, deciduous 21

Leaves with margin incurved, closely appressed, subpersistent

3c. phellos subsp. platyphyllum(p. 25)

21(20) Leaves broadly elliptic to obovate, not becoming ascending

3a. phellos subsp. phellos (v) 'diversicaule' (p. 25)
Leaves narrowly elliptic to oblong, spreading or becoming ascending

3a. phellos subsp. phellos (vi) 'tamanum' (p. 25)

22(8) Inflorescence-branching cymose; leaves always broadest at or below middle, rarely

petiolate , with margin plane or incurved , not minute and scale-like 23

Inflorescence-branching pseudo-dichotomous or lateral; leaves broadest below to above
middle, often petiolate, with margin plane or recurved or, if incurved, then minute and
scale-like 37

23(22) Leaves plane, oblong, with 3-7 basal veins becoming deflexed when withering; sepals 6-11

mm long 24

THE GENUS HYPERICUM L. 15

Leaves incurved or involute or conduplicate, or if plane then broadly ovate or 1-nerved,
becoming deflexed or not; sepals 2-6(-7) mm long 25

24(23) Leaves smooth, lustrous, the pairs very shortly united; stigmas narrow ... 10. simonsii(p. 34)
Leaves papillose, dull, the pairs united to form a cup-shaped sheath; stigmas capitate

11. papillosum(p. 34)

25(23) Leaves united only by interfoliar ridge, not becoming deflexed (except sometimes 12.

lycopodioides) 26

Leaves with bases of lamina united, often becoming deflexed 32

26(25) Leaves with midrib proximally prominent or wholly level with lamina; lamina condupli-
cate , apex usually acuminate to obtuse 27

Leaves with midrib proximally impressed; lamina incurved, apex usually obtuse to
rounded 28

27(26) Leaves coriaceous, markedly tetrastichous, apex acuminate to acute, margin plane

12. lycopodioides (p. 36)

Leaves chartaceous, weakly or not tetrastichous, apex acute to rounded, margin usually
undulate 13. thuyoides(p. 37)

28(26) Leaves sessile , elliptic-oblong to narrowly oblong, base cuneate to parallel 29

Leaves shortly petiolate or base angustate, if base narrowly cuneate, then lamina linear to

acicular 30

29(28) Leaves 5-7 mm long, elliptic or oblong to obovate; sepals 1-8-2-3 mm wide, apex plane

22. hartwegii(p. 51)

Leaves 2-4 mm long, narrowly elliptic or narrowly oblong to linear; sepals 0-7-1-7 mm
wide, apex incurved-cucullate 23. maguirei(p. 52)

30(28) Leaves subpapillose beneath, base cuneate to pseudopetiolate; sepals obtuse to rounded

19. sabiniforme(p. 45)

Leaves smooth beneath, base narrowly cuneate to angustate or parallel-sided; sepals acute
to obtuse 31

31(30) Plant a bushy or spreading shrub or small tree (0-1)0-3-6 m tall; leaves with glands not or

scarcely impressed; sepals usually plane 20. larici 'folium (p. 47)

Plant a pulviniform shrub 0-08-0-16 m tall; leaves with glands impressed; sepals markedly
ribbed 21 . martense(p. 51)

32(25) Flowers 18-40 mm in diam. ; leaf base rounded to cordate-amplexicaul 33

Flowers 6-12 mm in diam. ; leaf base parallel to truncate 35

33(32) Stem internodes smooth; leaves plane or slightly saccate, (2-)3-7 mm wide, margin not or

scarcely hyaline 16. cuatrecasii(p. 41)

Stem internodes with corky emergences, at least below upper leaves; leaves saccate or
incurved-cucullate, 0-7-3-5 mm wide, margin markedly hyaline 34

34(33) Leaves becoming reflexed, saccate; sepals 5-8 mm wide 14. goyanesii(p. 38)

Leaves spreading only, incurved-cucullate ; sepals 4-5 mm wide 15 . myricariifolium (p. 40)

35(32) Leaves becoming deflexed, broadly ovate to triangular-ovate or (lower) oblong, basal

veins 3(5) 17. quitense(p. 41)

Leaves at most spreading only, triangular-ovate or oblong to linear, basal veins 1(3-5) (18.
loxense) 36

36(35) Leaves broadly triangular-ovate to lanceolate, usually imbricate, base truncate to broadly

cuneate 18a. loxense subsp. aequatoriale (p. 44)

Leaves narrowly oblong or narrowly elliptic to oblanceolate or linear, usually spreading,
base narrowly cuneate to parallel 18b. loxense subsp. loxense (p. 45)

37(22) Leaves with parallel venation (5-7-veined) ; flowers 3-12 in condensed cyme ; shrub

61. cymobrathys(p. 103)

Leaves with pinnate venation or 1-nerved; flowers in lax cymes; suffrutices or herbs

[62-80. See Part 8]

38(5) Leaf venation parallel (5-7-veined) , veins unbranched ; flowers 3- 12 in condensed cyme

61. cymobrathys(p. 103)
Leaf venation flabellate or pinnate or 1-nerved, veins branched or not; flowers solitary

(except in 26. mexicanum) 39

16 NORMAN K. B. ROBSON

39(38) Leaves and usually sepals markedly incurved-cucullate; leaves distally broadened, not

linear 40

Leaves plane to incurved; leaf and sepal apices not cucullate or, if so, then lamina narrowly
elliptic or narrowly oblong to linear 45

40(39) Leaves with margin viscid, usually obovate; flowers often in dense cymes; stigmas

capitate 41

Leaves with margin not viscid, broadly elliptic to obovate-spathulate; flowers solitary;
stigmas narrow or, if capitate then leaves oblanceolate 42

41(40) Flowers solitary, 20-40 mm in diam., sometimes terminating clustered short shoots; leaves

7-10 mm long, always coriaceous; habit erect 25. gleasonii(p. 55)

Flowers in 3-13-flowered cymes or, if solitary, less than 20 mm in diam.; leaves up to 17
mm long, sometimes chartaceous and then habit decumbent and rooting

26. mexicanum(p. 56)

42(40) Leaves elliptic-ovate to elliptic-lanceolate, 3-7 mm broad venation flabellate; flowers

25-40 mm in diam 27. stuebelii(p. 58)

Leaves oblanceolate to narrowly oblong, 0-7-3 mm broad, venation pinnate or 1-nerved;
flowers 6-25 mm in diam. (petals unknown in 28.prietoi) 43

43(42) Sepals acuminate; leaves persistently imbricate-tetrastichous, markedly laterally com-
pressed distally; stigmas capitate 28. prietoi(p. 58)

Sepals acute; leaves eventually outcurving, not laterally compressed distally or, if so (29.
cassiopiforme in part) then stigmas narrow 44

44(43) Petals c. 14 mm long; styles longer than ovary; stigmas narrow; leaves oblanceolate (1: b =

c. 2.5) 29. cassiopiforme (p. 59)

Petals 5-9 mm long; styles 0-6-1 x as long as ovary; stigmas capitate; leaves narrowly
oblanceolate to linear-elliptic (l:b = 4-11) 30. decandrum(p. 59)

45(39) Leaves with venation flabellate, some laterals free or almost free from the base; styles stout

with stigma usually capitate 31 . pimeleoides(p. 60)

Leaves with venation pinnate, all laterals originating well above the base; styles and
stigmas various 46

46(45) Leaves petiolate or abruptly narrowed at the base ; stem sometimes apparently articulated 47
Leaves sessile , gradually narrowed or parallel-sided at the base ; stem not articulated 54

47(46) Styles 6-7 mm long; flowers 25-30 mm in diam.; leaves often spreading abruptly above

petiole 49. jaramiUoi(p. 86)

Styles 0-8-5 mm long; flowers up to 25 mm in diam. ; leaves outcurving or spreading from
the base 48

48(47) Leaves with midrib prominent or level with lamina beneath, glands usually visible beneath 49
Leaves with midrib impressed beneath , glands not visible beneath 53

49(48) Stigmas broadly capitate; leaves glaucous, dull 50. cardonae(p. 87)

Stigmas clavate or small ; leaves not glaucous, lustrous 50

50(49) Leaves with margins recurved (51 . caracasanum) 51

Leaves with margin or whole lamina incurved 52

51(50) Leaves9 x 4-5 mm or larger; petals c. 1-3 x as long as sepals; styles c. 1-3 x as long as ovary

51a. caracasanum subsp. caracasanum(p. 89)

Leaves 8 x 3-5 mm or smaller; petals 1-5-2 x as long as sepals; styles 2-2-5 x as long as
ovary 51b. caracasanum subsp. turumiquirense(p. 90)

52(50) Leaf lamina elliptic, 2-4 mm wide, plane with margin incurved; styles 5-6 mm long

52. ekmanii(p. 90)

Leaf lamina narrowly elliptic to linear, 0-5-2 mm wide, incurved to canaliculate; styles
3-5-4mmlong 53. pycnophyllum(p. 92)

53(48) Leaves plane or with margin subincrassate, narrowly ovate to narrowly oblong-elliptic;

sepalsl-5-4mmwide;styles2-5-4mmlong 55. ruscoides(p. 93)

Leaves incurved to canaliculate, very narrowly lanceolate to (usually) linear; sepals
0-5-1-5 mm wide; styles l-2(-3) mm long 59a. lancioides subsp. lancioides(p. 99)

54(46) Leaf and sepal apices incurved-cucullate; leaf lamina narrowly oblong or narrowly

elliptic to linear, narrowed at the base 55

THE GENUS HYPER1CUM L. 17

Leaf and sepal apices not incurved or cucullate or, if so, then leaves broadened at the base 60

55(54) Ovary ellipsoid to globose; styles 0-8-2(-3) mm long, 0-4-1-1 x as long as ovary, stigmas

broadly capitate ; sepals usually oblong to oblanceolate , not ribbed 56

Ovary ovoid; styles either 3-4-5 mm long, 1-3-1-5 x as long as ovary, stigmas scarcely to
narrowly capitate, or if styles shorter (1-5-2-5 mm long, 0-6-0-7 x as long as ovary) and
broadly capitate, then sepals lanceolate, ribbed 58

56(55) Stems erect or rarely decumbent; leaves 7-15 mm long; sepals 5-8 mm long; styles 3(4),

l-2(-3) mm long (59. lancioides) 57

Stems prostrate or ascending; leaves 4-7 mm long; sepals 3-4 mm long; styles (3)4-5,
0-8-lmmlong 60. selaginella(p. 102)

57(56) Flowers terminal, solitary, inflorescence-branching pseudo-dichotomous; habit bushy or

caespitose 59a. lancioides subsp. Iancioides(p. 99)

Flowers terminal and lateral in dense racemiform heads; habit bushy or
straggling 59b. lancioides subsp. congestifforum(p. 101)

58(55) Styles 3-4-5 mm long, 1-3-1-5 x as long as ovary; stigmas not or scarcely capitate; leaves

1-5-4 mm wide, margin not or scarcely hyaline 59

Styles 1-5-2-5 mm long, 0-6-0-7 x as long as ovary; stigmas broadly capitate; leaves
0-8-1 -5 mm wide, margin narrowly but distinctly hyaline 58. andinum(p. 97)

59(58) Sepals oblong-oblanceolate, not ribbed; leaves plane or with margin slightly incurved, 6-9

mm long, markedly tetrastichous, glaucous 56. Hanganaticum(p. 95)

Sepals ovate-lanceolate to oblong, ribbed; leaves with margin markedly incurved, 8-15
mm long, not or obscurely tetrastichous, lustrous 57. struthiolifolium(p. 96)

60(54) Styles 6-7-5 mm long; flowers 20-30 mm in diam.; leaves with midrib branched, lamina

elliptic to oblanceolate, plane or conduplicate, epidermis smooth 32. magdalenicum(p. 63)
Styles up to 5 mm long; flowers up to 20 mm in diam. ; leaves with midrib unbranched or, if
branched , then lamina narrowly oblong or epidermis papillose above 61

61(60) Leaves markedly tetrastichous, narrowly elliptic, papillose above; branches very strict,

parallel, with internodes 1-1-5 mm long 43. parallelum(p. 76)

Leaves not tetrastichous or if so , then linear , smooth , and branches not as above 62

62(61) Sepal margin green or, if hyaline, then inflorescence a racemiform synflorescence and

styles shorter than ovary with stigmas capitate 63

Sepal margin hyaline; inflorescence and styles not as above 72

63(62) Flowers solitary, not congested; styles 2-5-5 mm long, stigmas narrow 64

Flowers in congested heads or racemiform synflorescences; styles 1-3(4-5) mm long,
stigmas broadly capitate 70

64(63) Leaves 2-3 mm wide, narrowly elliptic, slightly incurved 33. valleanum(p. 64)

Leaves 0-4-2 mm wide, very narrowly oblanceolate to linear, markedly incurved to
conduplicate 65

65(64) Styles 2-5-3-5 mm long; leaves lustrous beneath 66

Styles 1-2 mm long; leaves dull beneath 69

66(65) Leaves becoming twisted , incurved but not carinate or conduplicate , margin very narrowly

hyaline, dorsally usually glandular when mature 67

Leaves remaining erect or becoming recurved, incurved to conduplicate, often at least
partly carinate , margin broadly hyaline , dorsally eglandular when mature 68

67(66) Sepals 7-11 mm long; flowers 20-30 mm in diam.; capsule c.5 mm long; leaves 6-12 x

0-8-1 -7 mm; stamens 60-75 34. sprucei(p. 65)

Sepals 4-7 mm long; flowers 10-18 mm in diam.; capsule 3-5-4 mm long; leaves 3-7-7 x
0-4-0-6 mm; stamens 30-55 35. aciculare(p. 66)

68(66) Leaves imbricate at first but not markedly tetrastichous, recurving, 5-8 mm long; flowers

15-20 mm in diam.; petals 8-12 mm long 39. recurvum(p. 68)

Leaves densely and persistently imbricate, 'winged'-tetrastichous, scarcely spreading,
9-14 mm long; flowers 20-25 mm in diam.; petals 15-18 mm long 40. wurdackii(p. 69)

69(65) Flowers 15-20 mm in diam. ; leaves 5-15 mm long, acute; stems woody, not rooting

38. costaricense(p. 70)

18 NORMAN K. B. ROBSON

Flowers 6-8 mm in diam.; leaves 3-5 mm long, subacute to obtuse; stems wiry, basally

rooting 39. bryoides(p. 71)

70(63) Leaves very narrowly elliptic, 15 x 3 mm or larger; sepals 9-11 mm long, coriaceous; styles

c.4-5 mm long 40. bolivaricum(p. 72)

Leaves linear, up to 14 x 1-5 mm; sepals 2-7 mm long, chartaceous to membranous; styles
0-6-3 mm long 71

71(70) Stems erect to decumbent; leaves 6-14 mm long; styles 1-5-3 mm long, longer than ovary

41.juniperinum(p. 72)
Stems prostrate; leaves 2-5 mm long; styles 0-6-0-8 mm long, shorter than ovary

42. prostratum(p. 75)

72(62) Leaves lustrous, margin markedly incurved, lamina twisting and sometimes recurving (44.

marahuacanum) 73

Leaves dull or with metallic sheen , margin plane to incurved , lamina recurving only 75

73(72) Stigmas small or slightly enlarged; ovary and capsule acute; inflorescence branching

pseudo-dichotomous or racemiform 74

Stigmas enlarged to capitate; ovary and capsule rostrate, inflorescence branching
pseudo-dichotomous to lateral-congested 44c. marahuacanum subsp. chimantaicum(p. 80)

74(73) Styles 4-5 mm long; sepals 7-9 mm long; inflorescence branching pseudo-dichotomous

44a. marahuacanum subsp. marahuacanum (p. 79)

Styles 2-3(-4) mm long; sepals 4-6 mm long; inflorescence branching lateral, racemi-
form 44b. marahuacanum subsp. strictissimum (p. 79)

75(72) Leaves plane or slightly incurved, oblong to oblong-linear or narrowly oblanceolate, the

tips not red when young 45. lancifolium(p. 80)

Leaves markedly incurved to canaliculate, linear to linear-acicular, the tips usually red
when young 76

76(75) Plant a prostrate shrublet with branches matted, rooting; leaves often secund, not

tetrastichous 46. horizontale(p. 81)

Plant an erect shrub with branches strict, not rooting; leaves never secund, usually
tetrastichous 77

77(76) Leaves with metallic sheen, markedly tetrastichous, erect to suberect 47. tetrastichum(p. 82)
Leaves dull, not tetrastichous (except sometimes in young parts or plants); usually
outcurving (48. strictumi) 78

78(77) Leaf and sepal apices acicular; sepals 1-3-2 mm wide, not ribbed; leaves 8-13 mm long;

branches strict, usually all from one stem 48a. strictum subsp. strictum(p. 84)

Leaf and sepal apices subacute; sepals 1-1-2 mm wide, clearly ribbed; leaves 5-7(8) mm
long; branches ascending, usually branched from the base

48b. strictum subsp. compactum(p. 85)

1 . Hypericum terrae-firmae Sprague & Riley

in Kew Bull. 1924: 12 (1924). Type: Belize, without precise locality, 1912? (fr), Peck
321 (K!, holotype; GH!, isotype).

H. styphelioides sensu Standley & Williams in Fieldiana Bot. 24 (7): 51 (1961); P.
Adams in Rhodora 64: 236 (1962), non A. Richard (1845).

Icon: Fig. 4.

Shrub or small tree, 1-2 m tall, erect, with branches strict, pseudo-dichotomous or
lateral. Stems orange-brown, 4-lined when young, soon terete, without corky
wrinkles, cortex exfoliating in strips; internodes 4-6 mm long. Leaves sessile, free
from the base, spreading to subimbricate and tetrastichous, deciduous at the base
without fading; lamina 16-30 x 4-6 mm, narrowly oblong to narrowly elliptic,
plane, not cucullate or carinate, concolorous, not or slightly glaucous, chartaceous to
thinly coriaceous; apex acute, base narrowly cuneate to subangustate, not sheathing,

* The leaves and petals in the marginal text figures are all respectively twice and one-and-a-half times natural size. In
the inflorescence diagrams O = open flower, = flower bud, x = vegetative bud, \ = continuing shoot.
Note. In part 3 (Robson, 1985) the magnifications in the legends to figs 9-24 should all have been reduced by half.

THE GENUS HYPERICUM L.

19

Fig. 4 H. terrae-firmae: (a) habit; (b) stem with leaf bases; (c) leaf scar; (d) leaf; (e) sepal; (f) petal;
(g) stamens (partly cut away) and ovary; (h) stamen; (i) anthers; (j) capsule (a, b = Vi; c x 1; d, j x 2; e-g
x 3;h x 8;i x 15). (a-d,g, j) Hunt 66; (e, f, h, \) Sutton et al. 193.

pairs free; basal veins 7, subparallel-sided, with short lateral branches, tertiary
reticulation not visible; laminar glands dense, not prominent. Inflorescence 1-
flowered, with pseudo-dichotomous branches from 1-2 nodes below; pedicel absent
or very short; upper leaves transitional. Flowers 40-60 mm in diam., stellate. Sepals
14-22 x 3-7 mm, narrowly oblong, acute; veins 15, dichotomising and reticulate
distally, with midrib prominent; glands linear. Petals bright yellow, 20-30 x 7-12
mm, c. 1-3 x sepals, oblanceolate; apiculus acute; glands linear, mostly uninter-
rupted. Stamens c. 250, longest 9-11 mm long, c. 0-3-0-4 x petals. Ovary (2-5-)3

20

NORMAN K. B. ROBSON

-4-5 x 2-2-5 mm, narrowly ovoid; styles 5, 5-7 mm long, 1-4-3 x ovary, erect,
outcurved below apex; stigmas small. Capsule 8-10 x 5-7 mm, broadly ovoid,
shorter than sepals. Seeds c. 1 mm long, not or shallowly carinate; testa finely
scalariform-reticulate .

In open pine or oak-pine forest or pine savannah on granite, often near streams;
lowland to 550 m.

Belize (Cayo, Belize). Map 1.

BELIZE. Belize: Colonel English Pine Ridge, Belize-Cayo road, 20.i.l958 (fl),
Gentle 9515 (DUKE, F, NY, S, US); Burrellboom, c. 9 m, 22. Hi. 1970 (fr), McDaniel
12853 (F). Cayo: Mountain Pine Ridge, San Miguel, 400 m, 2.vi.l979 (fl & fr),
Whitefoord 1935 (BM); Mountain Pine Ridge, Augustine, vii-viii.1936 (fl), Lundell
6617 (F, K, MICH, NY, US).

H. terrae-firmae, as previous authors have pointed out, is closely related to the
Cuban H. styphelioides, especially to forms of subsp. darense; but the thinner,
longer, acute leaves, which spread more or less widely from the base, and the large
flowers with longer, narrowly oblong sepals, allow the Belize plant to be accorded
specific rank. Its nearest ancestral taxon is in East Africa (H. revolutum subsp.
keniense (Schweinfurth) N. Robson), a subspecies of Sect. 1 Campy losporus.

2. Hypericum styphelioides A. Richard

in Sagra, Hist. Phys., Pol. Nat. He Cuba (Bot. - PI. Vase.): 237 (1845); Leon &
Alain, Fl. CubaS: 316, 1. 141 (1953); Lippoldin WissZ. Friedr. -Schiller Univ. Jena
(Math. -Nat. R.) 19: 378, f.2. (1970). Type: Cuba, [Pinar del Rio, near San Diego
de los Banos], Sagra s.n. (P, holotype).

Shrub 0-15-2 m tall, erect, with branches erect, strict, pseudo-dichotomous or
lateral. Stems orange-brown, 4-lined when young, soon terete, without epidermal
wrinkles, cortex exfoliating in strips or scales; internodes 2-6 mm long. Leaves
sessile, spreading from above the base to closely imbricate and tetrastichous,
deciduous at the base without fading; lamina 5-25 x 2-8 mm, narrowly obovate to
narrowly elliptic or narrowly oblong, plane or incurved, sometimes concave, not
carinate, concolorous, lightly to densely glaucous, thinly to thickly coriaceous; apex
shortly acuminate to acute, base cuneate to scarcely angustate, not or slightly
sheathing, pairs free; basal veins 5-9(-ll), subparallel to flabellate, branching
distally to form a lax reticulum, tertiary reticulum not visible; laminar glands dense,
prominent. Inflorescence l(2)-flowered, with pseudo-dichotomous branches from
1-2 nodes below, sometimes with additional branches from immediately lower node;
pedicel absent or very short; upper leaves transitional. Flowers 15-50(-70) mm in
diam., stellate. Sepals 5-15 x 2-5 mm, oblong or lanceolate to obovate, acute or
obtuse-apiculate to obtuse; veins 5-17, dichotomising, and reticulating distally, with
midrib prominent beneath; glands linear, sometimes distally punctiform. Petals
bright yellow, (8-)10-22 x 2-10 mm, c. 1-5 x sepals, oblanceolate; apiculus acute;
glands linear, uninterrupted. Stamens c. 70-200, longest 5-9 mm long, c. 0-3-0-5 x
petals. Ovary 2-4 x 1-5-3 mm, ovoid to rostrate-subglobose; styles (4)5, 3-5 mm
long, 1-2-1-5 x ovary, erect, outcurved below apex; stigmas small. Capsule (3-)4-8
x 2-7 mm, ovoid-subglobose, shorter than sepals. Seeds c. 1 mm long, not or
shallowly carinate; testa finely scalariform-reticulate.

In pine forest or pine savannah or often on sterile white sands, especially round
margins of lakes and pools; 0-800 m.

Cuba (Pinar del Rio, Las Villas, Oriente, Isla de Pinos). Map 1.

Three subspecies of//, styphelioides have been recognized. Subsp. darense, the most
similar to H. terrae-firmae, has a disjunct distribution (central Cuba, western Cuba,
Isla de Pinos), whilst the others are respectively eastern (subsp. moaense) and
western (subsp. styphelioides).

THE GENUS HYPERICUM L.

21

Map 1 1 . H. terrae-firmae . 2. H. styphelioides: a. subsp. clarense A , b. subsp. styphelioides O, c. subsp.
tnoaenseU.

2a. Hypericum styphelioides subsp. clarense Lippold

in Wiss. Z. Friedr. -Schiller Univ. Jena (Math. -Nat. R.) 19: 379, f. 2. (1970). Type:
Cuba, Las Villas, Santa Spiritus, Rio Gayabo, 7.xii.l911 (fl), Bro. Clemente 2884
(LS, holotype; NY!, isotype).

Hypericum incurvum Urban, Symb. Antill. 9: 405 (1925); Leon & Alain, Fl. Cuba 3:
317 (1953); Lippold in Wiss. Z. Friedr. -Schiller Univ. Jena (Math.-Nat. R.) 19:
380, f. 2. (1970). Type: Cuba, Isla de Pinos, Santa Barbara, Westport, 3.xii.l920
(fl), Ekman 12072 (S!, holotype).

Shrub 0-15-0-6(-l) m tall. Leaves 10-18 x 2-5-5 mm, narrowly elliptic to narrowly
oblong, thinly to thickly coriaceous, not or lightly glaucous, plane to concave or
incurved, spreading or outcurving; apex acute; basal veins 5(7). Flowers 15-30 mm
in diam. Sepals oblong to lanceolate, acute or obtuse, 5-veined; glands prominent.
Petals 7-16 mm long. Styles 5, 3-5 mm long.

Savannahs, fields, and coastal sands, sometimes on weathered dolomite; lowland to
montane.

In Las Villas, western Pinar del Rio and Isla de Pinos.

CUBA. Pinar del Rio: N. of San Diego de los Banos, 13. iv. 1900 (st), Palmer &
Riley 535 (NY, US); mountains near El Guama, 6.iii.l900 (fr), Palmer & Riley 160
(NY, US). Las Villas: Lomas de Banao, i.1920 (fl), Lund 123 (NY); Trinidad Mts,
Buenos Aires, near La Sabana, 5.viii.l936, Smith, Hodgson & Gonzales 3381 (F,
NY, S, US). Isla de Pinos: Siguanea, 26.ii.1916 (fl), N. L. Britton, E. G. Britton &
Wilson 14926 (F, NY, S, US); between Mine de Oro and Playa del Soldado,
12.i. 1956, Killip 45389 (US).

Subsp. clarense forms a cline from eastern Las Villas (where the flowers are larger
and the leaves are broader and flat) via eastern Pinar del Rio to Isla de Pinos (where
the flowers are smaller and the leaves narrow with an incurved margin and out-
curving towards the apex).

2b. Hypericum styphelioides A. Richard subsp. styphelioides

Shrub 0-3-2 m tall. Leaves 8-25 x 4-8 mm, oblanceolate to narrowly obovate,
plane, densely glaucous, thickly coriaceous, spreading to densely imbricate;
apex shortly acuminate; basal veins 7-9(-ll); glands not usually prominent. Flowers
15-50(-70) mm in diam. Sepals elliptic to obovate, acute or obtuse-apiculate to
obtuse, 9-17-veined. Petals 10-22 mm long. Styles 5, 3-5 mm long.

22 NORMAN K. B. ROBSON

In pine forest or savannah sand or in wet places; lowland.

In Isla de Pinos, Las Villas and Pinar del Rio.

CUBA. Pinar del Rio: El Sabalo, Finca Sabanalamar, near sea level, 22.xiii.1937
(fl), Killip 32263 (NY, US); Herradura, 1904 (fl), van Hermann 562 (BM, F, K, NY,
P, US); near Laguna Alcatraz Grande, 16.xii.1911 (fl), Shafer 11016 (NY, US). Las
Villas: Banao Mts., Sabana de la Gloria, 29.vii.1918 (fl), Leon & Roca 7928 (NY).
Isla de Pinos: San Pedro and vicinity, 12.ii.-22.ii.1916 (fl), N. L. Britton, E. G.
Britton & Wilson 14148 (F, NY, S, US); Los Indios, 4.U.1953 (fl), Killip 42603 (NY,
US).

Subsp. styphelioides forms two incompletely correlated clines eastward, showing
reduction in size and increasing leaf-imbrication. The only specimen seen from Las
Villas (Leon & Roca 7928) is somewhat intermediate between subsp. styphelioides
and subsp. moaense.

2c. Hypericum styphelioides subsp. moaense Lippold

in Wiss. Z. Friedr. -Schiller Univ. Jena (Math. -Nat. R.) 19: 379, f.2 (1970). Type:
Cuba, Oriente, Piano el Sierra de Moa, Bisse 11676 (JE, holotype; HAJB,
isotype).

Shrub 0-6-1-5 m tall. Leaves 7-ll(-16) x 2-6 mm, obovate (or rarely oblanceolate)
to narrowly elliptic, thickly coriaceous; slightly glaucous, cucullate or incurved,
usually densely imbricate; apex apiculate-obtuse to acute; basal veins 5-7; glands
prominent. Flowers 15-20 mm in diam. Sepals lanceolate, acute, 7-veined. Petals
8-12 mm long. Styles 5(4), c. 3 mm long.

In pine woods and open, dry or marshy habitats, on laterite or serpentine soils;
0-800 m.

Confined to Oriente.

CUBA. Oriente: Minas de Iberia, Taco Bay, in mountains, c. 800 m, 7-8. xii. 1914
(fl), Ekman 3787 (F, K, MICH, NY, S); Moa, near airfield, vii.1941 (fl), Howard
6020 (BM, NY, P, S, US); Maravi (near Barancoa), 25.xii.1914 (st), Ekman 4020

(NY,S,US).

The H. phellos group (Spp. 3-9) Fig. 1, p. 4

The most primitive form of 3 H. phellos has relatively large, marcescent leaves as in
10 H. simonsii. In the other forms of the species, however, the leaves are either (i)
smaller and more rigid, deciduous at the apparent base or persisting, or (ii) narrower
and more revolute, deciduous at the apparent base only after drooping but not
withering. The extreme variability of H. phellos in a relatively restricted area,
together with the absence of clear morphological disjunctions, makes it difficult to
recognize infraspecific taxa in this species and to differentiate it from its nearest
relatives. The form with marcescent leaves (3a subsp. phellos (i) 'marcescens')
differs from H. simonsii in having leaves free at the base with the apex acute to
acuminate and the margin recurved, as well as rounded sepals; but it intergrades with
both the form having thicker, narrow, drooping leaves with recurved margins and
densely corky internodes (13a subsp. phellos (ii) 'angustum') and the form with
smaller, chartaceous, spreading leaves, also with recurved margins, but with an
acuminate apex and sometimes only slightly corky internodes (3a phellos subsp.
phellos (v) 'diversicaule'). From (ii) 'angustum' two distinct trends are apparent in
subsp. phellos: to (iii) 'patens', in which the leaves are thicker with more recurved
margins and usually fall when spreading or soon after becoming appressed, and to
(iv) 'phellos' (including the type), in which the leaves are narrower and become
markedly appressed but not so thick. Variant (ii) 'patens' gives rise to the H. garciae
group (Spp. 4-6), with narrow revolute leaves, as well as 3b H. phellos subsp.
oroqueanum, with narrow continually appressed leaves, and 7 H. irazuense (Costa
Rica), whilst the (iv) 'phellos' trend continues to 8 H. stenopetalum, which has a
distribution overlapping that of//, phellos. On the other main division of//, phellos
subsp. phellos, (v) 'diversicaule' is related to 9 H. carinosum, which has smaller,

THE GENUS HYPERICUM L.

1-nerved leaves and smaller flowers, whereas (vi) 'tamanum', which differs from
'diversicaule' only in its relatively broader and more obtuse leaves and sepals and
more clearly differentiated lateral shoots, gives rise to a high-altitude condensed
form (3c subsp. platyphyllum) in which the leaves become incurved and remain
appressed until they are shed.

23

3. Hypericum phellos Gleason

in Bull. Torrey hot. Club 56: 106 (1929). Type: Colombia, Santander, Paramo de la
Puentes, above La Baja, 3700-3800 m, 25. i. 1927 (fl), Kiltip & Smith 18199 (NY!,
holotype; F!, GH!, K!, US! isotypes).

Shrub or small tree 0-3-4 m tall, erect, with branches strict, almost always lateral.
Stems yellow-brown, 4-lined and ancipitous when young, becoming terete, often
with transverse epidermal or corky wrinkles or emergences immediately below the
node or throughout the internode, cortex exfoliating in strips; internodes 1-5-3
mm long. Leaves sessile, spreading from the base or imbricate and tetrastichous,
deciduous near the base after fading or persistent; lamina 4-20 x (l-)2-6 mm,
narrowly to broadly elliptic or oblong to obovate or oblanceolate, plane or recurved,
not concave, not or scarcely carinate, concolorous, not glaucous, thickly to thinly
coriaceous; apex shortly acuminate to rounded, base cuneate to angustate, not
sheathing, free or pairs united to form a narrow interfoliar ridge; basal veins 3-7,
diverging, all or only the midrib branching, tertiary reticulation obscure or not
visible; laminar glands rather dense, impressed above, prominent beneath.
Inflorescence l(2-5)-flowered, terminal and on short lateral shoots, with extension
shoots usually from 2-3 nodes below; peduncle and pedicels 1-9 mm long; upper
leaves foliose or bracteose. Flowers 15-30 mm in diam., stellate. Sepals 4-9 x 1-5-3
mm, narrowly oblong to lanceolate or elliptic or rarely oblanceolate, acute or
acuminate to obtuse or rarely rounded; veins 5-7, branched distally, midrib not or
scarcely prominent; glands mostly linear or up to half punctiform. Petals bright
yellow to orange-yellow, sometimes margined red, 8-15 x 4-8 mm, c. 2 x sepals,
oblanceolate to narrowly obovate; apiculus acute; glands all linear or distally
interrupted. Stamens c. 40-200, longest 5-8 mm long, c. 0-5-0-7 x petals. Ovary
2-3-5 x 1-5-2 mm, ovoid; styles 3, (5-)6-9 mm long, 2-3 x ovary, spreading,
incurved distally; stigmas small or subcapitate. Capsules 4-5-5 x 3-5-5 mm,
subglobose to globose, shorter than sepals. Seeds c. 1 mm long, ecarinate; testa finely
scalariform.

On scrubby slopes and in moist woods immediately below them; 2600-4200 m.
Colombia (Santander, Norte de Santander, Cesar), Venezuela (Tachira). Map 2.

Hypericum phellos can be distinguished from its nearest relatives only by a combina-
tion of characters. Nearly always the stem internodes have corky ridges or emerg-
ences, especially below the node, and the leaves are deciduous. Where they axe
persistent, they are either marcescent with apex acute ('marcescens') or broad and
remaining appressed, thus apparently preventing the development of the corky
emergences (subsp. platyphyllum). The other recognizable subspecies (subsp. oro-
queanum) has leaves that remain appressed but eventually fall.

3a. Hypericum phellos subsp. phellos

H. tamanum Cuatrec. in Ciencia Mex. 4: 65 (1943). Type: Colombia, Norte de
Santander, Paramo de Tama, vicinity of La Cueva, 3100-3200 m, 27.x. 1941 (fl),
Cuatrecasas, Schultes & Smith 12647 (COL, holotype; BM!, F!, U!, US!, iso-
types).

H. thymifolium Cuatrec. in sched., non Kunth.

H. tachirense Steyerm. in sched.

Shrub or small tree 0-3-4 m tall, with shoots rounded to pyramidal or narrowly
cylindric, not or markedly differentiated into elongate main stems and short
laterals; internodes with slight or pronounced corky emergences. Leaves free,

5 1 1 1

Ill/

24

NORMAN K. B. ROBSON

Map 2 3. H. phellos (all subspecies) ; 10. H. simonsii A ; 11. H. papillosum O.

spreading then deciduous directly or after decurving or after deflexing and withering;
lamina 5-5-16 x 1-4-7 mm, oblanceolate to elliptic or oblong, acute to acuminate,
margin recurved to plane. Sepals 4-9 x 2-3 mm, narrowly oblong or elliptic to
lanceolate or ovate-lanceolate. Petals 10-15 x 4-8 mm. Stamens c. 40-200. Styles
5-9 mm long.

Colombia (Norte de Santander, Santander), Venezuela (Tachira); 2500-3900 m.

COLOMBIA. Norte de Santander: Paramo de Fontibon, 2600-2750 m, 15
-16.X.1941 (fl), Cuatrecasas, Schultes & Smith 12298 (BM, COL, GH, NY, U, US);
between Mutiscua and Pamplona, 3400 m, 23.ii.1927 (fl), Killip & Smith 19727 (GH,
S, US); Hoya del Rio Chitaga, Quebrada de Presidente, 3100-3300 m, 28.xi.1941
(fl), Cuatrecasas 13490 (COL, NY). Paramo de Tama, vicinity of the Cueva,
3100-3200 m, 27.X.1941 (fl), Cuatrecasas, Schultes & Smith 12647 (BM, F, US).
Santander: Paramo de Santurban entre Bucaramanga & Berlin, W. side, c. 3200 m,
3. i. 1960 (fl & fr), Barclay & Juajibioy 10422 (NY); Berlin to Picacho, 3058 m,
14.viii.1977 (fl), Renteria, Mantilla, Nino & Ortiz 555(3) (COL, MO); vicinity of La
Baja, 2700 m, 14-31. i. 1927, Killip & Smith 18756 (GH, NY, US).

VENEZUELA. Tachira: Paramo de Batallon, near La Grita, 3100 m, 8.iii.l979
(fr), Kieft 104 (BM, U, VEN); SE. of Paramo de Tama, Pata de Judio, near the
Colombia- Venezuela frontier, 2900-3000 m, 20.U968 (fl), Steyermark & Dunster-
ville 101230 (NY, VEN).

The following is a summary of the variation of H. phellos subsp. phellos. The
variation is continuous, thus preventing the recognition of these variants as taxa. The

THE GENUS HYPERICUM L. 25

names in single quotation marks are merely for reference and have no formal
nomenclatural status.

Variant 3a (i) 'marcescens': Stems not differentiated, internodal corky emergences
not pronounced; leaves eventually deciduous, drooping and marcescent, narrowly
elliptic to oblanceolate, acute to acuminate, margin recurved (Santander: Vetas, La
Baja).

Variant 3a (ii) 'angustum': Stems not differentiated, internodal corky emergences
evident; leaves eventually deciduous, drooping but not marcescent, oblanceolate to
linear, acute, margin recurved (Norte de Santander: Paramo de Fontibon, Mutis-
cua).

Variant 3a (iii) 'patens': Stems not differentiated, internodal corky emergences
evident; leaves deciduous when spreading, oblanceolate to elliptic, acute to obtuse,
margin recurved to plane (Santander: Paramo de Santurban, Norte de Santander:
Rio Chitaga, Tachira: Paramo de Batallon).

Variant 3a (iv) 'phellos': Stems not differentiated, internodal corky emergences
evident; leaves deciduous after becoming appressed, narrowly elliptic to oblan-
ceolate, acute, margin recurved to plane (Santander: La Baja, Paramo de Romeral).

Variant 3a (v) 'diversicaule': Stems often differentiated, internodal corky emerg-
ences evident; leaves deciduous when spreading, broadly elliptic to obovate, acute to
obtuse, margin recurved to plane (Norte de Santander: Paramo de Fontibon,
Mutiscua).

Variant 3a (vi) 'tamanum': Stems differentiated, internodal corky emergences
evident; leaves deciduous when spreading or ascending, narrowly elliptic to oblong,
subacuminate to acute, margin recurved (Norte de Santander and Tachira: Paramo
de Tama, Santander: Paramo de Almorzadero).

The type specimen of//, phellos apparently belongs to variant 3a (iv) 'phellos'.

3b. Hypericum phellos subsp. oroqueanum N. Robson, subsp. nov.

a subsp. phellos foliis ad caulem continue adpressis differt. Type: Colombia, Cordil-
lera Oriental, limites entre Los Departmentos Norte de Santander y Cesar, Jurisdic-
ciones, Cerro de Oroque, 3000-3700-3900 m, 22-27. vii. 1974, Garcia Barriga &
Jaramillo 20629 (COL!, holotype).

Small tree 4 m tall, with shoots rounded, not differentiated; internodes with
pronounced corky emergences. Leaves free, closely imbricate-appressed, tetra-
stichous, not or scarcely spreading, deciduous; lamina 5-5-8 x 1-6-2-2 mm,
narrowly elliptic-oblong to oblanceolate, acute, margin plane. Sepals c. 7 x 2-5 mm,
broadly elliptic. Petals 12-14 x 4-7 mm. Stamens c. 100. Styles 6-7 mm long.

Colombia (Norte de Santander/Cesar) ; 3700 m?
COLOMBIA. Known only from the type.

3c. Hypericum phellos subsp. platyphyllum (Gleason) N. Robson, stat.
nov.

Hypericum platyphyllum Gleason in Bull. Torrey hot. Club 56: 106 (1929) pro parte,
excl. spec. Killip & Smith 15616. Type: Colombia, Norte de Santander, Paramo de
Romeral, 3800-4200 m, 30.U927 (fl), Killip & Smith 18626 (NY!, holotype; A!,
COL,GH!,US!,isotypes).

Shrub 0-5-2 m tall, with shoots flattened terminally to rounded, not or slightly
differentiated; internodes smooth. Leaves united by interfoliar ridge, ascending to
closely imbricate, persistent; lamina 4-7 x 2-4 mm, broadly elliptic to obovate,
obtuse to rounded, margin recurved to incurved. Sepals 5-7 x 2-3 mm, narrowly
oblong to elliptic. Petals 8-14 x 4 mm. Stamens c. 80. Styles 5-5-7 mm long.

26

NORMAN K. B. ROBSON

Colombia (Norte de Santander, Santander), confined to the Paramos de Santurban
and de Romeral; 2700-4100 m.

COLOMBIA. Norte de Santander: Paramo de Santurban, extremo este, 3300
-3500 m, 27.vi.1940 (fl), Cuatrecasas & Garcia- Barriga 10297 (COL, F); Paramo de
Romeral, hoya del rio Cucutilla, 3600 m, 9.iv.l973 (veg), Cuatrecasas & Jaramillo
28751 (BM, COL, US). Santander: Paramo de Santurban, entre Berlin y Vetas,
3700-3800 m, 22.ix.1969 (fl), Cuatrecasas & Rodriguez 27897 (BM, COL, US);
Paramo de Romeral, 3800-4100 m, 29-30. i. 1927 (fl & fr), Killip & Smith 18580
(NY).

Killip & Smith 187716 (Paramo de Romeral) is intermediate between subsp.
platyphyllum and subsp. phellos. Gleason (1929) confused subsp. platyphyllum with
29 H. gleasonii (q.v.).

4. Hypericum garciae Pierce

in Bull. Torrey hot. Club 70: 174 (1943). Type: Colombia, Santander, Paramo del
Almorzadero, Peralonso, 3200-3300 m, 20.vii.1940 (fl), Cuatrecasas & Garcia-
Barriga 9935 (F!, holotype; NY!, US!, isotypes).

Hypericum chamaemyrtus sensu Gleason in Bull. Torrey hot. Club 56: 102 (1929),
non Triana & Planchon (1862).

Hypericum ericifolium Steyerm. in Fieldiana Bot. 28: 393 (1952) ['ericaefoUwn'].
Type: Venezuela, Tachira, Paramo de Tama, 2 km above Betania, 7 km above
Villapaez, 2500 m, 14.vii. 1944 (fi),Steyermark 57212 (F!, holotype; NY!, isotype).

Shrub 0-3-1 -2(-3) m tall, erect, with branches strict or ascending, lateral or
occasionally pseudo-dichotomous. Stems orange-brown to blackish, 4-lined and
ancipitous when young, the principal lines broader, eventually terete, without
epidermal emergences, cortex exfoliating in strips; internodes 1-4 mm long. Leaves
sessile, outcurving or ascending and subimbricate, tetrastichous, deciduous at
suprabasal articulation; lamina 4-10 x 0-5-2-3 mm, elliptic-oblong or narrowly
lanceolate to linear, broadly revolute and often concealing lower surface apart from
midrib, not carinate, concolorous, lucent above, papillose beneath, not glaucous,
coriaceous; apex acute, base cuneate, not sheathing, pairs united to form a narrow
interfoliar ridge; basal vein unbranched; laminar glands dense, slightly prominent.
Inflorescence l(2)-flowered, terminal and on short lateral shoots, occasionally with
pseudo-dichotomous branches; peduncle and pedicels 1-5-3 mm long, upwards
incrassate; upper leaves foliose. Flowers 17-20 mm in diam., stellate. Sepals 4-9 x
1-7-3 mm, lanceolate to ovate, acute to subacuminate; veins 5-7, obscure, not
visibly branching, midrib prominent; glands linear, distally punctiform. Petals bright
to deep yellow, sometimes tinged red, 8-13 x 3-6-5 mm, 1-5-2 x sepals, narrowly
obovate; apiculus acute; glands linear, interrupted distally. Stamens 40-70(7-100),
longest 5-8 mm long, c. 0-6-0-65 x petals. Ovary 1-5-2-5 x 1-2 mm, ovoid; styles
3(4), 3-5-7 mm long, 2-2-2-5(-3) x ovary, free, spreading-incurved; stigmas small
or slightly capitate. Capsule 4-5-5 x 3-3-5 mm, ovoid to subglobose, shorter than
sepals. Seeds c. 1-3 mm long, ecarinate; testa finely scalariform.

In paramo, usually on dry stony or sandy soil; 2400-3935 m.

Colombia (Norte de Santander to Meta), Venezuela (Tachira). Map 3.

COLOMBIA. Boyaca: Paramo de La Rusia, NW. to N. of Duitama, Serrania
Pena Negra, Hoya de la Laguna Agua Clara, 3935 m, 10.xii.1972 (fl), Cleef 6992
(COL, U); Vado Hondo, Siberia, between Pena de Arnical and Alto de Mogotas,
3290 m, 4.iv.l973 (fr & fr), Cleef '9332 (BM, COL, U). Cundinamarca: Municipio de
Tausa, paramo vecindad a la poblacion, 3200 m, 26.x. 1961 (fl), Huertas & Camargo
5285 (COL). Meta: Paramo de Sumapaz, Hoya El Nevado, Laguna El Sorbedero,
3550 m, l.ii.1972 (fl), Cleef 1489A (COL). Norte de Santander: Paramo de Hatico
between Toledo and Pamplona, c. 2900 m, 12-13.iii.1927 (fl), Killip & Smith 20661
(K, NY, US). Santander: Paramo del Almorzadero, 3600-3800 m, 28.xi.1941 (fl),
Cuatrecasas 13496 (F, NY, P); Paramo Rico, near Vetas, 3750-3850 m, 18.L1927
(fl), Killip & Smith 17659 (NY).

THE GENUS HYPERICUM L.

27

Map 3 4. //. garciae ; 5. H. acostanum

VENEZUELA. Tachira: Paramo de Tama, El Paramito, 2550 m, viii.1939 (fl),

H. garciae differs from the coriacous-revolute-leaved form of//, phellos ssp. phellos
('patens') principally by its narrower, closely revolute leaves, a character that
distinguishes it from all other species in sect. Brathys, except H. acostanum and //.
piriai. In addition, these species are unique in having the leaf-base incorporated in
the stem and forming a swelling, thus indicating that the leaf is not deciduous at the
base. The largest flowers and leaves in H. garciae occur in Boyaca, whence there is a
reduction trend along the Cordillera Oriental north-eastward to Tachira. As the
variation is continuous, H. ericifolium cannot be recognized as a distinct species.

5. Hypericum acostanum Steyerm. [in Acosta-Solis, Div. fitogeogr. y
formac. geobot. Ecuador: 103 (1968), nomen] ex N. Robson, sp. nov.

H. garciae Pierce af finis, sed foliis patulis persistentibus, pro ratione angustioribus,
uninervis, margine revolutis, sepalis angustioribus, petalis longioribus, inter alia
differt. Type: Ecuador, Loja, Canton Catachocha, Hacienda La Hamaca, Loma
Larga, 2200-2400 m, 15.iv.1944 (fl), Acosta Soils 7839 (F!, holotype; NY!, photo-
graph).

Shrub 0-1-0-8 m tall, erect, with branches strict, pseudo-dichotomous and lateral.
Stems orange-brown, 4- lined and ancipitous when young, the principal lines broader,
eventually terete, without epidermal emergences, cortex exfoliating in strips; inter-
nodes 2-5 mm long. Leaves sessile, outcurving or spreading but not imbricate,

28

NORMAN K. B. ROBSON

scarcely tetrastichous, persistent, not articulated but sometimes breaking off above
base; lamina 8-22 x 0-5-2-5 mm, narrowly oblanceolate to linear, broadly to
narrowly revolute, sometimes concealing lower surface apart from midrib, not
carinate, paler or ferrugineous and smooth beneath, lucent above, not glaucous,
coriaceous; apex acute, base parallel-sided, not sheathing, pairs united to form a
narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense, impressed.
Inflorescence 1 -flowered, terminal, often with pseudo-dichotomous branches;
pedicel 3-15 mm long, not upwards incrassate; upper leaves foliose. Flowers 20-25
mm in diam., stellate. Sepals 6-9 x 1-1-5 mm, narrowly triangular-lanceolate to
oblong-linear, acute, sometimes revolute above; veins 5-7, obscure, not visibly
branching, midrib slightly prominent; glands linear, distally punctiform. Petals
bright (?) yellow, tinged red in bud, 10-14 x 5-7 mm, c. 2 x sepals, obovate; glands
linear and striiform. Stamens c. 100, longest 4-5 mm long, c. 0-4 x petals. Ovary c. 2
x 1 mm, narrowly ovoid; styles 3, 3-3-5 mm long, c. 1-5 x ovary, divergent; stigmas
broadly capitate. Capsule 4-5 x 3-4 mm, ovoid-subglobose, shorter than sepals.
Seeds c. 1 mm long, ecarinate; testa finely scalariform.

On steep shrubby slopes of paramo; 2200-3300 m.

Ecuador (Loja, Morona-Santiago). Map 3.

ECUADOR. Loja: between San Pedro and Portovelo, slope towards Portovelo,
c. 2300 m, 13.X.1955 (fl & ir),Asplund 18133 (S); Celica-Guachanama road, Km 8 at
the Roldos memorial monument, 2700 m, 19. ii. 1985 (fl & fr), Marling & Anderson
2223 (BM, GB).

H. acostanum is clearly related to H. garciae, for which the nearest record is from
central Colombia (Meta).

6. Hypericum piriai Arechav.

in An. Mas. nac. Montevideo 3: 108 (1898), Fl. Uruguay 1: 108 (1898); L. B. Sm. in/.
Wash. Acad. Sci. 48: 314 (1958); Rodriguez Jimenez in Reitz, Fl. III. Catarin.,
Hipericdceas: 5, fig. 1 B (1980). Type: Uruguay, Mardonado, Pan de Azucar cerca
del Castillo de Piriapolis, xi.189- (fl & fr), Arechavaleta (MVM, holotype; GH!,
US!, photographs).

H. tenuifolium St.-Hil. in St.-Hil., Jussieu & Cambess., Fl. Bras. Merid. 1: 337
(1828); Walpers, Repert. Spec. nov. 1: 390 (1842); Dietrich, Syn. PI. : 1236 (1847);
Reichardt in Martius, Fl. Brasil. 12: 188 (1878), non Pursh (1814). Type: Brazil,
Sao Paulo, prope urbes Sancti Pauli, Villa do Castro et Curityba, iii.1820 (fl & fr),
St.-Hilaire 1174 (P!, holotype; US!, photograph).

H. hilaireanum L.B. Sm. in J. Wash. Acad. Sci. 48: 314 (1958); Angely, Fl. Anal.
Parana: 451 (1965). Type as for H. tenuifolium St.-Hil.

Subshrub or perennial herb 0-07-0-35 m tall, erect, with taproot, branches few, strict
or decumbent, lateral, from base or lower half of stem. Stems green, 4-lined above,
eventually terete, gland-dotted, with prominent glands along raised lines; cortex
eventually exfoliating in strips; internodes 3-21 mm long. Leaves sessile, outcurving
and deflexing, not tetrastichous, persistent, not articulated but occasionally breaking
off above base; lamina 10-40 x 1-2 mm, linear, revolute, concealing all or most of
lower surface apart from midrib, not carinate, paler beneath?, dull above,
glaucous, subcoriaceous; apex acute to acicular, base parallel, not sheathing, pairs
united to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands
dense, slightly impressed or not. Inflorescence l-18(-25)-flowered, terminal and
sometimes axillary, monochasial after first branching; pedicels 3-6 mm long; upper
leaves bracteose. Flowers 8-20 mm in diam., obconic to pseudotubular. Sepals 6-16
x 1-5-2-5 mm, linear-lanceolate to lanceolate or narrowly oblong, acute to acumin-
ate, margin slightly revolute or plane; veins 3-5, unbranched, midrib slightly
prominent or not; glands linear towards base or wholly punctiform. Petals bright (?)
yellow, not (?) tinged red in bud, 7-12 x 3-4-5 mm, 0-8-1-2 x sepals, obovate-
oblong; glands linear, distally punctiform. Stamens 25-60, longest 5-8(-10) mm, c.
0-75 x petals, 3(5)-fascicled, the double (antesepalous) fascicles sometimes incom-

THE GENUS HYPERICUM L.

pletely united, with filaments within each fascicle united shortly or usually to 0-7 of
their length. Ovary 1-5-4 x 1-1-5 mm, narrowly ovoid to narrowly ovoid-ellipsoid;
styles 3, 2-4 mm long, c. 1-1-3 x ovary, outcurving; stigmas subcapitate. Capsule
5-7 x 2-2-3 mm, ovoid-cylindric to ellipsoid, shorter than sepals. Seeds 0-8-1 mm
long, ecarinate; testa finely scalariform.

In dry, stony or damp grassland; lowland to 1000 m.

Brazil (Sao Paulo, Parana, Santa Catarina, Rio Grande do Sul), Uruguay. Map 4.

BRAZIL. Sao Paulo: Sao Paulo to Villa Emma, xii.1933 (fl), Brade 12978 (MO).
Parana: Municipio Guarapuava, estrada para Laranjeiras do Sul, 15. xi. 1957 (fl),
Hatschbach 4253 (US); Municipio Ponta Grossa, Parque Vila Velha, Furnas,
i.x.1965 (fl & fr), Hatschbach 12854 (K). Santa Catarina: Municipio Abelardo Luz,
8-12 km N. of Abelardo Luz, 900-1000 m, 15.xi.1964, Smith & Klein 13305 (NY,
US); Municipio Xanxere, 11 km N. of Abelardo Luz, 500-600 m, 25.xii.1956, Smith
& Klein 9231 (US). Rio Grande do Sul: Vila Manresa, prope Porto Alegre,
21. \.1951, Rambo 51325 (US); Porto Alegre, Morro da Policias, 21. ii. 1902 (fl & fr),
MalmeRegn. II 1408 (S).

URUGUAY. Mardonado: see type. Lavalleja: Co. Penitente, lO.i.1909 (fl),
fl?m>5533(US).

29

Map 4 6. H. piriai .

Despite the considerable distance between southern Ecuador and south-eastern
Brazil, there can be no doubt that H. piriai is a close relative of H. acostanum. The
character trends between H. garciae and H. acostanum are extended to H. piriai (for
example: tall tall shrub-dwarf shrub-subshrub to herb; leaves short, deciduous
-long, persistent; sepals short, broad-long, narrow, revolute-long to short, narrow,
revolute to plane). However, the flower shows new tendencies, viz. (i) towards
development of a cymose inflorescence and (ii) to become adapted to specialized
rather than open pollination. Unlike the other examples of this change in Hypericum
(H. elodes and the species of sect. 25. Adenotrias}, H. piriai shows intermediate
evolutionary stages and the specialization is less complex. Thus (i) the flower
apparently varies from obconic to pseudotubular and the stamen filaments within
each fascicle vary correspondingly from almost free to c. 0-7 united; (ii) the double
fascicles are often incompletely united; (iii) there are no fasciclodes to help open the
flower by expansion and no ligules on the petals to guide the tongues of foraging
insects.

30

NORMAN K. B. ROBSON

I agree with Rodriguez Jimenez (1980) that H. piriai cannot be divided into a
northern species (H. hilaireanum) and a southern one (H. piriai sensu stricto). The
more northern ones, suffruticose with fewer larger flowers and longer sepals, are the
nearest morphologically to H. acostanum.

9 9

\ o /

7. Hypericum irazuense Kuntze [Rev. Gen. PI: 61 (1891), in synon.;
Williams in Fieldiana Bot. 29: 355 (1961), nomen] ex N. Robson, sp. nov.

H. p Helios subsp. oroqueano N. Robson affinis, sed foliis persistentibus nervis haud
ramosis, differt; a H. stenopetalo Turcz. foliorum nervis basalibus haud ramosis
floribus maioribus, stylis longioribus, differt. Type: Costa Rica, Cartago, Volcan
Irazu, 3150 m, 25. vi. 1874 (fl), Kuntze 2359 (NY!, holotype; K!, isotype).

H. caracasanum var. scherzeri J. Zahlbr. , in sched.

H. silenoides sensu R. Keller in Engler & Prantl, Nat. Pflanzenfam. 3(6): 214 (1893),

in Bull. Herb. Boissier6: 258 (1898), in op. cit. II, 8: 177 (1908), in Engler & Prantl,

op. cit., 2nd ed. 21: 181 (1925), non Juss. (1804).

Icon: Fig. 5 A.

Shrub or small tree 0-4-5 m tall, erect, flat-topped, with branches strict, lateral and
sometimes pseudo-dichotomous, Stems yellow-brown, 4-lined and ancipitous when
young, becoming terete, with transverse corky emergences, cortex exfoliating in
strips; internodes 1-5-4 mm long. Leaves sessile, imbricate and tetrastichous at first,
spreading then becoming appressed, tardily deciduous near the base (usually after
withering) or persistent; lamina 10-15 x 1-5-3-5 mm, narrowly elliptic to oblan-
ceolate, plane to recurved, not carinate, concolorous, not glaucous, subcoriaceous to
coriaceous; apex acute, base cuneate, not sheathing, pair free but with incomplete
narrow interfoliar ridge; basal veins 3-5, parallel, inner pair (of 5) branching above,
tertiary reticulation not visible; laminar glands rather dense, impressed above, not
prominent beneath. Inflorescence 1 -flowered, terminal and on short lateral shoots,
with extension shoots from 2-3 nodes below; pedicel 2-5-4 mm long; upper leaves
bracteose. Flowers 25-30 mm in diam., stellate. Sepals 7-5-9 x 1-5-2 mm, linear-
lanceolate, acute; veins 5, unbranched, midrib not prominent; glands mostly linear,
punctiform near apex. Petals bright yellow to orange-yellow, not (?) tinged red,
13-18 x 7-10 mm, c. 2 x sepals, oblanceolate to obovate; apiculus acute; glands
linear, interrupted distally. Stamens c. 100, longest 6-9 mm long, 0-35-0-5 x petals.
Ovary 1-5-2-5 x 1-1-5 mm, ovoid; styles 3, 9-10 long, 4-6 x ovary, free, spreading,
distally incurved; stigmas small. Capsule 6-7 x 4-5 mm, broadly ovoid or ellipsoid
to globose, shorter than sepals. Seeds 1-1-3 mm long, scarcely carinate; testa finely
scalariform. 2n = 12.

On open paramo slopes or among bamboo (Chusquea); 2700-3730 m.

Costa Rica (San Jose, Cartago, Limon), on the cordillera and on Volcan Irazu and V.
Turrialba, and in adjacent Panama (Chiriqui). Map 5 (p. 32).

COSTA RICA. Cartago: Volcan Irazu, 3000-3300 m, 5.V.1928 (fl), Stork20ll (F,
MICH, NY); Finca Quemado to top of Volcan Turrialba, 2800-3200 m, lO.v.1971
(fl), Wilbur 14319 (DUKE); Cordillera de Talamanca, Cerro de la Muerte, Pan-
American Hwy., 5 km above Millsville, 3400-3500 m, 22.vii.1949 (fl), Holm & Iltis
458 (F, K, NY, P). Limon: Chirripo National Park, between Casa de Administration
and peak, c. 3400 m, 13. ii. 1983 (fl & fr), Garwoodetal. 1161 (BM). San Jose: Upper
Rio Talari, lower Valle de los Conejos and trail to Valle de los Leones, 3250-3450 m,
21-23.viii.1971 (fl & fr), Burger & Gomez P. 82848 (BM, DUKE, F); La Asuncion,
Cerro Sakira, 3400 m, 27.L1968 (fl), Wilbur & Stone 10047 (DUKE, F, MO).

PANAMA. Chiriqui: Cerro Fabrega and vicinity near Costa Rican frontier,
3150-3335 m, 7-8.iv.1976 (fl), Weston 10189 (MO).

H. irazuense has apparently been derived from H. phellos subsp. oroqueanum in a
similar way to the derivation of H. stenopetalum from H. phellos subsp. phellos, but it
can be distinguished from H. stenopetalum both morphologically and geographi-
cally.

THE GENUS HYPERICUM L.

31

Fig. 5 A. H. irazuense: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut
away) and ovary; (g) capsule. B. H. stenopetalum: (h) leaf; (i) sepal; (j) petal; (k) stamens (partly cut
away) and ovary (a x Vr, b x 1; d-g, i-k x 3; c, h x 4). A. Khan, Vickery & Tebbs 1398; B. Luteyn 6032.

The record of//, irazuense from Guatemala is apparently erroneous. A Guatema-
lan label was used for the Kew (K) specimen of Friedrichstal 1395, but the Chicago
(F) specimen of this collection has a similar label with 'Guatemala' crossed out and
replaced by 'Cartago'.

8. Hypericum stenopetalum Turcz.

in Bull. Soc. Nat. Moscou 31 (1): 385 (1858). Knuth in Reprium Spec. nov. Regni veg.
Beih. 43: 484 (1927); Gleason in Bull. Torrey Bot. Club 56: 103 (1929). Type:

32

NORMAN K. B. ROBSON

Map 5 7. H. irazuense A ; 8. H. stenopetalum .

Venezuela, Merida, Sierra Nevada, 2940 m, 1846 (fl), Funck & Schlim 1139 (LE,
holotype; BM!, P!, isotypes).

H. decorticans Planchon & Linden in Weddell, Chloris Andina 2: 272 (1861); Triana
& Planchon in Annls Sci. nat. (Bot.) IV, 18: 297 (1862), nomen.

H. meridense Steyerm. in Fieldiana Bot. 28 (2): 394. (1952). Type: Venezuela,
Merida, between San Jose and Beguilla, Paramo de Pozo Negro, 2590-3220 m,
3. v. 1944 (fl & fr), Steyermark 56272 (NY!, holotype; F!, isotype).

Icon: Fig. 5B.

Shrub or small tree 0-5-4 m tall, erect, branches strict or ascending, lateral or rarely
pseudo-dichotomous. Stems yellow-brown, 4-lined and ancipitous when young,
becoming terete, without or with weak corky emergences, cortex exfoliating in
strips; internodes 1-5-3 mm long. Leaves sessile, imbricate and tetrastichous at first,
becoming appressed, eventually deciduous near base or persistent; lamina 4-15 x
(1-)1 -5-4-5 mm, narrowly or rather broadly elliptic to oblanceolate, plane or
recurved or rarely concave to cucullate, not or scarcely carinate, concolorous, not
glaucous, subcoriaceous to coriaceous; apex shortly acuminate to subacute, base
cuneate to angustate, not sheathing, pairs free but with incomplete narrow interfol-
iar ridge; basal veins 3-5, diverging, all or only midrib branching, tertiary reticula-
tion obscure or not visible; lamina glands rather dense, impressed above,
prominent beneath. Inflorescence 1-flowered, terminal and sometimes on short
lateral shoots, with extension shoots usually from 2nd or 3rd node below; pedicel 1-9
mm long; upper leaves foliose or bracteose. Flowers 15-27 mm in diam., stellate.
Sepals 4-9 x 1-5-3 mm, narrowly oblong or elliptic to ovate-lanceolate or rarely
oblanceolate, acute or shortly acuminate to obtuse; veins 5-7, branched distally or
not, midrib not or scarcely prominent; glands mostly linear or up to half punctiform.
Petals bright yellow to orange-yellow, sometimes margined red, (6-)8-15 x 4-8
mm, c. 2 x sepals, oblanceolate to obovate; apiculus acute; glands all linear or
distally interrupted. Stamens 65-120, longest 5-8 mm long, 0-5-0-7 x petals. Ovary
2-3 x 1-2 mm, broadly ovoid; styles 3, (5-)6-9 mm long, 2-4 x ovary, free,
spreading, distally incurved; stigmas small or subcapitate. Capsule 4-6 x 3-5-5 mm,
broadly ovoid to globose, shorter than sepals. Seeds c. 1 mm long, ecarinate; testa
finely scalariform.

On open scrubby or stony paramo slopes and among shrubs or bamboos in the
tree/shrub zone immediately below them; 2580-4200 m.

THE GENUS HYPERICUM L.

Colombia (Santander), Venezuela (Tachira, Merida). Map 5.

COLOMBIA. Santander: [Paramo de Almorzadero] Peralonso, Timocato, 3600
m, 29. ix. 1969 (fl), Cuatrecasas & Rodriguez 27871 (BM, COL, US); Paramo de
Santurban, c. 3000 m, 27.viii.1948 (fl), Barkley & Araque M. 18sl69 (COL, US).

VENEZUELA. Merida: Paramo de Los Leones (La Lagunita, La Canada
Grande) al W. de Murucuba, 3400-3500 m, 31. v. 1930 (fl), Gehringer 159 (F, NY,
VEN); Paramo de San Jose, monte El Cupio, 3000 m, l.ii.1973 (fl & fr), Cuatrecasas,
Ruiz-Teran & Lopez-Figueras 28440 (BM, US). Tachira: Paramo de Portachuelo,
2860 m, 23.X.1978 (fl), Luteyn 6032 (BM, MO, NY, VEN).

Linden 446 (BM, K, W) is labelled 'Caracas'. In the absence of other records from
that area this probably merely indicates Venezuela.

33

9. Hypericum carinosum R. Keller

in Bull. Herb. Boissier, II, 8: 182 (1908), in Engler & Prantl, Nat. Pflanzenfam. 2nd
ed. 21: 181 (1925); Knuth in Reprium Spec. nov. Regni veg. Beih. 43: 483 (1927);
Gleason in Bull. Torrey hot. Club 56: 102-103 (1929). Type: Venezuela, Merida,
Paramo de Portachuelo, 2600-2700 m, 1846 (fl), Funck & Schlim 1459 (W!,
holotype; BM!, P!, isotypes).

H. nova species sensu Turcz. in Bull. Soc. Nat. Moscou 31 (2): 386 (1858); Knuth in

Reprium Spec. nov. Regni veg. Beih. 43: 484 (1927).
H. struthiolifolium sensu Trev., Hyper. Animad: 15 (1861) ['struthiolaefoliutn'] pro

parte, excl. typum.
H. thymifolium sensu Triana & Planchon inAnnls Sci. nat. (Bot.) IV, 18: 297 (1862)

pro parte, quoad syn. Trevirani.
H. stenoclados Cuatrec. in Ciencia Mex. 4: 64 (1943). Type: Colombia. Santander,

Paramo de Almorzadero, extremo sur, Peralonso, 3200 m, 19.vii.1940 (fl),

Cuatrecasas & Garcia Barriga 9919 (COL, holotype; F!, NY!, US! isotypes).

Shrub 0-6-C.2-5 m tall, erect, with branches strict, lateral, short (flowering) and long
(extension). Stems purplish-brown, 4-lined and ancipitous when young, eventually
terete, without epidermal emergences, cortex exfoliating in strips; internodes 2-9
mm long. Leaves sessile, spreading from the base or ascending but not imbricate or
tetrastichous, persistent; lamina 3-5-5 x 0-8-2 mm, narrowly elliptic or narrowly
oblong to oblanceolate-spathulate, plane, rarely apically subconcave, markedly
carinate, concolorous, not or slightly glaucous, chartaceous to subcoriaceous; apex
acute to subacute, base cuneate to angustate, not sheathing, pairs almost free
(interfoliar ridge rudimentary); basal vein 1, unbranched; laminar glands dense, not
impressed above, prominent beneath. Infloresence l-2(-3)-flowered, terminal and
on short lateral pinnate branches; peduncle and pedicels c. 2 mm long; upper leaves
bracteose. Flowers 8-13 mm in diam., stellate. Sepals 2-4-5 x 0-5-1-8 mm,
ovate-lanceolate to triangular-lanceolate, acute; veins 3(5), unbranched, midrib
distally incrassate; glands linear, interrupted distally. Petals bright (?) yellow, 4-8 x
2-3 mm, c. 2 x sepals, oblong-lanceolate to obovate; apiculus acute; glands linear,
interrupted distally. Stamens 40-80, longest 3-4 mm long, 0-5-0-6 x petals. Ovary
1-5-2 x 0-8-1 mm, ovoid; styles 3, 4-6(-7) mm long, c. 2-3 x ovary; stigmas small.
Capsule c. 4 x 3-5 mm, subglobose, shorter than sepals. Seeds not seen.

In thickets and woodland margins in the paramo; 2600-3200 (-4500) m.

Colombia (Norte de Santander, Santander), Venezuela (Merida). Map 6 (p. 37).

COLOMBIA. Norte de Santander: environs of Pamplona. 10.x. 1943 (fl), de
Garganta 626 (F). Santander: vicinity of Las Vegas, 2600-3600 m, 21-23.xii.1926,
Killip & Smith 15921 (GH, NY, US); vicinity of California, 2800 m, 11-27.U927,
Killip & Smith 16904 (F, GH, NY, US).

VENEZUELA. Merida: Paramo de Los Leones, 3400-4500 m, 31.V.1930 (st),
Gehringer 149 (VEN).

H. carinosum is related to H. phellos subsp. phellos 'diversicaule', and its area of
distribution is to the south-east of that of the latter. The two taxa, however, are quite

34

NORMAN K. B. ROBSON

distinct, H. carinosum being recognizable (inter alia) by the stem internodes without
corky emergences, the smaller, persistent, 1-nerved leaves, and the smaller flowers.
H. stenoclados has subacute, subcoriaceous leaves rather than the acute, char-
taceous leaves typical of//, carinosum, but these differences do not merit taxonomic
recognition.

10. Hypericum simonsii N. Robson, sp. nov.

H. phellos affinis, sed caulis haud suberosis, foliis persistentibus deflexis oblongis
apice rotundatis basi conjunctis venatione valde reticulata, sepalis subacutis vel
rotundatis, stylis (5-)6-9 mm longis, differt. Type: Colombia, Magdalena, Sierra
Nevada de Santa Marta, 1880 (fl), Simons s.n. (BM!, holotype).

H. stenopetalum [var.] (3 majus Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18:
297 (1862) ['major']. Types: Colombia, Magdalena, Santa Marta, source ['prov.']
of Rio Hacha, 3700-3800 m, vii.1844 (fl & fr), Purdies.n. (G, lectotype; GH!, K!,
syntypes); ibid., 1851-1852 (fl), Schlim 838 (G, syntype).

Icon: Fig. 6A.

Shrub 1-2 m tall, erect, with branches ascending, mostly lateral. Stems reddish-
brown, 4-lined when young, eventually terete, without epidermal wrinkles, cortex
flaking irregularly; internodes 2-10 mm long. Leaves sessile, spreading from the
base, not tetrastichous, becoming brown and deflexed, usually persistent until cortex
is shed; lamina (7-)8-16(-20) x (2-2-)3-5-2(-7) mm, narrowly elliptic to narrowly
oblong, plane, not concave or carinate, concolorous, pale green, chartaceous; apex
rounded, base cuneate, not sheathing, pairs united to form a narrow interfoliar
ridge; basal or near-basal veins 5-7, with branches and tertiary reticulation rather
conspicuous; laminar glands dense, visible beneath only. Inflorescence 1 (2)-
flowered, mostly terminal and lateral, occasionally pseudo-dichotomous; peduncle
and pedicels 4-7 mm long; upper leaves not transitional. Flowers 20-35 mm in
diam., stellate. Sepals 6-10-5 x 2-4-3-5 mm, lanceolate or narrowly elliptic to
narrowly oblong or oblanceolate, subacute to rounded; veins 5-7, not or obscurely
branched, midrib not prominent; glands distally punctiform. Petals bright yellow,
10-21 x 7-13 mm, c. 2 x sepals, oblanceolate; apiculus obtuse; glands distally
interrupted. Stamens c. 100, 'obscurely 5-fascicled', longest c. 1 mm long, 0-4-0-5 x
petals. Ovary 2-5-3-5 x 2-5 mm, ovoid; styles 3-4, 4-5-6 mm long, c. 2 x ovary,
spreading, distally incurved; stigmas narrowly capitate. Capsule c. 8 x 5 mm,
subglobose, shorter than sepals. Seeds not seen.

In paramo, bushy prairies, and thickets; 3250-4100 m.

Apparently confined to the Sierra Nevada de Santa Marta in Colombia (Magdalena).
Map 2 (p. 24).

COLOMBIA. Magdalena: near Rio Seville, 20.L1959 (fl), Barclay & Juajibioy
6557 (MO, NY); Sierra de Santa Marta, c. 48 km inland from Dibulla, c. 3850 m,
vii.1932 (fl), Seifriz 455 (US); Sierra de Santa Marta, SE. slopes, Hoya del Rio
Donachui, Meollaca, 3320-3260 m, 29.ix.1959, Cuatrecasas & Romero -Castaneda
24480 (COL, US); Sierra de Santa Marta, source of Rio Donachui, 4310 m,
22.V.1977, Starker White & Alverson 547A (NY); Sierra de Santa Marta, quebrada
from Laguna Rio Frio, 3250 m, 28.vii.1972, Kirkbride & Forero 1751 (BM, COL).

H. simonsii, which is apparently confined to the Sierra de Santa Marta massif, is
related to H. phellos (Venezuela-Colombia border area). It is morphologically more
specialized in that its leaves are united by an interfoliar ridge, but it is more primitive
in that they have 5-7 basal or near-basal veins.

11. Hypericum papillosum N. Robson, sp. nov.

H. simonsii N. Robson affinis, sed habitu humiliore densiore, foliis sepalisque
subtiliter papillosis, foliis angustioribus apice obtusis vel rotundatis basi breviter
vaginatis, stigmatibus valde capitatis, differt. Type: Colombia, Boyaca, Sierra

THE GENUS HYPERICUM L.

35

Fig. 6 A. H. simonsii: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut
away) and ovary; (g) capsule. B. H. papillosum: (h) habit; (i) leaf; (j) sepal; (k) petal; (1) stamens (partly
cut away) and ovary; (m) young capsule (a, h x l /r, b x 1 ; c-f x 4; g x 2; i-m x 3). A. Romero-Castaneda
4541; B: (h) Grubb, Curry & Fernandez-Perez 401; (i-m) Jaramillo Mejia 2728 A.

Nevada del Cocuy, above Laguna Grande, c. 3900 m, 14.viii.1957 (fl), Grubb, Curry
& Fernandez-Perez 401 (K!, holotype; US!, isotype).

Icon: Fig. 6B.

Shrub to c. 1m tall, erect, rounded, with branches ascending or spreading, mostly
lateral. Stems orange-brown, 4- lined and ancipitous when young, eventually terete,
without epidermal wrinkles, cortex flaking irregularly; internodes 2-9 mm long.
Leaves sessile, spreading from base or just above base, not tetrastichous, becoming
brown and deflexed, persistent until cortex is shed; lamina 10-22 x 4-7 mm,

36

NORMAN K. B. ROBSON

narrowly elliptic to narrowly oblong, plane, not concave or carinate, finely papillose
on both sides, concolorous, pale green, chartaceous; apex obtuse to rounded, base
cuneate, shortly sheathing at least when young, pairs united to form narrow
interfoliar ridge; basal or near-basal veins 3-7, with branches and tertiary reticula-
tion inconspicuous; laminar glands rather dense, visible on both sides or only
beneath. Inflorescence 1-flowered, mostly terminal and lateral, occasionally pseudo-
dichotomous; pedicel 4-6 mm long; upper leaves not transitional. Flowers c. 30 mm
in diam., stellate. Sepals 9-11 x 3-5 mm, oblong to lanceolate-oblong or oblan-
ceolate, subacute to obtuse, with epidermis papillose; veins 5, laterals branched,
midrib prominent; glands mostly punctiform. Petals bright yellow, 14-17 x 9-10
mm, c. 1-5-1-8 x sepals, oblanceolate; apiculus obtuse; glands striiform or puncti-
form. Stamens c. 100, c. 1 mm long, longest c. 0-5 x petals. Ovary c. 2-5 x 2 mm,
ovoid; styles 3, c. 5 mm long, c. 2 x ovary, spreading, distally incurved; stigmas
distinctly capitate. Capsule and seeds not seen.

On dry rocky slopes in paramo and subparamo; 3530-C.3900 m.

Colombia (Boyaca). Apparently confined to the Sierra Nevada del Cocuy and the
Paramo de Pisva/Paramo de Chita. Map 2 (p. 24).

COLOMBIA. Boyaca: Paramo de Pisva, road between Socha and La Punta, Km
61, 5-5 km E. of Los Pinos, 3680 m, 14.vi.1972 (fl), Clee/4541 (U); same road, Km
70, 1-5 km, W. of El Cadillal, 3530 m, 16.vi.1972 (fl), C/ee/4631 (U); Paramo de
Chita, vertiente oriental, sitio Piedras Negras, 3500-3700 m, 13.vii.1967 (fl), Jara-
millo & van der Hammen 2128 A (COL).

The papillose leaf- and sepal-epidermis (making them appear duller green) clearly
distinguish this isolated endemic of Dept. Boyaca from H. simonsii and, indeed,
from nearly all other species in sect. Brathys. It also differs from H. simonsii in the
obscure or almost invisible tertiary leaf -venation.

12. Hypericum lycopodioides Triana & Planchon

inAnnls Sci. not. (Bot.) IV, 18: 296 (1962). Type: Colombia, Cundinamarca, Andes
de Bogota, Cipaquira [Zipaquira], viii.1855, (fl), Triana 5464 (COL?, holotype;
BM!).

H. gyropodioides sensu Jackson & Hooker, Index Kew. 1: 1193 (1893), sphalm.

Shrub 0-8-2-5 m tall, erect, with branches strict to ascending, lateral, short (flower-
ing) and long, narrowly spiciform to cylindric (extension). Stems dark red-brown,
4- lined when young, eventually terete, without epidermal emergences, cortex
exfoliating in strips or small flakes; internodes 1-5-2-5 mm long. Leaves sessile,
imbricate to spreading, eventually reflexed, markedly terastichous; lamina 3-6 x
1-5-3 mm, oblong to elliptic or lanceolate, incurved-conduplicate, midrib usually
prominent beneath, margin narrowly hyaline and plane or undulate, not glaucous,
subcoriaceous; apex acute to acuminate, slightly cucullate, base broadly cuneate to
angustate, loosely clasping, pairs united to form narrow interfoliar ridge; basal vein
1, sometimes branched; laminar glands dense, not impressed or prominent, visible
on both sides. Inflorescence 1-flowered, terminal and on short lateral branches;
pedicels absent; upper leaves foliose, appressed to calyx. Flowers (15-)25-30 mm in
diam., stellate. Sepals (5-)6-7 x (2-)2-5-3 mm, ovate-oblong to ovate-lanceolate,
acute to obtuse, subcucullate, margin hyaline; veins 7, unbranched, midrib
scarcely prominent; glands linear, interrupted distally. Petals bright yellow, 13-16 x
9-10 mm, 2-2-2 x sepals, very obliquely obovate; apiculus subacute; glands
striiform and punctiform. Stamens c. 120, longest 6-8 mm long, c. 0-5 x petals.
Ovary 2-5-3 x 1-5-2 mm, ellipsoid; styles 3, 4-5 mm long, c. 1-4 x ovary,
spreading-incurved; stigmas small to narrowly capitate. Capsule c. 6 x 3-5 mm,
ellipsoid, shorter than sepals. Seeds 1-1-2 mm long, ecarinate; testa finely reticulate-
scalariform.

In subparamo and paramo, most frequently in thickets; 2850-4100 m.

THE GENUS HYPERICUM L.

37

Map 6 9. H. carinosum A ; 12. H. lycopodioides O; 13. H. thuyoides .

Colombia (Arauca, Boyaca, Cundinamarca) , on the humid side of the Cordillera
Oriental from Sierra Nevada del Cocuy to Paramo de Zipaquira. Map 6.

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Hoya San
Jose, 3450 m, 9.vi.l973 (fl), Cleef 10117 (U). Boyaca: Sierra Nevada del Cocuy,
Valle de San Jose, c. 4000 m, 10.ix.1957 (fl), Grubb, Curry & Fernandez-Perez 768
(COL, K, US); between Chita and Sacama, Quebrada del Curial (La Porquera),
3350 m, 15.ix.1969 (fl), Cuatrecasas & Laureano Rodriguez 27787 (BM, COL, US).
Cundinamarca: see type.

H. lycopodioides is morphologically and geographically intermediate between 10 H.
simonsii and 13 H. thuyoides. It is very similar to the latter, differing essentially in the
stiff, acute to acuminate leaves that give the shoots an Araucaria-like appearance.

13. Hypericum thuyoides Kunth

in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. PI. 5: 187, t. 456 (1822); R.
Keller in Bull. Herb. Boissier II, 8: 177 (1908), in Engler & Prantl, Nat. Pflanzen-
fam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey hot. Club 56: 101 (1929).
Type: Colombia, Tolima?, Andes de Quindiu, entre Cartago et Ibague, x.1805
(fl), Humboldt & Bonpland (P-HUM!, holotype; B-WILLD, P!, isotypes).

Brathys thuyoides (Kunth) Spach, Hist. not. veg. Phan. 5: 448 (1836), in Annls Sci.

nat. (Bot.) II, 5: 366 (1836).
H. hartwegii var. patens R. Keller in Bull. Herb. Boissier II, 8: 182 (1908). Type:

Colombia, Cundinamarca, mountains to the east of Bogota (fl), Holton 785 (K!).
H. genistoides Kunth in sched.

Icon: Humboldt, Bonpland & Kunth, Nova Gen. etSp. PI. 5: t. 456 (1822).

Shrub or small tree (0-5-)l-3(?-6) m tall, spreading, with branches ascending,
lateral, short (flowering) and long, narrowly spiciform to narrowly conical (exten-
sion), or rarely pseudo-dichotomous. Stems purplish-brown, 4-lined when young,
eventually terete, without epidermal emergences, cortex exfoliating in strips; inter-
nodes 1-5-3 mm long. Leaves sessile, spreading to recurved, tetrastichous, persis-
tent; lamina 2-8 x 1-2 mm, sometimes increasing in size along shoot, elliptic or
rhombic-elliptic or elliptic-oblong to oblanceolate-spathulate or narrowly oblong,
incurved-conduplicate, midrib prominent beneath or not, margin narrowly hyaline

38

NORMAN K. B. ROBSON

and usually undulate, not glaucous, thinly chartaceous; apex acute to obtuse,
cucullate, base cuneate to angustate or shortly pseudopetiolate, pairs united to form
marked interfoliar ridge; basal vein 1, branched or usually unbranched; laminar
glands dense, visible on both sides or above only. Inflorescence 1-flowered,
terminal and on short lateral branches, the whole shoot cylindric to conic and
sometimes racemiform due to retarded maturity of terminal flower, rarely with
pseudo-dichotomous branches from node below; pedicel amost absent; upper leaves
foliose. Flowers 20-30 mm in diam., stellate. Sepals 3-5 x 1-5-2-5 mm, oblong-
lanceolate or elliptic to obovate-elliptic, acute to obtuse, margin narrowly hyaline;
veins 9-11, not or scarcely branched, midrib apically incrassate; glands linear,
punctiform distally. Petals bright yellow to orange-yellow, 10-17 x 6-11 mm, 3-4 x
sepals, very obliquely obovate; apiculus obtuse; glands striiform with few distal dots.
Stamens 100-120, longest 5-6 mm long, c. 0-4-0-5 x petals. Ovary 2-3 x 1-2 mm,
ovoid to ellipsoid; styles 3, 4-7 mm long, c. 2 x ovary, spreading; stigmas scarcely to
narrowly capitate. Capsule c. 3-5 x 2 mm, ellipsoid, shorter than sepals. Seeds not
seen.

In scrub and thickets on slopes of the paramo; especially in boggy areas; 2500-
3600 m.

Colombia (Boyaca, Cundinamarca, Tolima?, Meta); mainly in the mountains
around Bogota. Map 6.

COLOMBIA. Boyaca: Sogamosa to Pajarito, near Km 268, c. 2500 m,
27.viii.1953 (fl), Langenheim 3601 (COL). Cundinamarca: Bogota to Choachi, 30
km, Paramo de Cruz Verde, 3050 m, 8.viii.l974 (fl), Grabandt & Idrobo 219 (COL);
Municipio de La Calera, Paramo de La Siberia, 3000-3500 m, 25.X.1953 (fl),
Humbert et al. 26890 (COL, P, S) Meta: Paramo de Sumapaz, Hoya Sitiales, 300 m
an al NW. de la Laguna La Primavera, 3580 m, 25.: 72 (fl), Cleef985 (COL, U).
Tolima?: see type.

H. thuyoides is a rather variable species with a restricted distribution in central
Colombia, records from Narino and northern Ecuador being errors for 20 H.
laricifolium (q.v.).* It is easily distinguishable from typical H. lycopodioides by the
thinner, usually undulate leaves, and from H . laricifolium by the denser habit and the
usually broader or spathulate, undulate leaves with midrib not impressed beneath.
15 H. myricariifolium, with which it has been confused, has smaller, thicker, broader
leaves and corky emergences near the top of the upper internodes, and the cortex
exfoliates in irregular flakes.

Some specimens are very close in form to H. lycopodioides (e.g. Humbert et al.
26890). From these there are morphological trends in two directions: i) towards
narrower leaves and narrower, spiciform shoots ('//. genistoides' in sched.) (e.g.
Langenheim 3601); ii) towards broader, shorter leaves and broader, pyramidal
shoots, thus approaching 14 H. goyanesii (e.g. Grabandt & Idrobo 219). Extreme
forms of trend i) have been confused with H. laricifolium, which has thicker, more
lucent leaves with midrib impressed.

14. Hypericum goyanesii Cuatrec.

[in Trab. Mus. nac. Cienc. not. Jard. hot. Madrid (Bot.) no. 27: cuadro 7 inter 54 et 55
(1934), nomen] in op. dr., no. 29: 9 (1935), op. cit. , no. 33: 83, f. 1A (1936). Type:
Colombia, Cundinamarca, Guasca, Los Gaques wood, 3000 m, 24. iv. 1932 (fl),
Cuatrecasas 3229 (MA, holotype; K!, isotype).

Shrub 1-3 m tall, spreading, with branches ascending to divergent or pendent,
lateral, short (flowering) and long (extension), not pseudo-dichotomous. Stems
reddish-brown, 4-lined and ancipitous when young, sometimes wrinkled, eventually
terete, cortex exfoliating in irregular flakes; internodes 2-8 mm long. Leaves sessile,
spreading to deflexed, tetrastichous; lamina 2-5-5 x 1-5-3-5 mm, increasing in size

* Knuth's record from Venezuela, Merida, Paramo de Timotes (John 847) (Reprium Spec,
nov. Regni veg. Beih. 43: 484 (1927)) must be a misidentification, but I have not seen the
specimen.

THE GENUS HYPERICUM L.

along shoot, oblong-ovate or ovate to triangular-ovate or subcircular, incurved-
conduplicate or saccate, with midrib slightly prominent beneath or not, margin
narrowly hyaline, not glaucous, chartaceous, deciduous almost at the base after
turning brown, leaving narrow basal ring; apex subacute or apiculate-obtuse to
rounded, base cordate-amplexicaul, pairs perfoliately united; basal vein 1,
sometimes with 1-2 pairs of near-basal branches, rarely with visible tertiary reticu-
lum; laminar glands dense to rather few, large, prominent on both sides.
Inflorescence 1 -flowered, terminal and on short lateral branches, the whole shoot
cylindric to conical, without pseudo-dichotomous branches; pedicel 1-3-5 mm long;
upper leaves foliose. Flowers 20-40 mm in diam., stellate. Sepals 5-8 x 2-3-5 mm,
the outer oblong to ovate, often cordiform, the inner oblong to lanceolate, acute or
obtuse to rounded, cucullate; margin narrowly hyaline; veins 7-11, branching and
anastomosing distally, midrib distally incrassate; glands linear, distally punctiform.
Petals bright yellow, 10-20 x 6-12 mm, 2-2-5 x sepals, obovate; apiculus obtuse;
glands striiform and punctiform. Stamens c. 120-150, longest 5-6 mm long, c.
0-4-0-5 x petals. Ovary 2-5-3 x 1-5 mm, narrowly ovoid; styles 3, 5-8 mm long, c.
2-2-5 x ovary, spreading-incurved; stigmas narrowly capitate. Capsule 5-6 x 4mm,
broadly ellipsoid to subglobose, equalling or exceeding sepals. Seeds c. 1 mm long,
ecarinate; testa finely scalariform-reticulate.

In degraded forest and scrub on open slopes and banks in the paramo and subpara-
mo, mainly near water and sometimes in pure stands (Lozano-C. & Schnetter
(1976)); 2730-3590 m.

Colombia (Cundinamarca, Meta). Map 7.

39

Map 7 14. H. goyanesii A ; 16. H. cuatrecasii ; 17. H. quitense .

40

NORMAN K. B. ROBSON

COLOMBIA. Cundinamarca: Paramo de Guasca, 3300 m, 16.xii.1938 (fl), Balls
5764 (BM, COL, K); Paramo de Zipaquira [Guerrero], between Zipaquira and
Pacho, 3100-3200 m, 16.vi.1940 (fl), Cuatrecasas 9526 (COL, F, P); Paramo de Cruz
Verde, fondo del valle al sur de La Viga, 3590 m, 28.viii.1972 (fl), Cleef3287 (BM,
COL, U). Meta: Macizo de Sumapaz, alredores de la Laguna La Guitarra, 3380
-3420 m, 4.vii.l981 (fl & fr), Diaz 2398 (COL).

H. goyanesii resembles H. thuy aides (which occurs in the same area) but differs in,
for example, the thicker, saccate, cordate-amplexicaul leaves that become deflexed
when withering, the spreading habit, and the exfoliation of the cortex in flakes rather
than strips. It is related to both H. thuyoides and 16 H. cuatrecasii.

15. Hypericum myricariifolium Hieron.

in Bot. Jahrb. 21: 321. (1895); R. Keller in Engler & Prantl, Nat. Pflanzenfam. 2nd
ed. 21: 181 (1925); Gleason in Bull. Torrey hot. Club 56: 102 (1929). Type:
Colombia, Cundinamarca, Paramo de Pasca, 3700-3800 m, vi.1868? (fl), Stubel
Colomb. 124 (Bt, holotype; F!, NY!, photographs).

H. myricariifolium Hieron. ex R. Keller in Bot. Jahrb. 42: 130 (1908), in Bull. Herb.

Boissier II, 8: 178 (1908). Type: as for H. myricariifolium Hieron.
H. lindenii sensu Cuatrec. in sched., non R. Keller.

Shrub up to 2 m tall, bushy or with branches ascending to spreading or prostrate and
matted, lateral, short (flowering) or long and narrowly conical (extension), rarely
pseudo-dichotomous. Stems orange-brown, 4-lined when young, sometimes wrink-
led or with corky emergences especially near flowers, eventually terete, cortex
exfoliating in irregular flakes; internodes 1-4-6 mm long. Leaves sessile, imbricate to
spreading, tetrastichous, cupressoid; lamina 1-3-5 x 0-7-2(-2-6) mm, triangular-
ovate to oblong-ovate, incurved-cucullate, with midrib plane or usually impressed
beneath, margin relatively broadly hyaline, not glaucous, coriaceous; apex subacute
to obtuse, base rounded to cordate-amplexicaul, pairs shortly united; basal vein 1,
unbranched, without visible tertiary reticulum; laminar glands dense to few, large,
prominent beneath or not. Inflorescence 1 -flowered, not terminal, on short lateral
branches, the whole shoot narrowly conical, racemiform, rarely with a pseudo-
dichotomous branch; pedicel 1-1-5 mm long; upper leaves foliose. Flowers 18-25
mm in diam., stellate. Sepals 4-5 x 1-7-2-7 mm, the outer ovate, sometimes
cordiform, the inner ovate-lanceolate to lanceolate-oblong, acute to rounded,
cucullate, margin narrowly to rather broadly hyaline; veins 9-11, usually unbran-
ched, midrib not prominent beneath; glands linear to punctiform. Petals bright (?)
yellow, 10-15 x 6-9 mm, c. 2-2-3 x sepals, obovate; apiculus rounded; glands
striiform and punctiform. Stamens 60-80, longest 5-6 mm long, c. 0-35-0-5 x petals.
Ovary 2-3 x 1-1-5 mm, narrowly ovoid-ellipsoid; styles 3, 5-7 mm long, c. 2-2-5 x
ovary, spreading-outcurved; stigmas narrowly capitate. Capsules 5x4 mm, broadly
ellipsoid, exceeding sepals. Seeds not seen.

On open slopes in the paramo; 3000-4120 m.

Colombia (Cundinamarca); in the higher paramos near Bogota. Map 8 (p. 44).
COLOMBIA. Cundinamarca: Cordillera Oriental S. of Usme, Paramo de Chisa-
ca, c. 4120 m, 9-ll.xi.1958 (fl), Barclay & Juajibioy 6178 (COL, MO, NY); Macizo
de Sumapaz, Andabobos, 3760-3720 m, 8.U969 (fl), Cuatrecasas & Jaramillo 27031
(BM, US); Paramo de Cruz Verde, W. slopes, 3150 m, 7.X.1938 (fl), Cuatrecasas 409
(COL, F, P).

H. myricariifolium was formerly though to be inseparable from H. thuyoides (see
Cuatrecasas, 1936: 83). However, it can be distinguished inter alia by the thick,
broad-based, cucullate leaves with a broad hyaline margin and large glands. Its
nearest relative is H. goyanesii, the area of which includes that of//, myricariifolium.
It is clearly a high-altitude derivative of//, goyanesii, and the altitudinal ranges of the
two taxa overlap from 3000-c. 3600 m; there is no evidence of hybridization between
them. They can be distinguished by the leaf shape (see key), which gives the young

THE GENUS HYPERICUM L.

shoots of H. myricariifolium a cupressoid appearance lacking in H. goyanesii, the
leaf aspect (leaves becoming reflexed in H. goyanesii but no more than spreading in
H. myricariifolium), and usually by the leaf size. It seems advisable, therefore, to
recognize them as species, despite their overlapping ranges.

16. Hypericum cuatrecasii Gleason

in RevtaAcad. Colomb. 5: 38 (1942), in Bull. Torrey hot. Club 70: 174 (1943). Type:
Colombia, Boyaca, Cordillera Oriental, between Arcabuco and Tunja, 2950 m,
5.viii.l940 (fl), Cuatrecasas 10439 (NY!, holotype; COL, F!, P!, U!, US!, iso-
types).

Shrub up to 2 m tall, spreading, with branches divergent, lateral, short (flowering)
and long (extension), or pseudo-dichotomous. Stems yellow to reddish-brown,
4-lined and ancipitous when young, eventually terete, without epidermal wrinkles or
emergences, cortex exfoliating in strips or flakes; internodes 2-5 mm long. Leaves
sessile, spreading to deflexed, obscurely tetrastichous, deciduous almost at the base
after turning brown, leaving narrow basal ring; lamina (2-)4-7 x (2-)3-7 mm,
gradually increasing in size along shoot, broadly ovate to subcircular, plane or
slightly saccate, with midrib prominent beneath, margin not or very narrowly
hyaline, not glaucous, coriaceous; apex acute to rounded, base cordate-amplexicaul,
shallowly sheathing, pairs perfoliately united; basal or near-basal veins 1-3, with
midrib branching, tertiary reticulum obscure; laminar glands dense to sparse, large,
not prominent. Inflorescence 1-flowered, terminal and on short lateral shoots, the
whole shoot cylindrical, sometimes with pseudo-dichotomous branches; pedicel
4-7 mm long; upper leaves foliose. Flowers c. 20 mm in diam. , stellate. Sepals 5-7 x
3-4 mm, the outer ovate, subcordiform, the inner ovate-oblong, subacute to
rounded, cucullate, margin narrowly to broadly hyaline; veins 7-11, obscurely
branching, midrib not prominent; glands linear, distally punctiform. Petals (bright?)
yellow, 9-12-5 x 6-5-7-5 mm, c. 1-8 x sepals, obliquely obovate; apiculus obtuse;
glands mostly striiform, distally punctiform. Stamens c. 120-150, longest 5-6 mm
long,c. 0-5 x petals. Ovary 2-5-3 x 1-5-2 mm, broadly ovoid; styles 3(4), 5-6-5 mm
long, 2 x ovary, spreading-incurved; stigmas scarcely to narrowly capitate. Capsule
c. 6 x 3-5 mm, ellipsoid, slightly exceeding sepals. Seeds not seen.

In paramo and degraded forests; 1990-3000 m.

Colombia (Boyaca, Santander), in the Paramo de Arcabuco. Map 7 (p. 39).

COLOMBIA. Boyaca: Sachica, 1990 m, viii.1964, Saravia 4425 (COL). Santan-
der: Cerro Berlin, between Arcabuco and La Palma, 2900 m, 28.iii.1973 (fr),
Cuatrecasas, Garcia-Barriga & Jaramillo 28668 (BM, COL, US).

H. cuatrecasii is most closely related to 14 H. goyanesii, from which it is separable by
the more spreading habit, and by leaf size and shape, the absence of pellucid leaf
margins, smaller flowers, and shorter styles. It appears to be restricted to extreme
eastern Boyaca as far as the border with Santander.

41

17. Hypericum quitense R. Keller

in Bull. Herb. Boissier II, 8: 182 (1908), in Engler & Prantl, Nat. Pflanzenfam. 2nd
ed. 21: 181 (1925); Gleason in Bull. Torrey hot. Club 56: 102 (1929). Type:
Ecuador, Azuay, near Cuenca, Surucucho, viii.1865 (fl), Jameson 128 (W!,
holotype; K!, isotype).

H. rimbachianum Diels in Biblthca hot. 116: 108 (1937). Type: Ecuador, Canar, N.
of Biblian, c. 3400 m, 13.viii.1933, Diels 497 (Bt, holotype).

Icon: Fig. 7.

Shrub or shrublet 0-3- l(-l-5) m tall, erect or spreading, with branches divergent,
lateral, short (flowering) and long (extension), not pseudo-dichotomous. Stems
reddish-brown, 4-lined and markedly ancipitous when young, eventually terete,
without wrinkles or corky emergences, cortex exfoliating in flakes; internodes 4-10

42

NORMAN K. B. ROBSON

Fig. 7 H. quitense: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut
away) and ovary; (g) capsule (a x 1/2; b, f, g x 4; c-e x 6). All Camp 4133.

mm long. Leaves sessile, spreading to deflexed, obscurely tetrastichous, turning
brown, deciduous with the cortex; lamina 4-9-5 x 2-5-5 mm, sometimes increasing
in size along shoot, oblong (lower) to broadly ovate, plane or slightly saccate or
conduplicate, midrib prominent beneath, margin narrowly hyaline, otherwise pale
green but not or only slightly glaucous, subcoriaceous; apex acute (lower) to obtuse,
base broadly cuneate, pairs shortly united; basal or near-basal veins 3(5), with
laterals and sometimes midrib branching, tertiary reticulum obscure or apparently
absent; laminar glands mainly peripheral, dense, rather large, not prominent.
Inflorescence 1 -flowered, terminal and on very short lateral shoots, the whole shoot
narrowly cylindrical, without pseudo-dichotomous branches; pedicel 3-7 mm long;
upper leaves foliose. Flowers 8-12 mm in diam. , stellate. Sepals 5 x 1-2 mm, elliptic
to narrowly oblong, the outer sometimes broader, apiculate-obtuse to acute, sub-
cucullate, margin narrowly hyaline; veins 3-5, not or scarcely branching, all promin-
ent; glands all linear or distally punctiform. Petals bright yellow, 5-6-5 x 3-4 mm,
1-5-2 x sepals, obovate; apiculus subacute to obtuse; glands few, striiform and

THE GENUS HYPERICUM L.

punctiform. Stamens 30-45, longest 3-4 mm long, c. 0-65 x petals. Ovary 1-5-2 x
0-7-1 mm, narrowly ovoid; styles 3, 3-4 mm long, c. 2 x ovary, spreading; stigmas
broadly capitate. Capsule 3-4-5 x 2-5-3-5, ellipsoid, equalling or slightly exceeding
sepals. Seeds c. 0-7 mm long, ecarinate, testa finely scalariform.

In damp meadows of the paramo; 2600-3800 m.

Ecuador (Bolivar, Tungurahua, Canar, Azuay). Map 7 (p. 39).

ECUADOR. Tungurahua: around Lago Pisayambo, 3800 m, 26.xii.1964 (fl),
Knight 268 (BM). Bolivar: Cordillera Occidental, Pucara de Telimbela, 2600-3000
m, 18.xi.1943 (ft),Acosta Solis 6825 (F). Canar: near Pimo, 3060-3120 m, 9.vii.l945
(fl), Camp E-4133 (BM, NY). Azuay: mountains above Sayausid, 3000-3200 m,
18.iii.1974 (fl & fr), Marling & Anderson 12630 (GB).

H. quitense is related to H. cuatrecasii but is much smaller in all parts.

18. Hypericum loxense Bentham

PL Hartweg.: 126 (1843); R. Keller in Bull. Herb. Boissierll,*: 177(1908),inEngler
& Prantl, Nat. Pfanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey hot.
Club 56: 102 (1929). Type: Ecuador, Loja, 'in montibus Loxa', viii.1842? (fl & fr),
Hartweg 720 (K!, holotype; BM!, F!, P!, isotypes).

Shrub orshrublet, 0-2-l(-l-5) m tall, erect or decumbent to prostrate, with branches
ascending to strict, lateral, short (flowering) and long (extension), not pseudo-
dichotomous. Stems reddish-brown, 4-lined and ancipitous when young, eventually
terete, without wrinkles or emergences, cortex exfoliating in irregular flakes or
strips; internodes 1-5-10 mm long. Leaves sessile, erect, imbricate and sometimes
markedly tetrastichous when young, spreading and turning brown, deciduous with
the cortex (or sometimes breaking off earlier); lamina 2-10 x 0-6-3 mm, not
increasing in size along shoot, narrowly oblong or oblanceolate or linear to triangu-
lar-lanceolate or triangular-ovate, incurved-conduplicate or subsaccate, midrib
prominent beneath or not, margin narrowly hyaline, not or slightly glaucous,
subcoriaceous to chartaceous; apex acute to obtuse, base narrowly cuneate or
parallel-sided to broadly cuneate or truncate, pairs shortly united; basal veins
1(3-5), with midrib sometimes branching, tertiary reticulum rarely present; laminar
glands dense to rather sparse, varying in size, prominent or not. Inflorescence either
(i) wholly 1 -flowered, terminal and on very short lateral shoots, the whole shoot
narrowly cylindric, with lateral branches sometimes also bearing flowering branches
but rarely with pseudo-dichotomous branches, or (ii) 1-c. 15-flowered in terminal
and sometimes lateral dichasia or mixed dichasia and pseudo-dichotomies; peduncle
and pedicels 2-9 mm long; upper leaves foliose, bracts gradually smaller. Flowers
6-10 mm in diam., stellate. Sepals 1-5-5-3 x 0-5-1-6 mm, elliptic or lanceolate to
narrowly oblong, the outer sometimes broader, apiculate-obtuse to acute, some-
times cucullate, margin narrowly hyaline; veins 3-5, not or scarcely branching, not
or only the midrib slightly prominent; glands all linear or distally punctiform. Petals
pale or bright yellow to orange, darker or reddish in bud, 3-6(-7-5) x 1-5-3 mm,
1-5-2 x sepals, obovate; apiculus obtuse or obsolete; glands few, striiform and
punctiform. Stamens 20-40, longest 2-4 mm long, c. 0-65 x petals. Ovary 0-7-1-5 x
0-5-1 mm, narrowly ovoid; styles 3(4), 1-3-3 mm long, 1-3-2 x ovary, spreading;
stigmas broadly capitate. Capsule 2-5-4 x 1-5-2-2 mm, ellipsoid or ovoid-
ellipsoid, equalling sepals. Seeds 0-7-0-8 mm long, ecarinate, testa finely scalar-
iform.

On dry slopes and in rocky places in the paramo; 2350-3800 m.

Ecuador (Cotopaxi, Chimborazo, Canar, Azuay, Loja), Peru (Cajamarca, La
Libertad). Map 8.

H. loxense is closely related to H. quitense, which although overlapping it in
distribution is apparently ecologically distinct. In addition, H. loxense has narrower
and/or smaller leaves and usually smaller flowers.
H. loxense comprises two populations of which subsp. aequatoriale is nearer to H.

43

44

NORMAN K. B. ROBSON

Map 8 15. H. myricariifolium
b. subsp. loxenseO.

I; 18. H. loxense: a. subsp. aequatoriale

quitense. The most primitive form of this subspecies is from Mt. Corazon (Cotopaxi),
where it is geographically isolated from the rest (Chimborazo to Loja). It is always
erect and has incurved-cucullate, relatively broad leaves. Subsp. loxense differs in
having relatively narrow, plane leaves and a usually laxer and more spreading habit.
Its most primitive form occurs in Loja, whence there is (i) a southern trend (with
disjunction) to Peru (Cajamarca, La Libertad) in which the leaves become smaller
and narrower, and (ii) a northern trend (Azuay, Canar, Chimborazo) towards
prostrate weak stems and somewhat smaller leaves. The occurrence of intermediates
in Loja prevents the recognition of these taxa as species.

18a. Hypericum loxense subsp. aequatoriale (R. Keller) N. Robson, stat.
nov.

H. loxense var. aequatoriale R. Keller in Bot. Jahrb. 42: 129 (1908), in Bull. Herb.
Boissier II, 8: 181 (1908). Type: Ecuador. Canar, between Azogues and Paramo
de Huairacaja, 2700-3000 m, ix-x.1905? (fl & fr), Lehmann 4795 (Bf, holotype,
F!, photograph; F!, K!, US!, isotypes).

Leaves usually erect and densely imbricate; lamina 2-5-5-2 x 1-2 mm, lanceolate
or rarely oblong to broadly triangular-ovate, plane to subsaccate, chartaceous to
subcoriaceous, apex acute to obtuse, base broadly cuneate to truncate, midrib
rarely prominent beneath. Inflorescence branches 1-3-flowered. Sepals 2-3 x
0-5-1-1 mm, narrowly oblong to lanceolate, acute to subacute.

THE GENUS HYPERICUM L. 45

Ecuador (Cotopaxi, Chimborazo, Canar, Azuay, Loja).

ECUADOR. Cotopaxi: Corazon, 3200 m, 22.vi.1876, Andre K. 436 (K). Chim-
borazo: Paramo de Cachenco, c. 3800 m, 27.vii.1959 (fl), Barclay & Juajibioy 8258
(NY). Canar: near Pimo, 3060-3120 m, 9.vii.l945 (fl), Camp E-4136 (BM, NY);
Tipococha, 3400 m, 9.vii.l939 (fl & fr), Penland 994 (F). Azuay: Benos, SW. of
Cuenca, c. 2700 m, 12. v. 1974 (fl), Marling & Anderson 14561 (GB); San Miguel to
Cuenca, 25 km, 3000 m, 15.ix.1969, B. & C. Maguire 61712 (BM, NY). Loja:
between Loja and Cuenca, c. 20 km N. of Saraguro, 3080 m, 21.X.1980 (fl & fr),
Croat 50854 (BM, MO).

18b. Hypericum loxense subsp. loxense

Leaves spreading or more rarely erect and subimbricate; lamina 4-10 x 0-6-3 mm,
narrowly oblong or narrowly elliptic or rarely oblanceolate to linear, plane to
incurved, chartaceous, apex acute to rarely subacute, base narrowly cuneate to
parallel-sided, midrib often prominent beneath. Inflorescence branches usually
dichasial or mixed, more rarely 1-flowered. Sepals 2-4-5(5-3) x 0-5-1-2(1-6) mm,
narrowly elliptic to oblong, acute.

Ecuador (Chimborazo, Canar, southern Azuay, Loja), Peru (Cajamarca, La Liber-
tad).

ECUADOR. Azuay: Paramo Portete, 3600m, 13.iii.1953 (fl), Prescott793 (NY).
Chimborazo: Urbina, towards Mt. Chimborazo, c. 3700 m, 27.vii.1939 (fl & fr),
Asplund 7913 (S). Canar: near El Tambo (c. 69 km S. of Sibombe), 2850-3000 m,
5.vii.l945 (fr), Camp E-4000 (BM, NY). Loja: Loja, S. of the town, c. 2350 m,
4.X.1955 (fl & fr), Asplund 17919 (S); between Cuenca and Loja, near Una, 1865 (fl
& fr), Jameson 127 (K, W).

PERU. Cajamarca: above Sunchubamba, 3680 m, 8.vi.l957 (fl), Ellenberg 1862
(U). La Libertad: Playapampa, c. 2700 m, 15-24.vi.1923 (fl), Macbride 4525 (NY).

The disjunct populations in Chimborazo and northern Azuay differ in their decum-
bent to prostrate habit, whereas the collection from Cajamarca differs in having
oblanceolate subacute leaves and somewhat larger flowers than normal.

19. Hypericum sabiniforme Trev.

Hyper. Animad. : 15 (1861); Triana & Planchon in Annls Sci. not. (Bot.) IV, 18: 296
(1862) ['sabinaeforme']. Type: Colombia, Boyaca, Prov. deTunja, Soata, 1300m,
iv.1843 (fl), Linden 1328 (BHU?, holotype; BM!, GH!, K!, P!, W!, isotypes).

Hypericum thymifolium sensu Turcz. in Bull. Soc. Nat. Moscou 31 (1): 386 (1858),

non Banks & Solander (1794) nee Kunth (1822).
Hypericum hartwegii sensu Triana & Planchon, torn. cit. : 295. (1862) pro parte, excl.

typum; R. Keller in Engler & Prantl, Nat. Pflanzenfam. 3 (6): 214 (1893) pro

parte, excl. typum, in op. cit., 2nd ed., 21: 181 (1925) pro parte quoad specim.

Colomb.
Hypericum lindenii R. Keller in Bull. Herb. Boissier II, 8: 183 (1908), in Engler &

Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey hot.

Club 56: 102 (1929). Type: Colombia (as for H. sabiniforme), Linden 1328 (W!,

holotype; BM!, GH!, K!, P!, isotypes).
Hypericum trianae N. Robson in Cleef , Veg. Pdram. Colomb. Cord. Or. : 164, 305

(1981), nomen.

Shrub 0-3-0-5 m tall, erect, bushy, with branches erect, strict, lateral or rarely
pseudo-dichotomous. Stems orange-brown, 4-lined when young, eventually terete,
cortex exfoliating in strips; internodes 1-5-4 mm long. Leaves sessile or with
pseudopetiole up to 1 mm long, outcurving to appressed, loosely to densely
imbricate, sometimes markedly tetrastichous, deciduous at or sometimes slightly
above the base; lamina 1-5-7 x 0-4-3 mm, ovate or elliptic to oblanceolate or
narrowly oblong, cucullate, incurved, with margin narrowly hyaline, midrib
impressed beneath, concolorous, glaucous, coriaceous, markedly papillose or sub-
papillose above, less so beneath; apex obtuse to rounded, base cuneate-

46

NORMAN K. B. ROBSON

pseudopetiolate to angustate or parallel, pairs united to form narrow interfoliar
ridge; basal vein 1, unbranched; laminar glands dense, large, not or slightly promin-
ent. Inflorescence 1-flowered, terminal and on lateral branches; pedicel 1-2 mm
long; upper leaves transitional or foliose. Flowers 5-20 mm in diam., stellate. Sepals
2-5 x 0-5-3 mm, ovate-oblong to oblong, obtuse to rounded, plane or cucullate,
margin narrowly hyaline; veins 3-9, unbranched, with midrib prominent beneath or
not; glands mostly linear to mostly punctiform. Petals bright (?) yellow, 4-11 x 2-6
mm, 2-2-5 x sepals, oblong-obovate; apiculus acute; glands distally interupted to
wholly punctiform. Stamens 60-125, longest 2-5-4-5 mm long, 0-4-0-65 x petals.
Ovary 1-5-2 x 0-8-1 mm, ovoid to ellipsoid-subglobose; styles 2, 1-5-5 mm long,
1-2 x ovary, spreading-outcurved; stigmas small. Capsule 3-5 x 2-5 mm,
ovoid-cylindric to ellipsoid, exceeding sepals. Seeds not seen.

In dry paramo; 2000-3870 m.
Colombia (Boyaca). Map 9.

COLOMBIA. Boyaca: Municipio de Susacon, paramo de Guantiva, 3140 m,
8.viii.l958 (fl), Jaramillo & Hernandez 965 (US); NW. of Belen, Quebrada Minas,
Hoya Clla. Larga, 3870 m, 25.ii.1972 (fl), CleeflSOl (COL; U); 1 km E. of Vado
Hondo, Valle de Rio Cusiara, 2915 m, 31.iii.1973 (fl). Cleef, Cuatrecasas & Jaramillo
9236B (COL); Paramo de Giiina, 3200 m, 17.vii.1940 (fl), Cuatrecasas & Garcia
Barriga. 9790 (F); Andes de Bogota, 2500-3000 m, (fl), Triana s.n. (BM).

The type of H. sabiniforme, which superficially resembles a species of the African
ericaceous genera Philippia or Blaeria, represents the extreme form of an eastward

Map 9 19. H. sabiniforme A ; 22. H. hartwegii O; 23. H. maguirei \25.H. gleasonii

THE GENUS HYPERICUM L.

dine. At the western end of the cline is a form with much larger ovate to elliptic
pseudopetiolate leaves, markedly papillose above, and much larger flowers. Varia-
tion along the cline, however, appears to be continuous. The western form is near the
broad-leaved form of//, thuy aides, but differs in having thicker, glaucous leaves with
undulate to papillose epidermis.

Wood 3682 (Cundinamarca, between Neusa and San Cayetano) is intermediate
between 13 H. thuyoides 'genistoides' (see p. 38) and 19 H. sabiniforme.

20. Hypericum laricifolium Juss.

in Annls Mus. Hist. nat. Paris 3: 160, t.16 f.l (1804); Choisy, Prodr. monogr. fam.
Hyperic. : 58 (1821), in DC., Prodr. syst. nat. regni veg. 1: 553 (1824); R. Keller in
Bull. Herb. Boissier 6: 257 (1898), in op. cit. II, 8: 176 (1908), in Engler & Prantl,
Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Kunth in Reprium Spec. nov. Regni veg.
Beih. 43: 483 (1927). Type: Peru, without precise locality or date (fl), /. deJussieu*
in Herb. A. L. de Jussieu 11800 (P- JUSS!, holotype; C!, F!, P!, isotypes).

H. acerosum Kunth in Humboldt, Bonpland & Kunth, Nova Gen. etSp. PI. 5: 187, t.

457 (1822). Type: Ecuador, Azuay, 'in Andium Assuay Quitensium', 3240 m,

vii.1805 (fl), Humboldt & Bonpland (P-HUM!, holotype; P!, isotype).
Brathys laricifolia (Juss.) Spach, Hist. nat. veg. Phan. 5: 366 (1836).
B. acerosa (Kunth) Spach, Hist. nat. veg. Phan. 5: 449. 1836, in Annls Sci. nat. (Bot.)

II, 5: 366 (1836).
Hypericum laricifolium var. acerosum (Kunth) Weddell, Chloris Andina 2: 271

(1857).
H. platypetalum Turcz. in Bull. Soc. Nat. Moscou. 31 (i): 388 (1858), non (Webb)

Steudel (1840). Type: Ecuador, Pichincha, Quito, 1836? (fl), Jameson 60 (LE,

holotype; BM!, K!, W!, isotypes).
H. racemulosum Turcz. in Bull. Soc. Nat. Moscou. 36 (1): 580 (1863). Type: Peru.

Amazonas, Chachapoyas, 1835 (fl), Mathews 1611 (LE, holotype; BM!, KW!, P!,

isotypes).
H. laricoides Gleason in Bull. Torrey hot. Club 56: 104. (1929). Type: Venezuela,

Merida, Paramo de Timotes, Paramos de Chachapos, 3800-4200 m, 24. i. 1928 (fl),

Pittier 12713 (US!, holotype; F!, K!, MO!, isotypes).

Icones: Cuatrecasas in Trab. Mus. Nac. Cienc. Nat. Madrid (Bot.), no. 27: t. 29
(1934); Humboldt, Bonpland & Kunth, torn. cit. : t. 457 (1822); Fig. 8A.

Shrub or small tree (0- 1-)0-3-3(-6) m tall, bushy or lax and spreading, with branches
erect to spreading, lateral, short (flowering) and long, narrowly to broadly conical
(extension). Stems 4-lined when young, eventually terete, blackish-brown, without
epidermal emergences, cortex exfoliating in strips or irregular flakes; internodes
1-5-5 mm long. Leaves sessile, spreading or erect, sometimes imbricate, not
markedly tetrastichous, persistent; lamina (2-)3-6(-8) x 0-3-2-2 mm, narrowly
elliptic or narrowly elliptic-oblong to acerose, conduplicate to incurved, with midrib
impressed beneath, margin narrowly hyaline and plane or subundulate, not
glaucous, chartaceous to coriaceous; apex subacute to obtuse, cucullate, base
narrowly cuneate to angustate, loosely clasping or not, pairs united to form narrow
interfoliar ridge; basal vein 1, not or obscurely branched; laminar glands dense,
scattered or in 1 row, impressed, usually visible beneath only. Inflorescence 1-
flowered, on short lateral branches, rarely branching pseudo-dichotomously; pedicel
1-5-5 mm long; upper leaves foliose. Flowers 15-25(-30) mm in diam., stellate.
Sepals 3-7 x 0-7-2-2 mm, oblong or elliptic to narrowly ovate or lanceolate,
subacute to acute, cucullate, margin hyaline; veins 7-9, not or scarcely branched,
midrib distally incrassate; glands linear, distally punctiform. Petals bright golden
yellow to orange-yellow, 7-14)-22) x 4-10 mm, 2-3 x sepals, very obliquely,
obovate; apiculus obtuse; glands striiform and punctiform. Stamens 70-110, longest
4-7 mm long, c. 0-5 x petals. Ovary 2-3-5 x 0-8-1 -5 mm, ellipsoid; styles 3, 4-9 mm
long, 2-4-5 x ovary, spreading; stigmas scarcely to narrowly capitate. Capsule 4-7 x

* The only specimen in Herb, de Jussieu that could be type material is labelled 'Perou, 1780,
Dombey.'

47

48

NORMAN K. B. ROBSON

Fig. 8 A. H. laricifolium: (a) habit; (b) stem with leaves (Ecuador); (c) stem with leaves (Venezuela); (d)
leaf; (e) sepal; (f) petals; (g) stamens (partly cut away) and ovary; (h) capsule. B. H. martense: (i) habit;
(j) stem with leaves; (k) leaf; (1) sepal; (m) petal; (n) capsule (a, i x Va; b, c, j x 2; d-h, k-n x 4). A.
Luteyn 6751 except (c) Luteyn 7583; B. Seifriz 421.

3-4 mm, ovoid to subglobose, equalling or exceeding sepals. Seeds 0-8-1 mm long,
ecarinate; testa finely scalariform.

In the subparamo and paramo, in open or more usually sheltered habitats, well-
drained or not; 2200-4300 m.

From western Venezuela (Lara, Trujillo, Merida, Zulia) along the Cordilleras

THE GENUS HYPERICUM L.

Central and Oriental of Colombia and Ecuador to central Peru (Huanuco, Ancash).
Map 10.

COLOMBIA. Antioquia: between Medellin and Retire, around Las Palmas, c.
2800 m, 16.x. 1947 (st), Gutierrez, Barkley & Rosano 24(F). Boyaca: Paramo La
Rusia, NNW. of Duitama, 3550 m, ll.ix.1969 (fl & fr), Cuatrecasas & Rodriguez
27730 (BM, COL, US). Caldas: Rio Otun, Nevado de Santa Isabel, Bagas de la
Laguna Negra, 3800-3750 m, 24.xi.1946, Cuatrecasas 23160 (F, P, U). Cauca:
Popayan, near Pitayo, 1841-1843 (fl), Hartweg 925 (BM, K, P, W); Paramo del
Purace, S. of the Volcan, San Francisco, 3450-3400 m, 23.vii.1943 (fl), Cuatrecasas
14571 (COL, F, P). Cesar: Municipio de Valledupar, Corregimiento de Manaure,
2800 m, n.d. (fl), Rodriguez s.n. (COL). Choco: Macigo de Tatama, valle El
Encanto, 3420 m, 13. ii. 1983 (fl), Cleef & Salamanca s.n. (COL). Cundinamarca:
Paramo de Guasca, vicinity of Bogota, 3150 m, 23.viii.1959 (fl), B. & C. Maguire
44029 (BM, NY); Paramo de Palacio, Lagunas de Buitrago, 3550 m, 27. iv. 1973 (fl),
Cleef '9577 B (COL). Magdalena: Sierra de Perija, plain between Cerro Venado and
Cerro Avion, 3270-3350 m, 8.xi.l959 (fl), Cuatrecasas & Romero-Castaneda 25128
(COL) . Narino : Volcan de Cumbal , via Laguna Bolsa , 2740-2980 m , 1 1 . ix . 1944 (fl) ,
Ewan 16139 (BM, S). Norte de Santander: Ocana to Pamplona, Serra PI. , 2400-3300
m, 30.viii.1878 (fl), Kalbreyerlll (K), Putumayo: Lagunda de la Cocha, Quebrada
de Santa Lucia, 2850 m, 8.U941 (fl), Cuatrecasas 11824 (COL, F). Risaralda:
Municipio de Pereira, vicinity of La Laguna de Otun, 4000-4300 m, 3.ii.l980 (fl),
Diaz, Valencia &Jaromillo 1738 (COL). Santander: Paramo de Santurban, Laguna
de Cunta, 3880 m, 21. i. 1927 (fl), Killip & Smith 17962 (COL, F). Tolima: Paramo de
Ruiz, 3200-3500 m, 16-17.xii.1917 (fl), Pennell 2999 (F, K, MO). Valle: Rio

49

Map 10 20. H. laricifolium: typical form , 'laricoides' O; 21. H. martense A.

50 NORMAN K. B. ROBSON

Bugalagrande, Barragan, Paramo de Bavaya, Corrales, 3550-3400 m, 10. iv. 1946
(fl), Cuatrecasas 20569 (BM, F, P, U, US).

VENEZUELA. Lara: between Buenos Aires and Paramo de las Rosas, 2285
-3290 m, 11. ii. 1944 (fl), Steyermark 55501 (F). Merida: Paramo de Mucuchies,
x.1865 (fl), Moritz 1166 (BM, K, P, S); Sierra Nevada de Merida, between La
Canada del Espejo and La Canada de Los Anteojos, Torre de Beffo, 4000-4130 m,
22. i. 1973 (fl), Cuatrecasas, Ruiz-Terdn & Lopez-Figueras 28570 (BM, US); Paramo
de Conejos, c. 8 km NE. of Merida, 3800-4100 m, 27.xii.1967, Wessels Boer 2135
(NY, U, YEN). Tachira: Paramo de La Negra, 3000 m, 13.xi.1939 (fl), Burrus &
Mutter 1019a (YEN). Trujillo: El Paramito, Jago to La Morita, hacia Tuname, over
3000 m, viii.1958 (fl), Aristeguieta & Medina 3488 (NY, YEN). Zulia: Zulia,
1983-1894 (fl), Mocquerys s.n. (NY, P).

ECUADOR. Azuay: 54 km from Cuenca on Suscal road, 3350 m, 13.ix.1969 (fl),
B. & C. Maguire 61696 (BM, NY). Bolivar: Cordillera Occidental, Gualicon Loma,
3240 m, 7. x.l 943, Acosta Soils 6263 (F). Canar: Biblian to Canar, north of Biblian, c.
3400 m, 23.iv.1968 (fl), Marling, Storm & Strom 8655 (GB, NY). Carchi: Paramo El
Angel, between El Angel and Tulcan, 3450-3500 m, 14.V.1973 (fl), Holm-Nielsen et
al. 5298 (AAU, F, S). Chimborazo: Cerro Chiguaza, c. 3200 m, 24.ix.1968 (fl), Lugo
475 (GB, NY). Cotopaxi: Cotopaxi, SW. slope, 3700m, 3.vii.l939 (fi),Asplund!516
(K, P, S). El Oro: NE. Zaruma, Tioloma, Hac. Ambocas, 3100 m, 30.viii.1947,
Espinosa 2034 (NY). Imbabura: Cotacache to Apuela, 21 km (Intec valley), 3300 m,
ll.viii.1976 (fl), 0llgaard & Balslev 8687 (AAU, BM, MO, U). Loja: 10 km south of
Loja, 2800 m, l-3.viii.1959 (fl), Hading 6239 (S). Napo: Cerro Antisana, Los
Corrales, near Papallacta, 3900 m, 21.vii.1960 (fl), Grubb et al. 218 (K, NY).
Pichincha: Mount Guamani, 3600-3800 m, 15.ix.1939 (fl), Asplund 8757 (K, P, S).
Tungurahua: between Ambato and Banos, 10.i.l981 (fl), D'Arcy 13975 (BM,
MO).

PERU. Amazonas: Chachapoyas, Cordillera Calla-Calla between Leimebamba
and Balsas, 2900-3000 m, 21.X.1963 (fl & fr), Ferreyra & Acleto 15302 (MO, NY).
Ancash: Cordillera Blanca, above Vicos, trail to Llacshacocha at Usacocha, 3350 m,
14.iii.1964 (fl), Hutchinson & Wright ^92 (F, K, MO, P, S). Cajamarca: 28 km N. of
Cajamarca towards Pedernal, 12 km W. to Hacienda Granja Porcon (SIP A), 3260
m, 6.vi.l963 (fl), D. & V. Ugent 5463 (GH, K, WIS). Huanuco: Mito, c. 2700 m,
S-22.ii.1922 (fl & fr), Macbride & Featherstone 1610 (F, S). La Libertad: Huama-
cucho, Pullac to Parcay, 69 km, above Rio Chuagon, 3920 m, 7.viii.l964 (fl),
Hutchinson, Wright & Straw 6188 (F, K, MO). Piura: Huancabamba, distr. Sondor,
2300 m, 21.vii.1975 (fl), Sagdstegui, Cabanillas & Dias 8214 (MO). San Martin:
Huallaga, valley of Rio Apisoncho 30 km above Jucusbamba, 3300 m, 5.viii.l965
(fl), Hamilton & Holligan 1090 (K, S).

H. laricifolium is closely related to H. thuyoides, and one collection from Cauca
(Fernandez Perez 7159) is intermediate in some characters. Nearly always, however,
H. laricifolium can be distinguished by the narrow rounded leaves with impressed
midrib and straight margin.

H. laricifolium is very variable, but the variation appears to be continuous and not
amenable to classification. From northern Ecuador and adjacent Colombia, where
the leaves are relatively broad, there are clines north and south towards shorter,
narrower, more densely imbricate leaves and smaller flowers with narrower, more
acute sepals:

(i) North-eastward along the Cordillera Central to Venezuela; the extreme form
from Boyaca north-eastward looks very different, with young shoots densely clothed
with appressed leaves ('H. laricoides'}. Both the typical form and 'H. laricoides'
occur in Boyaca and some parts of Cundinamarca, the latter favouring damper
habitats. In other parts of Cundinamarca (e.g. Paramo de Guasca) and in Caldas,
Risaralda, and Tolima, however, there is a continuous series of intermediate forms
that does not allow the recognition even of subspecies.

(ii) Southward through Ecuador to central Peru (Huanuco, Ancash) the leaves
become gradually narrower and smaller, but not markedly imbricate, except for
some populations from northern Peru.

THE GENUS HYPERICUM L.

21. Hypericum martense N. Robson, sp. nov.

H. laricifolio Juss. affinis, sed habitu multo humiliori caespitosa, ramification!
praecipue pseudo-dichotomo, foliis valde punctatis, sepalis valde nervatis, petalis
plerumque minoribus, staminibus paucioribus, stylis brevioribus, capsula angus-
tiora, differt. Type: Colombia, Magdalena, Sierra Nevada de Santa Marta, near
source of Rio Yebosimeina, 3900 m, 25. v. 1977 (fl), Starker-White & Alverson 586
(NY!, holotype; MO!, isotype!).

Icon: Fig. 8B.

Shrublet 0-08-0-16 m tall, forming clumps or cushions c. 15 mm in diam., with
branches strict, mostly pseudo-dichotomous, or the outer decumbent but not
rooting. Stems orange-brown, 4-lined and compressed when young, soon terete,
cortex exfoliating irregularly; internodes 1-4 mm long. Leaves sessile, erect to
closely imbricate, scarcely outcurving, not tetrastichous, persistent; lamina 4-7 x
0-6-0-8 mm, linear (upper) to acerose, incurved-canaliculate, cucullate, midrib not
impressed, margin narrowly hyaline, concolorous, both surfaces dull, glaucous,
subcoriaceous?; apex subacute to obtuse, base parallel, not clasping, pairs united to
form very narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense,
large, impressed beneath. Inflorescence 1 -flowered, with pseudo-dichotomous bran-
ches from node below; pedicel 3-4 mm long, slender; upper leaves not transitional.
Flowers 8-10 mm in diam. , obconic (?). Sepals 4-5 x 0-8-1-3 mm, narrowly oblong,
acute, incurved above; veins 5, unbranched, all prominent; glands mostly linear but
striiform to punctiform near hyaline margin. Petals 6-8 x 2 mm c. 1-5 x sepals,
oblong-obovate; apiculus acute; glands few, punctiform. Stamens c. 25, longest 4-5
mm long, c. 0-65 x petals. Ovary c. 1 x 0-5 mm, ellipsoid; styles 3, 2-2-5 mm long, c.
2-2-5 x ovary, outcurving to erect; stigmas broadly capitate. Capsule 4-5 x 2-2-5
mm, narrowly ellipsoid, obtuse, exceeding sepals. Seeds 1-2 mm long, ecarinate;
testa finely scalariform-reticulate.

In rocky paramo, muddy places in high-rainfall areas; 3000-3900 m.

Colombia (Magdalena); confined to the Sierra de Santa Marta. Map 10 (p. 49).

COLOMBIA. Magdalena: Sierra de Santa Marta, SE. slopes, Hoya del Rio
Donachui, Meollaca, c. 3400 m, 8.x. 1959, Cuatrecasas & Romero Castaneda 24644
(US); above San Miguel, c. 3000 m, vii.1932, Seifriz 421 (US).

H. martense is a high-altitude derivative of the 'laricoides' form of//, laricifolium. As
a result of its dwarf, caespitose habit, however, the non-pseudo-dichotomous parts
are much reduced, most of the clump being formed by pseudo-dichotomous (in-
florescence) branching. The branches of a pseudo-dichotomy, however, usually bear
lateral flowering branches.

22. Hypericum hartwegii Bentham

PI. Hartwegiana: 127 (1843) [' hartwegi']; Triana & Planchon in Annls Sci. nat. (Bot.)
IV, 18: 295 (1862) pro parte, quoad typum; R. Keller in Engler & Prantl, Nat.
Pflanzenfam. 3 (6): 214 (1893), in Bull. Herb. Boissierll, 8: 177 (1908), in Engler
& Prantl, op. cit. 2nd ed. 21: 181 (1925) pro parte, excl. pi. Colomb. Type:
Ecuador, Loja, mountains near Loja, Chiquiribamba, vii.1841 (fl), Hartweg 721
(K!, holotype; BM!, K!, P!, W!, isotypes).

Shrub 0-3-0-9 m tall, erect, with branches strict to spreading, lateral, short (flower-
ing) and long (extension) or rarely pseudo-dichotomous. Stems dark red-brown,
4-lined and ancipitous when young, eventually terete, without epidermal emerg-
ences, cortex exfoliating in strips; internodes 5-7 mm long. Leaves sessile, erect to
ascending, closely imbricate, tetrastichous, persistent; lamina 5-7 x 2-3 mm,
elliptic to oblong or obovate, plane, apically subconcave, not carinate but with
midrib incrassate distally and subimpressed proximally beneath, margin narrowly
hyaline, not glaucous, subcoriaceous; apex apiculate-obtuse to rounded, slightly
cucullate, base cuneate to parallel, scarcely clasping, pairs united to form very
narrow interfoliar ridge; basal vein 1, apparently unbranched; laminar glands quite

51

52

NORMAN K. B. ROBSON

dense, not impressed or prominent, visible on both sides. Inflorescence 1-flowered,
terminal and on short lateral branches; pedicel 1-2 mm long; upper leaves foliose.
Flowers 15-20 mm in diam., stellate. Sepals 4-6 x 1-8-2-3 mm, ovate to oblong-
elliptic, acute to subacute; veins c. 5, unbranched (?), midrib distally incrassate;
glands linear. Petals bright (?) yellow, 8-11 x 4-5 mm, 2-2-5 x sepals, obovate;
apiculus acute; glands striiform and punctiform. Stamens 45-50, longest 6-7 mm
long, c. 0-5 x petals. Ovary 2-2-5 x c.1-4 mm, ovoid-ellipsoid; styles 3, 5-6 mm
long, c. 2-5 x ovary, outcurving; stigmas small to subcapitate. Capsule and seeds not
seen.

Habitat and phenology not recorded.

Ecuador (Loja). Known from only one collection. Map 9 (p. 46).
ECUADOR. Loja: see type.

H. hartwegii appears to be a very local relict species, being mostly closely related to
H. lycopodioides (from east-central Colombia), which has spreading, incurved-
conduplicate leaves with margin sometimes undulate, oblique-obovate petals, and
more numerous stamens. See also 23 H. maguirei.

23. Hypericum maguirei N. Robson, sp. nov.

H. hartwegii Bentham affinis, sed foliis minoribus, dense imbricatis vaide cucullatis
costa media subtus impressa, floribus minoribus, differt. Type: Ecuador, Azuay,
5km past San Miguel on road to Cuenca, 3350m, 15.x. 1969 (fl), B. & C. Maguire
61707 (BM!, holotype; NY!, isotype).

Shrub 0-3-1 m tall, erect or spreading; branches strict or ascending, lateral, short
(flowering) and long (extension), narrowly spiciform to pyramidal, sometimes
secund. Stems dark red-brown, 4-6 lined when young, eventually terete, without
epidermal emergences, cortex exfoliating in strips or irregular flakes; internodes 1-3
mm, long. Leaves sessile, suberect to erect, densely imbricate, tetrastichous,
persistent; lamina 2-4 x 0-4-1-2 mm, narrowly oblong to narrowly elliptic or
linear, incurved, midrib impressed beneath, pale lucent green with margin narrowly
hyaline and plane, not glaucous, coriaceous; apex obtuse to rounded, markedly
cucullate, base narrowly cuneate to parallel-sided, not or scarcely clasping, pairs
united to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands
dense or in one row beneath, not impressed or prominent, sparser above. Inflores-
cence 1-flowered, terminal and on short lateral branches; pedicel 1-3 mm long;
upper leaves foliose. Flowers (5-)8-15 mm in diam., stellate (to obconic ?). Sepals
2-5 x 0-7-7-1 mm, acute, incurved-cucullate, margin hyaline; veins 3-5, unbran-
ched, midrib prominent beneath; glands linear, distally punctiform. Petals golden to
orange-yellow, 3-8 x 1-5-4 mm, 1-5-2 x sepals, obovate to oblong-obovate;
apiculus subacute to obsolete; glands striiform and punctiform. Stamens 25-40,
longest 2-6 mm long, 0-6-0-75 x petals. Ovary 1-2 x 0-5-1-2 mm, narrowly ovoid
to ellipsoid; styles 3, 2-5 mm long, 1-5-2-5 x ovary, spreading-incurved; stigmas
small. Capsule c. 3 x 2 mm, narrowly ovoid-ellipsoid, equalling sepals. Seeds not
seen.

On dry slopes with mostly low shrubs and ferns, and drier grassland; 2400-3350 m.

Southern Ecuador (Azuay, Loja, El Oro). Map 9 (p. 46).

ECUADOR. Azuay: Cuenca to Ona, Hacienda Pizhia, c. 2800 m, 23. ix. 1955 (fl),
Asplund 17766 (S). Azuay/Loja: Nudo del Cordillera Occidental y Cordillera
Oriental entre Ona y Rancho Ovejero, 2704-2800 m, l-2.viii.1959 (fl), Barclay &
Juajibioy 8452 (NY). El Oro: between Curtincapa and Guagra Uma, 13 km NE. of
Curtincapa, 2850 m, 16.viii.1943 (fl), Steyermark 53912 (F). Loja: about Ona, 2400
m, n.d. (fl), Jameson s.n. (K).

Larger specimens of H. maguirei at first sight resemble a small version of H.
lycopodioides, but differ from it in the sessile, markedly cucullate, imbricate leaves
with midrib impressed beneath, as well as in size of parts. It is most closely related to
H. hartwegii, which has larger, almost plane leaves and larger flowers. Although

THE GENUS HYPERICUM L.

these two populations are very close in distribution, they are morphologically so
distinct that they should be treated as separate species.

24. Hypericum magniflorum Cuatrec.

in Ciencia, Mex. 4: 64, f. 1. (1943); Steyermark in Fieldiana Bot. 28: 988 (1967).
Type: Colombia, Norte de Santander, Cordillera Oriental, Paramo de Tama, near
La Cueva, 27.X.1941 (fl), Cuatrecasas, Schultes & Smith 12634 (COL, holotype;
F!,GH!,P!,U!,isotypes).

Icon: Fig. 9.

Shrub 1-2-5 m tall, erect, with branches strict, mostly pseudo-dichotomous, some-
times lateral. Stems yellow-brown, 4-lined when young, without epidermal emerg-
ences, soon terete, cortex flaking irregularly; internodes 2-3 mm long. Leaves
sessile, spreading abruptly from above base, tetrastichous, becoming brown but

53

Fig. 9 H. magniflorum: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petals; (f) stamens (partly
cut away) and ovary; (g) capsule (a, b x l /2; c-g x 2). (b) Cuatrecasas & Rodriguez 27789; others
Steyermark 57378.

54

NORMAN K. B. ROBSON

not deflexed, persistent until cortex is shed; lamina 10-18 x 4-8 mm, elliptic to
oblanceolate-spathulate, plane, slightly cucullate, carinate, concolorous or paler
beneath, not glaucous, chartaceous; apex acute to obtuse-apiculate, base angustate,
sheathing, pairs united to form very narrow interfoliar ridge; basal veins 5-7, parallel
or almost so, branching distally, with tertiary reticulation sometimes visible beneath;
laminar glands dense above, sparse or absent beneath, slightly prominent. Infloresc-
ence 1-flowered, sometimes with several short flowering branches crowded together,
usually with pseudo-dichotomous branches from 1st or 2nd node below; pedicel up to
5 mm long; upper leaves transitional. Flowers c. 35 mm in diam., stellate. Sepals 9-11
x 3-5-5 mm, broadly oblong to obovate-spathulate, acute to obtuse; veins 11-19,
dichotomising distally, with midrib scarcely prominent beneath; glands punctiform
in upper third. Petals rich yellow, 16-22 x 12-14 mm, c. 2 x sepals, obovate-
oblanceolate; apiculus acute; glands punctiform distally. Stamens c. 80, longest 9-11
mm long, c. 0-5 x petals. Ovary 4-5 x 2-3 mm, broadly ellipsoid to subglobose;
styles 3, 5-9 mm long, 1-5-2 x ovary, erect, free, outcurved below apex; stigmas
slightly enlarged to broadly capitate. Capsule c. 7 x 5 mm, broadly ellipsoid, shorter
than sepals. Seeds not seen.

On rocky slopes of limestone outcrops; 3000-3800 m.

Eastern Colombia (Boyaca, Santander, Norte de Santander) and adjacent Vene-
zuela (Tachira). Map 11.

COLOMBIA. Boyaca: Paramo de Chita, cabeceras de Rio Casanare, 3080 m,
19.vi.1972 (fl), Cleef4743> (BM, U). Norte de Santander (see also type): Bucaraman-
ga, Cumbre of Paramo Mortino, Cucuta road, 3300 m, v.1948 (fl), Sandeman 6077

Map 11 24. H. magniflorum A ; 27. H. stuebelii ; 30. H. decandrum ; 61. H. cymobrathys O.

THE GENUS HYPERICUM L.

(K). Santander: Paramo de Santurban, c. 3000 m, 27.viii.1948 (fl & fr), Barkley &
Araquel8S155(F,COL).

VENEZUELA. Tachira: Paramo de Tama, near Colombo- Venezuelan bound-
ary, 3045-3475 m, 15.vii.1944 (fl). Steyermark 57378 (F, NY); NNE. slopes of Pata
de Judio, 2500-3100 m, 19.X.1978 (fl & fr), Luteyn 5945 (BM, VEN).

Though clearly related to H. terrae-firmae, H. magniflorum is constantly 3-styled.
The abrupt outward bend of the leaves is characteristic, as is the persistent leaf with a
sheathing narrowed base.

55

25. Hypericum gleasonii N. Robson, sp. nov.

H. magnifloro Cuatrec. affinis, sed foliis brevioribus, plusminusve imbricatis, inter-
dum viscidis, supra basin deciduis, stylis brevioribus stigmatibus manifeste capitatis,
differt. Type: Colombia, Santander, Paramo de Almorzadero, c. 32-34 km S. of
Chitaga, c. 3400 m, 12.V.1979 (fl), Luteyn 7653 (COL!, holotype; BM!, NY!,
isotypes).

H. platyphyllum Gleason in Bull. Torrey hot. Club 56: 106. (1929) pro parte, quoad
Killip & Smith 15616 pro parte.

Shrub 0-15-1-5 m tall, erect, with branches strict, mostly pseudo-dichotomous,
sometimes lateral. Stems yellow-brown, 4- lined when young, eventually terete,
without epidermal emergences, cortex flaking irregularly; internodes 2-3 mm long.
Leaves sessile, outcurved-ascending to imbricate-erect, tetrastichous, deciduous
above base before fading; lamina 7-10 x 3-6 mm, oblanceolate to obovate,
incurved, not cucullate or carinate, concolorous, not glaucous, subcoriaceous to
coriaceous; apex acute or apiculate to obtuse, base angustate, sheathing, pairs united
(at least when young) to form narrow interfoliar ridge; basal veins 5-7, flabellate,
branching distally (sometimes obscurely), reticulation obscure beneath; laminar
glands dense, not or slightly prominent, visible beneath; marginal glands usually
secreting viscous clear or white resin. Inflorescence 1 -flowered usually with pseudo-
dichotomous branches from 1st to 3rd node below and often with 1 or 2 lateral
branches immediately below these, forming flower cluster when young; pedicel 3-6
mm long; upper leaves transitional. Flowers 20-40 mm in diam., stellate. Sepals
6-10 x 3-5 mm, elliptic to oblanceolate, acute to apiculate or obtuse; veins 9-15,
sometimes dichotomising distally, with midrib scarcely prominent beneath; glands
punctiform in upper third to two-thirds. Petals rich yellow, (always?) tinged red
beneath, 12-20 x 10-11 mm, c. 2 x sepals, oblanceolate to obovate; apiculus obtuse
to rounded; glands distally punctiform. Stamens 80-100, longest 6-10 mm long, c.
0-5 x petals. Ovary 3-4 x 2-3 mm, broadly ellipsoid to ovoid; styles 3, 4-6 mm long,
1-25-1-5 x ovary, free, suberect to spreading; stigmas broadly capitate. Capsule 5-6
x 5-6 mm, subglobose, shorter than sepals. Seeds not seen.

In open paramo on fine talus or grassy slopes; 3200-3900 m.

Colombia (Norte de Santander, Santander). Apparently confined to the Paramo del
Almorzadero. Map 9 (p. 46).

COLOMBIA. Norte de Santander: Valley of rio Chitaga, 20 km SE. of Chitaga,
3475 m, 24.ix.1944 (fl & fr), St. John 207784 (COL, NY). Santander: Paramo del
Almorzadero, 3600-3800 m, 28. xi. 1941 (fl), Cuatrecasas 13495 (COL, GH, NY, P);
Paramo del Almorzadero, between Chitaga and Cerrito, 3900 m, 31.xii.1959-
l.i.1960 (fl), Barclay &Juajibioy 10389 (NY); Peralonso, Platera, 3400 m, 20.ix.1969
(fl), Cuatrecasas & Rodriguez 27828 (BM, COL, US); Peralonso, 3200 m,
19.vii.1940 (fr), Cuatrecasas & Garcia Barriga 9887 (COL).

H. gleasonii is a relict species intermediate between H. magniflorum and H.
mexicanum. It has larger flowers than H. mexicanum, and these are solitary, but it
shares with that species the deciduous leaves with viscous secretions. In H. gleasonii
the leaves are smaller than those of//, mexicanum (from Cundinamarca and Boyaca)

56

NORMAN K. B. ROBSON

but with a comparable erect habit; the form of//, mexicanum that grows in the same
area is a small-flowered shrublet with decumbent rooting stems.

Killip & Smith 15616 (from the Paramo de Las Vegas), which Gleason (1929)
placed in H. platyphyllum, is a puzzling collection. The BM and COL specimens
belong to H. mexicanum, but the NY specimen has larger flowers and tends towards
H. gleasonil.

26. Hypericum mexicanum L.

Diss. Hyperic. : 5, f. 2 (1776), Amoen. Acad. 8: 322, t.8, f.2 (1785); L. fil. , Suppl. PI. :
345 (1781); Lam., Encyl. meth. (Hot.) 4: 169 (1797); Gleason in Bull. Torrey hot.
Club 56: 103 (1929). Type: Colombia [Cundinamarca], Mutis in Herb. Linn. 943:
31 (LINN!, holotype; BM!, isotype).

Hypericum ? mexicanum sensu Choisy, Prodr. monogr. fam. Hyperic. : 60 (1821).

Hypericum mutisianum Kunth in Humboldt, Bonpland & Kunth, Nov. Gen. et Sp.
PI. 5: 185 (1822), nom. illegit.; R. Keller in Bull. Herb. Boissierll, 8: 178 (1908), in
Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Knuth in Reprium Sp.
nov. Regni veg., Beih. 43: 483 (1927). Type: Colombia, Cundinamarca, near Santa
Fe de Bogota, 2457 m, 1805 (fl), Humboldt & Bonpland s.n. (P-HUM!, holotype;
B-WILLD, isotype);

Brathys mutisiana (Kunth) Spach, Hist. not. veg. Phan. 5: 447 (1836); in Annls Sci.
nat. (Hot.) II, 5: 366 (1836).

Hypericum resinosum Bentham, PI. Hartweg.: 165 (1845); R. Keller in Bull. Herb.
Boissier II, 8: 177 (1908), in Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181
(1925). Type: Colombia. Cundinamarca, 1841-1843 (fl), Hartweg 924 (K!, holo-
type; BM!, P!, W!, isotypes).

H. mexicanum var. intermedium Kuntze, Rev. Gen. pi. 1: 59 (1891). Type: Vene-
zuela, Merida, Silla de Caracas, Kuntze s.n. (NY, holotype).

Shrub or shrublet 0-15-l-5(-2) m tall, erect or decumbent and sometimes rooting at
the base, with branches strict, pseudo-dichotomous and sometimes lateral especially
from short upper internodes. Stems yellow- to orange-brown, 4-lined when young,
eventually terete, cortex flaking irregularly; internodes 2-8 mm long. Leaves sessile,
outcurving-ascending to imbricate-erect, tetrastichous, deciduous above base usu-
ally without withering; lamina 7-17 x 3-10 mm, elliptic or oblanceolate to obovate,
plane to subconduplicate, not or scarcely cucullate, not carinate, concolorous, not
glaucous, coriaceous; apex obtuse or rounded, base cuneate to angustate,
sheathing, pairs united to form narrow interfoliar ridge; basal veins 5-7, flabellate,
branching and anastomosing distally, reticulation clearly to obscurely visible be-
neath; laminar glands dense to sparse, usually visible and sometimes slightly
prominent beneath; marginal glands secreting viscous clear or white resin. Inflores-
cence l-5(-13)-flowered, cymose, corymbose, often with 1-3 pairs of pseudo-
dichotomous branches from 1-3 nodes close below; peduncle and pedicels 1-6 mm
long; upper leaves transitional. Flowers 15-25 mm in diam., stellate. Sepals 5-9 x
2-4 mm, elliptic to oblanceolate-spathulate or obovate, acute to apiculate or obtuse;
veins 5-9, sometimes dichotomising distally, with midrib scarcely prominent be-
neath; glands punctiform distally. Petals golden yellow, (always?) tinged red be-
neath, 9-12 x 4-9 mm, 1-3-2 x sepals, obovate-oblanceolate to obovate-oblong;
apiculus obtuse to obsolete; glands punctiform distally. Stamens c. 50, longest 5-7
mm long, c. 0-5 x petals. Ovary 2-5-3 x 1-8-2 mm, broadly ellipsoid to ovoid; styles
3, 2-5-4 mm long, 1-1-2 x ovary, suberect; stigmas broadly capitate. Capsule 5-7 x
4-6 mm, broadly ellipsoid to subglobose, equalling or exceeding sepals. Seeds c. 0-7
mm long, ecarinate; testa finely scalariform.

In open paramo on fine talus or grassy slopes or in scrub; (1700)2400-4200 m.

Western Venezuela (Merida, Tachira) and north-eastern to central Colombia (Norte
de Santander to Valle de Cauca). Map 12.

COLOMBIA. Antioquia: entre Rio Negro y Sta Helena, 20650 m, vii.1944 (fl),
Garcia Barriga. 11094 (COL). Boyaca: Sierra Nevada del Cocuy, Quebrada de San

THE GENUS HYPERICUM L.

57

Map 12 26. H. mexicanum ; 28. H. prietoiO; 29. H. cassiopiforme .

Paulino proximo Alto Ritacuva, c. 3650, 3. v. 1959 (fl), Barclay & Juajibioy 7525
(COL, MO, NY); Paramo de Giiina, 3200 m, 17.vii.1940 (fl), Cuatrecasas & Garcia
Barriga. 9783 (COL). Cundinamarca: Paramo de Guasca, 2840 m, 15.xii.1938 (fl),
Balls 5700 (BM, COL, K); Sabana de Bogota, Municipio de Tenjo, Vereda de la
Punta, 2800-2670 m, 29.1.1966 (fl & fr), Cuatrecasas & Jaramillo 26755 (BM, COL,
US); Cerro de Monserrate, 3000 m, 10.iii.1966 (fl), Duque- Jaramillo 2877 (COL).
Meta: Macizo de Sumapaz, alredores de la Laguna La Guitarra, 3370-3400 m,
5.vii.l981, Diaz P. 2485 (COL). Norte de Santander: Paramo de Fontibon, 2700 m,
27.vii.1940 (fl), Cuatrecasas & Garcia Barriga. 10088 (NY, US). Santander: Paramo
de Santurban, 3600 m, 27.viii.1948 (fl), Araque & Barkley 18s004 (COL, F). Valle:
Rio Cali, Pichinde, 1700 m, vii.1938 (fl), Duque-Jaramillo 4107 (COL).

VENEZUELA. Merida: Silla de Caracas, Kuntze s.n. (NY). Tachira: Paramo de
Tama, 4000 m, 1937 (fl), Car dona 123 (US, YEN).

H. mexicanum is a misnomer, as the type comes from Cundinamarca and the species
as a whole is confined to Colombia and adjacent Venezuela. In attributing Mutis
collections to Mexico, Linnaeus made the same mistake here as he did in the case of
Vismia baccifera.

H. mexicanum differs from H. gleasonii in having larger, more spreading leaves
and smaller flowers, which are frequently in condensed cymes. The two species
remain distinct where their distributions overlap in the Paramo del Almorzadero.

Gleason (1929) confused the eastern form of H. mexicanum with H. platyphyllum,
which has penninerved (not flabellate-nerved) leaves and smaller stigmas and is
treated here as a subspecies of H. phellos.

58

NORMAN K. B. ROBSON

27. HypericumstuebeliiHieron.

in Bot. Jahrb. 21: 321 (1895); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908), in
Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925) Type: Peru, Amazonas,
Chachapoyas, between Pacasmayo and Moyobamba, near Mojon Cruz, 3300 m,
iv.-vi.1875? (fl), StubelPem21 (Bt; holotype; F!, GH!, NY!, photographs).

Hypericum stuebelii Hieron. ex R. Keller in Bot. Jahrb. 42: 129 (1908). Type: as for
H. stuebelii Hieron.

Shrub 0-3-1 m tall, erect, with branches strict or ascending, pseudo-dichotomous.
Stems reddish-brown, 4-lined when young, soon terete, cortex exfoliating in irregu-
lar flakes; internode 3-4 mm long. Leaves sessile, closely imbricate and tetrastichous
at first, gradually outcurving, deciduous above the base usually before fading; lamina
8-14 x 3-7 mm, broadly elliptic-ovate to elliptic-lanceolate, incurved-cucullate,
midrib prominent distally beneath, margin scarcely distinct, concolorous, yellow-
green, not glaucous, coriaceous; apex obtuse or apiculate to acute, base angustate,
sheathing, pairs united to form rather deep (0-4-1 mm) interfoliar ridge; basal or
near-basal veins 7-9, flabellate, not clearly branched, tertiary reticulum not visible;
laminar glands dense, not prominent, visible above and sometimes beneath. In-
florescence 1 -flowered, usually with pseudo-dichotomous branches from node be-
low; pedicel 4-6 mm long; upper leaves not transitional. Flowers 25-40 mm in diam. ,
stellate. Sepals 10-12 x (2-)3-6 mm, elliptic-ovate to lanceolate, obtuse to acute,
cucullate; veins 7-11, sometimes dichotomising distally, prominent beneath; glands
linear, punctiform in distal third. Petals bright? yellow, 15-20 x 5-10 mm, 1-5-1-7 x
sepals, obovate; apiculus subacute to obtuse; glands linear, interrupted distally.
Stamens 80-150, longest 7-10 mm long, c. 0-5 x petals. Ovary c. 4-5-5 x 2-5 mm,
ellipsoid; styles 3-4, c. 1 mm long, c. 1-7 x ovary, free, suberect to outcurved;
stigmas not or scarcely enlarged. Capsule c. 7-8 x 5 mm, broadly ellipsoid to
subglobose, shorter than sepals. Seeds not seen.

On open hillsides, (always ?) in damp or wet areas; 2400-3300 m.

Northern Peru (Amazonas, San Martin), almost confined to a small region round
Chachapoyas. Map 11 (p. 54).

PERU. Amazonas: Chachapoyas, 1-5 km W. of Molinopampa, Jalca zone,
2400-2450 m, 18.vii.1962 (fl), Wurdack 1353 (F, GH, K, NY, P, S, US); La Jalca
near Chachapoyas, 2700-3300 m, i.1930 (fl), Williams 7585 (F, NY); Chachapoyas,
Pass of Piscohuanuna, 2700 m, viii.1938 (fl), Sandeman s.n. (K); Cordillera of
Piscohuanuna between Chachapoyas and Moyobamba, xii.1868 (fl), Raimondi2096
(NY). San Martin: Bagazan, 1835 (fl), Mathews 1609 (K).

H. stuebelii appears to be a relict species with no ancestral relatives nearer than NE.
Colombia (H. magniflorum, H. pimeleoides). Its leaves vary from broad, obtuse, and
almost plane to narrow, acute, and incurved.

28. Hypericum prietoi N. Robson, sp. nov.

H. stuebelii Hieron. affinis, sed ramis strictioribus, foliis minoribus densissimis
manifeste quadrifariis, 1-nervis, parte distali obtuse carinata parte proximali subtus
costa media impressa, floribus minoribus, differt. Type: Ecuador, Azuay, Eastern
Cordillera, between Ona and the rio Yacuambi, 3000-3360 m, 10-19.ix.1945 (fr),
Prieto P-302 (NY!, holotype).

Shrub c. 0-3 m tall, erect; branches very strict, pseudo-dichotomous and lateral.
Stems reddish-brown, 4-6-lined when young, eventually terete, cortex exfoliating in
irregular flakes; internodes c. 2 mm long. Leaves sessile, very closely imbricate,
markedly tetrastichous, laterally compressed distally, not becoming outcurved,
deciduous above the base before fading; lamina 7-8 x 1-5-2 mm, oblanceolate,
distally obtusely carinate and cucullate, midrib impressed proximally beneath,
margin very narrowly hyaline, concolorous, lustrous yellow-green, not glaucous,
coriaceous; apex acute, base parallel-sided, scarcely sheathing, pairs united to form
shallow to rather deep (to 1-5 mm) interfoliar ridge; basal vein 1, unbranched;

THE GENUS HYPERICUM L.

laminar glands dense, not visible beneath. Inflorescence 1-flowered, with pseudo-
dichotomous branches from node below; pedicel 2-2-5 mm long; upper leaves not
transitional. Flowers stellate? Sepals 7-8 x 2 mm, lanceolate, acuminate, compress-
ed-cucullate; veins 3-5, unbranched, prominent beneath; glands linear. Petals and
stamens not seen. Ovary with styles 3, c. 4 mm long, suberect; stigma narrowly
capitate. Capsule 5 x 2-5 mm, broadly ellipsoid, shorter than sepals. Seeds 1-2 mm
long, ecarinate; testa finely scalariform.

On the crest of the Cordillera; 3000-3360 m.

Ecuador (Azuay). Map 12 (p. 57).

ECUADOR. Azuay: Eastern Cordillera between Ona and the rio Yacuambi,
3000-3360 m, 10-19.ix.1945 (fr), Prieto P-302 (NY).

H. prietoi, which has been collected once only, resembles H. stuebelii but has
smaller, laterally compressed, and markedly tetrastichous leaves and stricter
branches, some of which are lateral.

29. Hypericum cassiopiforme N. Robson, sp. nov.

H. stuebelii Hieron. affinis, sed ramis basalibus lateralibusque strictioribus, foliis
minoribus, oblanceolatis vel obovato-oblanceolatis, floribus minoribus, staminis
paucioribus, stylis brevioribus, differt. Type: Peru, Amazonas, Chachapoyas, 1-5
km W. of Molinopampa, 2400-2450 m, 18.vii.1962 (fl), Wurdack 1355 (US!,
holotype).

Shrub c. 0-3 m tall, erect, with branches strict, basal and lateral. Stems reddish-
brown, 4-lined when young, eventually terete, cortex exfoliating in irregular flakes;
internodes 2-4 mm long. Leaves sessile, imbricate-tetrastichous but not usually
laterally compressed distally, becoming outcurved, deciduous above the base before
fading; lamina 7-8 x 2-5-3 mm, oblanceolate, incurved-cucullate but scarcely
carinate, midrib slightly prominent to scarcely impressed beneath, margin narrowly,
hyaline otherwise concolorous, lustrous yellowish-green, not glaucous, sub-
coriaceous; apex subacute, base cuneate, broadening below, scarcely sheathing,
pairs united to form narrow interfoliar ridge; basal vein 1 with 2 obscure near-basal
branches; laminar glands dense, more visible above than beneath. Inflorescence
1-flowered; pedicel c. 3-5 mm long; upper leaves not transitional. Flowers c. 25 mm in
diam. , stellate. Sepals 8 x 1-8-2-3 mm, elliptic to oblong, acute, incurved-cucullate,
reddish; veins 5, unbranched, prominent beneath; glands linear, distally punctiform.
Petals bright yellow tinged red, c. 14 x 9 mm, c. 1-7 x sepals, obovate-oblanceolate;
apiculus acute; glands linear. Stamens c. 50, longest c. 10-11 mm long, c. 0-5 x
petals. Ovary c. 4 x 1-7 mm, ellipsoid; styles 3, c. 5 mm long, c. 1-2 x ovary, sub-
erect ?; stigmas narrow. Capsule and seeds not seen.

In the Jalca zone; 2400-2450 m.

Peru (Amazonas). Map 12 (p. 57).

PERU. Amazonas: 1-5 km W. of Molinopampa, 2400-2450 m, 18.vii.1962 (fl),
Wurdack 1355 (US).

H. cassiopiforme, like H. prietoi, is based on a single collection and appears to be
related directly to H. stuebelii. It is smaller, with smaller leaves and flowers and more
slender stems, and is intermediate in form between H. stuebelii and H. decandrum. It
differs from the latter by the broader leaves, the larger flowers with broader sepals
and the longer styles with narrow stigmas.

30. Hypericum decandrum Turcz.

in Bull. Soc. Nat. Moscou 31 (1): 389 (1858). Type: Ecuador [Pichincha (?)], Quito,
[1865?] (fl), Jameson 62 (LE, holotype; BM!, K!, W!, isotypes).

Hypericum gnidioides var. polytrichoides R. Keller in Bull. Herb. Boissier II, 8: 183
(1908). Type: Ecuador, Chimborazo, in pascuis montis Titaicun, 3900 m, xi.1859
(fr), Spruce 5599 (G!, holotype, F!, photograph; BM!, C!, GH!, K!, NY!, P!, W!,
isotypes).

59

60

NORMAN K. B. ROBSON

Icon: Fig. 10.

Shrub orshrublet (or perennial herb?) 0-1-0-6 m tall, erect or decumbent and rooting
with branches strict, basal (from decumbent and rooting portion) and pseudo-
dichotomous or sometimes lateral. Stems reddish- to yellowish-brown, 4-6-lined
when young, soon 2-lined, eventually terete, cortex exfoliating in irregular flakes;
internodes 1-5 mm long. Leaves sessile, erect and subimbricate to narrowly spread-
ing or slightly outcurving, deciduous above the base before fading; lamina 4-15 x
0-7-3 mm, narrowly oblanceolate to narrowly oblong, incurved-cucullate,
midrib distally prominent otherwise impressed or plane beneath, concolorous or
with margin rather narrowly hyaline, lustrous pale green, not glaucous, sub-
coriaceous; apex acute, base angustate, broadening below, not or scarcely sheathing,
pairs united to form narrow interfoliar ridge; basal vein 1, unbranched or with 1-2
pairs of obscure lateral branches; laminar glands dense, visible above only, or
obscure. Inflorescence 1 -flowered, with pseudo-dichotomous branches from node
below; pedicel 2-5-7 mm long; upper leaves not transitional. Flowers 6-20 mm in
diam., stellate. Sepals 3-5-9 x 1-3 mm, narrowly elliptic to narrowly oblong, acute,
compressed-cucullate, green (not reddish); veins 5-7, unbranched, sometimes
becoming prominent; glands linear, punctiform in distal third to half. Petals bright
yellow tinged red (? always), 5-9 x 2-3 mm, 0-9-1-2 x sepals, obovate-oblong to
oblanceolate-oblong; apiculus acute; glands absent. Stamens (5)10-45, longest
2-5-4 mm long, c. 0-5 x petals. Ovary 2-5-3 x 1-1-5 mm, ellipsoid; styles 3(4),
1-5-3 mm long, c. 0-5-1 x ovary, divergent; stigma narrowly to broadly capitate.
Capsule 3-5-5 x 2-3 mm, ellipsoid to cylindric-ellipsoid, shorter than sepals. Seeds
0-7-0-8 mm long, ecarinate; testa finely scalariform.

In grassy or shrubby slopes of the paramo and subparamo; (2200 ?) 2700-3930 m.

Ecuador (in scattered localities from Carchi to Loja), Peru (Amazonas). Map 11
(p. 54).

ECUADOR. Azuay: Loja to Cuenca, between Km 113 and 124, 2700-3000 m,
26.ix.1969 (fl & fr), B. & C. Maguire 44320 (BM, NY). Canar: Rivera NE. of
Pindilig, 2900 m, 13.xii.1980 (fl & fr), Holm-Nielsen, Jaramillo & Coello 29222
(AAU). Carchi: Nudo de Boliche, Voladero, 3800 m, 12.vi.1939 (fl & fr), Penland
900 (F, GH, NY). Chimborazo: Mt. Titaicun, 3900m, xi.1858 (fl) , Spruce 5599 (BM,
C, GH, K, NY, P, W). Loja: N. of Saraguro, 3045 m, 4.viii.l943 (fr), Steyermark
53743 (F, NY). Morona-Santiago: Paramo de Hatanga, Km 36 on road Sigsig-
Gualaquiza, E. of pass, 3300 m, 14.xii.1980 (fl), Holm-Nielsen, Jaramillo & Coello
29386 (AAU). Napo: Valle de Chalupas near junction of Rio Chalupas and Rio
Aguas Buenas, 3200 m, 20.U980 (fl), Holm-Nielsen 21014 (AAU). Pichincha?:
Quito, 1865? (ft), Jameson 62 (BM, K, W). Tungurahua: Cordillera de Llanganates,
near junction of rio Golpe and rio Sangarinas (Desaguadero), 26. xi. 1939 (fl & fr),
Asplund9989(S).

PERU. Amazonas: Cerro San Marino, Amaybamba, 3100 m, Hi. 1919 (fr), Bues
567 (US).

H. decandrum is most frequent towards the south of Ecuador and absent or rare in
some central parts of that country. It is related to H. cassiopiforme but differs in the
longer, often narrower, less closely imbricate leaves and the smaller flowers with
shorter, capitate styles. It is quite variable, the most reduced, weaker-stemmed form
from the extreme south of its range being considerably different from the sturdy
northern plants.

31. Hypericum pimeleoides Planchon & Linden ex Triana & Planchon

in Annls Sci. nat. (Bot.) IV, 18: 299 (1862); Gleason in Bull. Torrey hot. Club 56: 103
(1929). Type: Colombia, Norte de Santander, Prov. de Ocana, Cerro Pelado, 2270
m, ii.1846-1852 (fl), Schlim 443 (sphalm. 243) (COL?, holotype; BM!, K!, KW!,
P!,US!,isotypes).

Hypericum caracasanum var. ocanense R. Keller in Bull. Herb. Boissier II, 8: 183
(1908). Type: as for H. pimeleoides (G!, holotype).

THE GENUS HYPERICUM L.

61

Fig. 10 H. decandrum: (a) habit, large form; (b) habit, small form; (c) stem with leaves; (d) leaf; (e)
sepal; (f) petal; (g) stamens (partly cut away) and ovary; (h) capsule (a, b x VT, c-h x 5). (b) Espinosa
923; others B. & C. Maguire 61708.

Shrub 0-4-1-5 m tall, erect, with branches strict, pseudo-dichotomous and occa-
sionally lateral. Stems yellow- to orange-brown, 4-lined and compressed when
young, the subfoliar ridge broad, soon terete, cortex exfoliating in strips; internodes
3-6 mm long. Leaves sessile or with broad petiole to l-5(-2) mm long, densely
imbricate and scarcely outcurving to laxly imbricate and narrowly spreading,
markedly tetrastichous, deciduous above petiole without fading; lamina 10-24 x
3-5-10-5 mm, broadly to narrowly elliptic, plane, not or scarcely cucullate, midrib
not prominent beneath, margin not distinct or rarely indurated, concolorous,
glaucous, coriaceous; apex acute or subapiculate to obtuse, base cuneate to angus-
tate, sheathing or not, pairs united to form narrow interfoliar ridge; basal or
near-basal veins (3)5-9, flabellate, branching and often visibly anastomosing distal-
ly, tertiary reticulation obscure or apparently absent; laminar glands dense, often

62

NORMAN K. B. ROBSON

obscure beneath. Inflorescence 1 -flowered, with pseudo-dichotomous branches from
one or two nodes below; pedicel 4-12 mm long, incrassate upwards; upper leaves not
transitional. Flowers 18-35 mm in diam., stellate to subcyathiform. Sepals (7)10-14
x (2)3-8 mm, elliptic-subcircular to elliptic-oblong or rarely oblong, acute to
subacute; veins (5)7-11, dichotomising distally, with midrib not or slightly promin-
ent; glands linear, distally punctiform. Petals bright yellow, 10-22 x 4-10 mm, c.
1-3-1-5 x sepals, obovate; apiculus acute; glands linear, distally punctiform. Sta-
mens c. 40-90, longest 5-10 mm long, c. 0-5 x petals. Ovary (2-5)3-4 x 2-2-5 mm,
ovoid-ellipsoid to subglobose; styles 3-4, (2)3-7 mm long, 1-1-75 x ovary, outcurv-
ing to diverging, stout; stigmas scarcely to broadly capitate. Capsule 5-8 x 4-6 mm,
cylindric-ellipsoid, equalling sepals. Seeds c. 1-2 mm long, ecarinate; testa finely
scalariform.

In open paramo; 3000-4300 m.

Colombia (Cesar/Norte de Santander to Boyaca). In the Cordillera Oriental from
Cerro de Oroque to Vado Hondo. Map 13 (excluding Cundinamarca records, which
belong to 55 H. ruscoides).

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Agua
Tendida, 3100 m, 10. vi. 1973 (fl), Cleef 10114 (U). Boyaca: Sierra Nevada del Cocuy,
near El Playon, c. 3800 m, 10.ix.1957 (fl), Grubb, Curry & Fernandez-Perez 111
(COL, K, US); Paramo de Chita, cabeceras de Rio Casanare, Km 93 de la carretera a
La Punta, 3080 m, 19.vii. 1972 (fl), Cleef 4144 (BM, U); Pena de Arnical, N. de Vado
Hondo, 3600 m, 6.iv. 1973 (fl), Cleef '9418 (U). Cesar/Norte de Santander: 20 km al S.

Map 13 31. H. pimeleoides

wurdackii O.

I; 32. H. magdalenicum A; 33. H. valleanum A; 34. H. sprucei ; 37. H.

THE GENUS HYPERICUM L.

de Abrego, Las Jurisdicciones (Cerro de Oroque), 3700-3900 m, 19-21.V.1969 (fl &
fr), Garcia- Barriga & Jaramillo 19744 (COL). Norte de Santander: Paramo de Las
Jurisdicciones, SW. of Ocana, 3000-3500 m, 8.vii.l984 (fl), Wood 4526 (K).

H . pimelioides shows a continuous morphological trend southwards from a tall form
in the Cerro de Oroque with stout branches, large, sessile, sheathing leaves, and
large flowers with long styles and scarcely broadened stigmas, to a relatively low
form in the Vado Hondo area with slender branches, small, petiolate, not sheathing
leaves, and small flowers with short styles and broad stigmas. In the Cerro de Oroque
population the large sessile or subsessile leaves have up to 9 free basal veins and a
thin margin, whereas in the Vado Hondo population the smaller petiolate leaves
have only one vein (the midrib); but 1-2 pairs of midrib branches originate near the
base, so that the venation is reduced-flabellate. This is in contrast to the leaf venation
of plants in the Sierra Nevada del Cocuy and elsewhere in Boyaca that belong to H.
cardonae. These have rather similar leaves, but they are relatively narrower and less
crowded, the margin is not or only slightly indurated, and the midrib branches,
where visible, arise at least one third of the distance from the base.

H. pimeleoides appears to be related to H. magniflorum and H. stuebelii because of
its primitively broad-based leaves with flabellate venation and large flowers with 3-5
styles. It is the basic species to which the remainder of sect. Brathys is related,
respectively through 32 H. magdalenicum and 49 H. jaramilloi.

63

32. Hypericum magdalenicum N. Robson, sp. nov.

H. pimeleoidei Planchon & Linden ex Triana & Planchon af finis, sed foliis semper
sessilibus angustioribus basi haud vel vix vaginatis venatione pinnata, sepalis plerum-
que angustioribus, ovario angustiori, stigmatibus haud vel vix capitatis, differt.
Type: Colombia, Magdalena, Sierra Nevada de Santa Marta, quebrada from Laguna
Rio Frio, 3250 m, 28.vii.1972 (fl), Kirkbride & Forero 1748 (COL!, holotype; NY!,
isotype).

Shrub (0-2-)0-7-l-5 m tall, erect, with branches strict, pseudo-dichotomous and
sometimes with 2-3 laterals closely beneath flower. Stems orange-brown, 4-lined and
compressed when young, the subfoliar ridges broad, soon terete, cortex exfoliating
irregularly; internodes 2-4 mm long. Leaves sessile, densely imbricate, erect or
outcurving, markedly tetrastichous, deciduous above base without fading; lamina
8-18 x 2-6-2 mm, broadly to rather narrowly elliptic or oblanceolate, plane to
incurved-conduplicate, not or slightly cucullate, midrib plane or slightly impressed
beneath, margin not distinct, epidermis sometimes undulate, concolorous, some-
times glaucous, thinly to thickly coriaceous; apex acute, base narrowly cuneate,
scarcely sheathing, pairs united to form narrow interfoliar ridge; basal vein 1 , with c.
3-4 pairs of ascending lateral branches, sometimes branching distally, not prominent
or impressed beneath, tertiary reticulation not visible; laminar glands dense,
sometimes visible beneath. Inflorescence 1 -flowered, with pseudo-dichotomous
branches from node below; pedicel 4-5 mm long, not incrassate upwards; upper
leaves not transitional. Flowers 20-30 mm in diam., stellate. Sepals (7-)9-ll x
(2-)2-5-3 mm, elliptic, acute, veins c. 9-11, obscurely reticulating distally, with
midrib not or slightly prominent; glands linear, punctiform near apex. Petals deep
yellow, (10-)14-17 x (5-)9-ll mm, 1-5-2 x sepals, obovate; apiculus acute; glands
linear, distally interrupted to punctiform. Stamens c. 100-120, longest c. 5-8 mm
long, c. 0-5 x petals. Ovary 2-5-3 x 1-5-2 mm, ellipsoid to subglobose; styles 3,
6-7-5 mm long, 2-5-3-5 x ovary, spreading-incurving; stigmas narrow to narrowly
capitate. Capsule (6-)8-9 x 4-6 mm, ellipsoid, shorter than sepals. Seeds c. 1-2 mm
long, ecarinate; testa finely scalariform.

In paramo forest, scrub or damp grassland; 2400-4875 m.

Colombia (Magdalena), Venezuela (Zulia and Merida). In the Sierra Nevada de
Santa Marta and the Sierra de Perija, with one probably conspecific record from the
Paramo del Molino. Map 13.
COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, alredores de cabeceras

64

NORMAN K. B. ROBSON

de Rio Sevilla, 3320-3410 m, 22.1.1959 (fl & fr), Barclay & Juajibioy 6620 (COL,
MO, NY); Sierra Nevada de Santa Marta, valley of Rio Yebosimeina, 3500-4000 m,
22.V.1977 (fl & fr), Starker White & Alverson 547 (COL, MO, NY); Sierra de Perija,
E. of Manaure, quebrada de Floridablanca, 2700-2800 m, 9-12.xi.1959, Cuatre-
casas & Romero Castaheda 25171 (COL).

VENEZUELA. Merida: Paramo del Molino, 2600 m, 19.U922 (fl), John 940
(YEN). Zulia: Perija, Cerro Fetari, iv.1952 (st), Urbano 2 (YEN).

H. magdalenicum differs from the sessile-leaved form of H. pimeleoides by (i) the
leaves with pinnate venation but with a scarcely sheathing base and (ii) the narrower
petals and sepals and usually shorter styles.

H. magdalenicum, which has a pivotal position in relation to over half the
remaining species of sect. Brathys (Spp. 38-53), is rather variable. The nearest form
to H. pimeleoides (from Sierra de Santa Marta, Rio Sevilla) has relatively broad,
plane leaves with conspicuous gland dots beneath. Most other forms from Sierra de
Santa Marta, as well as those from Sierra de Perija, have narrower, incurved-
conduplicate leaves in which the gland dots are inconspicuous or superficially
invisible (except in some Sierra de Perija populations). Of these, the population
from the Rio Donachui area has stiffly erect, relatively thick stems and markedly
imbricate leaves (tending towards 43 H. parallelum and 40 H. bolivaricum), whereas
the other populations have more divergent, relatively thin stems and less markedly
imbricate, though distinctly tetrastichous leaves (tending towards 50 H. lancifolium
and its relatives) .

33. Hypericum valleanum N. Robson, sp. nov.

H. sprucei N. Robson af finis, sed foliis latioribus, nee profunde incurvatis nee
cucullatis, manifeste glanduloso-punctatis, pedicello 3-4 mm longo, stylis crassius-
culis, differt. Type: Colombia, Valle de Cauca, Los Farallones, filo de la cordillera,
matorrales de paramo en el cerro La Torre, c. 3750 m, 10.x. 1944 (fl), Cuatrecasas
177861 (F!, holotype & isotype).

Shrub c. 1 m tall, erect, with branches strict, pseudo-dichotomous and lateral. Stems
orange-brown, compressed? when young, soon subterete to terete, cortex exfoliat-
ing irregularly; internodes 1-2 mm long. Leaves sessile, subimbricate, narrowly
spreading and twisting, subtetrastichous, deciduous above base without fading;
lamina 11-17 x 2-3 mm, narrowly elliptic, shallowly incurved, not cucullate, midrib
impressed, margin very narrowly hyaline, concolorous, not glaucous, coriaceous;
apex sharply acute, base angustate, not sheathing, pairs united to form very narrow
interfoliar ridge; basal vein 1, unbranched; laminar glands dense, large, visible
beneath. Inflorescence 1 -flowered, with pseudo-dichotomous branches from node
below; pedicel stout, 3-4 mm long; upper leaves not transitional. Flowers c. 20 mm in
diam.?, stellate. Sepals 7-10 x 2-2-8 mm, narrowly elliptic to lanceolate, acute to
acuminate, incurved above, margin narrowly hyaline, veins 5, the outer branched,
midrib not or scarcely prominent; glands linear, distally punctiform. Petals bright (?)
yellow, 10-12 x 5-6 mm, c. 1-2-1-4 x sepals, obovate; apiculus sharply acute;
glands striiform to punctiform. Stamens c. 60 ?, longest 5-6 mm long, c. 0-5 x petals.
Ovary c. 2 x 1 -5 mm, ellipsoid; styles 3, c. 3 mm long, c. 1-5 x ovary, stout, suberect;
stigmas narrow. Capsule and seeds not seen.

In paramo; 3750 m.

Colombia (Valle de Cauca). Known only from the type collection. Map 13 (p. 62).

COLOMBIA. Valle de Cauca: Cordillera Occidental, Los Farallones, cerro La
Torre, c. 3750 m, 10.x. 1944 (fl), Cuatrecasas 17861 (F).

H. valleanum is intermediate in form and distribution between the more primitive
form of H. magdalenicum (with broad punctate leaves) and H. sprucei, differing
from the former by its shorter styles and narrower leaves, and from the latter by its
broader, thicker, and densely punctate not cucullate leaves and its stout styles and
generally stronger habit. It appears to be a relict population.

THE GENUS HYPERICUM L.

34. Hypericum sprucei N. Robson, sp. nov.

H. aciculari Kunth af finis, sed caulibus crassioribus, foliis latioribus et saepissime
longioribus, coriaceis, sepalis petalisque longioribus, staminibus numerosioribus,
differt. Type: Ecuador, Tungurahua, [Volcan] Tungurahua, 2400-3000 m, xi.1857
(fl), Spruce 5110 (BM!, holotype; c!, GH!, K!, P!, S!, isotypes).

Hypericum struthiolifolium sensu R. Keller in Bull. Herb. Boissier II, 8: 180 (1908)
[ l struthiolaefolium']\ Gleason in Bull. Torrey hot. Club 56: 102, 105 (1929), pro
parteexcl. typum.

Icon: Fig. 11.

Shrub 0-2-2 m tall, erect to ascending, with branches strict, lateral or more rarely
pseudo-dichotomous. Stems orange-brown, 4-lined and ancipitous when young,
eventually terete, cortex exfoliating irregularly; internodes 1-5-7 mm long. Leaves
sessile, densely to loosely imbricate, erect, spreading and twisting, scarcely tetra-
stichous, deciduous above base without fading; lamina 6-12 x 0-8-1-7 mm, very
narrowly oblanceolate to linear, incurved-cucullate, midrib impressed beneath,
margin very narrowly hyaline, concolorous, not glaucous, coriaceous; apex acute,
base parallel, not sheathing, pairs united to form narrow interfoliar ridge; basal vein
1, unbranched; laminar glands rather dense, few or sometimes none visible beneath.
Inflorescence 1 -flowered, rarely with pseudo-dichotomous branches from node
below; pedicel 1-5-3 mm long, not incrassate upwards; upper leaves not transitional.
Flowers 20-30 mm in diam. , stellate. Sepals 7-11 x 1-5-2-5 mm, lanceolate, acute to
acuminate, incurved above, margin hyaline, veins 5, midrib not prominent; glands
linear, distally punctiform. Petals bright to deep yellow, 10-15 x 5-7 mm, c. 1-4 x
sepals, oblong-oblanceolate; apiculus sharply acute to apiculate; glands striiform to
punctiform, sparse. Stamens 60-75, longest 5-9 mm long, 0-5-0-6 x petals. Ovary
2-3-5 x 1-5-2 mm, ellipsoid-subglobose; styles 3(4), (2-5)3-4-5 mm long, (1-3)1-5
-2 x ovary, outcurving, slender; stigmas narrowly or not capitate. Capsule c. 5 x 3
mm, broadly ellipsoid to subglobose, shorter than sepals. Seeds c. 0-8 mm long
(immature?), ecarinate; testa finely scalariform.

Open, dry or damp paramo; 2725-4000 m.

Ecuador (Carchi to Loja, apparently absent from Cotopaxi and Bolivar), Peru
(Piura). Map 13 (p. 62).

ECUADOR. Azuay: c. 10 km NE. of Sevilla de Oro, 3000-3200 m, 13. ix. 1976
(fl), 0llgaard & Balslev 9449 (AAU, BM, MO); 'Oriente' border, Eastern Cordil-
lera between Ona and rio Yacuambi, 3000-3360 m, 10-19.ix.1945 (fl & fr), Prieto
P-297 (BM, NY). Canar: between Biblian and Canar, c. 3350 m, 21.ix.1955 (fl),
Asplund 17665 (S). Carchi: Base of Volcan Chiles, km34-36onTulcan-Maldonado
road, 3900-4050 m, 19.V.1973 (fl), Holm-Nielsen etal. 5888 (AAU, F, GB, MO, S,
U). Chimborazo: Atilio, c. 3700 m, 29.L1968 (fl), Marling, Storm & Strom 6705
(GB). Imbabura: Ibarra to Mariano Acosta, at pass (Hacienda Yura Cruz),' 3600
-3800 m, 10.viii.1976 (fl & fr), 0llgaard & Balslev 8664 (AAU, BM, MO, U). Loja:
Loja - Zamora road, 1 km east of pass (Km 12), 2725-2750 m, 17.iv.1973 (fl),
Holm-Nielsen et al. K. 3698 (S). Morona-Santiago: Gualaceo-General Plaza
(Limon) road, 3000-3100 m, 20.ic.1967 (fl), Sparre 18778 (S). Napo: Cordillera de
Los Llanganati, Chihuila Sacho o Ainchilibi, c. 3750 m, 25-29. viii. 1959, Barclay &
Juajibioy 8996 (MO, NY); Sucumbios 'Playon de San Francisco', Camino al Cerro el
Mirador, 3600-3800 m, 29.xii.1980 (fl), Jaramillo & Coelle 3903 (AAU, QCA).
Pichincha: 15-4 km west of Papallacta, 3980 m, 26.iii.1972, MacBryde & Dwyer 1185
(MO). Tungurahua: Llanganati Mts, Lake Aucacocha, 3700 m, vii.1969, Edwards
58(K); Paramo of Minza Chica, 3800 m, 4.iv.l939 (fl), PenlandlQl (F, GH). Zamora
- Chinchipe: Loja to Zamora, 2800 m, 18.ix.1961, Dodson & Thien 685 (BM).

PERU. Piura: above Huancabamba, 3000 m, iv.1912 (fl), Weberbauer 6133 (F,
GH, S).

H. sprucei provides a morphological and geographical link between H. valleanum
(Colombia) and three species in southern Ecuador and Peru (Spp. 35-37). From H.

65

A

66

NORMAN K. B. ROBSON

Fig. 11 A. H. sprucei: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut
away) and ovary; (g) capsule. B. H. valleanum: (h) habit. C. H. aciculare: (i) habit (a, h, i x V^; b x 2;
c-g x 3). A. 0llgaard & Balslev 8454; B. Cuatrecasas 17861; C. Hartweg 719.

valleanum it differs in the slender styles and the shorter, narrower leaves. For
differences between it and the southern species, see their accounts below.

H. sprucei varies clinally from north to south, the plants from Carchi being most
similar to H. valleanum. Towards the southern end of its range (Azuay to Piura), the
leaves become slenderer and more flexuous, and the habit low and multi-stemmed.
In Chimborazo the leaves and flowers of some plants are relatively small, thus
indicating a trend towards 35 H. aciculare.

35. Hypericum aciculare Kunth

in Humboldt, Bonpland & Kunth, Nova Gen. et Spec. PI. 5: 190 (1822); Choisy in
DC., Prodr. syst. nat. regni veg. 1: 553 (1824); Jameson, PI. Aequator. 1: 108
(1865); R. Keller in Bull. Herb. Boissier II, 8: 176 (1908) ? (see below), in Engler

THE GENUS HYPERICUM L.

& Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925). Type: Ecuador, Loja, locis
temporis prope Loxam (Regni Quitensi), 1908 m, viii.1805, Humboldt & Bon-
plands.n. (PI, holotype; F!, US!, photographs)

H. struthiolifolium Juss. in Annls Mus. Hist. not. Paris 3: 160 (1804) ['struthiolae-

folium'] pro parte, quoad var. innom. (p. 161, t. 16 f. 2 b).
H. struthiolifolium [var.] |3 sensu Choisy, Prodr. monogr. fam. Hyperic.: 59 (1821)

['struthiolaefolium'] pro parte, quoad spec. Peruv.
H. struthiolifolium [var.] (3 minutum Choisy in DC. , Prodr. syst. nat. regn. veg. 1: 533

(1824) ['struthiolaefolium']. Type: Peru, Dombey s.n. (G, holotype; P!, isotype).
Brathys acicularis (Kunth) Spach, Hist. nat. veg. Phan. 5: 366 (1836), in Annls Sci.

nat. II, 5: 366 (1836).
H. laricifolium sensu Bentham, PI. Hartweg. : 126 (1843-1844).

Shrub 0-3-2 m tall, erect and bushy to decumbent and slender, with branches strict to
ascending, lateral and frequently pseudo-dichotomous. Stems orange-brown, 4-lined
and ancipitous when young, eventually terete, cortex exfoliating irregularly;
internodes 1-7 mm long. Leaves sessile, subimbricate or not, suberect, outcurving
and usually twisting, not or scarcely tetrastichous, deciduous above base without
fading; lamina 3-5-7 x 0-4-0-6 mm, very narrowly elliptic to linear-acicular,
incurved, slightly cucullate, midrib impressed beneath, margin very narrowly
hyaline, concolorous, not glaucous, subcoriaceous to chartaceous; apex acute, base
parallel-sided, not sheathing, scarcely broadening but pairs forming narrow inter-
foliar ridge; basal vein 1, unbranched; laminar glands dense to very sparse, visible
beneath. Inflorescence 1 -flowered, terminal, with pseudo-dichotomous branches
from node below and terminating short unbranched lateral shoots; pedicel 1-5-2-5
mm long, not incrassate upwards; upper leaves not transitional. Flowers 10-17 mm
in diam., stellate. Sepals 4-7-5 x 0-7-1-5 mm, narrowly lanceolate, acute, incurved
above, margin hyaline, veins (3)5, midrib not prominent but whole sepal often
becoming ribbed; glands linear, distally punctiform. Petals yellow or apricot to
orange, sometimes tinged red, 6-10 x 3-7 mm, c. 1-5 x sepals, oblong-obovate,
apiculus sharply acute; glands striiform to punctiform. Stamens 30-55, longest 3-5
mm long, c. 0-5 x petals. Ovary 1-5-2-5 x 1-1-5 mm, ellipsoid-subglobose; styles 3,
2-5-3(4) mm long, 1-2-2 x ovary, outcurving-ascending; stigmas narrow. Capsule
3-5-4 x 2-5-3-5 mm, broadly ellipsoid, shorter than sepals. Seeds 1-2 mm long,
ecarinate; testa finely scalariform.

Dry, open scrub on slopes of paramo, dense moist forested slopes, and wet
sphagnum; 1800-3700 m.

Ecuador (Azuay, Loja), Peru (Piura, Amazonas). Map 14 (p. 68).

ECUADOR. Azuay: Km 85 on Pan-American Highway north of Loja, 2850-2950
m, 3.V.1973 (fl), Holm-Nielsen et al. 4808 (AAU, F, GB, S, U); Rio Collay south of
El Pan, Huagrarancha slopes, 2650-3290 m, 6.vii.l943 (fl), Steyermark 53372 (F);
*30-50 km S. of Cuenca, Paramo de Tinajillas; 3300-3450, 17.iii.1945, Camp
E-2229 (BM, NY);* Rancho Ovejero, c. 3000 m, 1-2. viii. 1969 (fl). Barclay &
Juajibioy 8484 (NY). Loja: between La Toma and Loja, 1800-2600 m, 4.ix.l923 (fl),
Hitchcock 21431 (GH, NY, US);* 7 km from Loja on road to Zamora, 2350 m,
8.ix.l959 (fl), B. & C. Maguire 61667 (BM, NY);* W. slopes of Cerro Villonaco,
2200-2250 m, 10.iv.1974 (fl & fr), Hading & Anderson 13351 (GB, NY).

PERU. Piura: Huancabamba to Cuello del Indio, Mitopampa, 2650 m, 22.vii. 1975
(fl), Sagdstegui, Cabanillas & Dios 8253 (MO). Amazonas:* Chachapoyas, 3-6 Km
W. of Molinopampa, 2200-2450 m, 19.vii.1962 (fl), Wurdack 1403 (F, K, NY, S,
US). *Pajonales, Pozuro, 3000 m, vii.1863 (fl), Pearce 252 (BM, K).

H. aciculare differs essentially from H. sprucei in habit and size of parts. It most
resembles some populations of the latter from Chimborazo, from which it is
separated by a disjunction of merely c. 100 km. Other, less similar populations of H .
sprucei, however, are found in the same area as H. aciculare, which occurs in two
ecologically somewhat distinct forms with widely overlapping areas of distribution. If

67

Wet habitat form, see p. 68.

68

NORMAN K. B. ROBSON

Map 14 35. H. aciculare A ; 36. H. recurvum .

these forms prove to be genetically distinct, it may be possible to recognize them as
subspecies, as they are almost always distinguishable from one another. Plants on
open or forested paramo slopes are bushy with branching mostly pseudo-
dichotomous, whereas those in marshes and seepages are slender with branching
mostly lateral (records marked*).

Keller (1908 supra) keys out H. aciculare with H. nitidum Lam. as having connate
styles. If his material did indeed belong to this species, then he must have interpreted
it wrongly.

36. Hypericum recurvum N. Robson, sp. nov.

H. sprucei N. Robson af finis, sed foliis aureo-marginatis apice longe acuminatis,
recurvascentibus haud torquescentibus, petalis angustioribus, differt; a H. aciculari
Kunth isdem characteribus differt. Type: Peru, Amazonas, Chachapoyas, middle
eastern Calla-Calla slopes, near Kms 416-419 of Leimebamba-Balsas road, 2900
-3100 m, 9.vii.l962 (fl & fr), Wurdack 1303 (K!, holotype; F!, GH!, NY!, P!, S!, US,
isotypes).

Shrub 0-15-0-7 m tall, many-stemmed, erect or decumbent and rooting, with
branches strict, pseudo-dichotomous and sometimes lateral. Stems orange- to red-
dish-brown, 6-lined when young, eventually terete, cortex exfoliating in irregular
flakes; internodes 1-3 mm long. Leaves sessile, imbricate-suberect but not markedly
tetrastichous at first, eventually becoming markedly outcurved but not twisted,
deciduous above the base after turning brown; lamina 5-8 x 0-8-1-2 mm, subulate,
distally conduplicate and cucullate, otherwise incurved with midrib slightly
prominent proximally to slightly impressed beneath, margin narrowly glaucous,
chartaceous; apex acicular, base broadened, markedly sheathing, pairs united to
form narrow interfoliar ridge ; basal vein 1 , unbranched ; laminar glands rather dense ,

THE GENUS HYPERICUM L.

visible beneath when young. Inflorescence 1-flowered, with pseudo-dichotomous
branches from node below; pedicel 3-5-5 mm long, not incrassate upwards; upper
leaves not transitional. Flowers 15-20 mm in diam. , stellate. Sepals 7-9 x 1-1-5 mm,
narrowly-lanceolate-acerose, long-acuminate, cucullate, with golden-hyaline mar-
gin; veins 3-5, unbranched, not prominent; glands linear, interrupted in distal 1/4.
Petals golden yellow, 8-12 x 2-5-4 mm, 1-2-1-4 x sepals, narrowly obovate-
oblong; apiculus acicular; glands linear, distally interrupted. Stamens c. 60, longest
6-8 mm long, c. 0-7 x petals. Ovary 2-5-3 x 1-1-2 mm, narrowly ellipsoid; styles 3,
4-5 mm long, c. 1-7 x ovary, outcurved-ascending; stigmas narrow to scarcely
capitate. Capsule 3-5 x 2-3 mm, ellipsoid, shorter than sepals. Seeds 0-7-0-8 mm
long, ecarinate; testa finely scalariform.

On open hillsides in exposed and (in Pasco) boggy areas; 2700-3100 (7-3300) m.

Peru (Amazonas (Chachapoyas) , Junin, Pasco), apparently almost restricted to the
Cerro Calla-Calla. Map 14.

PERU. Amazonas: Chachapoyas, Cordillera Calla-Calla between Leimebamba
and Balsas, 2900-3000 m, 21.viii.1963 (fl), Ferreyra & Acleto 15284 (NY); between
Leimebamba and Balsas, 2700 m, l.vi.1963 (fl & fr), Lopez, Sagdstegui & Callantes
4429 (NY); Cerro Calla-Calla, E. side, 19 km above Leimebamba on Balsas road,
3100 m, 4.vU964 (fl), Hutchinson & Wright 5505 (F, NY, UC, US); Jalca de
Calla-Calla, 3100 m, 7.V.1970 (fl & fr), Sagdstegui 7451 (F, NY, YEN); Chachapoyas
to Moyobamba, La Jalca, 2700-3300 m, 20.U930 (fl), Williams 7583 (F, NY);
Chachapoyas, 1826 (fl & fr), Mathews 2130 (BM, K, P). Junin: Huancayo, Quebrada
Occopilla, ii.1948 (fl), Soukoup 3636 (MO). Pasco: Oxapampa, Cordillera Yana-
chaga, Cerro Pajonal 12 km SE. of Oxapampa, 2700-2800 m, 7.x. 1982 (fr), Foster
8992 (BM, MO).

H. recurvum is clearly related to the southern, low, many-stemmed form of H.
sprucei, differing from it mainly in the often glandular, acuminate leaves, which
becomes recurved, not twisted, and are suffused with a golden tint (at least when
dried). The longer styles and narrower, acuminate sepals distinguish H. recurvum
from H. andinum, which has an overlapping area of distribution.

37. Hypericum wurdackii N. Robson, sp. nov.

H. recurvo N. Robson affinis, sed foliis longioribus, carinatis, densissime imbricatis,
valde tetrastichis alas formantibus, floribus maioribus, differt; a H. acicular e Kunth
isdem characteribus differt. Type: Peru, Amazonas, Chachapoyas, summit of Cerro
Malcabal (Cerro Tumbe), 3-6 km SW. of Molinopampa, 2850-2900 m, 20.vii.1962
(fl & fr), Wurdack 1426 (K!, holotype; F!, GH!, MICH!, NY!, P!, S!, US, isotypes).

Shrub 0-15-0-3 m tall, several-stemmed, erect, with branches strict, pseudo-
dichotomous and sometimes lateral. Stems orange-brown, completely enclosed by
leaves when young, eventually terete, cortex exfoliating irregularly; internodes 1
mm long. Leaves sessile, imbricate, markedly tetrastichous with the vertical rows
winglike, not or scarcely outcurving, not twisted, deciduous above the base after
turning brown; lamina 9-14 x 1-2 mm, linear-acerose, incurved-conduplicate,
cucullate, carinate wholly or only at apex and base, margin narrowly golden-hyaline
especially towards base, otherwise concolorous, yellow-green, not glaucous, char-
taceous; apex sharply acute, base broadened, markedly sheathing, pairs united to
form deep interfoliar ridge ; basal vein 1 , unbranched ; laminar glands dense , invisible
below in mature leaf. Inflorescence 1-flowered, with pseudo-dichotomous branches
from node below; pedicel 2-3 mm long, not incrassate upwards; upper leaves not
transitional. Flowers 20-25 mm in diam., stellate. Sepals 10-12 x 3-4 mm, lan-
ceolate-ace rose, sharply acute, not cucullate, with golden-hyaline margin; veins 7,
unbranched, not or scarcely prominent; glands linear, punctiform in upper half.
Petals golden (?) yellow, 15-18 x 7-8 mm, c. 1-5 x sepals, narrowly obovate;
apiculus acute; glands linear, distally interrupted. Stamens c. 100, longest 8-9 mm
long, c. 0-5 x petals. Ovary c. 2-5 x 1-5 mm, ellipsoid; styles 3, 4-5 mm long, 1-7-2
x ovary, outcurved-ascending; stigmas narrow. Capsule c. 6 x 3-5 mm, ellipsoid,
shorter than sepals. Seeds not seen.

69

70

NORMAN K. B. ROBSON

No habitat data known; 2850-2900 m.

Peru (Amazonas-Chachapoyas). Known only from type collection. Map 13 (p. 62).
PERU . Amazonas : Chachapoyas , summit of Cerro Malcabal (Cerro Tumbe) , 3-6
km SW. of Molinopampa, 2850-2900 m, 20.vii.1962 (fl & fr), Wurdack 1426 (F, GH,
K, MICH, NY, P, S, US).

This bizarre species, which resembles a large Cassiope tetragona in habit, is clearly
related to the low, many-stemmed form of H. sprucei from Piura. Although it is
larger in all parts than H. recurvum, it can be interpreted as carrying the trends in leaf
form and arrangement of that species a stage further. The leaves are even more
closely imbricate, so that the tetrastichy characteristic of the young shoots of H.
recurvum is very marked in H. wurdackii.

38. Hypericum costaricense N. Robson, sp. nov.

H. sprucei N. Robson af finis, sed foliis subcoriaceis saepe glaucis, apice haud vel vix
cucullatis, floribus minoribus, sepalis apice acutissimis, stylis brevioribus, stigmati-
bus anguste vel late capitatis, capsula ovoidei-ellipsoideis, differt. Type: Costa Rica,
Alajuela, upper slopes of Volcan Poas, c. 2700m, I.ix.l968(fl&fr), Wilbur &Stone
10634 (BM!, holotype; DUKE!, NY!, US!, isotypes).

Shrub or shrub let c. 1 - 1 m tall , erect or ascending , forming dense rounded clumps ,
with branches strict, pseudo-dichotomous and lateral. Stems orange-brown, 4-
angled and (?) ancipitous when young, eventually terete, cortex exfoliating in strips;
internodes 2-4 mm long. Leaves sessile, imbricate to suberect, spreading to
outcurving and twisting, markedly tetrastichous when young, deciduous above
base without fading; lamina 5-15 x 0-6-1 mm, linear-elliptic to linear, incurved-
canaliculate, not or scarcely cucullate, midrib impressed beneath, margin narrowly
hyaline, concolorous, lucent or often glaucous, subcoriaceous; apex acute to sub-
acute, base parallel-sided, scarcely broadening, pairs forming narrow interfoliar
ridge; basal vein 1, unbranched; laminar glands dense, visible beneath. Inflores-
cence 1 -flowered, terminal, with pseudo-dichotomous branches from node below;
pedicel 3-4 mm long, rather slender, not incrassate upwards; upper leaves not
transitional. Flowers 15-20 mm in diam., stellate. Sepals 5-8 x 0-8-1-6 mm,
narrowly oblong to narrowly triangular-lanceolate, sharply acute, margin hyaline,
veins 5(7), unbranched, midrib or all sometimes prominent; glands linear, distally
punctiform. Petals bright yellow to orange-yellow, (6-5-)8-13 x 3-4 mm, c. 1-3-1-5
x sepals, oblong-obovate; apiculus sharply acute; glands linear, distally punctiform.
Stamens c. 40-50, longest 5-6 mm long, c. 0-65 -0-75 x petals. Ovary c. 1-5-2 x
1-1-5 mm, ellipsoid; styles 3, 1-5-2 mm long, 0-7-1-5 x ovary, erect; stigmas
narrowly to broadly capitate. Capsule 4-5 x 2-5-3 mm, ovoid-ellipsoid, obtuse to
rounded, shorter than sepals. Seeds 0-7-1 mm long, ecarinate; testa finely scalar-
iform-reticulate.

In open paramo on rocky, often volcanic slopes; (900-)2440-3700 m.

Costa Rica (San Jose, Alajuela, Cartago, Limon, Puntas Arenas), Panama (Chiri-
qui, Bocas del Toro), Colombia (Bolivar, Antioquia). Map 15.

COSTA RICA. Alajuela: upper slopes of Volcan Poas, c. 2700 m, l.ix.1968 (fl &
fr), Wilbur & Stone 10634 (BM, DUKE, GH, MO, NY, US). Cartago: Asuncion of
Cerro de la Muerte, c. 3150 m, 17.V.1971 (fl), Wilbur 14563 (DUKE, MICH, MO).
Limon: Chirripo National Park, between Casa da Administration and peak, c. 3400
m, 13.ii.1983 (fl & fr), Garwood et al. 1162 (BM). Puntarenas/Bocas del Toro
(Panama): Cordillera de Talamanca, Cerro Echandi, on the International border,
3050-3160 m, 22.viii.1983 (fl), Davidseetal. 2389 (BM, MO).

PANAMA. Bocas del Toro: Valle de Silencia, 2440 m, 8.viii.l979 (fl & fr),
Antonio 1547 (BM, MO). Chiriqui: Cerro Fabrega and vicinity, near Costa Rican
frontier, 3150-3335 m, 7-8.iv.1976 (fl), Weston 10149 (MO).

COLOMBIA. Antioquia: Paramo Frontino, near Llano Grande, 3450 m,
25. x.l 976 (fl & fr), Boeke & McElroy 229 (NY), Bolivar/ Antioquia: Paramo de
Chaquiro, 900-960 m, 23.ii.1919 (fl & fr), Pennell 4285 (GH, NY).

THE GENUS HYPERICUM L.

71

Map 15 38. H. costaricense ; 39. H. bryoides O; 44. H. marahuacanum: a. subsp. marahuacanum
subsp. strictissimum A, c. subsp. chimantaicum D.

H. costaricense is related to 34 H. sprucei, having similar markedly incurved leaves
that twist when long enough to do so, but they are not cucullate and no more than
subcoriaceous. Moreover, the flowers are smaller, with shorter styles and capitate
stigmas.

In a specimen from the highest point of the Pan-American Highway on the
Talamanca Range (J. & C. Taylor 11757 (NY)) the leaves are broader, the flowers
larger, and the styles longer than in the other specimens. It seems probable that the
suggestion on the label is correct, i.e. that it is the hybrid H. irazuense x H.
costaricense. The two Colombian collections belong to H. costaricense despite the
disjunct distribution and the lower altitude of Pennell 4205, which is a dwarf shrub
(0-1-0-15 m tall) with numerous lateral branches, leaves relatively long (c. 10 mm),
recurving markedly and twisting, flowers that are relatively small for the species but
nevertheless fall within its range of variation, and stigmas that are broadly capitate. It
resembles a dwarf version of the (primitive) Panamanian form of//, costaricense and
appears to be a reduction along a different line from that leading to H. bryoides. The
other Antioquia collection (Boeke & McElroy) is taller but otherwise similar.

39. Hypericum bryoides Gleason

in Bull. Torrey hot. Club 56: 102 (clav.), 103 (1929). Type: Colombia, Santander,
Paramo de Santurban, 3000 m, 18. ii. 1927 (fl), Killip & Smith 19533 (NY!,
holotype; GH!, US!, isotypes).

Suffrutex 0-09-0-15 m tall, erect or ascending from creeping branching and rooting
base, forming dense Polytrichum-like rounded clumps, with branches strict, pseudo-
dichotomous and lateral. Stems orange-brown, 4- angled and(?) ancipitous when
young, soon terete, cortex exfoliating irregularly; internodes 1-3-5 mm long. Leaves
sessile, imbricate, eventually outcurving and slightly twisting, tetrastichous, de-
ciduous above base without fading; lamina 3-5 x 0--0-8 mm, linear-subulate,
incurved-canaliculate, cucullate, midrib impressed beneath or not, margin narrowly
hyaline, concolorous, not glaucous, chartaceous; apex subacute to obtuse, base
gradually broadened and sheathing, pairs forming relatively broad interfoliar ridge;
basal vein 1, unbranched; laminar glands dense, visible beneath. Inflorescence
1 -flowered, terminal, with pseudo-dichotomous branches from node below; pedicel
c. I mm long, relatively stout; upper leaves not transitional. Flowers 6-8 mm in
diam., obconic (?). Sepals 3-5 x 0-7-1-3 mm, narrowly oblong to narrowly

72

NORMAN K. B. ROBSON

oblong-lanceolate, acute, incurved above, margin hyaline, veins 3-5, unbranched,
midrib or all prominent; glands all linear or distally punctiform. Petals bright yellow,
3-5-6-5 x 1-5-3 mm, c. 1-2-1-4 x sepals, obovate; apiculus acute or obsolete;
glands striiform. Stamens 20-35, longest 2-5-3 mm long, c. 0-75 x petals. Ovary
0-8-1 x 0-6-0-8 mm, ellipsoid; styles 3, 1-1-2 mm long, c. 1-2 x ovary, erect;
stigmas narrowly to broadly capitate. Capsule 4-4-5 x 2-2-5 mm, cylindric-
ellipsoid, subacute to obtuse, slightly exceeding sepals. Seeds not seen.

Rocky places in open paramo; 3000-4200 m.

Colombia (Norte de Santander, Santander, Boyaca). Map 15 (p. 71).

COLOMBIA. Boyaca: Sierra Nevada del Cocuy, Alto La Cueva, 3850 m,
5.vU973, Cleef 10000 (BM, COL, U). Norte de Santander: Paramo de Romeral,
3800-4200 m, 30.U927 (st), Killip & Smith 18631 (F, GH, K, NY, S, US); Paramo
de Santurban, 5 km E. of Berlin, 3200 m, 4.vii.l984 (fl), Wood 4504 (K). Santander:
Paramo de Almorzadero, 3500-3700 m, 20.vii.1940 (fl), Cuatrecasas & Garcia
Barriga 9993 (F), 9995 (COL), 9996 (NY).

H. bryoides is a reduced suffruticose version of the more specialized form of H.
costaricense , differing from it essentially in size of parts. The dense moss-like clumps
expand by the creeping and rooting of the outer shoots.

40. Hypericum bolivaricum N. Robson, sp. nov.

H. magdalenico N. Robson affinis, sed foliis angustissime ellipticis basi angustatis
saepe terminantibus, petalis brevioribus, staminibus paucioribus, stylis brevioribus
stigmatibus manifeste capitatis, differt. Type: Colombia, Bolivar, headwaters of Rio
Sinu, 1918 (fl), Pennell4183 (NY!, holotype).

Shrub, with branches strict, lateral. Stems orange-brown?, 4-angled when young, the
subfoliar ridges broad, eventually terete, cortex exfoliating in scales; internodes
1-5-2-5 mm long. Leaves sessile, subimbricate at first, spreading and twisting,
scarcely tetrastichous, deciduous above base without fading; lamina 15-20 x 3-3-5
mm, very narrowly elliptic, incurved, not cucullate, midrib not prominent beneath,
margin very narrowly hyaline, not glaucous, coriaceous; apex acute, pungent, base
angustate, scarcely sheathing, united to form narrow interfoliar ridge; basal vein 1,
unbranched; laminar glands rather sparse, not or scarcely visible beneath; marginal
glands dense, small. Inflorescence 1 -flowered, probably with pseudo-dichotomous
branches from node below; pedicel c. 2 mm long; upper leaves not transitional.
Flowers c. 15 mm ? in diam., stellate? Sepals 9-11 x 2-3 mm, oblanceolate-oblong
to narrowly oblong, acute, margin narrowly hyaline; veins 5-7, unbranched, not
prominent; glands linear, punctiform in upper two-fifths. Petals yellow, c. 12 x 6
mm, c. 1-1 x sepals, obovate-oblong; apiculus acute; glands mostly striiform to
punctiform. Stamens 34, longest 5-6 mm long, c. 0-5 x petals. Ovary 3x2 mm,
narrowly ellipsoid; styles 3, 4-5 mm long, 1-5 x ovary, slightly outcurved; stigmas
rather broadly capitate. Capsules and seeds not seen.

No habitat or altitude cited.

Colombia (Bolivar). Known from only one collection. Map 16 (p. 74).
COLOMBIA. Bolivar: headwaters of Rio Sinu, 1918 (fl), Pennell4783 (NY).

Although only one specimen of H. bolivaricum has so far been studied, its taxonomic
position is clear and it is quite distinct from its nearest relatives. Morphologically and
geographically it is intermediate between the long-leaved form of H. magdalenicum
and the H. juniperinum group, having the thickly coriaceous leaves of the former but
the 1-nerved, twisting leaves and predominantly lateral flowers of the latter.

41. Hypericum juniperinum Kunth

in Humboldt, Bonpland & Kunth, Nova Gen. etSp. PI. 5: 189 (1822); Bentham, PI.
Hartweg.: 165 (1845). Type: Colombia, Cundinamarca ?, 'in frigidis Andium
Nova-Granatensium', vii.1805 (fl & fr), Humboldt & Bonpland (P-HUM!, holo-
type; P! (F!, photograph), GH!, isotypes).

THE GENUS HYPERICUM L.

Brathys juniperina L. f.,Suppl. PL: 268 (1781). Type: Colombia, Cundinamarca, 'in
Nova Granada', Mutis (LINN!, holotype; BM!, MA!, US!, isotypes).

H. brathys Sm., Plant. Ic. Ined. Herb. Linn. 2: 41, t. 41. (1790), nom. illegit.; Lam.,
Encycl. meth. (Bot.) 4: 152 (1797); Choisy, Prodr. monogr. fam. Hyperic.: 59
(1821); Kunth in Humboldt, Bonpland, Kunth, Nova Gen. et Spl. PI. 5: 188
(1822); Choisy in DC., Prodr. syst. nat. regni veg.: 1: 554 (1824); Bentham, PL
Hartweg. : 165 (1845); Weddell, Chloris Andina 2: 270 (1857); Treviranus, Hyper.
Animad.: 15 (1861); Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18: 292
(1862); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908), in Engler & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 181 (1925); Knuth in Reprium Spec. nov. Regni veg.
Beih. 43: 484 (1927); Gleason in Bull. Torrey hot. Club 56: 102 (1929); Cuatrec. in
Trab. Mus. nac. Cienc. nat. Jard. hot. Madrid (Bot.) 33: 85 (1936). Type: as for
Brathys juniperina Linn. f.

H. brathys [var.] |3 juniperinum (Kunth) Choisy in DC., Prodr. syst. nat. veg. 1: 554
(1824); Weddell, Chloris Andina 2: 270 (1857); Knuth in Reprium Spec. nov.
Regni veg., Beih. 43: 483 (1927).

Brathys juniperina (Kunth) Spach, Hist. nat. veg. Phan. 5: 450 (1836), in Annls Sci.
nat. (Bot.) II, 5: 366 (1836), non Linn. f. (1781).

H. pseudobrathys Turcz. in Bull. Soc. Nat. Moscou 31 (1): 387 (1858) ['pseudo-
brathys'] pro parte, quoad lectotypum et Linden 1242. Type: Venezuela, Merida,
'in Sierra Nevada', 3000 m, 1846 (fl), Funck & Schlim 1140 (LE, lectotype;
BM!, P! (GH!, photograph), W!, syntypes).

H. jahnii R. Keller in Bot. Jahrb. 58: 196 (1923), in Engler & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey hot. Club 56: 103
(1929); Steyermark in Fieldiana Bot. 28: 988 (1957). Type: Venezuela, Tachira,
Paramo de Batallon, 3000 m, iii.1911 (fl), John 2 (Bf, holotype; NY!, US!,
isotypes, F!, GH!, NY!, photographs).

H. struthiolifolium var. congestiflorum sensu Knuth in Reprium Spec, nov Regni
veg. Beih. 43: 484 (1927) ['struthiolaefolium'] pro parte, quoad Funck & Schlim
1140.

H. holtonii Gleason in Bull. Torrey hot. Club 56: 104 (1929) ['holtonf]. Type:
Colombia, Cundinamarca, 'in montibus juxta Bogotam', 10. xi. 1852 (fl & fr),
Holton 787 (NY!, holotype; GH!, K!, isotypes).

H. strictum var. reductum Gleason in Bull. Torrey hot. Club 56: 104 (1929). Type:
Colombia, Santander, west slope of the Paramo de San turban towards Tona, 3000
m, 18.ii.1927 (fl & fr), Killip & Smith 19534 (NY!, holotype).

Shrub or shrublet 0-2-2-5 m tall, erect or rarely decumbent and rooting at the base,
with branches strict, lateral or sometimes pseudo-dichotomous. Stems orange-
brown, 4- angled and ancipitous when young, soon 2-lined, eventually terete, cortex
exfoliating irregularly; internodes 1-5-10 mm long. Leaves sessile, not or scarcely
imbricate, erect to spreading and twisting, not or scarcely tetrastichous, deciduous
above base without fading; lamina 6-14 x 0--1-5 mm, linear to acicular, rarely
cucullate, midrib impressed beneath, margin narrowly hyaline, concolorous, rarely
glaucous, coriaceous to subcoriaceous; apex acute, base parallel-sided, scarcely
broadening, pairs forming narrow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense to sparse, sometimes visible beneath. Inflorescence 1-flowered,
terminal (usually) and on up to 12 short lateral shoots, sometimes with pseudo-
dichotomous branches from terminal node; pedicel 0-5-3 mm long, not incrassate
upwards; upper leaves transitional. Flowers 4-12 mm in diam., stellate to obconic.
Sepals 2-5-7 x 0-5-2 mm, narrowly lanceolate, acute, veins 5, unbranched, midrib
not prominent, margin hyaline; glands linear, sometimes distally punctiform. Petals
bright to pale yellow, (3-)4-9 x l-5-3(-3-5) mm, 1-3-2-1 x sepals, narrowly
oblong-obovate to obovate; apiculus acute to apiculate; glands mostly linear to
mostly punctiform. Stamens 20-75, 3-5 mm long, longest 0-5-0-75 x petals. Ovary
0-8-3 x 0-6-1-8 mm, broadly ellipsoid or broadly ovoid to globose; styles 3-5(6),
1-3 mm long, 0-5-1-5 x ovary, ascending; stigmas narrowly to broadly capitate.
Capsule 2-5-4 x 2-3 mm, ovoid-subglobose to globose, shorter than sepals. Seeds
0-6-1-2 mm long, ecarinate; testa finely scalariform.

73

9 9

74

NORMAN K. B. ROBSON

Map 16 40. H. bolivaricum O; 41. H. juniperinum .

In open forest, forest margins and damp or shaded areas in paramo and subparamo;
(1800) 2200-3800 m.

Venezuela (Trujillo, Merida, Tachira), Colombia (Magdalena and Norte de Santan-
der to Putumayo). Map 16.

COLOMBIA. Antioquia: Paramo de Urrao, 3500 m, ix.1960 (fl), Espinal & Perez
258 (COL); 1 km norte de Santa Rosa de Osos, c. 2600 m, 25.ix.1948 (fl & fr), Perez
& Barkley 18A058 (COL, NY); Paramo Frontino, near Llano Grande, 3450 m,
25.V.1976 (fl & fr), Boeke & McElroy 219 (MO, NY). Boyaca:* between Soata and
Cocuy, Paramo del Alto del Canutal, 3300-3400 m, 8.ix.l938 (fl & fr), Cuatrecasas
1197 (F); Sierra Nevada del Cocuy, between Praton and Cobugon, c. 3100 m,
15.viii.1957 (fl), Grubb, Curry & Fernandez-Perez 507 (COL, K); Guican, Hoya del
rio Tabor, en direccione a Ritacuva, 3570-3530 m, 18.ix.1969 (fl), Cuatrecasas &
Rodriguez 27833 (BM, COL, US); Paramo de la Rusia, NW.-N. de Duitama, cerca
del puente de rio Surba, 3240 m, 9.xii.l972 (fl & fr), Clee/6925 (COL, U); carreteria
Sogamoso-Pajarito, Km. 57-73, 2850-3010 m, 20.ix.1969 (fl), Diaz 196 (COL).
Caldas: cabeceras del rio Otun, hacia el Nevado de Santa Isabel, Bagas de la Laguna
Negra (Quebrada de la Leona), 3800-3750 m, 24.xi.1946 (fl), Cuatrecasas 23159
(BM, F, P, U). Cuaca: Valley of Rio Cofre, near Gabriel Lopez, c. 3100 m, 14.x. 1961
(fl & fr), Cuatrecasas & Willand 26431 (COL); Paramo de Guanaco, 1843 (fl),
Hartweg 926 (BM, K, P, W);* Macizo Colombiano, Valle de Las Papas, alredores de
Valencia, Los Andes, margens del ri Sucubun, 2910 m, 11. ix. -1.x. 1958, Idrobo,
Pinto & Bischler 3725 (COL, P). Cundinamarca: Macizo de Bogota, El Retire,
Bogota, 2600-2700 m, (v.1946 (fl & fr), Schultes 7004 a (COL, F, GH); Bogota to
Choachi, 27 km W. of Choachi, 3260 m, 4.1.1976 (fl), Luteyn 4730 (BM, COL, MO,
NY); Paramo de Guasca, 2840 m, 15.xii.1938 (fl), Balls 5680 (BM, COL, F, K);
Paramo de Sibate, 3570 m, 22.xi.1981 (fr), Inglesias 19 (COL); entre Bogota y La
Calera, 2650-3000 m, 27.xi.1947 (fl), Barkley, Garcia-Barriga & Vanegas 17C814
(COL, US). Magdalena: Sierra Perija, near Laguna el Juncos, above 'Africa', c.
2200 m, 16.xii.1944 (fl & fr), Haught4503 (F, K). Meta: Paramo de Sumapaz, Hoya
El Nevado, Laguna La Guitarra, 3460 m, 21.1.1972 (fr), C/?/824 (COL, U). Norte
de Santander: Paramo del Hatico (between Toledo and Pamplona), c. 2900 m,
12-13.iii.1927 (fl), Killip & Smith 20662 (GH, NY). Putumayo: between Laguna La
Cocha and Paramo de Tabano, c. 2800-3000 m, l.i.1946 (fl), Schultes & Villareal
7834a (F). Santander:* Bucaramanga (Cumbre to Cucuta), 3300 m, v.1948 (fl),

* Robust, large-flowered form (= H. pseudobrathys Turcz. sensu stricto).

THE GENUS HYPERICUM L.

Sandeman 6067 (BM, K); Paramo de Almorzadero, 3600-3800 m, 28.xi.1941 (fl),
Cuatrecasas 13493 (COL, F, NY); limites entre los Departamentos Santander y
Boyaca, Corregimiento de Virolin, Finca 'La Sierra', 2500-2600 m, 18.vi.1976 (fl),
Lozano, Torres & Diaz 2636 (COL). Tolima: 'Rosalito', near Paramo de Ruiz,
2800-3100 m, 15-17.xii.1917 (fl & fr), Pennell2956 (F, GH, K, MO, NY).

VENEZUELA. Merida: Paramo de La Negra above Bailadores, 3100 m,
13.ii.1939 (ft), Alston 7024 (BM, S, U);* Paramo de Muruti, x.1865 (fl), Moritz 1165
(BM, GH, K, P, S) ; Paramo La Negra between Tovar and La Grita, 3000 m, 7.x. 1965
(fl), Breteler 4619 (K, MO, NY, S, U, YEN). Tachira: Paramo Zumbador, 14 km S.
of El Cobre, 2500 m, 31.iii.1974 (fl), Gentry, Morillo & Morillo 11068 (GH, MO,
NY, VEN); Dtto. Uribante, a las cabeceras del rio Uribante, 51 km al SW. de
Pregonero, 2700-2900 m, 28.ix.1981 (fl), Steyermark & Manara 125420 (BM, VEN).
Trujillo: Alredores de Guirigay, hacia Laguna La Parida, 3300 m, viii.1958 (fl),
Aristeguieta & Medina 3537 (NY, VEN).

The variation in H. juniperinum is rather complex. Spach and Triana & Planchon
distinguished a plant with larger, always 3-styled, sometimes solitary flowers and
fruits and more appressed leaves (based on Kunth's H. juniperinum) from the typical
small-flowered, spreading-leaved H. brathys Sm. (an illegitimate name) with 3-5
styles. Such a large-flowered plant can be recognized in western Venezuela (Trujillo,
Merida) and in isolated Colombian localities in Santander, Boyaca, Cundinamarca,
Antioquia, and Cauca, but it is linked to the typical 'H. brathys' by intermediates.
Indeed Kunth's type of H. juniperinum has flowers somewhat intermediate in size.

In Cauca there is a separate apparently continuous morphocline in which the styles
are always three in number. This cline links the tall erect plants with crowded
subappressed leaves and larger flowers (i.e. H. juniperinum sensu Spach and Triana
& Planchon) with smaller, spreading to decumbent plants with stems rooting, longer
internodes, smaller, closely appressed leaves, and smaller flowers. In its extreme
expression, the latter form approaches H. prostratum, but can be distinguished from
it by the larger leaves and flowers and the longer styles. Specimens exemplifying this
end of the morphocline include Idrobo, Pinto & Bischler3725 (COL, P), 3717 (COL)
and Barclay & Juajibioy 5784 (COL, MO, NY), 5809 (COL, MO, NY), all from
Valle de Los Papas, Los Andes, near Valencia.

75

42. Hypericum prostratum Cuatrec.

in Brittonia 11: 166 (1959). Type: Colombia, Cundinamarca, Paramo de Sumapaz,
3800 m, 10.vi.1952 (fl), K0ie 4621 (US!, holotype; C!, isotype).

Shrub or shrublet with shoots 0-1-0-3 m long, wiry, decumbent to prostrate and
rooting at the base, the branches strict to spreading, lateral (irregularly pinnate), not
pseudo-dichotomous. Stem orange-brown, 4-lined when young, soon 4-angled and
ancipitous, eventually terete, cortex exfoliating irregularly; internodes 2-3 mm long.
Leaves sessile, appressed at first, eventually spreading and twisting, not tetra-
stichous, deciduous above base without fading; lamina 2-5 x 0-5-0-8(-l-2) mm,
linear to acicular, cucullate, midrib impressed beneath, margin broadly hyaline or
golden, concolorous, not glaucous, subcoriaceous, apex acute, base parallel to
cuneate, not broadening, pairs forming narrow interfoliar ridge; basal vein 1,
unbranched; laminar glands dense and prominent above, sparse or not visible
beneath. Inflorescence 1 -flowered, terminal and sometimes on up to 6 short lateral
shoots, without pseudo-dichotomous branches; pedicel almost absent or up to 2 mm
long, not incrassate upwards; upper leaves not transitional. Flowers 4-8 ,m in diam. ,
stellate to obconic. Sepals 4-5, 2-3-5 x 0-5-1 mm, lanceolate, acute, veins 5,
unbranched, midrib not prominent; glands linear. Petals 4-5, bright yellow, 3-6 x
1-5-2 mm, 1-5-2 x sepals, oblong-obovate; apiculus acute; glands mostly linear.
Stamens 7-10, longest 2-5-3-5 mm long, c. 0-65 x petals. Ovary 0-8-1 x 1 x 0-8 mm,
subglobose; styles 3(4), 0-6-0-8 mm long, c. 0-8 x ovary, suberect; stigmas capitate.
Capsule 2-2-5 x 1-5-2 mm, ellipsoid-subglobose to globose, shorter than to slightly
exceeding sepals. Seeds 0-5-0-6 mm long, ecarinate; testa finely scalariform.

In open paramo, usually in damp areas; 3200-4200 m.

76

NORMAN K. B. ROBSON

Colombia (Cundinamarca, Meta, Boyaca, Santander). Map 17.

COLOMBIA. Boyaca: Paramo de Pisva, carretera Socha-La Punta, Km 61.5, 6
km al E. de Los Pinos, Alto de Granados, 3570 m, ll.vi.1972 (fl), C/ee/4374 (BM,
COL, U); Paramo de La Rusia, NW.-N. de Duitama, Aislada, 3575 m, 6.xii.l972 (fl
& fr), Cleef6147 (COL, U); between Soata and Cocuy, Paramo del Alto del Escobal,
3750 m, 8.ix.l938 (fl), Cuatrecasas 1224 (F). Cundinamarca: Bogota-Usme road,
Paramo de Chisaca, between Kms 28 and 33, 3375-3450 m, 17.V.1979 (fl), Luteyn
7770 (BM, COL, MO, NY); Paramo de Cruz Verde, aprox. 5 km al ENE. de la
Laguna El Verjon, 3360 m, 2.V.1972 (fl), Cleef337l (BM, COL, U); entre Cogua y
San Cayetano, Laguna Verde y alredores, 3600 m, 12.xi.1972 (fl), C/ee/6260 (BM,
COL, U). Meta: Paramo de Sumapaz, Hoya El Nevado, Laguna La Guitarra y
alredores, 3460 m, 21. i. 1972 (fl), Cleef834 (BM, COL, U). Santander: Paramo de
Almorzadero, 3500-3700 m, 20.vii.1940 (fl), Cuatrecasas & Garcia Barriga 9998
(COL, US).

H. prostratum continues one of the reduction trends in H. juniperinum, differing
from the decumbent Cauca plants in (for example) its prostrate habit, shorter and
initially strongly appressed leaves, and smaller flowers and fruits. The direction of
the trend, however, has been reversed, i.e. the westernmost specimens (from
Cundinamarca and Meta) are nearest to the above-mentioned Cauca populations of
H. juniperinum both morphologically and geographically.

43. Hypericum parallelum N. Robson, sp. nov.

H. magdalenico N. Robson af finis, sed foliis angustioribus glandulis prominentibus
costa media haud ramosa, floribus minoribus stylis brevioribus sepalis glandulose-
punctatis, differt; a H. pimelioidei Planchon et Linden ex Triana et Planchon inter
alia ramis strictissimis foliis angustioribus valde tetrastichis differt. Type: Colombia,
Norte de Santander/Cesar, Linea divisoria entre los Deptos. Santander del Norte y
Cesar, 20 km al sur de Abrego, Las Jurisdicciones (Cerro de Oroque), 3700-3960 m,
19-21. v.1969 (fl), Garcia- Barriga &Jaramillo 19749 (COL!, holotype).

Shrub 0-3-0-4 m tall, erect, with branches very strict, crowded, parallel, pseudo-
dichotomous, corymbiform. Stems orange-brown becoming blackish, 4-lined but not
compressed when young, the subfoliar ridges broad, soon (?) terete, cortex exfoliat-
ing irregularly; internodes 1-1-5 mm long. Leaves sessile, densely imbricate, slightly
outcurving, markedly tetrastichous, soon deciduous above base without fading;
lamina 6-11 x 1-4-1-7 mm, narrowly elliptic, plane to incurved, cucullate, midrib
slightly impressed beneath, margin narrowly hyaline, concolorous, papillose above,
lucent beneath, not glaucous, coriaceous, apex acute, base narrowly cuneate,
scarcely sheathing, pairs united to form narrow interfoliar ridge; basal vein 1,
apparently unbranched; laminar glands dense, prominent beneath. Inflorescence
1-flowered, with pseudo-dichotomous branches from node below; pedicel 2-2-5 mm
long, slightly incrassate upwards; upper leaves not transitional. Flowers c. 10-12 mm
in diam. (?), stellate (?). Sepals 6-7 x 1-5-2 mm, lanceolate, acute, veins 5,
unbranched, the midrib not prominent; glands linear proximally, punctiform distal-
ly. Petals bright yellow, c. 9-10 x 2-5 mm, c. 1-3 x sepals, obovate; apiculus acute;
glands linear, distally punctiform. Stamens c. 40-50, longest 4-5 mm long, c. 0-5 x
petals. Ovary c. 2 x 1 mm, ellipsoid; styles 3, 2-2-5 mm long, c. 1-1-3 x ovary,
spreading; stigmas broadly capitate. Capsule 5-5-5 x 2-5-3 mm, ellipsoid, acute,
shorter than sepals. Seeds 0-8-0-9 mm long, ecarinate; testa scalariform.

Paramo?; 3700-3960 m.

Colombia (N. de Santander/Cesar); known only from Las Jurisdicciones, Cerro de
Oroque. Map 17.

COLOMBIA. N. de Santander/Cesar: Jurisdicciones, Cerro de Oroque, '3000
-3700-3900 m', 22-27.vii.1974, Garcia- Barriga & Jaramillo 20659 (COL).

H. parallelum is clearly derived from the form of H. magdalenicum with small,
coriaceous, densely imbricate leaves, but differs in the markedly tetrastichous,
cucullate leaves with conspicuous gland dots and the very strict branching.

THE GENUS HYPERICUM L.

77

Map 17 42. H. prostratum ; 43. H. parallelum A ; 45. H. lancifolium O.

44. Hypericum marahuacanum N. Robson, sp. nov.

H. magdalenico N. Robson affinis, sed foliis minoribus acicularibus minus coriaceis,
costa media baud ramosa, sepalis angustioribus stylis gracilioribus brevioribus, inter
alia differt; a H. lancifolio Gleason foliis angustioribus lucentibus torquescentibus
margine valde incurvatis differt. Type: Venezuela, Amazonas, Dept. Atabapo,
Cerro Marahuaca, cumbre Seccion suroriental, 2685 m, 15. i. 1981, Maguire, Steyer-
mark etal. 65629 (YEN!, holotype; MO!, NY!, isotypes).

Icones: Fig. 12A-C.

Shrub 0-3-1-5 m tall, erect, with branches strict to very strict, mostly pseudo-
dichotomous to all lateral. Stems orange-brown, 4- angled and ancipitous when
young, the subfoliar ridges broad, soon terete, cortex exfoliating irregularly; inter-
nodes 0-7-2 mm long. Leaves sessile, densely imbricate, erect to suberect, outcurv-
ing, twisting slightly or not, subtetrastichous to strictly tetrastichous, deciduous
above base without fading; lamina 5-12 x 0-6-1 mm, linear, incurved, not cucullate,
midrib impressed beneath, margin narrowly hyaline, concolorous, polished, not
glaucous, coriaceous; apex acute, often slightly pungent, base parallel-sided, not
sheathing, pairs united to form narrow interfoliar ridge; basal vein 1, without
reticulation; laminar glands in single irregular row, visible beneath or not. Inflores-
cence 1-flowered, sometimes with pseudo-dichotomous branches from node below;
pedicel 2-3 mm long, not incrassate upwards; upper leaves not transitional. Flowers
10-15(-20?) mm in diam., stellate. Sepals 4-9 x 0-6 mm, narrowly lanceolate or
narrowly oblong-lanceolate, acute, veins (3)5-9, unbranched, midrib impressed;
glands linear, becoming prominent (sepals ribbed). Petals deep yellow to orange-
yellow, 6-11 x 2-5-4 mm, c. 1-2-1-7 x sepals, obovate (?) to oblanceolate; apiculus
sharply acute; glands linear, usually distally interrupted. Stamens 30-50, longest 3-9
mm long, 0-45-0-8 x petals. Ovary 1-2-3 x 1-1-5 mm, narrowly ellipsoid or
narrowly ovoid-ellipsoid to ellipsoid-subglobose, acute to rostrate; styles 3(4), 2-5
mm long, 1-5-2 x ovary, outcurved to erect or ascending; stigmas narrow to
narrowly capitate. Capsule 3-5-5 x 2-2-5 mm, narrowly ovoid to ovoid-ellipsoid,
shorter than sepals. Seeds 1-1-2 mm long, ecarinate; testa finely scalariform.

Along shaded (?) streams, in swampy areas, and in drier places and on escarpment;
2500-2800 m.

Colombia (Boyaca), Venezuela (Merida, Amazonas, Bolivar). Map 15 (p. 71).

O O

78

NORMAN K. B. ROBSON

Fig. 12 H. marahuacanum. A. subsp. marahuacanum: (a) habit; (b, b') stem with leaves (two aspects);
(c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut away) and ovary; (g) capsule. B. subsp. strictissimum:
(h) habit. C. subsp. chimantaicum: (i) habit (a, h, i x l /2\ b-g x 3). A. Maguire etal. 65629; B. Fosberg
22209; C. Huber & Steyermark 7014.

H. marahuacanum sensu lato comprises four geographically isolated populations,
which form three taxa. Although geographically separate, the morphological varia-
tion among them is almost continuous, and so the appropriate rank would seem to be
subspecies.

Subsp. marahuacanum (Amazonas: Cerro de Marahuaca) has longer and usually
broader sepals, longer petals, and nearly always longer styles than the other
subspecies, the inflorescence branching is pseudo-dichotomous, and the ovary and
capsule are acute and the stigmas narrow. Subsp. chimantaicum (Bolivar: Chimanta
Massif, Roraima) has a rostrate ovary and capsule and broad to capitate stigmas,
whereas in subsp. strictissimum (Merida, Boyaca) the inflorescence branching is
wholly lateral.

THE GENUS HYPERICUM L. 79

44 a. Hypericum marahuacanum subsp. marahuacanum
Icon: Fig. 12 A.

Branches strict, mostly paired and pseudo-dichotomous, rarely lateral. Leaves
twisting slightly, strictly tetrastichous; lamina 7-10 x 1 mm, with glands visible
beneath. Inflorescence branches pseudo-dichotomous; pedicel 2-3 mm long. Flow-
ers 10-15(7-20) mm in diam. Sepals 7-9 x 1-3-2 mm, narrowly lanceolate or
narrowly oblong-lanceolate, veins 5-9. Petals 10-11 x c.3 mm, c. 1-2-1-3 x sepals,
obovate (?). Stamens c. 50, longest 7-9 mm long, 0-7-0-8 x petals. Ovary 2-5-3 x
1-3-5 mm, narrowly ellipsoid to narrowly ovoid-ellipsoid, acute; styles 4-5 mm long,
1-5-2 x ovary, outcurved; stigmas narrow to narrow capitate. Capsule c. 5 x 2 mm,
narrowly ovoid.

Along shaded (?) steams and on escarpment; 25-2800 m.

Venezuela ( Amazonas) .

VENEZUELA. Amazonas: Departamento Atabapo, Cerro de Marahuaca, parte
central de la Meseta S.-E., Quebrada Yekuana, afluente del rio Negro, 2560 m,
10-12.X.1983, Steyermark 129511 (BM, VEN); Cerro de Marahuaca, al NE. de, y
casi contigua con, Cerro Duida, c. 2750 m, 2 & 9.H.1975, Tillett, Colveeetal. 752-335
(VEN).

Subsp. marahuacanum, apparently confined to the Cerro Marahuacana, is the only
member of the genus recorded from that eminence. It is most closely related to the
narrow-leaved form of H. magdalenicum from the Cerro de Perija (Zulia), differing
from it in the smaller, narrower leaves and the usually smaller flowers with narrower,
ribbed sepals and shorter, more slender styles.

44 h. Hypericum marahuacanum subsp. strictissimum N. Robson, subsp.
nov.

a subsp. marahuacano ramulis plerumque lateralibus, foliis angustioribus, floribus
minoribus, sepals 3-5-venatis, petalis quam sepalis l-7plo longioribus, stylis 2-5-3
mm longis, inter alia differt. Type: Colombia, Boyaca, Socha, valley of Rio
Chicamocha, 2700-2750m, 8.xi.l944 (fl), Fosberg 22209 (US!, holotype; NY!, P!,
isotypes).

Icon: Fig. 12B.

Branches strict to very strict, some paired and pseudo-dichotomous or all lateral.
Leaves twisting slightly or not, markedly tetrastichous; lamina 5-11 x 0-5-0-8 mm,
with glands relatively few, mostly obscure. Inflorescence branches short, lateral,
from up to c. 6 nodes below, sometimes with pseudo-dichotomous branches from
uppermost node; pedicel c. 2 mm long. Flowers c. 10-12 mm in diam. Sepals 4-6 x
0-7-1-5 mm, narrowly lanceolate, veins (3)5. Petals 7-9 x 3-4 mm, c. 1-7 x sepals,
obovate (?). Stamens c. 40, longest c. 3-4 mm long, c. 0-45 x petals. Ovary 1-2-1-5
x 1 mm, ellipsoid-subglobose; styles 2-5-3 mrn long, c. 2 x ovary, suberect; stigmas
scarcely enlarged. Capsule 4x2-5 mm, ovoid, acute, shorter than sepals.

Paramo and subparamo, in dry stony places; 2700-3250 m.

Colombia (Boyaca), Venezuela (Merida).

COLOMBIA. Boyaca: Carretera Socha-Los Pinos, Km 41, 2 km al N. del Alto
Los Pinos y 4 km al ENE. de Socha, 2970 m, 22.V.1973 (fl), Cleef9891 (COL, U);
Paramo de Belen, c. 14 km N. of Belen, near 259 km marker (from Bogota), c. 3250
m, 9.V.1959 (fl), Barclay & Juajibioy 7668 (NY).

VENEZUELA. Merida: Paramo de Las Coloradas (entre Santa Cruz de Mora y
El Molino), San Rafael, 2950-2950 m, 9.H.1973 (fl), Cuatrecasas, Ruiz-Terdn &
Lopez-Figueiras 28529 (BM, US); Sierra Nevada, i.1865 (fl), Moritz 1602 (BM).

Subsp. strictissimum, which comprises two geographically distinct populations,
differs from subsp. marahuacanum in having narrower leaves, smaller flowers which
terminate lateral branches as well as main ones, and shorter styles. When well

80 NORMAN K. B. ROBSON

developed (as in the type collection), the strict lateral branches gives subsp.
strictissimum the appearance of some species of Erica.

44c. II \ perk um marahuacanum subsp. chi man talcum N. Robson, subsp.
nov.

a subsp. marahuacano caulibus gracilioribus, nonnihil magis divergentibus, foliis
haud vel vix punctata, pedicellis longioribus, floribus minoribus interdum ramulis
brevibus lateralibus terminantibus, stylis brevioribus anguste capitatis, capsula
elliptica, differt. Type: Venezuela, Bolivar, Chimanta Massif, Central Seccion, E.
branch of headwaters of Rio Tirica, 2121 m, 12. ii. 1955 (fr), Steyermark & Wurdack
7581 (YEN!, holotype; BM!, MO!, NY!, isotypes).

Icon: Fig. 12C.

Branches strict to ascending, paired and pseudo-dichotomous, rarely lateral. Leaves
twisting slightly, subtetrastichous; lamina 8-12 x 0-6-0-8 mm, with glands few,
obscure or apparently absent. Inflorescence sometimes with 2-4 short axillary shoots
immediate below, pseudo-dichotomous; pedicel c. 3 mm long. Flowers c. 10 mm in
diam. Sepals 4-6 x 0-6-1 mm, linear-lanceolate, acute, veins 5-7. Petals 6-8 x
2-5-3 mm, c. 1-5 x sepals, oblanceolate. Stamens c. 30?, longest c. 4-5 mm long,
0-6-0-7 x petals. Ovary c. 1-5-2 x 1 mm, ovoid-ellipsoid, rostrate; styles 2-3-5 m
long, c. 1-5 x ovary, erect to ascending; stigmas narrowly capitate. Capsule 3-5-5 x
2-2-5 mm, ovoid-ellipsoid, acute to subrostrate.

In swampy open savanna; 2121-2200 m.

Venezuela (Bolivar: Macizo de Chimanta and Roraima).

VENEZUELA, Bolivar: Distrito Piar, Macizo de Chimanta, E., Apacara-tepui,
sector Norte de Macizo, c. 2200 m, 30.i-l.ii.1983 (fl), Steyermark, Huber & Carreno
128351 (BM, VEN), Huber & Steyermark 7014 (BM, VEN); Cimo del Roraima,
i.1977 (fl), Delascio & Brewer 4794 (VEN).

All the characters by which subsp. chimantaicum differs from subsp. marahuacanum
are apomorphic (advanced). It is the easternmost taxon of those in the H.
magdalenlcum affinity and is as isolated as subsp. marahuacanum.

45 . Hypericum lancifolium Gleason

in Bull. Torrey hot. Club 56: 103 (clav.), 105 (1929). Type: Colombia, Santander,
vicinity of La Baja, 2200-2600 m, 14.U928 (fl), Kilttp & Smith 17138 (NY!,
holotype; US!, isotype).

H. struthiolifolium sensu Knuth in Reprium Spec. nov. Regni veg., Beih. 43: 484
(1927) ['struthiolaefolium']; Cuatrec. in Trab. Mus. nac. Cienc. nat. Jard. hot.
Madrid (Bot.) 33: 83 (1936), non Juss. (1804).

H. weberbaueri sensu Gleason in Bull. Torrey hot. Club 56: 102, 103, 105 (1929) pro
parte, quoad specim. Colomb. cit., non R. Keller (1908).

H. lancioides sensu Steyerm. in Fieldiana Bot. 28: 988 (1957).

Shrub 0-2-1-5 m tall, erect, with branches strict, pseudo-dichotomous and some-
times lateral. Stems orange- to reddish-brown, 4-lined and ancipitous when young,
the subfoliar ridges broad, soon terete, cortex exfoliating irregularly; internodes
1-5-3-5 mm long. Leaves sessile, subimbricate to narrowly spreading, tetras-
tichous, deciduous above base without fading; lamina 6-16 x 0-8-3 mm, narrowly
oblong to very narrowly oblanceolate, plane to slightly incurved, not or scarcely
cucullate, midrib not or scarcely impressed distally, margin narrowly hyaline,
concolorous, undulate-papillose on both sides or sublucent beneath, sometimes
glaucous, subcoriaceous; apex sharply acute, base angustate or parallel-sided, not
sheathing, pairs united to form very narrow interfoliar ridge; basal vein 1, occa-
sionally with 1-2 pairs of ascending lateral branches, not branching distally, not
prominent beneath, tertiary reticulation absent; laminar glands dense, frequently
prominent beneath. Inflorescence 1-flowered, with pseudo-dichotomous branches

THE GENUS HYPERICUM L.

from node below; pedicel 2-4 mm long, rather slender; upper leaves not transitional.
Flowers 10-20 mm in diam., stellate. Sepals 6-10 x 1-5-3 mm, narrowly to
acuminate, incurved above, veins 3-5, unbranched or laterals branched, midrib not
prominent; glands sometimes linear to striiform towards base, otherwise (or wholly)
punctiform. Petals pale yellow, 8-12 x 3-5 mm, 1-1-1-3 x sepals, narrowly
obovate; apiculus acute; glands punctiform. Stamens c. 30-70, longest 5-7 mm long,
0-5-0-75 x petals. Ovary 2-3 x 1-2 mm, ellipsoid; styles 3, 2-4 m long, 1-1-75 x
ovary, outcurving; stigmas scarcely to broadly capitate. Capsule 4-7 x 2-5-3 mm,
ellipsoid, acute, shorter than sepals. Seeds 0-7-0-8 mm long, ecarinate; testa finely
scalariform-reticulate.

In paramo; c. 2600-3700 m.

Colombia (Boyaca, Santander, N. de Santander), Venezuela (Tachira, Apure).
Map 17 (p. 77).

COLOMBIA. Boyaca: Paramo de Chita, cabeceras del rio Casanare cerca la
Lagunas de Ocubi, 3080 m, 23.V.1973, Cleef 9903 (U). Norte de Santander:
Pamplona, cerro al NE., 2700 m, 26.vii.1940, Cuatrecasas & Garcia Barriga 10218
(COL, F, US). Santander: Paramo de la Puentas, above La Baja, 3500-3700 m,
25.1.1927, Killip & Smith 18187 (F, GH, K, NY, S, US).

VENEZUELA. Apure: Paramo de Pata de Judio, en la f renter a Colombo-
Venezolana, 30 km al sur de San Vicente de la Revancha, SE. del Paramo de Tama,
19.L1968, Steyermark & Dunsterville 10122 (NY, VEN). Tachira: Paramo de Tama,
cerca de la frontera Colombo- Venezolana, 3000-3200 m, 20-23. v. 1967, Steyermark
& Dunsterville 98752 (VEN).

H. lancifolium is closest to the Duriamena population of//, magdalenicum, differing
in the more slender stems, the narrower smaller, thinner leaves, and the smaller
flowers with shorter styles. The typically plane narrow leaves are distinctive.

H. lancifolium has usually been labelled H. weberbaueri, but it differs from that
Peruvian species (= H. struthiolifolium) in several respects, e.g. the plane (not
incurved) leaves with prominent glands. These two species are not closely related.

The Venezuelan population has shorter, narrower, outcurving leaves and smaller
flowers with fewer stamens and shorter ovary, styles, and capsule. In addition, its
height is usually less and its sepals usually narrower. The leaves are rarely plane as in
H. lancifolium, always more or less incurved. It thus forms a transition to 46 //.
horizontal.

81

46. Hypericum horizontale N. Robson, sp. nov.

H. lancifolio Gleason affinis, sed habitu prostate vel adscendenti, foliis secundis
apice acicularibus, floribus minoribus, stigmatibus vix vel anguste capitatis, capsula
minori, inter alia differt. Type: Colombia, Santander, Paramo de Las Vegas,
3700-3800 m, 20-21. xii. 1926, Killip & Smith 15617 (BM!, holotype; A!, GH!, K!,
US!,isotypes).

Shrublet, up to c. 0-05 m tall, matted, with branches up to c. 0-3 m long, prostrate
with apex ascending, rooting, branching mainly pseudo-dichotomous. Stems orange-
brown, 4-angled and ancipitous when young, soon 2-lined, eventually terete, cortex
exfoliating irregularly; internodes 1-3 mm long. Leaves sessile, not or scarcely
imbricate, ascending to outcurving and secund, not tetrastichous, deciduous above
base without fading; lamina 5-9 x 0-5-1-3 mm, linear, incurved-canaliculate, not
cucullate, midrib impressed dorsally, margin narrowly hyaline, concolorous when
mature, often red-tipped when young, sometimes sublucent, not glaucous, sub-
coriaceous; apex acicular, base parallel-sided, scarcely broadened, pairs forming
narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense to sparse,
visible on both sides. Inflorescence 1 -flowered, terminal, and sometimes also on up to
9 short, paired lateral shoots, usually with pseudo-dichotomous branches from
terminal node; pedical 1-2 mm long or almost absent; upper leaves not transitional.
Flowers 8-12 mm in diam., stellate (? or shallowly obconic). Sepals (3-5-)4-6
- x 1-1-5 mm, narrowly oblong to narrowly or broadly lanceolate, distally incurved,

82

NORMAN K. B. ROBSON

acute, margin hyaline, tip sometimes reddish, veins 5, unbranched, prominent;
glands linear towards base, punctiform or absent distally. Petals bright (?) yellow,
5-7 x 2-5-3-5 mm, 1-1-1-4 x sepals, obovate;apiculusapiculate; glands striiform to
punctiform. Stamens c. 25, longest 3-4 mm long, c. 0-6 x petals. Ovary 1-5-2 x
1-1-5 mm, ellipsoid-subglobose to globose; styles 3-4, 1-5-2-5 mm long, 1-1-3 x
ovary, outcurving; stigma scarcely to narrowly capitate. Capsule 3-4 x 2-2-5 mm,
cylindric-subglobose, obtuse to rounded, shorter than sepals. Seeds 0-6-0-8 mm
long, ecarinate; testa finely scalariform.

In dry, open, sandy areas; 2800-4500 m.

Colombia (Norte de Santander, Santander). Map 18.

COLOMBIA. Norte de Santander: Paramo de Santurban, en route from Tona to
Mutiscua, 4200-4500 m, 19. ii. 1927 (fl & fr), Killip & Smith 19564 (BM, GH).
Santander: Paramo de Las Vetas, a 2 km del paraje de Berlin, c. 2800 m, 18.xii.1948
(fl & fr), Molina & Barkley 18S5410 (COL, F); idem, c. 3450 m, 3.vi.l960, Barclay &
Juajibioy 10445 (MO, NY); Paramo de Almorzadero, 4200 m, 9.ix.l955 (fl),
Vareschi 4070 (YEN p.p.); idem, auf der Passhohe, 3850 m, 24.vii.1952, Schneider
1232 (S).

Although quite different in habit, H. horizontal is clearly related to the Venezuelan
population of H. lancifolium. Its distinct distribution and morphological differences,
taken together, indicate that it should be treated as a separate species.

47. Hypericum tetrastichum Cuatrec.

in Ciencia Mex. 23: 144, f. 3A-G (1964). Type: Colombia, Santander, Municipio de
Onzaga, vereda de Chaguaca, alto de la Laguna de Los Bobos, en el filo divisorio
Santander-Boyaca, 3800 m, 7.viii.l958 (fl), Jaramillo-Mejia, Herndndez-
Camacho & van der Hammen 922 (US, holotype; COL, isotype); same data,
Jaramillo-Mejia, Herndndez-Camacho & van der Hammen 918 (COL!, topotype).

Shrub or shrublet 0-05-1 m tall, erect and densely caespitose, with branches strict,
pseudo-dichotomous or (mostly) lateral. Stems orange-brown to blackish, 4-lined
and compressed when young, the subfoliar ridges broad, soon or eventually terete,
cortex exfoliating in strips or irregularly; internodes 0-5-6 mm long. Leaves sessile,
erect, densely imbricate, markedly tetrastichous, the longer eventually spreading or

Map 18 46. H. horizontale A; 47. H. tetrastichum .

THE GENUS HYPER1CUM L.

outcurving, not twisting, deciduous above the base without fading; lamina 6-13 x
0-6-1-5 mm, linear-acicular, carinate to incurved-canaliculate, cucullate or not,
midrib prominent to impressed dorsally, margin narrowly (golden-) hyaline, some-
times red-tipped or red-tinged when young, otherwise concolorous, dull above, dull
to sublucent beneath, not glaucous (at least when mature), subcoriaceous to
coriaceous; apex acuminate to subacute, base broadened or parallel-sided, not
sheathing, pairs united to form narrow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense to sparse or often absent beneath. Inflorescence 1-flowered,
terminal, sometimes with short pseudo-dichotomous branches from node below;
pedicel almost absent or up to 3 mm long, rather slender; upper leaves not transitional.
Flowers 7-15 mm in diam., obconic. Sepals 4-8 x 1-2-2 mm, narrowly oblong to
lanceolate, acutely acuminate to acute, incurved above, sometimes outcurving,
often red-ripped or red-tinged, margin broadly hyaline, veins 3-5, unbranched,
midrib or all becoming prominent (ribbed); glands linear, distally sometimes inter-
rupted to punctiform. Petals bright yellow, 6-11 x 2-5-5-5 mm, 1-2-1-3 x sepals,
oblong-obovate to obovate; apiculus acute to acuminate; glands interrupted to
punctiform. Stamens 25-40, longest 3-5-5 mm long, 0-5 x petals. Ovary 1-2 x
0-7-1-3 mm, ellipsoid-subglobose to subglobose; styles 3(4), 1-2-5 mm long, 1-1-25
x ovary, outcurving to spreading; stigmas narrowly to broadly capitate. Capsule
3-5-5-5 x 2-5-3-5 mm, cylindric-ellipsoid to ellipsoid-subglobose, obtuse to round-
ed, shorter than sepals. Seeds 0-6-0-8 mm long, ecarinate; testa finely scalariform.

Wet or dry, often exposed areas in paramo and subparamo; 2800-4160 m.

Colombia (Boyaca, Santander, Venezuela (Merida). Map 18.

COLOMBIA. Boyaca: Municipio Villa de Leyva, Parque Nacional Santuario de
Iguaque, 3700 m, 20.vii.1979, Melampy 161 (COL); Municipio de Duitama, Paramo
de La Rusia, 3400-3500 m, 28.xi.1978 (fl), Diaz P. 1243 (COL); Paramo de
Guantiva, 3100 m, 26.vi.1964, Espinal 1733 (COL); paramos al NW. de Belen,
vereda S. Jose de la Montana, Alto de las Cruces y alredores, Cabeceras Q. El Toral,
c. 3950 m, 24.ii.1972 (fl), Clee/1136 (COL, U). Santander: Paramo Ruso, 3150 m,
12.vii.1968 (fl), Barclay & Mullen 38C137 (COL).

VENEZUELA. Merida: Sierra Nevada de Merida, margen N. de la Laguna verde
(al pie del Pico Humboldt), 4100 m, fl. 15-18.ii.1966, Schulz, Rodriguez &
Ramirez 358 (U).

H. tetrastichum is closely related to H. lancifolium and H. strictum, differing
essentially from the latter in its markedly tetrastichous foliage and in the length of the
styles. The leaves appear pungent but may not be rigid when alive.

In some Boyaca localities, H. tetrastichum occurs in two apparently distinct forms:
(i) leaves and flowers larger; leaves glandless beneath, apex acuminate-acicular
(damp habitats); (ii) leaves and flowers smaller; leaves glandular beneath, apex
acute (dry habitats). Until more information is available on the nature of this
variation (genetical or environmental), it seems preferable not to give form (ii) any
taxonomic recognition. Specimens of this form include:

Paramo de La Rusia, NW.-N. de Duitama, Laguna Negra, 3725 m, 14.xii.1972
(fl), C/ee/7208 (COL, U), 7208A (COL); Paramos al NW. de Belen, cabeceras
Quebrada Minas, Hoya Laguna El Alcohol, 3900 m, 26.ii.1972 (fl), Cleefl844 (U).

The drawing of the ovary accompanying the description of H. tetrastichum by
Cuatrecasas depicts the styles as shorter than the ovary. However, my observations
indicate that they are as long as or longer than the ovary, and suggest that the ovary
depicted by Cuatrecasas had begun to enlarge after fertilization.

83

Q Q

48. Hypericum strictum Kunth

in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. PI. 5: 190 (1822); Gleason in
Bull. Torrey hot. Club 56: 104 (1929) pro parte excl. vars. Type: Columbia,
Cundinamarca, prope Santa Fe de Bogota, 2880 m, 1805 (fl), Humboldt &
Bonpland (P!, holotype; US!, photograph).

Shrub 0-2-1 m tall, erect, with branches strict, pseudo-dichotomous or mostly

84

NORMAN K. B. ROBSON

lateral, sometimes decumbent and rooting at the base. Stems orange-brown, 4-
angled and ancipitous when young, then 2-lined, subquadrangular, eventually
terete, cortex exfoliating in strips; internodes 1-7 mm long. Leaves sessile, imbri-
cate, outcurving or spreading but not twisting, not or obscurely tetrastichous,
deciduous above base without fading; lamina 5-13 x 5-6-1-3 mm, linear-acicular,
incurved-canaliculate, cucullate or not, midrib impressed dorsally, margin narrowly
hyaline, sometimes red-tipped when young, otherwise concolorous, dull to sub-
lucent on both sides, glaucous when young, coriaceous to chartaceous; apex acute to
subacuminate, base parallel-sided, pairs forming narrow interfoliar ridge; basal vein
1, unbranched; laminar glands absent or rarely sparse and visible beneath. Inflores-
cence 1-flowered, terminal and sometimes on short to long lateral branches from up
to 5 nodes below (racemiform), sometimes with pseudo-dichotomous branches from
node below; pedicel 2-5 mm long, rather slender to rather stout, not incrassate
upwards; upper leaves not transitional. Flowers c. 12-20 mm in diam., stellate to
obconic. Sepals 4-5-10 x 1-2 mm, lanceolate to narrowly oblong, acute to acutely
acuminate, veins 3-5, unbranched, not or scarcely prominent but glands often
depressed ('sepals ribbed'); margin broadly hyaline; glands linear, distally inter-
rupted. Petals bright yellow, 6-11 x 3-7 mm, 1-1-1-3 x sepals, oblong-obovate to
obovate; apiculus acute to apiculate; glands interrupted, distally subpunctiform.
Stamens 35-50, longest 4-7 mm long, c. 0-65 x petals. Ovary 1-3-2 x 1-1-5 mm,
ellipsoid to ovoid-ellipsoid or subglobose; styles 3, 2-3 mm long, 1-1-5 x ovary,
ascending; stigmas narrowly to broadly capitate. Capsule 4-6 x 2-5-3-5 mm,
ellipsoid, acute to obtuse, snorter than sepals. Seeds 0-6-0-8 mm long, ecarinate;
testa finely scalariform-reticulate.

In exposed areas in paramo and subparamo, and open grassland; 2120-4000 m.
Colombia (Santander, Boyaca, Cundinamarca, Meta). Map 19.

H. strictum is closely related to H. tetrastichum, differing from it in the leaves, which
soon cease to be clearly tetrastichous as a result of stem internode elongation and leaf
outcurving. In addition the leaf apex is never so sharply acuminate as it is in the
longer-leaved form of//, tetrastichum, and the sepal form and glands are distinct. H.
strictum also has a different, though overlapping distribution, being centred in
Cundinamarca and showing trends north-east and south-west from there along the
Cordillera Oriental. Some Boyaca specimens verge toward H. tetrastichum morpho-
logically; others are quite distinct from that species and, indeed, form a derivative
subspecies of H. strictum.

48a. H. strictum subsp. strictum

H. struthiolifolium var. strictum (Kunth) Choisy in DC. , Prodr. syst. nat. regni veg. 1:
553 (1824) ['struthiolaefotium']', Weddell, Chloris Andina 2: 271 (1857); Triana &
Planchon in Annls Sci. nat. (Bot.) IV, 18: 294 (1862).

Leaves 8-13 mm long, outcurving, with apex subacuminate to acute, not cucullate,
rarely red-tipped when young, dull, usually eglandular beneath. Stems simple,
usually erect. Sepals 5-10 x 1-3-2 mm, lanceolate, acutely acuminate to acute,
usually outcurving, rarely red-tipped; veins 5; glands adjacent to midrib often
depressed. Petals 8-11 mm long. Stamens c. 50, longest 4-7 mm long. Ovary 1-5-2
mm long, ellipsoid to ovoid-ellipsoid. Seeds 0-7-0-8 mm long.

In dry rocky paramo and open grassland.

Colombia (Boyaca, Cundinamarca, Meta).

COLOMBIA. Boyaca: Rio Grande, 2 km N. of Cdmbita, 3445 m, l.ix.1944 (fl),
St. John 20695 (NY, US); NW. of Belen, Hoya El Pulpito, 2 km ESE. of Laguna
Grande, 3805 m, 6.V.1973 (fl), Cleef9715 B (COL). Cundinamarca: Represa del
Neusa, 3350 m, 22.vi.1957 (fl), Barclay 4157 (COL); Zipaquira - Pacho, Paramo
Alto, ll.iii.1951 (fl), Romero-Castaneda 2282 (COL, F); Paramo de La Calera,
3000-3300 m, 25.ii.1950 (fl & fr), Philipson, Idrobo & Fernandez 2440 (BM, COL,
F); Bogota, paramo above El Chic6, 3150-3300 m, ll.vii.1943 (fl), Fosberg &

THE GENUS HYPERICUM L.

85

Map 19 48. H. strictum: a. subsp. strictum , b. subsp. compactum ('gracile' form) O, ('compactum'
form) .

Villareal 20585 (COL, NY, US). Meta: Paramo de Sumapaz, Cerro Nevado del
Sumapaz, 3720 m, 31.1.1972 (fl), Clee/1535 (U).

H. strictum subsp. strictum can be recognized by its dull, outcurving, usually
apparently eglandular leaves and the lanceolate sepals that are not or scarcely ribbed
but often have the two linear glands on either side of the midrib impressed.

48b. H. strictum subsp. compactum (Triana & Planchon) N. Robson,
comb, et stat. nov.

H. pseudobrathys Turcz. in Bull. Soc. Nat. Moscou 31 (1): 387 (1858) [pseudo-

brathys'] pro parte, quoad spec. Funck & Schlim 1279, Linden 1313.
H. struthiolifolium [var.] y compactum Triana & Planchon [in Weddell, Chloris

Andina 2: 271 (1857) [ l struthiolaefolium\ 'confertum'], nomen] in Annls Sci. nat.

(Bot.) IV, 18: 294 (1862). Type: Colombia, Santander, prov. de Pamplona, San

Urban, 2570 m, 1846 (fl), Funck & Schlim 1279 (COL?, holotype; BM!, P!,

isotypes).
H. brathys sensu Trev., Hyper. Animad.: 15 (1861) pro parte, quoad spec. Linden

1313.
H. struthiolifolium [var.] E gracile Triana & Planchon [in Weddell, Chloris Andina 2:

271 (1857), nomen] in Annls Sci. nat. (Bot.) IV, 18: 294. (1862), ['struthiolae-

folium'] nom. illegit., pro parte, quoad spec. Linden 1313.
H. laricifolium var. glaucum R. Keller in Bull. Herb. Boissierll, 8: 182 (1908). Type:

Colombia, Boyaca, prov. Tunja, Paramo de Coati, pres de Serinsa, 2870 m,

1842-1843 (fl), Linden 1313 (W!, holotype; GH!, KW!, P!, W!, isotypes).
H. strictum var. compactum (Triana & Planchon) Gleason in Bull. Torrey hot. Club

56: 104(1929).
H. graciliforme N. Robson in Cleef, Veg. Pdram. Colomb. Cord. Or.: 304 (1981),

nomen.

Stems usually branched from the base, ascending. Leaves 5-7(-8) mm long,
outcurving or spreading with apex subacute, cucullate, markedly red-tipped or
-tinged when young, dull to sublucent, often glandular beneath. Sepals 4-5-5-5(-6)
x l-l-2(-l-5) mm, triangular-lanceolate to narrowly oblong, acute to subacute, not
outcurving, red-tipped or -tinged; veins 3(-5), all impressed. Petals 6-8(-10) m

86

NORMAN K. B. ROBSON

long. Stamens 30-40, longest 4-5 mm long. Ovary 1-3-2 mm long, ellipsoid-
subglobose to subglobose. Seeds 0-6-0-7 mm long.

Wet areas in paramo and subparamo, often with Espeletia spp.

Colombia (Boyaca, Santander).

COLOMBIA. Boyaca: Paramo de Santa Rosa, entre Santa Rosa de Viterbo y
Cerinza, 2800 m, 17.vii.1940 (fr), Cuatrecasas & Garcia Barriga. 9721 (COL, K, NY,
P); Paramo de Guantiva, Alto de Canutes, 3200-3400 m, 3.X.1940 (fl & fr),
Cuatrecasas 10329 (COL, F, US). Paramo de La Rusia, NW.-N. de Duitama,
Aislada, 3490 m, 8.xii.l972 (fl & fr), Cleef 6835 (U). Santander: Paramo de
Santurban, near Vetas, 3950-4160 m, 17.U927 (fl), Killip & Smith 17567 (A, BM,
K, US).

Subsp. compactum differs from subsp. strictum in the subacute to acute, cucullate,
usually glandular leaves and the markedly ribbed sepals, both leaves and sepals being
frequently red-tipped or red-tinged when young. It comprises two forms, which are
geographically almost distinct and differ mainly in habit. The form in Santander
('var. compactum'), which is confined to the Paramo de Santurban, has relatively
stout stems with crowded branches that at higher altitudes (3570-4160 m) form a
compact, caespitose clump or prostrate mat. The leaves are relatively stout, often
broader towards the tip, and remain erect and distinctly tetrastichous, and the sepals
are rarely red-tipped. Most of the Boyaca and Cundinamarca plants have taller,
relatively slender, more loosely branched stems forming lax bushes ('var. gracile'),
and the sepals are usually red with a hyaline margin.

The 'gracile' form has been confused with the superficially similar Peruvian -
Ecuadorian H. aciculare, from which it differs by its oblong (not lanceolate) sepals,
which (like the leaves) are nearly always tipped or tinged reddish, become markedly
ribbed and have relatively broad hyaline margins. In addition, the styles are capitate
and the seeds only about half as long, and the leaves become outcurved, not twisted.

49. Hypericum jaramilloi N. Robson, sp. nov.

H. caracasano Willd. affinis, sed foliis et floribus maioribus, sepalis longioribus,
petalis latioribus, staminibus numerosioribus, stylis longioribus, stigmatibus angus-
tioribus, differt. Type: Colombia, linea divisoria entre los Deptos. Santander del
Norte y Cesar, entre Abrego y Las Jurisdicciones (Cerro de Oroque), 3440-3750 m,
19-21. v. 1969 (fl), Garcia-Barriga & Jaramillo Meija 19810 (COL!, holotype).

Shrub 1-1-5 m tall, erect, with branches strict, pseudo-dichotomous or occasionally
lateral. Stems orange-brown, 4-angled when young, the subfoliar ridges broad,
eventually terete, cortex exfoliating in strips; internodes 3-9 mm long. Leaves with
distinct petiole 1-3 mm long, densely imbricate at first, soon outcurving then
usually widely spreading above petiole, subtetrastichous, deciduous above petiole
without fading; lamina 9-15)-20) x 2-5-5(-6-5) mm, usually narrowly elliptic,
plane to subincurved, not cucullate, midrib not prominent beneath, margin not
distinct, concolorous, glaucous, coriaceous; apex acute, base angustate, incurved
but not sheathing, pairs united to form narrow interfoliar ridge; basal vein 1, with
3-4 pairs of ascending main laterals, obscurely branching distally, tertiary reticula-
tion absent; laminar glands dense, sometimes obscure. Inflorescence 1 -flowered,
with pseudo-dichotomous branches from node below; pedicel 5-13 mm long, slightly
incrassate distally; upper leaves broader but not transitional. Flowers 25-30 mm in
diam. , stellate. Sepals 9-14 x 3-6 mm, broadly to narrowly obovate or oblanceolate
to elliptic-oblong or narrowly obovate or oblanceolate to elliptic-oblong or narrowly
elliptic, acute to subacuminate, veins 5-7, unbranched, with midrib not or slightly
prominent; glands linear to punctiform. Petals bright yellow, 12-17 x 6-10 mm,
1-2-1-5 x sepals, obovate; apiculus acute; glands linear, distally punctiform.
Stamens c. 70-80, longest 6-8 mm, c. 0-5 x petals. Ovary 3-5-4 x 2 mm,
cylindric-ellipsoid; styles 3, 6-7 mm long, c. 1-5 x ovary, distally outcurved; stigmas
scarcely capitate. Capsule c. 8 x 4-5 mm, cylindric-ellipsoid to cylindric, shorter
than sepals. Seeds not seen.

THE GENUS HYPERICUM L.

Paramo; 2600-3960 m.

Costa Rica (Limon), Colombia (Magdalena, Cesar/Norte de Santander). Map 20.

COSTA RICA. Limon: Cordillera de Talamanca, Cerro Kamuk massif, between
Cerro Dudu and Cerro Apri, 2900-3100 m, 23 & 26.iii. 1984 (fl), Davidse, Herrera &
Warner 25880 (MO).

COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, transecto del Alto Rio
Buritaca, 2900 m, 5.viii.l977 (ft), Jaramillo etal. 5371 (BM, COL); loc. cit., 3300 m,
9.viii.l977 (fl), Jaramillo etal. 5454 (COL, U). Cesar/Norte de Santander: Cerro de
Oroque, Las Jurisdicciones, 3700-3900 m, 22-27. vii. 1974 (fl), Garcia Barriga &
Jaramillo 20598 (COL); Cerro de Oroque, 20 km al S. de Abrego, 3700-3960 m,
19-21. vii. 1974 (fl), Garcia Barriga & Jaramillo 19728 (COL).

H. jaramilloi is related on the one hand to H. pimeleoides and on the other to both H.
cardonae and H. ruscoides. Its area of distribution overlaps that of H. pimeleoides in
the Cerro de Oroque, where it can be distinguished by (i) the narrower spreading
leaves with narrow (and usually longer) petioles and pinnate venation and (ii) the
narrower sepals. From both H. cardonae and H. ruscoides it can be distinguished by
size of parts (it resembles a larger, more primitive version of the latter), in particular
by the longer styles.

87

50. Hypericum cardonae Cuatrec.

in Revta Acad. colomb. Cienc. exact, fis. nat. 6: 62 (1944), in Trab. Com. Bot. etc.
Colomb.: 36 (1944). Type: Venezuela, Tachira, Paramo de Tama, 3100-3300 m,
vii.1939 (fl), Cardona 303 (YEN!, holotype; F!, isotype).

H. caracasanum sensu Turcz. in Bull. Soc. Nat. Moscou 31 (1): 389 (1858); Knuth in
Reprium Spec. nov. Regni veg. Beih. 43: 483 (1927) pro parte, quoad spec, merid. ,
et auct. plur.

H. caracasanum subsp. cardonae (Cuatrec.) N. Robson in Cleef, Veg. Pdram.
Colomb. Cord. Or.: 305 (1981), comb, illegit. sine basionym.

Shrub or shrublet 0-2-1 m tall, erect or rarely decumbent and rooting at the base,
with branches strict, pseudo-dichotomous or lateral. Stems yellow- to orange-brown,
4-lined and ancipitous when young, the subfoliar ridges broad, soon terete, cortex
exfoliating in strips; internodes 3-7 mm long. Leaves with petiole 1-2-5 mm long,

Map 20 49. H. jaramilloi A ; 50. H. cardonae ; 51. H. caracasanum: a. subsp. caracasanum , b. subsp.
turumiquirense D .

NORMAN K. B. ROBSON

persistently imbricate-tetrastichous to widely spreading from above petiole, de-
ciduous above petiole without fading; lamina 7-15 x 2-5-7 mm, broadly to narrowly
elliptic or narrowly oblong, plane to incurved or subconduplicate, not cucullate,
midrib not or slightly prominent beneath, margin plane to incrassate, concolorous,
often glaucous to blue-green, coriaceous; apex acute to subrounded, base angustate
to cuneate, the petioles not sheathing but united to form narrow interfoliar ridge;
basal vein 1, with (l-)2-3 pairs of ascending lateral branches (often obscure), rarely
visibly branching, not impressed beneath, tertiary reticulation not visible; laminar
glands dense, visible beneath or not. Inflorescence 1-flowered, sometimes with
several flowering branches clustered, with pseudo-dichotomous branches from one
or two nodes below; pedicel 4-6 mm long, not incrassate distally; upper leaves not
transitional. Flowers 12-18 mm in diam. , stellate or obconic. Sepals 6-12 x 2-4 mm,
broadly to narrowly elliptic or oblong-elliptic to oblong-spathulate, acute to sub-
acute ; veins 5-7 , sometimes branched distally, with midrib not or slightly prominent;
glands mostly linear, distally punctiform. Petals bright yellow to orange-yellow, 8-11
x 3-6 mm, c. 1-3 x sepals, oblanceolate to obovate; apiculus acute; glands mostly
linear, distally punctiform. Stamens 40-60, longest 4-7 mm long, c. 0-5 x petals.
Ovary 2-5-3 x 1-5-2 mm, ovoid-ellipsoid to ellipsoid-subglobose; styles 3, 3-5 mm
long, 1-1-6 x ovary, divergent to suberect; stigmas broadly capitate. Capsule 4-8 x
2-4 mm, cylindric-ellipsoid, equalling or shorter than sepals. Seeds 1-1-2 mm long,
ecarinate; testa finely scalariform.

In humid (shaded or open) places in the paramo; 2500-4160 m.

Costa Rica (Cartage, San Jose), Venezuela (Trujillo, Merida, Tachira, Apure),
Colombia (Norte de Santander, Santander, Boyaca, Arauca). Map 20 (p. 87).

COSTA RICA. Cartago: Cordillera de Talamanca, 'Cerro de la Muerte', Pan-
American Highway 5 km above Millsville (8 km above Nivel), 3400-3300 m, vii.1949
(fl), Holm & Iltis 454 (F, GH, K, MO, NY, P, U). San Jose: Valle de los Conejos
(upper Rio Talari) and trails to Cerro Cirripo and Valle de los Lagos, 3400-3820 m,
22.viii.1971 (fl), Burger 8281 (BM, DUKE, F).

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, 3350 m,
13.iii.1973 (fl), Cleef9l82D (COL). Boyaca: paramos al NW. de Belen, cabeceras
Quebrada Minas, Hoya Laguna El Alcohol, 3800 m, 26.ii.1972 (fl), CleeflSlQ (U);
Cocuy Mts, valley of Quebrada Playitas above Guican, 3800 m, 24. vi. 1984 (fl),
Wood 4453 (K). Norte de Santander: Paramo de Tama, alredores de la Cueva, 3000-
3200 m, 28.x. 1941 (fl), Cuatrecasas, Schultes & Smith 12691 (COL). Santander:
Paramo de Santurban near Vetas, 3950-4160 m, 17.L1927 (fl), Killip & Smith
21 167 (NY).

VENEZUELA. Apure: Paramo de Pata de Judio, 30 km al S. de San Vicente de
Revancha, 3000-3200 m, 19.U968 (fl), Steyermark & Dunsterville 101140 (NY,
VEN). Merida: Paramo de Los Leones (La Lagunita, La Canada Grande), W. of
Mucuruba, 3500 m, 31.V.1930 (fl & fr), Gehringer 138 (F, GH, NY, VEN); Villa de
Caracas, Las Lagunetas, xi. (fl & fr), Moritz 1167 (BM, GH, K, KW, S). Tachira:
Paramo El Batallon, 3200-3600 m, 3.x. 1956 (fl), Vareschi549l (VEN); Hoya del Rio
Tachira, cabeceras de la Quebrada de El Reposo, 2800-3000 m, 16.L1973 (fr),
Cuatrecasas, Ruiz-Terdn & Lopez-Figueras 28330 (BM, US); Paramo de Tama,
cerca la frontera Colombo- Venezolana, 3000-3200 m, 20-23.V.1967 (fl), Steyermark
& Dunsterville 98735 (VEN).

Some forms of H. cardonae from Merida are rather similar to H. caracasanum, but
the capitate stigma and the thicker, glaucous leaves with glands obscure or invisible
will easily distinguish it. There is a southward reduction trend in habit and in leaf
shape, plants from the Paramo de Tama to Belen being dwarfer (almost suffruticose)
and having smaller leaves, with thickened margin, obtuse to subrounded apex,
and a longer thinner petiole. The Costa Rican population is more similar to
plants from Merida, but shows a similar reduction tendency to that of the southern
plants.

For differences between H. cardonae and H. caracasanum, see the description of
the latter species. The reduced form of//, pimeleoides can be distinguished from that
of H. cardonae by the broader leaves with flabellate venation visible beneath.

THE GENUS HYPERICUM L.

5 1 . Hypericum caracasanum Willd .

Sp. PL 3: 1458 (1802); Choisy, Prodr. monogr. fam. Hyperic. 58 (1821); Kunth in
Humboldt, Bonpland & Kunth, Nova Gen. et Spec. PL 5: 186 (1822); Choisy in
DC. , Prodr. syst. nat. regni veg. 1: 554 (1824); Turcz. in Bull. Soc. Nat. Moscou 31
(1): 389 (1858); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908); in Engler &
Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Knuth in Reprium Spec. nov.
Regni veg. Beih. 43: 483 (1927) pro parte, excl. spec, merid.; Gleason in Bull.
Torrey hot. Club 56: 103. (1929); Steyerm. & Huber, PL Avila: 493, t. 152B
(1978). Type: Venezuela, Distrito Federal, Silla de Caracas, i.1800 (fl), Humboldt
& Bonpland 649 (B, holotype, F!, US!, photographs; P!, isotype).

Shrub or shrublet 0-5-1-8 m tall, erect, with branches strict, pseudo-dichotomous or
lateral. Stems yellow- to orange-brown, 4-6-lined and compressed when young, the
subfoliar ridges broad, soon or eventually terete, cortex exfoliating in strips;
internodes 1-4 mm long. Leaves subsessile to broadly or narrowly petiolate with
petiole up to 1-5 mm long, densely imbricate, tetrastichous, ascending to outcurv-
ing, deciduous above petiole without fading; lamina 5-14(-17) x 2-7 mm,
narrowly elliptic or ovate-elliptic to oblong, plane to incurved or slightly cucullate,
midrib slightly prominent to impressed beneath, margin plane to recurved or slightly
incrassate, paler beneath or concolorous or with margin pinkish, not glaucous,
coriaceous; apex acute to subacute, base angustate, the petioles not or scarcely
sheathing but united to form narrow interfoliar ridge; basal vein 1, with 1-3(4) pairs
of ascending lateral branches, not visibly branching, not impressed beneath, tertiary
reticulation not visible; laminar glands dense, usually visible beneath. Inflorescence
1-flowered, with pseudo-dichotomous branches from node below; pedicel 2-5-5-5
mm long, not or scarcely incrassate upwards; upper leaves not transitional. Flowers
12-20 mm in diam., stellate. Sepals 5-11 x 2-4-5 mm, broadly or narrowly elliptic or
oblong-elliptic to obovate-spathulate or oblanceolate, acute to subacute; veins 3-7,
sometimes branched distally, with midrib slightly prominent; glands linear, distally
punctiform. Petals bright? to deep yellow, 8-13 x 3-5-6 mm, 1-4-2 x petals,
oblanceolate; apiculus acute; glands mostly linear, distally interrupted to punct-
iform. Stamens c. 50-70, longest c. 4-7 mm long, c. 0-5 x petals. Ovary 2-3-5 x 2
mm, ovoid-ellipsoid; styles 3, 4-5 mm long, c. 1-5-2-5 x ovary, diverging and
sometimes incurved; stigmas narrow or clavate. Capsule (5)6-8 x (2)3-4 mm,
cylindric-ellipsoid, equalling sepals. Seeds c. 1-2 mm long, ecarinate; testa finely
scalariform.

In humid or shaded habitats in the paramo and subparamo; (1600) 2000-2765 m.
Venezuela (Distrito Federal, Miranda, Aragua, Anzoategui, Sucre). Map 20 (p. 87).

H. caracasanum and H. cardonae are vicarious species which differ in leaf shape,
thickness and glandularity, and in stigma shape, but H. caracasanum is more
restricted in morphological variation and distribution than is its vicariad. In H.
caracasanum the leaf glands are always visible and the stigma is not or scarcely
enlarged. It appears to be directly related to H. jaramilloi.

The extreme easternmost population of H. caracasanum, in the Cerro de Turumi-
quire (Sucre), has smaller, more crowded leaves and somewhat smaller flowers than
the rest of the species, thus showing a trend in morphology as well as geography
towards H. ekmanii from Hispaniola. The variation between the Sucre population
and the rest is apparently almost continuous, and therefore subspecies is the
appropriate rank for the two populations.

51a. H. caracasanum subsp. caracasanum

Brathys caracasana (Willd.) Spach, Hist. nat. veg. Phan. 5: 448 (1836) in Annls Sci.
nat. (Bot.) II, 5: 366 (1836).

Icones: Steyerm. & Huber, FL Avila: t. 152B (1978); Fig. 13 A.

Shrub or shrublet 0-5-1 m tall. Stem internodes 2-4 mm long. Leaves subsessile or
with petiole up to 1-5 mm long; lamina 7-14(-17) x (2-)3-5-7 mm, narrowly elliptic

89

90

NORMAN K. B. ROBSON

to oblong. Pedicels 4-5-5-5 mm long. Flowers 15-20 mm in diam. Sepals 8-11 x
2-4-5 mm, broadly or narrowly elliptic or oblong-elliptic to obovate-spathulate,
veins 5-7. Petals bright? yellow, 11-13 x 5-6 mm, c. 1-4 x sepals. Ovary 2-3 x
1-5-2 mm; styles 4-4-5 mm long, c. 1-5-2 x ovary.

In humid habitats in the paramo; (1600)2000-2763 m.

Venezuela (Distrito Federal, Miranda, Aragua, Anzoategui).

VENEZUELA. Anzoategui: Rio Mantiales E. of Bergantia, Cerro Peonia above
Santa Cruz, 2350 m, 21.iii.1945 (fl), Steyermark 61667 (F, NY, S, VEN). Aragua:
prope Colonia Tovar, 1854-1855 (fl), Fendler40 (GH, K, MO, P). Distrito Federal:
Silla de Caracas, Cordillera de Avila, between Los Venados and Pico Oriental,
1675-2640 m, 27-28. ii. 1944 (st), Steyermark 55644 (F, NY). Miranda: Pico de
Naiguata, above Los Chorros, 2200-2675 m, 16-17.vi.1945 (fl), Steyermark 62943
(F, NY).

51b. H. caracasanum subsp. turumiquirense (Steyerm.) N. Robson, stat.
nov.

H. caracasanum var. turumiquirense Steyerm. in Fieldiana Bot. 28: 393 (1952). Type:
Venezuela, Sucre, Cerro Turumiquire, north-facing slopes, 2360-2500 m,
6.V.1945 (fl), Steyermark 62579 (F!, holotype; NY!, US!, isotypes).

Shrub 0-8-1-8 m tall. Stem internodes 1-2 mm long. Leaves with petiole 1-1-5 mm
long; lamina 5-8 x 2-3-5 mm, ovate-elliptic to elliptic. Pedicels 2-5-3 mm long.
Flowers 12-18 mm in diam. Sepals 5-6 x 2-3 mm, obovate-oblong to oblanceolate-
spathulate, veins 3-5. Petals deep yellow, 8-12 x 3-5-4-5 mm, 1-5-2 x sepals.
Ovary c. 2 x 1 mm; styles 4-5 mm long, 2-2-5 x ovary.

On steep north-facing sandstone slopes in subparamo vegetation; 2360-2910 m.

Venezuela (Sucre). Confined to the Cerro Turumiquire.
VENEZUELA. Sucre: Cerro de Turumiquire, 1925 (fl), Tate 329 (US).

52. Hypericum ekmanii A. H. Liogier

in Phytologia 47: 183 (1980). Type: Dominican Republic, Peravia, San Jose de Ocoa,
2000 m, 4.vi.l978 (fl & fr), A & P. Liogier 27672 (SDM, holotype; GH!, isotype).

Icon: Fig. 13B.

Shrub up to 0-75 m tall, erect, forming thickets, with branches ascending, pseudo-
dichotomous and sometimes lateral. Stems orange-brown, 4-lined and compressed
when young, the subfoliar ridges broad, soon 2-lined, eventually terete, cortex
exfoliating at first in strips between subfoliar ridges; internodes 2-4 mm long. Leaves
with short petiole (0-5-1 mm long), densely spreading or imbricate, tetrastichous,
deciduous above petiole without fading; lamina 6-10 x 2-4 mm, elliptic, plane to
slightly concave, scarcely cucullate, midrib not or slightly prominent beneath,
margin incurved, not incrassate, concolorous, not glaucous or slightly so beneath,
subcoriaceous; apex acuminate to apiculate, base angustate to cuneate, the petioles
not sheathing but united to form very narrow interfoliar ridge; basal vein 1, with 1
short near-basal pair of branches or unbranched; laminar glands dense, visible on
both sides. Inflorescence 1 -flowered, with pseudo-dichotomous branches from node
below; pedicel 3-10 mm long, not incrassate upwards; upper leaves not transitional.
Flowers c. 12-15 mm in diam., stellate. Sepals 5-6 x 1-2-5 mm, unequal to equal
narrowly oblong to oblanceolate-spathulate, acute to subacuminate; veins 3, un-
branched, with midrib not prominent, glands linear, punctiform in distal 1/4. Petals
deep? yellow, 8-10 x 2-4 mm, c. 1-5 x sepals, oblong-oblanceolate; apiculus
subacute; glands nearly all punctiform. Stamens c. 80, longest 5-6 mm long, c. 0-6 x
petals. Ovary 2x1 mm, narrowly ovoid; styles 3, 5-6 mm long, 2-5-3 x ovary,
diverging-incurved; stigmas clavate. Capsule 3-6 x 2-3 mm, ellipsoid, shorter than
sepals. Seeds 1-2 mm long, ecarinate; testa finely ribbed-scalariform.

On rocky slopes in cloud forest; 1300-2550 m.

THE GENUS HYPERICUM L.

91

Fig. 13 A. H. caracasanum subsp. caracasanum: (a) habit; (b) leaf; (c)- sepal; (d) petal; (e) stamens
(partly cut away) and ovary. B. H. ekmanii: (f) habit; (g) stamens (partly cut away) and ovary. C. H.
pycnophyllum: (h) habit. D. H. millefolium: (i) habit (a, f, h, i x Vi; b x 2; c-e, g x 3). A. Davidse &
Gonzalez 19550; B. Ekman 11718; C. Liogier 21790; D. Holdridge 1776.

Dominican Republic (Sierra de Ocoa). Map 21 (p. 92).

DOMINICAN REPUBLIC. Azua: Sierra de Ocoa, San Jose de Ocoa, S. slope of
Tetero de Majia, c. 2550 m, l.ii.1929 (fl & fr), Ekman H. 11718 (S, US).

Ekman (on the back of the label) states that the species is quite common in 'these
mountains' (i.e. Sierra de Ocoa) and occurs at 1300 m near Bejucal. The gap in
distribution between H. ekmanii (Hispaniola) and its nearest relative, H. caracasa-
num subsp. turumiquirense (NE. Venezuela), is likely to have resulted from long-
distance dispersal and therefore cannot be adduced to support the theory that North
and South America were originally linked via the Greater Antilles and the Yucatan
Peninsula (see p. 9).

92

NORMAN K. B. ROBSON

Map 21 52. H. ekmanii ; 53. H. pycnophyllum A ; 54. H. millefolium

53. Hypericum pycnophyllum Urban

Symb. Antill. 7: 523 (1913). Type: Dominican Republic, La Vega, Loma Rosilla,
1700 m, 6.vii.l912 (fl), Fuertes 1749 (Bf, holotype; NY!, P!, W!, isotypes).

Icon: Fig. 13C.

Shrub 0-2-1 m tall, erect, with branches very strict, pseudo-dichotomous and lateral.
Stems reddish-brown, 2-lined and compressed when young, the subfoliar ridges
broad, very soon terete, appearing articulated when old, cortex exfoliating at first in
strips between subfoliar ridges; internodes 1-1-5 mm long. Leaves petiolate with
short petiole (c. 0-5 mm long), densely imbricate, markedly tetrastichous, deciduous
above petiole without fading or with attached cortex below after fading; lamina 3-9
x 0-5-2 mm, very narrowly elliptic to linear, incurved to canaliculate, cucullate,
midrib impressed beneath, margin narrowly hyaline, dull above, lucent beneath, not
glaucous, coriaceous; apex acute, base angustate to above 'articulation' then
broadening, pairs forming narrow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense, obscure, visible above only. Inflorescence 1 -flowered, terminal
and on lateral branches, sometimes with single or pseudo-dichotomous branches
from node below; pedicel absent or up to 3 mm long, not or slightly incrassate
upwards; upper leaves not transitional. Flowers 10-20 mm in diam., stellate to
obconic. Sepals 5-6(-7) x 1-2 mm, subequal, lanceolate to oblong-lanceolate,
acute to acuminate, cucullate, with margin narrowly hyaline; veins 3-7, unbranched,
midrib not prominent; glands linear, uninterrupted. Petals deep? yellow to orange-
yellow, 7-14 x 3-5-7 mm, c. 1-6 x sepals, oblanceolate; apiculus acute; glands
linear, distally interrupted. Stamens 55, longest 4-5-6-5 mm long, c. 0-6 x petals.
Ovary c. 2 x 1 mm, ovoid; styles 3, 3-5-4 mm long, 1-5-2 x ovary, diverging-
incurved; stigmas subclavate. Capsule 5-7 x 2-5-3 mm, narrowly ovoid, shorter
than sepals. Seeds c. 1 mm long, ecarinate; testa finely ribbed-scalariform.

In open Pinus occidentalis forest and among rocks at higher altitudes; 1700-3175 m.

Dominican Republic (Santiago, La Vega, San Juan, Azua). Map 21.

DOMINICAN REPUBLIC. Azua: El Pico del Yaque, 3125 m, 3.U944 (st),
Jimenez 232 (US). La Vega: Sabana Alte, c. 2700 m, 16.x. 1929 (fl & fr), Ekman H.
13793 (S). San Juan: Pico Duarte, 3170m, 10-14.vii.l976(fl), A & P. Liogier 25040
(NY). Santiago: Mention, top of Cerro Lucio, c. 2525 m, 12. vi. 1929 (fl & fr), Ekman
H.12850 (C, F, K, NY, S, US).

The specimens from higher altitudes have shorter leaves. Hypericum pycnophyllum
is apparently more widespread than H. ekmanii and links it morphologically with H.
millefolium.

THE GENUS HYPERICUM L.

54. Hypericum milk-folium Urban & Ekman

in Ark. Bot. 20 A, no. 15: 79 (1926). Type: Haiti, Massif de Selle, Morne de la Selle,
Morne Emerillon, 2000 m, i.1925 (fl & fr), Ekman H. 3126 (Bt, holotype; K!,
NY!,S!,isotypes).

Icon: Fig. 13D.

Shrub up to c. 2 m tall, erect, juniperoid, with branches strict, pseudo-dichotomous
(extension) and shorter, lateral (flowering). Stems 4-6-lined and ancipitous when
young, persistently 4-angled and ancipitous until cortex exfoliates at first in strips
between subfoliar ridges, then terete, fragile, green becoming pale grey; internodes
1-2 mm long. Leaves sessile, imbricate to curved-ascending, tetrastichous, eventu-
ally deciduous with attached cortex below after (?) fading; lamina 0-7-2 x 0-8-1
mm, lanceolate-triangular, incurved, markedly cucullate, midrib impressed be-
neath, margin narrowly hyaline, dull above, lucent or glaucous beneath,
coriaceous; apex acute, pungent, base broad, subamplexicaul, pairs united to form
narrow interf oliar ridge ; basal vein 1 , unbranched ; laminar glands obscure . Infloresc-
ence 1-flowered, terminal and on numerous short lateral branches (branching or
not), occasionally with pseudo-dichotomous branches from node below; pedicel
absent or up to 0-5 mm long, incrassate upwards; upper leaves not transitional.
Flowers 5-7 mm in diam., obconic. Sepals 3-3-5 x 1 mm, lanceolate, acute and
incurved-cucullate to pungent; veins 5, unbranched, midrib prominent; glands
linear, uninterrupted. Petals deep? yellow, 6-7 x 2-5-3 mm, 2 x sepals, oblan-
ceolate; apiculus very acute; glands linear, distally subpunctiform. Stamens c. 50,
longest 4-5 mm long, 0-6-0-7 x petals. Ovary 1-5x1 mm, broadly ovoid; styles 3,
4-5 mm long, c. 3 x ovary, diverging-incurved; stigmas narrow. Capsule 4x2 mm,
ovoid, exceeding sepals. Seeds not seen.

Among steep limestone rocks; 1750-2350 m.

Haiti (Massif de la Selle). Map 21.

HAITI. Quest: Massif de la Selle, Morne la Selle, Bouche Bouqui, 1750 m,
15.X.1943 (fl & fr), Holdridge 1776 (BM, MICH, NY, US); Croix des Bouquets,
Morne Badeau, c. 2200 m, 24.ii.1927 (fl), Ekman H. 7679 (F, S, US), Petionville, top
of Morne Cabaio, 2350 m, 10.iv.1927 (fl), Ekman H. 7979 (C, S).

Hypericum millefolium, with its ericoid habit, is at first glance quite distinct from any
other species in sect. Brathys. Closer investigation, however, shows it to be similar to
the smaller form of H. pycnophyllum. Its triangular leaves and numerous fragile
flowering stems, nevertheless, make it easily recognizable.

55. Hypericum ruscoides Cuatrec.

in Ciencia Mex. 4: 63 (1943). Type: Colombia, Putumayo, Paramo de San Antonio
del Bordoncillo, between El Encano and Sibundoy, 3250 m, 4. i. 1941 (fl), Cuatre-
casas 11720 (COL, holotype; F!, P!, US!, isotypes).

Shrub 0-1-1-5 m tall, erect or with stems sometimes decumbent and rooting at the
base, with branches strict, pseudo-dichotomous and sometimes lateral. Stems yel-
low-brown, 4-lined and compressed when young, the subfoliar ridges broad, even-
tually terete, cortex exfoliating in strips between subfoliar ridges; internodes 2-5-6
mm long. Leaves petiolate with petiole 1-5-3-5 mm long, ascending to spreading,
tetrastichous, deciduous above petiole without fading; lamina 8-18 x 3-6(-8) mm,
narrowly ovate or lanceolate to narrowly oblong-elliptic or the uppermost elliptic,
plane or slightly conduplicate, not or rarely carinate, slightly cucullate, midrib not or
slightly impressed beneath, margin not distinct or rarely subincrassate, concolorous,
sometimes ferrugineous, lucent or dull and undulate-papillose on both sides,
sometimes glaucous, coriaceous; apex acute to apiculate, base angustate, the
petioles not sheathing but united to form narrow interfoliar ridge; basal vein 1, with
2-4 pairs of obscure ascending branches, rarely visibly branched or reticulating;
laminar glands dense, sometimes visible on either or both sides. Inflorescence
1-flowered, with pseudo-dichotomous branches from node below; pedicel 4-8 mm

93

94

NORMAN K. B. ROBSON

long; upper leaves often broader but not transitional. Flowers 15-25 mm in diam.,
stellate. Sepals 5-10 x 1-5-4 mm, obovate-spathulate to narrowly oblong (unequal
to subequal), acute to obtuse; veins 5, branched or not, midrib not prominent; glands
linear, distally punctiform. Petals deep yellow to orange-yellow, 8-15 x 4-6 mm, c.
1-5 x sepals, obovate-spathulate; apiculus subacute to obtuse; glands linear, distally
punctiform. Stamens 30-60, longest 4-6-5 long, c. 0-5 x petals. Ovary 2-3 x 1-5-2
mm, ovoid; styles 3, 2-5-4 mm long, c. 1-3 x ovary, suberect to divergent; stigmas
broadly capitate. Capsule 4-8 x 3-3-5 mm, cylindric-ellipsoid to ellipsoid, shorter
than sepals. Seeds c. 1-5 mm long, ecarinate; testa finely scalariform.

In paramo; 2450-3800 m.

Colombia (Cordilleras Oriental, Central and Occidental from Boyaca and Cundina-
marca to Putumayo), Ecuador (Pichincha and Napo: Volcan Cayambe). Map 22.

COLOMBIA. Boyaca: Villa de Leiva, Parque Igaque, Quebrada Carrizal, 3400 m,
25.xi.1984 (fl & fr), Wood 4621 (K). Caldas: Navarco, 3000 m, 12.ix.1951 (fl),
Romero-Castaneda 3021 (COL). Cauca: Cordillera Central, Paramo de las Moras,
3800-3700 m, 19.ii.1969 (fl), Cuatrecasas 27424 (BM, COL, US). Cundinamarca:
Municipio de Fomeque, paramo de Chingaza, cerros vecinos a la Laguna y limites
con el Depto. del Meta, 3300 m, 2.xi.l966 (fl), Huertas & Camargo 6572 (COL).
Huila: Cordillera Oriental, 20 km SE. of Gigante, 20.ix.1944, Little 8687 (COL).
Meta: Paramo de Sumapaz, Cerro Nevado de Sumapaz, 3725 m, 29. i. 1972 (fl), Cleef
1206 (BM, COL, U). Narino: Pasto a Sibundoy, c. 3000 m, 5.i. 1952 (fl), Fernandez &
Knoth 1023 (COL). Putumayo: Valle de Sibundoy, Sibundoy, 2800 m, 30.vi.1953
(fl), Schultes & Cabrera 20138 (GH, NY). Tolima: Cordillera Central, Municipio de

Map 22 55. H. ruscoides A ; 60, H. selaginella O.

THE GENUS HYPERICUM L. 95

Roncevalles, paramo de Normandia, 3200 m, 8.xi.l980 (fl), Camargo 7508 (COL).
Valle: Cordillera Occidental, Los Farallones, cerro Alto de Buey, 3500-3600 m,
ll.x.1944, Cuatrecasas 17917 (BM, F, US), 17940 ^COL).

ECUADOR. Pichincha: Cordillera Oriental, Laguna de Chiqui, NE. of
Cayambe, 3800 m, 4.vi.l944 (fl), Acosta Soils 8151 (F). Napo: ENE. of Cayambe
Mtn, 3210 m, 6.xii.l961, Cazalet & Pennington 5522 (K, NY).

Hypericum ruscoides is related to H. jaramilloi, being intermediate between it and
H. lancioides. It can easily be confused with reduced forms of//, pimeleoides in the
area where the ranges of these species overlap, but can be recognized by the usually
narrower and less crowded leaves with longer, narrower petioles, acute apex, and no
free veins, i.e. the venation is pinnate.

H. ruscoides appears to be rare in the Cordillera Occidental. Two collections from
Valle de Cauca, Los Fardallones, cerro Alto del Buey (Cuatrecasas 17917, 17940)
have thinner, narrower, more shortly petiolate leaves than usual, and are nearest in
form to H. lancioides. Cuatrecasas has distinguished them as 'var. occidental on the
labels, and if these characters are constant in Cordillera Occidental populations,
then this population may warrant recognition as a subspecies.

56. Hypericum llanganaticum N. Robson, sp. nov.

H. struthiolifolio Juss. af finis, sed statura alta, ramis haud radicantibus, foliis planis
vel incurvatis, manifeste tetrastichis, plusminusve glaucis, sepalis oblongo-
oblanceolatis haud porcatis, differt. Type: Ecuador, Napo, Llanganati, paramo SE.
of Chosa Aucacocha, between Aucacocha and Pan de Azucar, 3800-3900 m,
15.V.1982 (fl), 0llgaard, Holm-Nielsen etal. 38464 (AAU!, holotype).

Shrub 0-4-2 m tall, erect to decumbent but not rooting, with branches strict,
pseudo-dichotomous or lateral. Stems orange-brown, 4-lined and ancipitous when
young, eventually terete, cortex exfoliating in irregular flakes; internodes 1-2 mm
long. Leaves sessile or rarely subsessile, erect to narrowly spreading, imbricate-
tetrastichous, deciduous above the base after fading; lamina 7-13 x 1-5-4 mm,
narrowly elliptic to linear-oblong, plane or slightly incurved, scarcely to markedly
cucullate, midrib impressed beneath, margin not or very narrowly hyaline, concol-
orous, rather glaucous, coriaceous; apex acute, base angustate, not sheathing,
pairs united to form rather shallow interfoliar ridge; basal vein 1, (always?) unbran-
ched; laminar glands dense, visible above and rarely also beneath; marginal glands
rather dense. Inflorescence 1-flowered, with pseudo-dichotomous branches from
node below; pedicel 3-4 mm long; upper leaves not transitional. Flowers c. 20-25
mm in diam., stellate. Sepals 7-5-9 x 1-8-2-3 mm, oblong-oblanceolate, acute, not
or slightly incurved-cucullate; veins 5, branched, not or scarcely prominent beneath;
glands striiform to punctiform. Petals (bright?) yellow, 12-16 x 4-10 mm, c. 1-5-1-8
x sepals, oblong-oblanceolate; apiculus acute; glands linear to striiform. Stamens c.
80-90, longest 6-10 mm long, 0-5-0-7 x petals. Ovary c. 3 x 2-5 mm, broadly ovoid;
styles 3, 3-6-4-3 mm long, 1-2-1-4 x ovary, divergent; stigmas not or narrowly
capitate. Capsule c. 5-5 x 3-5 mm, ovoid-pyramidal, shorter than sepals. Seeds not
seen.

In grassy paramo with scrub or reed forest; 3250-4200 m.

Ecuador (Tungurahua, Napo). Confined to the Cordillera de Los Llanganates. Map
23 (p. 96).

ECUADOR. Napo: Cordillera de Los Llanganati, Chihuila o Ainchilibi, 3750 m,
25-29.viii.1959 (fl), Barclay & Juajibioy 8996 (MO, NY); junction of Rio Chalupas
and Rio Agua Buenas, E. end of upper Chalupas valley, 3250 m, ll.viii.1979 (fl),
Holm-Nielsen 19074 (AAU). Tungurahua: Cordillera de Llanganates, near Las
Torres, 3700-3800 m, 22.xi.1939 (fl), Asplund 9878 (S); Llanganati Mts., Lake
Aucacocha, 3700 m, vii.1969 (st), Edwards 35 (K).

H. llanganaticum comprises a relict population intermediate in form and distribution
between H. ruscoides and H. struthiolifolium. Asplund 9878 differs from the other
specimens in its broader leaves and thus approaches H. ruscoides, differing from it

96

NORMAN K. B. ROBSON

Map 23 56. H. llanganaticum D; 57. H. struthiolifolium

essentially only in the narrower leaves with a slightly cucullate apex and narrower
stigmas. The remaining specimens are nearer to H. struthiolifolium but differ from it
in having more crowded tetrastichous leaves (which are dull blue-green or glaucous,
not lucent) and unribbed, oblanceolate to oblong sepals.

57. Hypericum struthiolifolium Juss.

in Annls Mus. Hist. nat. Paris 3: 161, t. 16, f.2 (1804) ['struthiolaefolium']; Choisy,
Prodr. monogr. fam. Hyperic. : 58 (1821), in DC. , Prodr. syst. nat. regni veg. 1: 553
(1824); R. Keller in Bull. Herb. Boissier II, 8: 176 (1908), in Engler & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 181 (1925). Type: Peru, without precise locality or date
(fl & fr), Dombey s.n. (P-JUS!, holotype; F!, P!, isotypes).

Brathys struthiolifolia (Juss.) Spach, Hist. nat. veg. Phan. 5: 449 (1836)
['struthiolaefolia'], in Annls Sci. nat. II, 5: 366 (1836).

H. struthiolifolium var. genuinum Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18:
293 (1862) ['struthiolaefolium'], pro parte excl. spec. cit.

H. weberbaueri R. Keller in Bot. Jahrb. 42: 129 (1908), in Bull. Herb. Boissier, II, 8:
178 (1908), in Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason
in Bull. Torrey hot. Club 56: 102, 103, 105 (1929), pro parte quoad typum. Type:
Peru, Junin, Tarma, Huacapostana, 3000-3100 m, n.d. (fl & fr), Weberbauer 2093
(Bf, holotype; F!, GH!, NY!, US!, photographs; G!, isotype).

Shrub up to c. 0-5 m tall, erect or sometimes decumbent and rooting, with branches
strict, pseudo-dichotomous or sometimes lateral. Stems orange-brown, 4-lined and
ancipitous when young, eventually terete, cortex exfoliating in irregular flakes;
internodes 2-7 mm long. Leaves sessile, suberect to spreading or slightly outcurving,
not or scarcely imbricate, deciduous above the base after fading; lamina 8-15 x
1-5-2-5 mm, very narrowly elliptic to linear-oblong, incurved- cucullate, midrib

THE GENUS HYPERICUM L.

mostly impressed beneath, margin not or slightly hyaline, concolorous, lucent, not
glaucous, coriaceous; apex acicular to subacute, base angulate, not sheathing,
pairs united to form shallow interfoliar ridge; basal vein 1, occasionally with 1-2
pairs of obscure branches; laminar glands rather dense, visible above but not
beneath. Inflorescence 1-flowered, with pseudo-dichotomous branches from node
below; pedicel 5-9 mm long; upper leaves not transitional. Flowers 15-25 mm in
diam., stellate. Sepals 5-8 x 1-3 mm, ovate-lanceolate to oblong, acute, incurved-
cucullate; veins 5-9, not branched, prominent beneath; glands linear, punctiform in
distal third. Petals bright yellow, (8-)10-15 x 3-7 mm, c. 1-5 x sepals, oblong-
oblanceolate to oblong-obovate; apiculus acute; glands striiform. Stamens 60-75,
longest 5-9 mm long, 0-5-0-6 x petals. Ovary 2-5-3-5 x 1-5-2 mm, broadly ovoid;
styles 3(4), 3-4 mm long, 1-3-1-5 x ovary, spreading; stigmas not or narrowly
capitate. Capsule 4-5 x 2-2-5 mm, ellipsoid, snorter than sepals. Seeds c. 0-7 mm
long, ecarinate; testa finely scalariform.

On open hillsides among shrubs; 3000-3700 m.

North and Central Peru (Amazonas, Pasco, Junin, Huancavelica, Cuzco). Map 23.
PERU. Amazonas: Chachapoyas, Pass of Piscohuanuna, 2700 m, viii.1938 (fl),
Sandeman 239 (K); Bagua, Cordillera Colan E. of La Peca, 2880 m, 8.viii.l978 (fl),
Barbour 3230 (MO). Cuzco: Urubamba, Puyupatamarca area, 3200 m, 29.iii.1894
(fl), Vargas 2762 (F). Huancavelica: Tayacaja, between Marcavalle and Huachocol-
pa, 3600 m, 24.iv.1963 (fl & fr), Rovar4225 (NY). Junin: Chilifruta to Huancayo, 11
km, 3700 m, 30.x. 1968 (fl), B. & C. Maguire 61657 (BM, NY). Pasco: Oxapampa,
Cordillera Yanachaga 12 km SE. of Oxapampa, 2700-2800, 7.x. 1982 (fl), Foster
9048 (BM, MO).

H. struthiolifolium was one of the first species in sect. Brathys to be described, and
the name has frequently been misapplied. The species is confined to north and
central Peru, where it is apparently relatively uncommon. It differs from H.
llanganaticum in the less crowded, more incurved, lucent (not glaucous) leaves and
the usually smaller flowers with sepals that are ribbed and usually broader below the
middle. Also, its decumbent stems sometimes root. Its distribution overlaps con-
siderably that of the derivative H. andinum, but these species remain distinct.

97

58. Hypericum andinum Gleason

in Torreya 29: 137 (1929). Type: Bolivia, Mt. Tunare, 1891 (fl & fr), Bang 1026 (NY!,
holotype; BM!, F!, GH!, K!, MO!, US!, W!, isotypes).

H. struthiolifolium var.parviflorum R. Keller in Bull. Herb. Boissierll, 8: 182 (1908)
['struthiolaefolium']. Type: Bolivia, La Paz, Larecaja, Majabaga, Catarguata,
2800-3200 m, X.1858-V.1859 (fl & fr), Mandon 790 (G!, holotype; BM!, GH!, K!,
NY!, P!,S!,W!, isotypes).

Shrub or shrublet 0-1-0-5 m tall, erect or decumbent and rooting, bushy with
branches strict, pseudo-dichotomous or lateral. Stems reddish to yellowish-brown,
4-angled when young, soon 2-lined then terete, cortex exfoliating in irregular flakes;
internodes 1-4 mm long. Leaves sessile, suberect to outcurved, subimbricate to
spreading, deciduous above the base after fading; lamina 3-5-10 x 0-8-1-5 mm,
narrowly oblong-elliptic to linear, incurved-cucullate, midrib mostly impressed
beneath, margin narrowly hyaline, otherwise concolorous, not glaucous, sub-
coriaceous to chartaceous; apex acute to rounded, base cuneate to parallel, not
sheathing, pairs united to form shallow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense, impressed, visible beneath. Inflorescence 1-flowered, with
pseudo-dichotomous branches from node below; pedicel 3-6 mm long; upper leaves
not transitional. Flowers 10-18 mm in diam., stellate. Sepals 3-5-7 x 0-8-2 mm,
broadly to narrowly lanceolate, subacuminate to acute, cucullate, with hyaline
margin; veins (3)5(7), unbranched, prominent; glands linear, punctiform, in distal
third. Petals golden yellow, 6-10 x 3-5 mm, c. 1-5 x sepals, narrow obovate to
obovate-oblong; apiculus acute to obtuse or obsolete; glands shortly striiform to
punctiform. Stamens 20-50, longest 3-6 mm long, c. 0-5 x petals. Ovary 1-8-2-3

98

J

NORMAN K. B. ROBSON

(-3-3) x 1-1-2 mm, narrowly ovoid; styles 3, 1-5-2-5 mm long, 0-6-0-7 x ovary,
free, spreading; stigmas broadly capitate. Capsule 4-6 x 1-2-3 mm, ovoid, shorter
than sepals. Seeds not seen.

On open or stony grassy slopes and Sphagnum bogs, sometimes near the snowline;
2400-4200 m.

Peru (La Libertad, Junin, Huancavelica, Cuzco, Puno), Bolivia (La Paz, Cochabam-
ba, Santa Cruz). Map 24.

PERU. Cuzco: Paucartambo Valley, 26 km from Cuzco to Pilcopata, 3425 m,
27.ix.1968 (fl), B. & C. Maguire 61564 (BM, NY). Huancavelica: Tayacaja, between
Colcabamba and Paucarbamba, Arriba de la Hda. Jocas, 3900 m, 20. iv. 1954, Tovar
2020 (NY). Junin: Huancayo, ii.1939 (fl), Soukup 389 (F). La Libertad: Huamachu-
co, Jalca de Huaguil, 3950 m, 24.vi.1958 (fl), Lopez & Sagdstegui 2736 (NY). Puno:
Ayapata, vi.1854 (fl & fr), Lechler 1817 (K, P, W).

BOLIVIA. Cochabamba: Road to Chapare [from Cochabamba], Km 104, 3100
m, 23.xii.1926, Steinbach 650 (F, MO, NY, S, U, US, WIS). La Paz: Sorata, 3900 m,
ii.1886 (fl & fr), Rusby 1350 (MICH, NY). Santa Cruz: Comarapa, Cerro de San
Mateo, 3400 m, 25.x. 1928 (fl & fr), Steinbach 8535 (BM, F, GH, K, MO, NY, S, U).

Although the area of H. andinum overlaps considerably that of H. struthiolifolium,
the former can be distinguished by the shorter capitate styles and the narrower,
usually less acute, recurving leaves with markedly hyaline margins.

59. Hypericum lancioides Cuatrec.

in Ciencia Mex. 4: 65 (1943). Type: Colombia, Putumayo, Paramo de San Antonio
del Bordoncillo, between El Encano and Sibundoy, 3250 m, 4. i. 1941, Cuatrecasas
11721 (COL, holotype; F!, US!, isotypes).

Shrub c. 0-1-1 m tall, erect and bushy or decumbent and caespitose, branches
strict, pseudo-dichotomous and lateral, or spreading (very rarely pinnate). Stems
orange-brown, 6-lined and compressed when young, the subfoliar ridges broad,
eventually terete, cortex splitting between and below subfoliar ridges; internodes
0-5-4(-6*) mm. Leaves sessile or rarely narrowing to a pseudopetiole up to 2 mm
long, ascending to subimbricate, sometimes tetrastichous, deciduous above base (or
pseudopetiole) after twisting spirally; lamina 7-15 x l-l-4(-4*) mm, very narrowly
elliptic-oblong to linear incurved to canaliculate or rarely subconduplicate,
markedly cucullate, midrib impressed beneath, margin distinct, narrowly to rather
broader hyaline, concolorous, lucent beneath, undulate-subpapillose above,
glaucous or not, coriaceous to somewhat succulent; apex acute, base angustate to
parallel-sided, not or scarcely sheathing, pairs united to form narrow interfoliar

Map 24 58. H. andinum A .
See note on p. 99.

THE GENUS HYPERICUM L.

ridge; basal vein 1, unbranched; laminar glands rather sparse, usually visible above
only but sometimes biseriate beneath. Inflorescence l(3)-flowered, with pseudo-
dichotomous branches from node below and sometimes short, often congested,
lateral branches (racemiform) ; pedicel absent or up to 4 mm long; upper leaves not
transitional. Flowers 10-16 mm in diam., stellate. Sepals (5-)6-8 x 0-5-1-5 mm,
oblanceolate-oblong to oblong or rarely linear, unequal to subequal, acute, cucul-
late, margin broadly hyaline; veins 3-5, unbranched or outer branched, all not or
slightly prominent; glands linear, punctiform in upper third. Petals yellow, 8-11 x
2-5-4 mm, c. 1-5 x sepals, narrowly obovate-oblong; apiculus acute to acuminate;
glands linear, distally punctiform. Stamens 25-40, longest 3-5-5 mm long, c. 0-5 x
petals. Ovary 1-5-2-5 x 0-8-1-5 mm, ellipsoid to subglobose; styles 3(4), l-2(-3*)
mm long, 0-5-0-9(-l-l*) x ovary, outcurved to suberect; stigmas broadly capitate.
Capsule 4-5 x 1-5-2-5 mm, ovoid-ellipsoid to cylindric-ellipsoid, shorter than
sepals. Seeds 0-8-1 mm long, ecarinate; testa finely scalariform.

In damp (sometimes marshy) or sheltered areas of the open paramo; 2870-4700 m.

Western Venezuela (Merida), Colombia (Santander to Narino) and northern
Ecuador (Carchi to Cotopaxi). Map 25.

H. lancioides differs from H. ruscoides especially in its narrower, 1 -veined, sessile
leaves, narrower sepals and shorter styles. One collection from Colombia, Putumayo
(Schultes & Villareal 7821), has relatively broad, petiolate leaves and longer styles
(see characters marked*), and Cuatrecasas 17861 (Valle, Los Farallones, cerro La
Torre) is even closer to H. ruscoides and could be of hybrid origin.

H. lancioides is variable and comprises two subspecies, one (subsp. lancioides)
with its most primitive form in southern Colombia and extending from there north to
Cundinamarca and south to Cotopaxi, has a simply pseudo-dichotomous inflores-
cence, whereas the other (subsp. congestiflorum) occurs from Cauca north-east to
Merida and has lateral flowers crowded below the terminal flower. Intermediate
specimens between the subspecies occur in Cauca.

99

59a. Hypericum lancioides Cuatrec. subsp. lancioides

H. struthiolifolium var. genuinum Triana & Planchon in Annls Sci. not. (Bot.) IV, 18:
293 (1862) ['struthiolaefolium'] pro parte, quoad spec. cit.

Map 25 59. H. lancioides a. subsp. lancioides A , b. subsp. congestiflorum A

100

NORMAN K. B. ROBSON

Fig. 14 A. H. lancioides subsp. lancioides: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) stamens (partly cut
away); (f) capsule. B. H. lancioides subsp. conges tiflorum: (g) habit. C. H. selaginella: (h) habit; (i) stem
with leaves; (j) leaf; (k) sepal; (b) petal; (m) stamens (partly cut away) and ovary; (n) capsule (a, g, h x
1/2; b-f x 3; i-m x 4). A. B. & C. Maguire 61808; B. Cuatrecasas et al. 28120; C. Barclay & Juajibioy

7407.

H. strictum sensu Gleason in Bull. Torrey hot. Club 56: 102 (1929) pro parte, quoad

spec. cit.
H. lancioides forma pygmaea Cuatrec. in sched.

Icon: Fig. 14 A.

Shrub 0-2-1 m tall, erect, bushy, with branches strict, pseudo-dichotomous and
lateral. Leaves shortly petiolate or sessile, usually spreading, sometimes tetra-
stichous, very narrowly elliptic or oblanceolate to linear. Inflorescence branches
pseudo-dichotomous with flowers solitary, terminal and sometimes on well-

THE GENUS HYPERICUM L. 101

developed branches from lower nodes. Sepals oblong to linear, with veins promin-
ent.

3000-4100 m.

Southern Colombia (Cundinamarca and Meta southwards) and Ecuador (Carchi to
Cotopaxi).

COLOMBIA. Cauca: Cordillera Central, Paramo de las Moras, 3800-3700 m,
12.xii.1969 (fl & fr), Cuatrecasas 27419 (BM, COL, US). Cundinamarca: Cordillera
Oriental S. of Usme, paramo de Chisaca above Laguna Grande and L. Negra,
9-ll.xi.1958 (fl), Barclay & Juajibioy 6191 (COL, MO, NY). Huila: Municipio de
La Plata, Inspection de Policia de Santa Leticia, Paramo de La Candelaria, 2370 m,
15.vii.1975 (fl), Diaz, Lozano & Torres 595 (COL). Narino: Province de Pasto,
Fuguerres, 300 m, v.1853 (fl), Triana 5464 (BM, COL, K, MEXU, NY, P, W);
Volcan El Galeras, c. 3200-3500 m, 4.vi.l946, Schultes & Villareall96Q (COL, F).
Putumayo: Pasto to Sibundoy, Paramo de San Antonio, 2790-2880 m, 13.iii.1953 (fl
& fr), Schultes & Cabrera 18828 (COL, GH, NY, U). Valle: Cordillera Occidental,
Los Farrallones, cerro de La Torre, La Laguna, 3500-3550 m, l.viii.1946 (fl),
Cuatrecasas 21870 (BM, F, P, U, US).

ECUADOR. Carchi: Tulcan to Maldonado, c. 43 km, SW. slopes of Volcan
Chilas, 3740 m, 6.viii.l976 (fl), 0llgaard & Balslev 8549 (AAU, BM). Cotopaxi:
Ruminahui, 3850 m, 17.ii.1947 (fl & fr), Aubert de La Rue s.n. (P). Imbabura: Lago
San Marcos, Cayambe, 3360 m, 26.xi.1961 (fl), Cazalet & Pennington 5349 (K).
Napos: Laguna Porjuantag, ENE. of Volcan Cayambe, 3750 m, 10. ii. 1953 (fl),
Prescott 347 (NY). Pichincha: Paso de Guamani, 35-37 km ESE. of Quito, head-
water of Rio Papallacta, 23.V.1947 (fl), Fosberg 27609 (NY, US); Cordillera Orien-
tal, environs of Laguna de Chiqui, 3800 m, 4.vi.l944 (fl & fr), Acosta Solis 8176 (F).

59b. Hypericum lancioides Cuatrec. subsp. congestiflorum (Triana & Plan-
chon) N. Robson, comb, et stat. nov.

H. struthiolifolium var. congestiflorum Triana & Planchon [ex Weddell, Chloris
Andina 2: 271 (1857) l struthiolaefolium\ nomen] inAnnls Sci. nat. (Bot.) IV, 18:
294 (1862) pro parte, excl. syn. ; Knuth in Reprium Spec. nov. Regni veg. Beih. 43:
484 (1927) pro parte, excl. spec. Funck & Schlim 1140. Type: Venezuela, Merida,
Sierra Nevada de Merida, 2870 m, 1842 (fl), Linden 432 (COL?, holotype; P!,
isotype).

Icon: Fig. 14B.

Shrub 0-1-0-5 m tall, erect and bushy or decumbent and caespitose with branches
strict, lateral (very rarely pinnate) , spreading. Leaves sessile, ascending to imbricate,
tetrastichous, linear. Inflorescence branches elongate, sympodial (from terminal
node) and very short, lateral, with flowers solitary or in triads in a congested
spiciform thyrse, immediately below terminal flower or with several intervening
sterile nodes. Sepals oblong-lanceolate, with veins not or slightly prominent.

2870-4500 m.

Western Venezuela (Merida), north-eastern and central Colombia (Santander to
Cauca).

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Plan de S.
Jose, 3625 m, 9.vi.l973 (fl), Cleef 10051 (U). Boyaca: Paramo de Chita, cabeceras
del Rio Casanare, Hoya Laguna Chorros Blancos, 3340 m, 24.V.1973 (fl), Cleef
9931A (COL, U). Caldas: Paramo del Quindio, 4100-4300 m, 15-20.viii.1922 (fl),
Pennell & Hazen 9874 (GH, K, S). Cauca: Paramo de Purace, 3500 m, 26.V.1944 (fl),
Killip & Lehmann 38548 (COL). Cundinamarca: Paramo de Palacio, c. 600 m al
NW. de Mina de Cal, 3520 m, 13.V.1972 (fl), Cleef 3733 (BM, COL, U); Lagunas de
Buitrago, 3555 m, 27.iv.1973, Cleef 9588A (COL). Meta: Paramo de Sumapaz,
Cerro Nevado de Sumapaz, 3615 m, 29.U972, Cleef 1269 (COL, U). Risaralda:
Municipio de Pereira, alredores de la Laguna Otun, 4000-4300 m, 3.ii.l980 (fl),
Diaz, Valencia & Jaromillo 1754 (COL). Santander: Paramo del Almorzadero,

102

NORMAN K. B. ROBSON

3500-3700 m, 20 vii.1940 (fl), Cuatrecasas & Garcia Barriga 9998 (COL, F). Tolima:
Paramo de Herves, Eresno, 3600 m, 14.vii.1939 (fl), Hanbury-Tracy 603 (K).

VENEZUELA. Merida: Paramo de Timotes, 3200 m, 25.U939 (fl), Alston 6616
(BM, NY); Quebrada de Saisay, 3880 m, 10.iv.1930, GehringerSl (NY, YEN).

In Venezuela there is a tendency in subsp. congestiflorum towards regular pyramidal
branching, shorter glandular-punctate leaves, and smaller flowers, sometimes with 4
styles. All these characters foreshadow H. selaginella.

60. Hypericum selaginella N. Robson, sp. nov.

H. lancioidei subsp. congestiflorae (Triana & Planchon) N. Robson affinis, sed
habitu nano prostratoque, ramificatione pinnato, foliis brevioribus dense imbricatis,
flores minoribus, stylis (3)4-5 brevioribus, differt. Type: Colombia, Boyaca, Cordil-
lera Oriental, Sierra Nevada del Cocuy, Alto Ritacuva, Station 16, c. 4400 m,
ll-29.iv.1959 (fl & fr), Barclay & Juajibioy 7407 (COL!, holotype; MO!, NY!,
isotypes).

H. selaginoides N. Robson in Cleef, Veg. Pdram. Colomb. Cord. Or.: 305 (1981),
nomen.

Icon: Fig. 14C.

Shrublet up to 0-15 m tall, prostrate or ascending, forming tufts up to 150 mm in
diam., with branches pinnate, spreading. Stems orange-brown, 6-lined and com-
pressed when young, the subfoliar ridges broad, eventually terete, cortex splitting
between and below subfoliar ridges; internodes c. 0-5 mm long. Leaves sessile,
densely to loosely imbricate, tetrastichous, deciduous above base or more usually
only with the cortex after fading; lamina 4-7 x 0-5-0-8 mm, linear, incurved-
canaliculate, cucullate, midrib impressed beneath, margin broadly hyaline, con-
colorous, sublucent beneath, sublucent to dull above, sometimes subglaucous,
coriaceous; apex acute to subacute, base parallel-sided, not sheathing, pairs united
to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense
above, rather sparse or absent beneath. Inflorescence 1-flowered, with pseudo-
dichotomous branches from node below and also short congested lateral branches
(racemiform) ; pedicel absent or up to 2 mm long; upper leaves not transitional.
Flowers c. 8 mm in diam., stellate. Sepals 3-4 x 0-8-1-2 mm, oblong to triangular-
lanceolate (equal), acute, cucullate, margin broadly hyaline; veins 3(5?), unbran-
ched, slightly prominent; glands mostly linear. Petals bright yellow, tinged red
outside, 4-5-6 x 2-2-5 mm, 1-5 x sepals, obovate-oblong; apiculus very short or
almost absent; glands linear, distally punctiform. Stamens c. 13, longest 3-4 mm
long, c. 0-65 x petals. Ovary 1-5-2-5 x 1-5-2 mm, subglobose; styles (3)4-5, 0-8-1
mm long, 0-4-0-5 x ovary, outcurved; stigmas broadly capitate. Capsule c. 3 x 3
mm, cylindric-globose to globose, shorter than sepals. Seeds 0-7-0-8 mm long,
ecarinate; testa finely scalariform.

In dry and stony or dampish paramo or superparamo; 3300-4340 m.

Colombia (Boyaca, Arauca, Cundinamarca), in the Sierra Nevada del Cocuy,
Paramo de Pisva and Macizo de Sumapaz. Map 22 (p. 94).

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Cabeceras de la Quebrada, El
Playon, Patio Bolos, 2-5 km S. of Alto La Plaza, 4340 m, 9.iii.l973 (fl & fr), Cleef
8944 A (COL). Boyaca: Sierra Nevada del Cocuy, Valle de los Corallitos, c. 4300 m,
31.vii.1957 (fl), Grubb, Curry & Fernandez-Perez 178 (COL, K, US); Paramo de
Pisva, Socha to La Punta Km 61.5, 6 km al NE. de Los Pinos, 3600 m, 10.vi.1972 (fl &
fr), Cleef 4328 (BM, COL, U). Cundinamarca: Macizo de Sumapaz, Cuchilla La
Rabona, 3900-3950 m, 17.vii.1981 (fl), Diaz etal. 2871 (COL).

In H. selaginella the reduction trends of H. lancioides are continued. The different
characters between it and H. lancioides subsp. congestiflorum (dwarf prostrate habit,
smaller and more densely imbricate leaves, smaller flowers, shorter and usually more
numerous styles) are clear-cut except for a few collections of the latter taxon from
Merida, which are somewhat intermediate in habit and size of parts.

THE GENUS HYPERICUM L.

103

61. Hypericum cymobrathys N. Robson, sp. nov.

H. terrae-firmae Sprague & Riley affinis, sed habitu humiliori, foliis brevioribus
crassioribus, interdum leviter cucullatis, supra basin deciduis, floribus minoribus in
cymes 3-12-floris dispositis, 3-stylis, inter alia differt. Type: Colombia, Boyaca, La
Uvita, by road from Chita, 3000-3050 m, 16.ix.1969 (fl), Cuatrecasas & Rodriguez
27805 (COL!, holotype; BM!, US!, isotypes).

Shrub 0-5-1 m tall, erect, with branches strict, pseudo-dichotomous or occasional-
ly lateral. Stems orange-brown to purplish-brown, 4-lined and markedly ancipitous
when young, soon 4-angled, cortex exfoliating in irregular flakes; internodes 2-5 mm
long. Leaves sessile, spreading from above base, tetrastichous, deciduous from
above base without fading; lamina 9-18 x 3-8 mm, oblong to elliptic, plane, slightly
cucullate, midrib prominent towards the base beneath, concolorous, glaucous,
coriaceous; apex acute to obtuse, base cuneate to angustate, sheathing, saccate,
pairs united to form narrow interfoliar ridge; basal veins 5-7, subparallel, apparently
unbranched, tertiary reticulum not visible; laminar glands dense, prominent to
impressed. Inflorescence 3-12-flowered, crowded (monochasial)?, with pseudo-
dichotomous branches from node below; peduncle and pedicels 3-5 mm long,
sometimes incrassate upwards; upper leaves intermediate, bracts narrowly elliptic-
oblong, acute. Flowers c. 15 mm in diam., stellate or cyathiform. Sepals 6-8 x
2-2-2-4 mm, narrowly lanceolate, acute; veins 5(7), unbranched, with midrib slightly
prominent; glands linear, distally striiform. Petals bright yellow, 8-9 x 3-5 mm, c.
1-2 x sepals, narrowly obovate; apiculus acute; glands linear, interrupted distally.
Stamens c. 100, longest 4-5 mm long, c. 0-5 x petals. Ovary 2x1-5 mm, broadly
ellipsoid; styles 3, 3 mm long, 1-5 x ovary, outcurving; stigmas narrow. Capsule c. 5
x 3 mm, broadly ellipsoid, shorter than sepals. Seeds 1-2-1-3 mm long, carinate;
testa finely scalariform.

Shrubby woodland on dry slopes; 2970-3500 (-3900?) m.

Colombia (Boyaca). Apparently confined to a small area of the Cordillera Oriental
around Guantiva and Chita. Map 11 (p. 54).

COLOMBIA. Boyaca: Socha to Los Pinos, Km 41, 2970 m, 22.V.1973 (fl), Cleef
9871 (BM, U); Paramo de Guantiva, 3000-3900 m, 23.vii.1952 (fl), Schneider 1225
(S); Municipio de Tola, Cerro los Arcos, alredores de Lago de Tota, 3050 m, i.1976
(fr), Aguirre & Rangel 391 (COL).

Although the cymose inflorescence of H. cymobrathys is reminiscent of H. mexica-
num, its flowers and leaves are very different. Its relationships appear to be with H.
terrae-firmae, although the much smaller flowers, 3-12-flowered inflorescence, and
smaller leaves make it superficially unlike it. Indeed, it appears to be an isolated
relict species at the base of a clade that includes H. chamaemyrtus, H. gnidioides, and
the H. gentianoides group, as well as H. denticulatum and its relatives (see part 8).

3. Acknowledgements

I am especially grateful to Dr Bassett Maguire (NY), who has helped my studies of tropical American
Hypericum in many ways; I have also received valuable assistance from Dr Antoine Cleef (U), Dr Jose
Cuatrecasas (US), Dr Richard Howard (A), Dr Alicia Lourteig (P), Dr James Luteyn (NY), Dr Benjamin
011gaard (AAU), Dr Peter Raven (MO), Dr Peter Stevens (A, GH), Dr Julian Steyermark (YEN), Dr
Robert Wilbur (DUKE), and John Ironside Wood (British Council, Bogota). My thanks are also due to
the directors of the following herbaria for the loan of specimens: Arnold Arboretum, Harvard University
(A); Herbarium Jutlandicum, Aarhus (AAU); Botanical Museum, Copenhagen (C); Institute de Ciencias
Naturales de la Universidad Nacional, Bogota (COL); Duke University, Durham, North Carolina
(DUKE); Botanical Museum Goteborg (GB); Gray Herbarium, Harvard University (GH); Royal Botanic
Gardens, Kew (K); Missouri Botanical Garden, St. Louis (MO); New York Botanical Garden (NY);
Museum National d Histoire Naturelle, Paris (P); Naturhistoriska Riksmuseet, Stockholm (S); Institute
for Systematic Botany, University of Utrecht (U); Institute Botanico, Caracas (YEN); Naturhistorisches
Museum, Vienna (W). I must also thank Miss Joan Malcolm for help with plotting distributions, and Dr
Richard Pohl (ISC) and Dr Chris Humphries (BM) for providing photographs.
With regard to the production of this paper, I am again deeply indebted to Mrs Margaret Tebbs for

104 NORMAN K. B. ROBSON

drawing the plates and figures; I am also very grateful to Miss Maria Duda for typing most of the
manuscript. Finally I would like to thank various colleagues for helpful discussions and suggestions, in
particular Arthur Chater, who commented on the typescript, and Bob Press for improving my marginal
illustrations.

4. References

Cleef, A. M. 1981. The vegetation of the paramos of the Colombian Cordillera Oriental. Thesis, University

of Utrecht.
Cuatrecasas, J. 1936. Resumen de mi actuation en Colombia con motive del II centenario del nacimiento

de Mutis. Trab. Mus. nac. Cienc. not., Madr. (Bot.) 33: 83-85.
Gleason, H. A. 1929. Hypericum, subsect. Eubrathys, in northwestern South America. Bull. Torrey hot.

Club 56: 100-107.
Howarth, M. K. 1981. Palaeogeography of the Mesozoic. In L. R. M. Cocks (Ed.), The evolving Earth:

197-220. London.

Kimura, Y. 1951. Hypericaceae. In T. Nakai & M. Honda, Nova flora japonica 10. Tokyo.
Lozano-C., G. & Schnetter, R. 1976. Estudios ecologicos en el Paramo de Cruz Verde, Colombia. II. Las

comunidades vegetales. Caldasia 11 (54): 53-68.
Melville, R. 1981. Vicarious plant distributions and the palaeography of the Pacific region. In G. Nelson &

D. E. Rosen (Eds), Vicariance biogeography: a critique. New York.
Owen, H. G. 1976. Continental displacement and expansion of the earth during the Mesozoic and

Cenozoic. Phil. Trans. R. Soc. A, 281: 223-292.
Raven, P. H. & Axelrod, D. J. 1974. Angiosperm biogeography and past continental movements. Ann.

Mo. hot. Gdn 61: 529-673.
Robson, N. K. B. 1977. Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric classification. Bull.

Br. Mus. nat. Hist. (Bot.) 5: 291-355.
1981 . Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus. Bull. Br. Mus. nat.

Hist. (Bot.) 8: 55-226.

1985. Studies in the genus Hypericum L. (Guttiferae). 3. Sections 1. Campylosporus to 6a.

Umbraculoides. Bull. Br. Mus. nat. Hist. (Bot.) 12: 163-325.
Rodriguez Jimenez, C. 1980. Hipericaceas. In R. Reitz, Flora illustrada catarinense. Itajai, Santa Catarina.
Rosen, D. E. 1976. A vicariance model of Caribbean biogeography. Syst. Zool. 24: 431-464.
1985. Geological hierarchies and biogeographical congruence in the Caribbean. Ann. Mo. hot. Gdn.

72: 636-659.
Standley, P. C. & Williams, L. 0. 1961. Flora of Guatemala, Guttiferae. Fieldiana Bot. 24 (7): 36-61.

5. Systematic index

Accepted names are in roman and synonyms in italic; new names are in bold, as are principal references.
An asterisk (*) denotes a figure.

Brathys acerosa (Kunth) Spach

(= 20) 47
acicularis (Kunth) Spach (= 35)

67
caracasana (Willd.) Spach

(= 51a) 89
juniperina (Kunth) Spach (= 41)

73

juniperina L. f. (= 41) 73
laricifolia (Juss.) Spach (= 20)

47
mutisiana (Kunth) Spach (= 26)

56
struthiolifolia (Juss.) Spach

(=57)96
thuyoides (Kunth) Spach (= 13)

37
Hypericum sect. 29. Brathys (Mutis

ex L.f.) Choisy 12

acerosum Kunth (= 20) 47
aciculare Kunth (35) 66*, 68*
iK-iist anum Steyerm. ex N.

Robson (5) 27*
andinum Gleason (58) 97, 98*
bolivaricum N. Robson (40) 72,

74*

brathysSm. (= 41)73
brathys sensu Trev. pro parte

(= 48b) 85
brathys var. juniperinum (Kunth)

Choisy (= 41) 73
bryoides Gleason (39) 71*
caracasanum Willd. (51) 89
caracasanum sensu Turcz. pro

parte (= 50) 87
caracasanum subsp. cardonae

(Cuatrec.) N. Robson (= 50)

87

caracasanum subsp. caracasanum

(51a)87*,89,91*
caracasanum subsp.

turumiquirense (Steyerm.) N.

Robson (51b) 87*, 90
caracasanum var. ocanense R.

Keller (= 31)60
caracasanum var. scherzeri (= 7)

30
caracasanum var. turumiquirense

Steyerm. (= 51b)90
cardonae Cuatrec. (50) 87*
carinosum R. Keller (9) 33, 37*
cassiopiforme N. Robson (29)

57*, 59
chamaemyrtus sensu Gleason

(=4)26
costaricense N. Robson (38) 70.

71*

THE GENUS HYPERICUM L.

105

cuatrecasii Gleason (16) 39*, 41
cymobrathys N. Robson (61)

54*, 103
decandrum Turcz. (30) 54*, 59,

61*
decorticans Planchon & Linden

(= 8)32
ekmanii A. H. Liogier (52) 90,

91*, 92*

ericifolium Steyerm. (= 4) 26
genistoides Kunth (= 13)37
garciae Pierce (4) 26, 27*
gleasonii N. Robson (25) 46*, 55
gnidioides var. polytrichoides R.

Keller (= 30)59
goyanesii Cuatrec. (14) 38, 39*
graciliforme N. Robson (= 48b)

85
gyropodioldes sensu Jackson &

Hooker (= 12)36
hartwegii Bentham (22) 46*, 51
hartwegii sensu Triana &

Planchon pro parte (= 19)45
hartwegii var. patens R. Keller

(= 13)37

hilaireanum (= 6) 28
holtonii Gleason (= 41) 73
horizontale N. Robson (46) 81,

82*

incurvum Urban (= 2a) 21
irazuense Kuntze ex N. Robson

(7) 30, 31*, 32*
jahniiR. Keller (= 41)73
jaramilloi N. Robson (49) 86, 87*
juniperinum Kunth (41) 72, 74*
lancifolium Gleason (45) 77*, 80
lancioides Cuatrec. (59) 98
lancioides sensu Steyerm. (=45)

80
lancioides subsp. congestiflorum

(Triana & Planchon) N.

Robson (59b) 99*, 100*, 101
lancioides subsp. lancioides (59a)

99*, 100*
lancioides forma pygmaea

Cuatrec. (59a) 100
laricifolium Juss. (20) 47, 48*,

49*
laricifolium sensu Bentham

(=35)67
laricifolium var. acerosum

(Kunth) Weddell (= 20) 47
laricifolium var. glaucum R.

Keller (= 48b) 85
laricoides Gleason (= 20) 47
littdeniiR. Keller (= 19)45
lindenii sensu Cuatrec. (= 15) 40
llanganaticum N. Robson (56)

95, 96*

loxense Bentham (18) 43
loxense subsp. aequatoriale (R.

Keller) N. Robson (18a) 44*
loxense var. aequatoriale R.

Keller (= 18a)44
loxense subsp. loxense (18b) 44*,

45

lycopodioides Triana & Plauchon

(12)36
magdalenicum N. Robson (32)

62*, 63
magniflorum Cuatrec. (24) 53*,

54*

maguirei N. Robson (23) 46*, 52
marahuacanum N. Robson (44)

77
marahuacanum subsp.

chimantaicum N . Robson (44c)

71*, 78*, 80
marahuacanum subsp.

marahuacanum (44a) 71*, 78*,

79
marahuacanum subsp.

strictissimum N. Robson (44b)

71*, 78*, 79
martense N. Robson (21) 48*,

49*, 51

meridense Steyerm. (= 8) 32
mexicanum L. (26) 56, 57*
? mexicanum sensu Choisy

(= 26) 56
mexicanum var. intermedium

Kuntze (=26) 56
millefolium Urban & Ekman (54)

91*, 92*, 93

mutisianum Kunth (=26) 56
myricariifolium Hieron. (15) 40,

44*
myricariifolium Hieron. ex R.

Keller (= 15)40
papillosum N. Robson (11) 24*,

34,35*
parallelum N. Robson (43) 76,

77*

phellos Gleason (3) 23, 24*
phellos subsp. oroqueanum N.

Robson (3b) 25
phellos subsp. phellos (3a) 23
phellos subsp. platyphyllum

(Gleason) N. Robson (3c) 25
pimelcoides Planchon & Linden

ex Triana & Planchon (31) 60,

62*

piriai (6) 28, 29*
platypetalum Turcz. (= 20) 47
platyphyllum Gleason (= 3c) 25
platyphyllum Gleason pro parte

(= 25)55

prietoi N. Robson (28) 57*, 58
prostratum Cuatrec. (42) 75, 77*
pseudobrathysTurcs. ( 41)73
pseudobrathys Turcs. pro parte

(= 48b) 85
pycnophyllum Urban (53) 91*,

92*
quitense R.Keller (17) 39*, 41,

42*

racemulosum Turcz. (= 20) 47
recurvum N. Robson (36) 68*
resinosum Bentham (= 26) 56
rimbachianum Diels (= 17)41
ruscoides Cuatrec. (55) 93,

94*

sabiniforme Trev. (19) 45, 46*
selaginella N. Robson (60) 94*,

100*, 102
selaginoides N . Robson ( = 60)

102
silenoides sensu R. Keller (= 7)

30
simonsii N. Robson (10) 24*, 34,

35*
spruce! N. Robson (34) 62*, 65,

66*

stenoclados Cuatrec. (= 9) 33
stenopetalum Turcz. (8) 31*, 32*
stenopetalum var. majus Triana

& Planchon (= 10) 34
strictum Kunth (48) 83
strictum sensu Gleason pro parte

(= 59a) 100
strictum subsp. compactum

(Triana & Planchon) N.

Robson (59b) 85*
strictum subsp. strictum (48a) 84,

85*
strictum var. compactum (Triana

& Planchon) Gleason (= 48b)

85
strictum var. reductum Gleason

(=41)73

struthiolifolium Juss. (57) 96*
struthiolifolium Juss. pro parte

(=35)67
struthiolifolium sensu Knuth

(= 45)80
struthiolifolium sensu R. Keller

pro parte ( 34) 65
struthiolifolium sensu Trev. pro

parte (= 9)33
struthiolifolium var. compactum

Triana & Planchon (= 48b) 85
struthliolifolium var.

congestiflorum Triana &

Planchon (= 59b) 101
struthiolifolium var.

congestiflorum sensu Kunth

pro parte (= 41)73
struthiolifolium var. genuinum

Triana & Planchon (= 57) 96
struthiolifolium var. genuinum

Triana & Planchon pro parte

(= 59a) 99
struthiolifolium var. gracile

Triana & Planchon (= 48b) 85
struthiolifolium var. minutum

Choisy (= 35)67
struthiolifolium var. parviflorum

R. Keller (= 58)97
struthiolifolium var. P sensu

Choisy pro parte (= 35) 67
struthiolifolum var. strictum

(Kunth) Choisy (= 48a) 84
stuebelii Hieron. (27) 54*, 58
stuebelii Hieron. ex R. Keller

(= 27) 58

styphelioides A. Richard (2) 20
styphelioides sensu Standley &

Williams (= 1) 18

106

NORMAN K. B. ROBSON

styphelioides subsp. clarense

Lippold (2a) 21*
styphelioides subsp. moaense

Lippold (2c) 21*, 22
styphelioides subsp.

styphelioides (2b) 21*
tachirense Steyerm. (= 3a) 23
tamanum Cuatrec. (= 3a) 23
tenuifolium St.-Hil. (= 6) 28

terrae-firmae Sprague & Riley

(1)18, 19*, 21*

tetrastichum Cuatrec. (47) 82*
thuyoides Kunth (13) 37*
thymifolium Cuatrec. (= 3a) 23
thymifolium sensu Planchon &

Triana pro parte (= 9) 33
thymifolium sensu Turcz. (= 19)

45

trianae N. Robson (= 19) 45
valleanum N. Robson (33) 62*,

64,66*

weberbaueri R. Keller ( = 57) 96
weberbaueri sensu Gleason

(= 45) 80
wurdackii N. Robson (37) 62*,

69

Studies in the genus Hypericum L. (Guttiferae)

N. K. B. Robson

When completed in nine parts, this series of papers will constitute a monograph of
Hypericum, St John's wort. This ancient genus, with an almost world-wide distribu-
tion, is of great scientific interest, as well as of considerable importance in horticul-
ture and, to a lesser extent, pharmacy. Parts 1 (1977) and 2 (1981) are introductory,
and the detailed taxonomy begins with part 3 (1985), in which the first seven sections
(sects l-6a) are treated. The present paper (part 7) contains the majority of sect. 29,
the remainder of which will be included in part 8, along with accounts of sects 30 and
31. Subsequent papers (parts 4-6) will contain treatments of sects 7-28, and the
concluding paper (part 9) will contain addenda, corrigenda, a revised sectional key
and enumeration, and a continuous numerical series of species.

Bulletin British Museum (Natural History): Botany Series

Vol. 5 No. 6 Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric
classification. N. K. B. Robson. 1977. Pp. 291-355, 9 figs. 7.25

Vol. 8 No. 2 Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the
genus. N. K. B. Robson, 1981. Pp. 52-226, 73 figs, 8 tables. 23.50

Vol. 12 No. 4 Studies in the genus Hypericum L. (Guttiferae). 3. Sections 1.
Campylosporus to 6a. Umbraculoides . N. K. B. Robson. 1985. Pp. 163-325, 24
figs, 34 maps. 565 08003 2. 28.80

Titles to be published in Volume 16

Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)

ByN. K. B.Robson

The lichen genus Ramalina in Australia
By G. N. Stevens

An annotated list of vascular plants collected in the valleys south of
Mt Everest

By G. Miehe

An illustrated catalogue of type specimens in the Greville diatom herbarium

By D.M.Williams

Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester

Bulletin of the

British Museum (Natural History)

The lichen genus Ramalina in Australia
G. Nell Stevens

Botany series Vol 16 No 2 25 June 1987

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

Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum's resources. Many of the papers are
works of reference that will remain indispensable for years to come.

Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW75BD,
England.

World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)

British Museum (Natural History), 1987

The Botany series is edited in the Museum's Department of Botany
Keeper of Botany: Mr J. F. M. Cannon

Editor of Bulletin: Mr J. R. Laundon

Assistant Editor: Dr A. J. Harrington

Editor's Assistant: Miss M. J. Short

ISBN 565 07016 9
ISSN 0068-2292

British Museum (Natural His
Cromwell Road
London SW7 5BD

^mg*^ #

Botany series

Vol 16 No 2 pp 107-223

Issued 25 June 1987

The lichen genus Ramalina in Australia

PRESE
G. Nell Stevens

Department of Botany, University of Queensland, St Lucia, 4067, Australia

Contents

Synopsis 108

Introduction 108

The Australian environment 110

Historical understanding of Ramalina and Ramalinaceae 115

Collectors of Australian Ramalinae 117

Materials and methods 120

Results 121

Morphology 121

Anatomy 126

Chemistry 132

Correlation between secondary-product chemistry and ecogeography

in the Ramalina subfraxinea complex 134

Distribution patterns in relation to climatic indices 135

Biogeography 140

Palaeobiogeography 142

Taxonomy 147

Taxonomic ranks 147

Hybridization 148

Description of the genus Ramalina 149

Enumeration of the Australian taxa 150

Key to Ramalina in Australia 150

1 . R. aUstraliemis 152

2. R. caespitella 155

3. R. canariensis 156

4. R.exiguella 158

5. R.filicaulis 160

6. R.fimbriata 161

7. R. glaucescens 163

8. R.litorea 169

9. R. peruviana 170

10. R.reducta 175

11. R.tenella 176

12. R. unilateralis 177

13. R. whinrayi 179

14. R.celastri 180

15. R.inflata 185

16. R.fissa 193

17. R.nervulosa 195

18. R.pacifica 201

19. R. subfraxinea 203

20. R.tropica 210

Appendix. Disposition of exluded Ramalina taxa 217

Acknowledgements 217

References 218

Index .. 222

Bull. Br. Mus. not. Hist. (Bot.) 16 (2): 107-223 Issued 25 June 1987

108 G. NELL STEVENS

Synopsis

This paper presents a revision of the lichen genus Ramalina in Australia. Of the 60 Ramalina taxa
previously listed for Australia in catalogues only nine of those names are retained, six with unchanged
status. Twenty-eight taxa belonging to 20 species are recognized for Australia. Two are new species: R.
filicaulis and R. tropica. A new variety R. subfraxinea var. norstictica and a new subspecies R. inflata subsp.
australis are described, and four names, R. celastri subsp. ovalis, R. inflata subsp. per pus ilia, R. subfraxinea
var. confirmata, and R. subfraxinea var. leiodea are new combinations.

The taxonomically important features of this genus (viz. morphology, anatomy, and chemistry) are
discussed. In delimiting taxa, the morphological plasticity shown by this genus has been taken into
consideration so that a wide range in infraspecific variation has been accepted for some of the taxa. The
distribution patterns of the Australian Ramalinae are shown to closely follow the distribution of Australian
phanerogam groups, which are controlled by the environmental parameters of rainfall and temperature;
this enables the Ramalinae to be conveniently divided into three response groups (viz. megatherm,
mesotherm, and micro therm) as used to divide the Australian phanerogams. On the basis of species
diversity it is hypothesised that the genus had its origin in west Gondwanaland (South America and Africa)
and then spread to the east. It is suggested that subsequent speciation occurred within Australia.

Introduction

The genus Ramalina has been studied in the northern hemisphere for over two hundred years
and several hundred taxa have been named. In the southern hemisphere very little research has
been undertaken and no revisional work had been carried out on the Australian Ramalinae
before the present study. During the course of previous research into the taxonomy and ecology
of twenty-four lichen genera found on eastern Australian mangroves (Stevens, 1978) it became
apparent how necessary it was for a revision of the Australian representatives of the genus to be
undertaken.

This present work provides a taxonomic revision of the Australian Ramalinae as well as an
ecological survey and a biogeographic assessment of the genus. Research into the Australian
taxa in this genus also contributes to the overall understanding of the biogeography of allied
Ramalina species which occur in the other Gondwanaland regions of Africa, India, and South
America.

The genus is regarded as a difficult one and its morphological plasticity is renowned. Howe
(1913-14) aptly summed up the dilemma when he stated 'there is little difficulty in recognizing
the characteristics of the genus Ramalina. The species on the other hand, have presented a most
difficult problem, caused largely no doubt by the innumerable intergrades that present them-
selves, and by the unwarranted description of new species'.

This problem was encountered by the author when confronted by the record of 60 different
Ramalina taxa reported for Australia (Wetmore, 1963; Weber & Wetmore, 1972) in catalogues
which listed all taxa recorded for Australia in papers published from 1804 to 1972. It was
apparent quite early in the study that many of these names were incorrect; some were synonyms,
others misapplication of the names of northern hemisphere species which they superficially
resembled.

Other recent mention of Australian Ramalinae occurs in an unpublished key to the genus
Ramalina by Dahl (1971) and in Filson & Rogers (1979), but in both of these accounts little
preliminary research had been carried out before naming the taxa. Most of the existing literature
summarizing Australian material was therefore of little help in the present study.

There are few modern publications recording research which include the southern hemi-
sphere Ramalinae. Krog & Swinscow (1974, 1975, 1976) have published detailed work on the
East African Ramalinae. Rundel (1978a) has dealt with the Ramalina usnea complex in North
and South America and in a paper on the lichens of Tristan da Cunha, J0rgensen (1977)
discussed six species of Ramalina. No other publications discuss this genus in detail; however, a
series of papers by Osorio published from 1967 to 1983 list the lichens found in Uruguay,
Paraguay, Brazil, and Argentina and these include some species of Ramalina. An account of the
lichens of New Zealand (Galloway, 1985) which includes the genus Ramalina lists only nine
species.

RAMALINA IN AUSTRALIA

109

During the present research four months were spent in Europe and U.S.A. visiting herbaria
(BM, FH, H, HAW, O, S, UPS and US) and becoming familiar with northern hemisphere
Ramalina species. This enabled an insight to be gained into the reason why Australian species
were often given the names of the northern hemisphere species. The type material of species
which occur in other southern continents was also examined where possible.

In this study the approach has been to reinforce the taxonomic revision of the genus by
incorporating ecological data in order to explain phenotypic variation, and to enable the
delimitation of the taxa to be made more confidently. The extreme morphological variation
within the species was a major problem. The extent of such variation in Australian material had
not previously been gauged, nor was there any conception of the evolutionary relationships that
might exist between the different taxa.

Many thousands of specimens therefore needed to be examined before any decisions could be
made on the delimitation of species. Examination of too few specimens makes it impossible to
understand the breadth of morphological plasticity exhibited by a single species, and the early
taxonomy of Australian Ramalinae was based on single or at most several thalli which were
usually collected by someone other than the taxonomist.

Collection of Ramalina material by the author began in 1975, when the lichens on mangroves
were studied (Stevens, 1978). Since then collecting trips have covered parts of New South
Wales, Victoria, Tasmania, Western Australia, and Northe i Territory and innumerable
collecting forays were made in many areas of Queensland. After several expeditions inland it
was found that the genus Ramalina was restricted to a zone varying within 100-200 km of the
coastline, thus eliminating the need to collect in the arid interior (Fig. 1).

Observation of the different ecological habitats occupied by any one species was made to
enable an appreciation of the variation that occurs in the morphology of a species under different
environmental pressures. As well as sampling different habitats in the one region, thalli were

Fig. 1 Ramalina distribution in Australia.

110 G. NELL STEVENS

collected (wherever possible) from latitude to latitude in order to assess if a continuum existed
between the different morphologies found over a wide geographical range.

To a large extent this research has been carried out on personal collections of fresh material,
supplemented by collections made by present day Australian lichenologists, material available
from herbaria in each of the Australian states, and to a far lesser extent on old collections held in
overseas herbaria. The latter proved to be of little use as only vague information concerning the
location of the collection was ever recorded, e.g. New Holland. In contrast the Australian
material was well documented on the whole but the specimens remained unnamed or bore
doubtful identifications.

All of the Australian types were examined and, where possible, compared with fresh material
from the type location. In this way the amount of variation that occurred within the particular
species could be assessed. It was found in some instances that the type represented one extreme
in a large population of morphotypes.

An attempt has been made to show possible evolutionary relationships between some of the
taxa, based on morphological and/or chemical evidence associated with geographical patterns of
distribution, as suggested by Imshaug & Brodo (1966).

This revision has treated all Australian material as belonging to the genus Ramalina Ach. , as
observations during the present study did not substantiate separation into the two genera
Ramalina Ach. and Fistulariella Bowler & Rundel. No species belonging to any other divisions
within the Ramalinaceae (Bowler, 1981) occur in Australia. A representative collection has
been deposited in BM.

The Australian environment

Although it is beyond the scope of this study to provide a detailed discussion of the geography,
climate, botany, and geology of Australia, a general outline is presented to enable a better
appreciation of the factors which are influencing Ramalina distribution (Fig. 1).

(1) Geography

The area of Australia is approximately 8,700,000 km 2 , larger than all of western Europe, and its
immense coastline measures some 36,700 km. In total it comprises the Australian mainland, the
large island of Tasmania, and numerous small islands around its coastline. It is washed by three
oceans, the Pacific, Southern, and Indian, and four seas, the Tasman, Timor, Coral, and
Arafura. It extends from approximately 930'S (Dauan Island in Torres Strait) to 43S (southern
Tasmania) and includes Macquarie Island at 55S; its climate varies from equatorial through
subtropical in the north to warm temperate, cool temperate to sub-antarctic in the south.

Overall the landmass is low: exceeding 1000 m in only very limited areas, most of these being
close to the eastern coastline, predominantly in a north-south alignment through Queensland,
New South Wales, and Victoria. This continuation of the Eastern Highlands and Great Dividing
Range from north to south creates a fairly uniform topography which is conducive to species
dispersal over a wide geographical area without creating any substantial barrier to the distri-
bution, which could lead to endemism.

In the south this mountain range turns west through the Victorian Alps to the Grampians of
western Victoria. Farther south the highland chain is apparent as islands in Bass Strait and in
Tasmania. During times of low sea-level in the Pleistocene glacial period, these islands and
Tasmania were joined to the mainland. Two endemic species have evolved, probably as a result
of isolation of these islands from the continent. Several well separated parts of the mountain
chain have individual mountains which exceed 1500 m elevation. These are found in Tasmania,
the Victorian Alps, the Snowy Mountains, the New England Tableland in New South Wales,
and Bellenden Ker Range in Queensland.

The majority of the western and southern Australian land mass is a vast flat peneplain with an
elevation between 200 m and 500 m. In the north-west area the highest peak in the Hamersley
Ranges exceeds 1200 m in height, and in central Australia the Macdonnell and Musgrave Ranges
reach a similar height. Much of the interior of Australia is arid sand-ridge desert; the arid zone

RAMALINA IN AUSTRALIA 111

extends to the western and southern coasts between the moist south-west corner of Western
Australia and the more fertile south-east of Australia. Extensive salt lakes occur in the south and
south-western regions of the arid interior.

(2) Climate

The general climatic characteristics of Australia are dictated by its latitudinal position lying
astride the mid-latitude high pressure belt. The prevailing wind patterns and general aridity over
continental Australia reflect the dominating control of eastward-moving anticyclonic cells which
track between 37 -38S in summer and between 29-32S in winter. Seasonal change is linked
with these sun-controlled shifts in the paths of the anticyclonic cells and associated movements
of the south-easterlies and the inter-tropical convergence zone which influence the climate of
eastern and northern Australia, and the westerlies which influence the climate of southern
Australia and Tasmania (Nix, 1981).

Summer rainfall predominates in the north, winter rainfall in the south, and along the eastern
coast a more uniform distribution of rainfall is common (Figs 2A & B). The eastern coastline of
Australia receives a substantial rainfall, which ranges from over 2000 mm in north Queensland
to 741 mm in Westernport Bay, Victoria. The western coastline receives far less precipitation
per annum, partly due to the lack of any mountain ranges close to the coast. The annual
precipitation ranges between 230 mm and 840 mm per annum with the highest recordings in the
southwest corner where winter rains occur. Winter rainfall is predominant as far north as 24S,
then summer rainfall becomes dominant in the monsoonal area.

More than 80% of the continent has at least three months each year which are without
effective precipitation (Nix, 1981). Two-thirds of Australia has less than 500 mm rainfall per
annum and one-third less than 250 mm. Most of the coastline represents a moist rim around an
otherwise dry continent; however parts of this coastline in South and Western Australia are also
extremely arid, and no Ramalinae occur there. The absence of Ramalinae in the arid inland is
possibly due to a combination of inhibiting factors such as high summer temperatures, low
rainfall, and lack of a suitable substrate.

In tropical Australia the highest relative humidity occurs during the rainy summer season
while in the temperate region relative humidity is generally highest in winter and lowest during
the summer. The relative humidity variation during the day closely follows the diurnal variation
of temperature, being highest with low temperatures and lowest with high temperatures.

Mean minimum temperatures occur in July throughout Australia. Freezing temperatures and
frosts occur regularly below the 5C isotherm and the frost-free limit coincides approximately
with the 10C isotherm. The ameliorating influence of the surrounding oceans is evident in the
coastal areas of southern Australia and Tasmania. Rainfall is the dominant mode of precipi-
tation, with sleet and snow confined to the higher mountains of southeastern Australia on the
mainland and in Tasmania.

The greater part of the Australian land mass receives over 3000 hours of sunshine per year
while the east coast is reported to receive between 2500 and 3000 hours, indicating that
cloudiness is higher near the coast.

The Queensland coast as far south as Mackay and the Western Australian coast as far south as
Carnarvon are subject to tropical cyclones, normally between December and March, which
cause very heavy rainfall and accompanying strong winds. Sea surface temperatures around
Australia range from 15-30C in summer to 12-26C in winter.

(3) Botany

Specht (1970) arbitrarily divided the Australian vegetation into structural formations based on
foliage cover and life form of the upper stratum (Table 1). The structure of the community and
the species therein reflect the amount of moisture available to the plants e.g. tall dense plant
communities are found in high rainfall areas whereas more stunted open communities occur in
drier regions. Edaphic factors also influence the eventual structural formation.

The majority of Ramalina collection locations lie in those plant communities listed on the top
left hand side of Table 1 , representing those which occur in relatively high rainfall areas and have

112

G. NELL STEVENS

Fig. 2 A. Summer seasonal rainfall (November-April) equalled or exceeded 8 years in 10; B. Winter
seasonal rainfall (May-October) equalled or exceeded 8 years in 10 (adapted from Nix, 1981).

RAMALINA IN AUSTRALIA

Table 1 Structural formations in Australian plant communities (modified from Specht,

113

1970).

Foliage projective cover of tallest stratum

Life form of

tallest stratum

100-70%

70-50%

50-30%

30-10%

<10%

Trees >30 m

Tall ctosed-

Tall

Trees 10-30 m

forest
Closed-
forest

open-forest
Open-forest

Open-forest

Woodland

Open-
woodland

Trees < 10m

Low closed-
forest

Low
open-forest

Low
open-forest

Low
woodland

Low open-
woodland

Shrubs >2 m

Closed
scrub

Open-scrub

Open-scrub

Tall
shrubland

Tall open-
shrubland

Shrubs 0-25-2 m

Heathy

Closed-
heathland

Open-
heathland

Open-
heathland

Low
shrubland

Low open- -
shrubland

Chenopodiaceous

Low
shrubland

Low
shrubland

Low open-
shrubland

Shrubs <0-25 m

Dwarf open-
heathland

Dwarf open-
heathland

Hummock grasses

Hummock
grassland

Open hummock
grassland

a relatively high Foliage Projective Cover (FPC) e.g. (1) Closed-forest, (2) Tall open-forest, (3)
Open-forest, and (4) Open-scrub. These structural formations are made up of overstorey and
understorey vegetation, and depending on the amount of canopy cover, vary in FPC from 100%
to 30%. They can be briefly described as follows:

(a) Closed-forest (also referred to as rain-forest, it can also describe some mangrove forests).
This community comprises many different tree genera and species which alter in composition
with change in latitude. It is found in disjunct pockets from Cape York to Tasmania and grades
from tropical mesophyll vine forests (with leaf size 45-180 cm 2 ) to subtropical mesophyll and
notophyll vine forest (with leaf size 20-25-45 cm 2 ) to warm temperate notophyll vine forest to
cool temperate microphyll moss forest (with leaf size 2-25-20-25 cm 2 ).

Closed-forest communities occur in humid areas where rainfall exceeds 1500 mm per annum;
overstorey FPC in such a formation approaches 100%. These formations reach their best
development on the basaltic soils of the Eastern Highlands. Several species of Ramalina grow in
this habitat, two species are specific to the rain-forest canopy, and another three species are
found in both closed-forest and open-forest communities.

These closed-forests can contain trees which exceed 30m in height, but in monsoonal
northern Australia and coastal Queensland, low closed-forests occur which are composed of
trees reaching less than 10 m in height. During the wet season these forests have a FPC of 100% ,
but during the dry winters the FPC may be as low as 70% as some of the trees are semi-
deciduous. In inland Queensland the low closed-forest is the only formation which supports any
Ramalina species.

(b) Tall open-forest (with sclerophyllous understorey with or without tree ferns) occurs in
sub-humid sites of eastern, south-eastern and south-western Australia. Overstorey FPC in such
a community varies from 70% to 50%. The dominant tree species in the overstorey is
Eucalyptus, whereas the low tree/tall shrub stratum (less than 10 m high) contains 18 genera of
angiosperms (Specht, 1981a). Several species of Ramalina have been found growing on twigs in
the understorey, but rarely do species of Ramalina grow on eucalypts.

(c) Open-forest (with either heathy sclerophyll or grassy understorey). This formation is the
most widespread in Australia, occurring in the north, east, and southern parts of the continent;
each area experiences quite different rainfall patterns ranging from summer rains to summer-
winter uniform rainfall, to winter rains respectively. The genus Eucalyptus dominates these

114 G. NELL STEVENS

communities, although species of Casuarina and Acacia are also plentiful. The overstorey FPC
for open-forest communities varies between 70-30% . No Ramalina species grow in the tropical
open-forests, but occur on Casuarina and Acacia in the temperate communities,
(d) Mallee open-scrub (with savannah-chenopod understorey) . This structural formation is
common in the drier Mediterranean climate of southern Australia where species of Eucalypt
dominate, but other genera, including Acacia are scattered between the mallee eucalypts.
Overstorey FPC in these areas can be as low as 30%. The relatively low rainfall (250-800 mm
p. a.) occurs in winter, followed by hot dry summers. Four species of Ramalina have been
collected in this formation.

Serai plant communities exist along the coastal fringe of Australia; these are coastal dune
communities and saline coastal communities, the latter made up of salt-marsh, mangrove
vegetation, and marine meadow (Specht, 19810); only the coastal dune community and the
mangroves act as phorophytes for Ramalina species. Such communities usually produce a series
of structural formations, influenced by micro-habitat conditions; the overstorey FPC therefore
differs in different parts of the community.

(A) Coastal dune community: In the tropics, lowland rain-forest abuts the open coastline in
areas where mangroves are not dominant; in the subtropics the sand dune vegetation comprises
various shrub species ofBanksia integrifolia, Callitris columellaris , and Casuarina equisitifolia as
the dominant tree species. Along the southern coast-line, the foredunes are vegetated by heathy
shrubs; Pimelea serpyllifolia, Leptospermum laevigatum, Leucopogon parviflora, Monotoca
elliptica, Bursaria spinosa, and Acacia sophorae. Several Ramalina species occur in this
community, the species varying with change in latitude and longitude.

(B) The mangroves (saline coastal community): This vegetation occupies the muddy areas at
the mouths of rivers and bays along the eastern, north-western, and northern coastlines of
Australia, with pockets in South Australia. The structure of such communities varies from
closed-forest to woodland to tall shrubland. The major concentration of mangrove vegetation
(approx. 28 species) is confined to the tropics gradually dwindling in species numbers in the
subtropics and eventually being reduced to one species, Avicennia marina, towards the
temperate zone. Both FPC and the amount of rainfall vary considerably depending on the
latitude of the mangrove community.

Those mangrove species which act as phorophytes to the Ramalinae are Rhizophora stylosa,
Rhizophora apiculata, Ceriops tagal, Bruguiera gymnorhiza, Avicennia marina, Aegiceras
corniculatum, Lumnitzera racemosa, and Excoecaria agallocha.

(4) Geology

The saxicolous species of Ramalina are totally dependent on the presence of rock outcrops

within their distribution range. Corticolous species, however, are only indirectly influenced

through the presence or absence of particular plant communities which grow on the different soil

types.

The four main rock types involved in the areas where Ramalina species occur are (1) basaltic
rocks (2) metamorphics and fine grained sedimentary rocks (3) sandstones and (4) granites.

The major areas of basalt are Tertiary in age, and are confined to the Eastern Highlands. Most
of these elevated areas are forested with either rain-forest or tall open-forest. Additional areas
of basalt of Quaternary age are found in north-east and central-east Queensland, supporting low
closed-forests, and western Victoria at lower altitudes on the coastal plain which supports
extensive areas of open-woodland and grassland.

In eastern Australia the metamorphic rocks and many fine-grained, compacted sedimentary
rocks which are associated with them are of Palaeozoic age, and have been folded and elevated,
forming mountain ranges and hilly country in the Eastern Highlands. They have been intruded
by granite batholiths, which also make up large areas, having been revealed by prolonged
erosion e.g. New England, Murrumbidgee, Kosciusko, and many other masses in the eastern
states. These are areas of high elevation subject to cold winters.

The sandstones which are most important are those sub-horizontal deposits of Mesozoic age

RAMALINA IN AUSTRALIA 115

forming for example, much of the surface of the Sydney Basin, Moreton Basin, and the
Carnarvon Range. The Grampians are composed of dipping sandstones of Palaeozoic age.

In South Australia and Western Australia only a few species of Ramalina occur on trees
growing on soils derived from old granites and metamorphic rocks (schists and quartzites).
There are very few occurrences of Ramalinae in areas of calcareous rocks of the Tertiary basins
and Pleistocene dune deposits.

Historical understanding of Ramalina and Ramalinaceae

(1) THE GENUS RAMALINA

The genus Ramalina was first described by Acharius (1810) when he distinguished it from

Parmelia by its cartilaginous thallus and apothecial characters.

Acharius (1810) listed ten species (together with their forms) as belonging to the genus,
namely Ramalina homalea, R. linearis, R. complanata, R. peruviana, R. polymorpha, R.
fraxinea, R. fastigiata, R. scopulorum, R. farinacea, and R. pollinaria. Except for the first, all of
these species have been retained in the genus to the present day.

Many more taxa have been added to this original number in the last 175 years, some being
removed later into separate genera because of differences found in the anatomy, the spore shape
and size, or the colour of pycnidia. Montagne (1852) examined the cortex of R. homalea Ach.
and found that the hyphae were arranged perpendicular to the surface without any mechanical
support tissue in the cortex; he therefore erected a new genus Desmazieria Mont, to incorporate
all Ramalina species possessing this anatomical feature, using R. homalea as the type.

Massalongo (1854) erected the genus Cenozosia Massal. based on R. inanis Mont, as the type
and characterized by its fistulose spongy thallus with a single layer of branching cells in the
cortex. This monotypic genus has been retained as a distinct taxon up to the present.

By using the colour of spermatia, as well as anatomical and thallus differences, Stizenberger
(1862) divided the genus Ramalina into three parts (i) Cenozosia (Massal.) (type R. inanis Mont.)
as a species with thallus hollow, spermatia black (ii) Desmazieria (Mont.) (type R. ceruchis Ach.)
for those species with simple cortex, thallus interior cottony and spermatia black and (iii)
Euramalina Stizenb. (type R. scopulorum Retz.) for species having a double cortex, thallus
interior cottony, and spermatia pale or colourless.

Nylander (1870) also used colour of spermogonia conceptacles for initial division within the
genus, separating species with (1) totally black spermatia (2) partially black or (3) pale or
colourless spermatia. Within group (1) he then used differences in anatomical structure of the
cortex to separate R. inanis Mont. (= Cenozosia Massal.) from the rest. He placed five other
species in another group which corresponds to Desmazieria Mont. , namely R. ceruchis Ach. , R.
combeiodes Nyl. , R. homalea Ach. , R. flaccescens Nyl. , and R. testudinaria Nyl.

Ramalina melanothrix Laurer was separated from the rest because of its two-layered cortex
(= Euramalina Stizenb.); its present position is in the genus Trichoramalina Rundel & Bowler.
Group (2) contained only R. carpathica Korber, a species which Bowler & Rundel (1977) placed
in the genus Fistulariella Bowler & Rundel. Group (3) contained 56 species which today belong
in three genera, Ramalina Ach. , Niebla Rundel & Bowler, and Fistulariella Bowler & Rundel.

Vainio (1890) also laid stress on anatomical structure of the cortex, accepting section
Euramalina Stizenb. as having a chondroid cortex made up of longitudinal hyphae, and sections
Desmazieria (Mont.) and Cenozosia (Massal.) as having a cortex made up of transverse hyphae.
Vainio divided Euramalina Stizenb. into two stirps based on thallus structure (1) Fistularia
Vainio species with inflated hollow branches and (2) Myelopoea Vainio species with a con-
tinuous arachnoid or cottony medulla. He used branch shape to subdivide stirps Myelopoea into
series Teretiusculae Vainio (containing species with terete or angular branches) and series
Compressiusculae Vainio (containing species with flattened branches rarely two-edged).

Hue (1901) followed Vainio in this division of Ramalina when he divided the genus into three
groups on morphology: (A) Teretiusculae Vainio (B) Compressiusculae Vainio and (C) Fistulo-

Steiner (1904) based his divisions of Ramalina on the arrangement of cortical tissue external to

116 G. NELL STEVENS

the algal layer. His section Corticatae (outer layer consisting of more or less transversely
arranged conglutinated hyphae) was equivalent to Desmazieria Mont., his section Bitectae
(outer layer as above overlying a tissue of longitudinally arranged conglutinated hyphae) was
similar to Euramalina Stizenb. He made a third division Ecorticatae (cortex composed of a layer
of longitudinally oriented conglutinated hyphae which formed a closed ring). Ramalina arabum
(Ach.) Meyen & Flotow is the type for Ecorticatae. Choisy (1954) commented that this thallus
structure was similar to Alectoria Ach.

Howe (1913-14) used spore characters to distinguish the sections he created within the genus
Ramalina as he thought this represented a more natural classification. Section Ellipsosporae R.
H. Howe contained species with ellipsoid or oblong spores (9-20 x 3-5-7-5 pm) and section
Fusisporae R. H. Howe contained species with long fusiform spores (16-35 x 3-7 /xm) and
section Bistortae R. H. Howe contained a species with sigmoid spores.

Each of these sections he divided into series and these series correspond in the main with
previous divisions, e.g. Howe's series Desmazieria (Mont.) = section Corticatae Steiner = genus
Niebla Bowler & Rundel. Series Myelopoeae = part of Myelopoea Vainio, series Fistulariae =
stirps Fistularia Vainio = genus Fistulariella Bowler & Rundel. He made three other series,
Tenuicorticate, Ciliate, and Fusisporae. This classification is not used.

Du Rietz (1926) relegated the genus Desmazieria Mont, to subgenus Desmazieria (Mont.)
(containing R. ceruchis, R. homalae, etc.) and Euramalina Stizenb. to subgenus Euramalina
(Stizenb.), which he divided into section TenuicorticataeR. H. Howe and section Bitectae Steiner.
Section Tenuicorticatae contained species without a chondroid element and this section he
further divided into subsection Solidae Du Rietz (species with a solid thallus, containing R.
evernioides, etc.) and subsection Tubulosae Du Rietz (species with inflated fistulose thallus: R.
inanisMont. = CenozosiaMassal.).

Du Rietz maintained section Bitectae Steiner (= Euramalina Stizenb.) but divided it into two
subsections (a) subsection Myelopoea Vainio thallus solid, and (b) subsection Fistularia Vainio
thallus inflated, fistulose. These extra subsections make the classification complicated although
the basic criteria used for division are sound.

Zahlbruckner (1907) combined the sections of both Stizenberger (1862) and Steiner (1904)
and made three divisions in the genus Ramalina: (i) section Ecorticatae Steiner (ii) section
Corticatae Steiner (which incorporated Cenozosia Massal. and Desmazieria Stizenb.) and (iii)
section Euramalina Stizenb. (= Bitectae Steiner). Zahlbruckner subdivided Euramalina
Stizenb. using the Vainio names Fistularia Vainio and Myelopoea Vainio and series Teretius-
culae Vainio and Compressiusculae Vainio.

Present day taxonomists are still divided in their acceptance of which species should be placed
in the genus Ramalina sens. str. Bowler & Rundel (1977) established the new genus Fistulariella
Bowler & Rundel for those species originally placed in Ramalina stirps Fistularia Vainio. They
placed emphasis on the hollow, usually perforate thallus as traits warranting genus status, yet
Australian material belonging to stirps Fistularia Vainio present a broad range of morphology
varying from almost totally compressed to totally fistulose. It is therefore regarded here that the
trait of inflation of the thallus should have no greater emphasis placed upon it than the roundness
of branches.

Rundel & Bowler (1978) proposed a new generic name Niebla to replace the genus name
Desmazieria Mont, as they found the former name was a homonym for the earlier described
genus Desmazeria Dumortier (Poaceae), and 13 taxa were transferred by them from Desma-
zieria and Ramalina into Niebla Rundel & Bowler.

Of the various divisions put forward up to the present day, the taxonomic system used by
Vainio would most suitably classify the Australian material, e.g. the Australian Ramalinae
would broadly divide into three groups: (1) fistulose thalli (2) terete and solid thalli and (3)
compressed and solid thalli.

(2) THE FAMILY RAMALINACEAE

Agardh (1821: 93) erected the family Ramalinaceae ['Ramalineae'], in which he placed the genus

Ramalina. Since that time the genus has been alternatively included with Usnea and other

RAMALINA IN AUSTRALIA 117

fruticose genera in the family Usneaceae, or isolated in the family Ramalinaceae. Table 2 shows
the various classifications which record Ramalina in Ramalinaceae (with or without other
members of Usneaceae) and the diversity of opinions as to the position of the genus.

Apparently Zahlbruckner (1907) and Smith (1921) did not regard septate spores nor a cortex
with mechanical tissue sufficiently different to separate Ramalina from other genera in the
family Usneaceae; but Watson (1929) regarded spore septation as most important and placed
Ramalina in the Ramalinaceae.

With modern day improvement in techniques of microscopy and chemistry together with a
better understanding of the fungal/algal composition, the natural relationships which exist
between the genera are being more fully understood.

Follmann & Huneck (1969) placed three genera in the family Ramalinaceae based on their
chemotaxonomy and anatomy (viz. Ramalina, Ramalinopsis, and Desmazieria). Poelt (1974)
also regarded these three genera as forming a natural entity and placed them in Ramalinaceae,
but he added a fourth genus which was not closely related, e.g. Speerschneidera. Culberson &
Culberson (1970) pointed out that chemical evidence supported segregation of Ramalinaceae as
a separate family as no species in the Usneaceae proper produces orcinol meta-depsides, a
category of substances richly represented in Ramalina.

Henssen & Jahns (1973) have kept the genus Ramalina in the family Ramalinaceae, because
of its two-celled spores and exobasidial conidiophores, and because of the specific lichen
substances it contains. They stated that the structure of the apothecium of Ramalina resembles
that of the Parmeliaceae in that there is a more or less complete algal layer beneath the
hymenium, but the Ramalinaceae differ from the characteristic cupular exciple of the
Parmeliaceae in that the algal layer is next to, not separated from the loosely interwoven
subhymenium.

Keuk (1979) divided the genera in Ramalinaceae into two groups based on anatomical and
chemical differences, and colour of pycnidial walls (a) Ramalina, Ramalinopsis, and Tricho-
ramalina and (b) Niebla (= Desmazieria Mont, and Cenozosia Massal.).

The most modern division within the family Ramalinaceae has been based on cortical
diversity. Bowler (1981) defined seven genera in the Ramalinaceae: Ramalina Ach., Fistu-
lariella Bowler & Rundel, Niebla Rundel & Bowler, Ramalinopsis (Zahlbr.) Follm. & Huneck,
Trichoramalina Rundel & Bowler, Dievernia M. Choisy, and Cenozosia Massal. Of these, the
genus Ramalina contains the largest number of taxa.

It can be seen from Table 2 that all recent authors who have specifically dealt with the question
of relationships of genera within the two families Ramalinaceae and Usneaceae, have con-
sidered them to be distinct morphologically, chemically, and taxonomically.

Collectors of Australian Ramalinae

Because the early explorers in Australian waters sailed in the temperate latitudes, it was only to
be expected that the first Ramalina collected in Australia would be a temperate region species;
this collection was made by Labillardiere, a botanist on the French ship 'Recherche' which sailed
along the Great Australian Bight and visited Tasmania in 1791 (Ducker, 1979). This first
specimen, held at H, (H-NYL 37226) was annotated 'Ramalina leiodea Nyl. Labillardiere, C.
van Dieman'. This was an erroneous identification which has now been determined as R. inflata
J.D. Hook. & Taylor.

In 1802, Robert Brown in his voyages with Matthew Flinders collected R. inflata from Port
Jackson (Sydney, N.S.W.), but this specimen was determined as Lichen fastigiatus Ach. by
Crombie (1880).

When J. D. Hooker accompanied Sir James Ross in his Antarctic expedition in the ships
'Erebus' and 'Terror' in 1839, he collected the type of R. inflata J.D. Hook. & Taylor on the
Lord Auckland Islands and from New Zealand he collected the type of R. geniculataL D. Hook.
& Taylor. They also visited Tasmania and collected material which was later published in Flora
Tasmaniae (1859). In this work Churchill Babington cites R. tasmanica Nyl. as a new species but

118

G. NELL STEVENS

Table 2 The classification of genera within the two families Ramalinaceae and Usneaceae as proposed by
various lichenologists from 1814 to the present day.

Author

Date Class/Order

Tribe/Family

Genus

Acharius, E.

1814 Class

Homothalami

Order Primus Scutellati

Order Secundus Peltati

Ramalina, Alectoria, Collema
Usnea, Cornicularia

Agardh, C.

1821

Ramalineae

Ramalina

Eschweiler, F. G.

1824

Usneaceae Usnea, Ramalina, Alectoria

Cornicularia, Dactylina,
Desmaziera, Endocena, Evernia,
Everniopsis, Letharia,
Neuropogon, Oropogon,
Siphula, Thamnolia.

Montagne, J. F. C. 1852

Tr. Parmeliacea
subtribe Usnea

Ramalina, Usnea

Nylander, W. 1857 Lichenacei

Ser. Ramalodei Tr. Ramalinei

Tr. Usneei

Ramalina, Alectoria, Evernia,

Dufourea, Dactylina
Usnea, Neuropogon, Chlorea

Stizenberger, E.

1862

Class
Homothalami

Tr. Usneaceae
Ramalineae

Usneaceae

Ramalina, Dactylina, Cetraria,
Evernia, Dufourea
Usnea, Alectoria, Oropogon,
Argopsis

Vainio, E. A.

1890

Order
Cyclocarpeae

Tr. Parmelieae

Ramalina, Usnea, Alectoria,
Evernia

Hue, A. M.

1901

Series
Cyclocarpae

Radiatae
Tr. Ramlineae
Tr. Usneae
Tr. Alectoreae

Ramalina, Dufourea
Usnea
Alectoria

Zahlbruckner, A.

1907

Order

Usneaceae

Ramalina, Usnea, Alectoria,

Cyclocarpeae

Oropogon, Dufourea, Evernia,
Everniopsis, Endocena,
Letharia, Thamnolia,
Siphula, Dactylina.

Smith, A. L.

1921 subseries Usneaceae

Cyclocarpineae

Ramalina, Usnea, Alectoria,
Evernia, Letharia,
Everniopsis, Oropogon,
Thamnolia, Siphula, Dactylina,
Dufourea, Endocena

Watson, W.

1929 Order

Parmeliales

Ramalinaceae Ramalina

Usneaceae Usnea, Alectoria, Letharia,

Evernia, Everniopsis, Dactylina,
Dufourea, Thamnolia, Siphula,
Endocena

Rasanen, V. J. P. B. 1943 Order

Cyclocarpeae

Usneaceae Ramalina, Usnea, Alectoria,

Evernia, Letharia,

Everniopsis, Dufourea,
Dactylina, Oropogon, Ramalea

Choisy, M.

1957

Ramalinaceae Ramalina

Usneaceae Usnea, Desmazieria, Cenozosia

RAMALINA IN AUSTRALIA

119

Table 2 - cont.

Author

Date Class/Order Tribe/Family

Genus

Follmann, G. &
Huneck, S.

1969 Ramalinaceae

Ramalina, Ramalinopsis,
Desmazieria

Henssen, A. &
Jahns, H. M.

1973 Order Ramalinaceae
Lecanorales
suborder
Lecanorineae

Ramalina

Poelt, J.

1974 Order Ramalinaceae
Lecanorales
suborder Usneaceae
Lecaniorieae

Ramalina, Ramalinopsis,
Desmaziera, Speerschneidera,
Usnea, Alectoria, Bryopogon,
Cornicularia, Evernia,
Everniopsis, Himantormia,
Letharia, Sulcaria, Neuropogon,
Oropogon

Bowler, P. A.

1981 Ramalinaceae

Ramalina, Fistulariella,
Trichoramalina, Niebla,
Dievernia, Cenozosia,
Ramalinopsis

examination of this material during the present study has shown this specimen to be yet another
morphotype of the very variable R. inflata.

The Reverend Dr. W. Woolls, a school master at Parramatta, N.S.W., in 1832, collected in
that area, and one Ramalina specimen called R. gracilis Nyl. (MEL 9432) is actually R. exiguella
Stirton.

During the period 1842-46, J. P. Verreaux (a French ornithologist) who was sent to Australia
by the Museum of Natural History in Paris, collected widely in Tasmania and mainland
Australia. In his botanical collections were several Ramalina species which he had obtained
during that period, either by collecting them himself or by purchase of material from others
(Lamy, in litt.). Because of this practice of obtaining specimens from other collectors there is
uncertainty about the locality given by Nylander for two 'type' specimens he records as collected
by Verreaux from Swan River, Western Australia, these are R. australiensis Nyl. and R.
confirmata Nyl. Both of these species grow in eastern Australia, and the Western Australian
types are the only material collected from that area. Nylander himself appeared unsure of the
locality of R. australiensis as he noted: 'in Australia (prope Swan River, ni fallor) legit Verreaux
1846'. Verreaux collections held at PC indicate he travelled north as far as Moreton Bay,
Queensland, and collected several Ramalina species from mangroves.

The first woman recorded as a lichen collector was Amalie Dietrich, who collected lichens in
Queensland for a wealthy Hamburg merchant named Godeffroy, for his private museum. Some
of these collections are now housed at Miinchen. They are collections of Ramalina species from
the Rockhampton area.

Friedrich Ludwig Leichhardt travelled through Queensland on his way to Port Essington
(Darwin), 1844-45. He collected a Ramalina (now R. filicaulis N. Stevens) along the way but
gave no exact location. As the occurrence of this species is rare today, more information about
the locality would have been helpful in the present study.

Ferdinand von Mueller, Victorian Government Botanist (1853-96), received lichen speci-
mens from various collectors in Australia and these specimens were sent to overseas lichenol-
ogists for determination, e.g. Anton Krempelhuber, Jean Miiller [Arg.], and James Stirton.

Daniel Sullivan, a headmaster, collected lichens for von Mueller, one being the 'type' of R.
glaucescens Krempelh., from Mt Ararat, western Victoria, and another was R. leiodea var.
fastigiatula Mull. Arg. 'type' from the Grampians. Another Ramalina which Sullivan collected
from Mt Ararat area (MEL 9471) now bears the name R. fimbriata Krog & Swinscow, a species
common in East Africa. Charles French, a plant propagator at Melbourne Botanical Gardens

120 G. NELL STEVENS

(Filson, 1976), collected the type of/?, inflata var.fissa Mull. Arg. from King Island, Bass Strait.
The type of/?, lacerata Mull. Arg. was collected at Eucla, Western Australia, by J. Oliver.

Collectors in New South Wales for the Victorian Botanical Department included T. White
who collected the type of R. myriodada Mull. Arg. from Twofold Bay, New South Wales.
Charles Knight, a New Zealand surgeon, collected lichens in the Sydney area; he named one
specimen R. subgeniculata C. Knight, which was subsequently changed to R. knightiana Zahlbr.
Knight also named R. minuscula var. alba a specimen collected by James Keys at Mt Perry,
Queensland. This name was published by Bailey (1886), but no description was given and later
John Shirley (1888) described it.

Messrs. Pentzke and Hartmann were two collectors in Queensland for the Victorian Botanical
Department (Wilson, 1889). Pentzke collected the type material of R. farinacea var. nervulosa
Mull. Arg. from Daintree River, north Queensland, and C. H. Hartman of Toowoomba
collected the type material of R. geniculata var. compacta Mull. Arg.

Collections made by F. M. Bailey, Government Botanist in Queensland, John Shirley, a
school inspector, and Mrs M. Thozet, were sent to James Stirton, W. Leighton, Charles Knight,
or Jean Miiller [Arg.] for identification (Wilson, 1889). It was Stirton who named the two
Ramalina types R. exiguella Stirton and R. perpusilla Stirton. Miiller Arg. described a
Rockhampton specimen collected by Mrs Thozet as R. farinacea var. squarrosa Mull. Arg. The
type of R. farinacea var. dendroides Mull. Arg. was collected by E. Forde from the Hunter
River, New South Wales. Both of these taxa have now been found to be morphotypes of R.
peruviana Ach.

The Reverend F. R. M. Wilson, from Kew, Victoria, collected prolifically (1877-97) in
eastern Australia, from Tasmania to Brisbane, and more particularly along Port Phillip Bay and
Gippsland, Victoria (Filson, 1976). He collected the type of/?, calicaris var. australica Rasanen
from Barwon Heads. Wilson himself described several taxa of Ramalina, e.g. R. unilateralis F.
Wilson, R. brevis F. Wilson, and R. brevis var. brevissima F. Wilson.

More recent collectors of Ramalinae in Australia are J. H. Willis, Assistant Government
Botanist at the National Herbarium Melbourne (Filson, 1976), A. C. Beauglehole, who
collected Ramalinae in South Australia and Victoria, J. S. Whinray, who made an extensive
survey of the lichen flora of the Bass Strait Islands, and G. C. Bratt and J. A. Cashin, whose
collections of Tasmanian lichens have proved invaluable in the present study.

Other present day lichenologists who have collected Ramalina specimens which have been
used in the present study are A. Archer (N.S.W.), J. A. Elix (A.C.T.), R. Filson (Vic.), G.
Kantvilas (Tas.), R. W. Rogers (Qld.), N. Sammy (W.A.), C. Scarlett (Qld), and R. Seppelt
(S.A.).

Materials and methods

The source of material for this research was threefold, (a) Personal collections (numbering over
10,000 specimens) , the majority from Australia, with small collections from the Pacific Islands of
Fiji, New Caledonia, Rarotonga, Tahiti, New Hebrides, New Zealand, and Hawaii, (b)
Herbarium material from institutes, as acknowledged, (c) Herbarium and living material from
private collections, as acknowledged.

Most of the specimens examined during this study were tested by thin-layer chromatography
using the techniques of Culberson (1972). Solvents B and C proved to be the most useful solvents
to separate the acids found in the various species of Ramalina. The chemistry of type material
was also checked. Examination of the surface features of the thallus (e.g. pseudocyphellae,
soralia, and soredia shapes) and spore size and shape, was carried out by use of the Scanning
Electron Microscope (SEM) using either a Cambridge or a Phillips 505. In order to observe the
shape and thickness of the hyphae composing the outer cortex and the supportive tissue thin
sections were examined by TEM, using an AEI Corinth 275 operated at 60 kV. Preparation of
thallus material involved fixation of the tissue in glutaraldehyde, post fixation in osmium
tetroxide, dehydration and then infiltration and embedding with Spurrs medium. The results
were not always successful as the tissue tended to pull away from the medium along the outer
cortex junction.

RAMALINA IN AUSTRALIA 121

All climatic data were derived from Meterological Bureau records (Climatic averages of
Australia, 1975).

Results

Morphology

(a) Soralia and soredia: Soralia are defined herein as those areas of thallus where clusters of
soredia are found, and the term soredia is applied to the more or less spherical bodies composed
of algae surrounded by short hyphae which act as vegetative diaspores. These vegetative
reproduction bodies are produced by 11 of the Australian Ramalinae, and their constancy in
occurrence enables this attribute to be used taxonomically.

The position and form of the soralia are also useful criteria in the separation of each sorediate
taxon. Round or ellipsoid soralia are produced by R. pacifica, R. caespitella, R. nervulosa var.
nervulosa, R. nervulosa var. luciae, and R. nervulosa var. dumeticola (Plate 1, fig. 3); these occur
marginally and laminally. Ramalina peruviana and R. tenella produce small, punctiform soralia
(Plate 1, figs 1, 2) which are mainly apical in R. tenella and marginal or lateral in R. peruviana.
The soralia found in the other four taxa are produced in several ways, e.g. (i) by disintegration of
the lower cortex (R. fimbriata, Plate 1, fig. 5) (ii) by the partial separation, both marginally and
apically, of the upper and lower cortices (R. canariensis) (iii) by lack of a continuous lower
cortex (R. reducta, Plate 1, fig. 4) and (iv) by the splitting or disintegration of parts of the lower
cortex at intervals along the branches (R. unilateralis , Plate 1, fig. 6). The soralia terminology
used by Du Rietz (1924) and Beltman (1978) was not generally applicable to the Ramalinae.

The composition of the soredia was examined under SEM; the algal cells were not discernable
but the fragments of hyphae which make up the outer surface of the soredium were observed to
differ in form in several of the taxa. The northern hemisphere species R. capitata is regarded as
having soredia covered by an 'epicortex-like' layer (James, pers. comm.) (Plate 2, fig. 1), and
such a structure was looked for in the Australian sorediate taxa. Four species were found to
possess some semblance of an outer covering (Plate 2, figs 2, 3, 4, 5). Whether this layer should
be compared to the 'epicortex' which occurs in some taxa in the Parmeliaceae as described by
Hale (1973, 1981) is debatable; but it can be likened to the 'Kittsubstanz' (cementing substance)
mentioned by Peveling (1970). Such a layer may act as a protection against wetting of the
soredia.

The Australian taxa which produce soredia with this outer covering are R. caespitella, R.
reducta, R. tenella, and R. fimbriata, and even in these only a minority of the soredia show it
clearly. The soredia of R. caespitella are small, spherical, and have a partially smooth surface
(Plate 2, fig. 2). Soredia of R. reducta are irregular in shape and size but fusion of the external
hyphal segments was apparent (Plate 2, fig. 3). Ramalina tenella soredia are small but well
defined, and some have a distinct outer covering (Plate 2, fig. 4). Ramalina fimbriata soredia are

Plate 1 (overleaf) SEM photographs showing soralia shapes in some of the Australian Ramalinae. Fig. 1
Apical, punctiform soralia - R. tenella. Scale 10 mm = 530 /urn (x 19). Fig. 2 Marginal/lateral,
punctif orm soralia -R. peruviana. Scale 10mm = 150^un(x 65). Fig. 3 Round and ellipsoid soralia-/?.
nervulosa var. dumeticola. Scale 10 mm = 220 /u,m (x 44). Fig. 4 Exposed lower surface with chondroid
strands of cortical tissue across area of soralia -R. reducta. Scale 10 mm = 270 /u-m (x 36). Fig. 5 Patches
of exposed medulla by disintegration of the under side-/?, fimbriata. Scale 10mm = 200//,m(x 51). Fig.
6 Lower surface split apart and eroded at intervals - R. unilateralis. Scale 10 mm = 530 /am (x 19).

Plate 2 (overleaf) SEM photographs showing soredia forms in some of the Australian Ramalinae. Scale
indicated by black bar. Fig. 1 R. capitata - showing smooth outer layer partially covering the soredium.
Scale 10 mm = 22 /urn (x 440). Fig. 2 R. caespitella - showing portion of soredia covered by a smooth
layer. Scale 10 mm = 22 /urn (x 440). Fig. 3 R. reducta - showing soredia with smooth surface (x 364).
Fig. 4 R. tenella - showing thin layer partially enclosing the soredium (x 1236). Fig. 5 R. fimbriata -
showing thin layer totally enclosing soredium (x 958). Fig. 6 R. unilateralis - showing soredium with a
'woolly' appearance; no outer layer present (x 1527). Fig. 7 R. nervulosa var. dumeticola - showing
soredia with irregular surface (x 873). Fig. 8 R. canariensis - showing soredia with a 'woolly'
appearance; no outer layer present (x 909).

122

G. NELL STEVENS

Plate 1

RAMALINA IN AUSTRALIA

123

Plate 2

124 G. NELL STEVENS

large and spherical, and the formation of an external layer could be discerned on many of the
soredia at various stages of development; the most advanced layer formation is shown in Plate 2,
fig. 5. All of these taxa are regarded as having granular soredia.

The soredia produced by R. unilateralis (Plate 2, fig. 6) and by R. canariensis (Plate 2, fig. 8)
are small in size and are composed of both thick and thin pieces of hyphae; there is no external
fusion of the hyphae so the outer surface remains loosely interwoven giving a woolly appearance
to the soredia (Plate 2, fig. 6). The soredia are farinose. The four taxa which belong in the R.
farinacea complex (R. pacifica, R. nervulosa var. nervulosa, R. nervulosa var. luciae, and R.
nervulosa var. dumeticola; Plate 2, fig. 7; Plate 3, figs 1, 2, 3) also produce soredia which have an
irregular surface made up of separate hyphal pieces which are not fused (Plate 3, fig. 2), and the
soredia are farinose. Ramalina peruviana produces soredia which have an irregular surface
(Plate .3, fig. 4).

Fibrils often arise from mounds of farinose soredia (Plate 1 , fig. 2) and these features have the
same structure as the soredia (Plate 3, fig. 7).

(b) Apothecia: The production of apothecia as a sexual means of reproduction occurs in all but
four of the Australian Ramalinae (i.e. seven of the 1 1 sorediate taxa also produce apothecia , but
only rarely, and often the mature ascospores are few and difficult to find). The esorediate taxa
generally produce numerous apothecia.

The position of the apothecia on the branches can be of taxonomic importance, being
terminal, subterminal, marginal, lateral, or laminal. Terminal refers to the apothecia at the end
of a branch. Subterminal refers to apothecia positioned near the end of the branch, with the
branch extending beyond the apothecium, forming a short attenuate branchlet or spur, which is
usually bent at an angle in relation to the main branch. If the spur grows long it makes the branch
appear geniculate. Marginal refers to apothecia on the edges of compressed branches, and
laminal refers to apothecia occurring on the surface of the thallus. If branches are subterete to
terete the apothecia occur laterally. When laminal apothecia occur specifically at the base of
bifurcating branches they are referred to as positioned at the axil of the branch.

Apothecial shape varies from concave to plane to convex; in some species the apothecia
remain concave to maturity, in others the apothecia are concave to plane when immature, but
are markedly convex at maturity. The thalline margin is usually distinct and entire, occasionally
becoming incised at maturity; when the disc becomes convex the margin is often obscured.

The size of the apothecia varies greatly from 0-2 mm to 5-0 mm with exceptions to 10 mm.
Most apothecia are shortly stalked, but sessile apothecia are found in R. australiensis and R.
filicaulis. The colour of the disc is usually yellow-green and pruinose; however, taxa in the R.
celastri complex have been found with orange coloured discs.

The apothecial tissue comprises a colourless hymenium formed of asci and paraphyses. The
asci are clavate to subcylindrical and lie amongst sparsely branched paraphyses over a developed
hypothecium. This hypothecial layer is usually brownish-yellow as is the epithecium [as used in a
lichenological context - Henssen & Jahns (1973)].

(c) Spores: The number of spores per ascus is eight, usually distichous in arrangement (Plate 5 ,
fig. 5), hyaline, 1-septate, varying considerably in size and shape, being broadly ellipsoidal to
fusiform and either straight to slightly curved or reniform (Plates 4 & 5).

Within a single ascus spores can vary from straight to curved, a feature also noted by Krog &

Plate 3 SEM photographs showing soredia forms and, surface features in some of the Australian
Ramalinae. Scale indicated by black bar. Fig. 1 R. nervulosa var. luciae - showing soredia with irregular
surface (x 750). Fig. 2 R. nervulosa var. nervulosa - showing fragments of hyphae making up the outer
surface, without a surface layer ( x 945). Fig. 3 R. pacifica - showing soredia with an irregular surface ( x
525). Fig. 4 R. peruviana - showing soredium with an irregular surface (x 1300). Fig. 5 Punctiform
pseudocyphellae-^?. subfraxinea \ar. confirmata (x 75). Fig. 6 Enlargement of section of Fig. 5 (x 750).
Fig. 7 Fibril produced from a mound of soredia - R. peruviana. Scale 10 mm = 30 /xm (x 341). Fig. 8
Tuberculate pseudocyphellae - R. subfraxinea var. subfraxinea. Scale 10 mm = 30 /xm (x 341).

RAMALINA IN AUSTRALIA

125

126 G. NELL STEVENS

Swinscow (1975) and Landron (1972) (Plate 4, fig. 4); therefore the shape of spores was not used
as a taxonomic character. The size of spores has also been found to be of limited taxonomic value
as most mature spores of the Ramalina species lie within the size range (8-)10-16 x 3-5-6 /xm
(Plate 4, figs 2, 3, 4, 5, 6). The relationship between the width and length of spores, as suggested
by Krog & Swinscow (1974) was investigated in this study to see if this could be a taxonomic tool
for separation of closely related taxa; however exceptions could always be found in the spore
ratios which negated the usefulness of this characteristic (Plate 5, figs 2, 3, 4).

Although the spores are 2-celled, some of the spores in the size range 14-16 x 4/nm appear 3
or 4-celled. These additional divisions are not produced by distoseptae so must be dense
cytoplasmic strands (which take up the cotton blue stain) across the cell which produce a 'mock
septation'.

The spores of each taxa examined under SEM showed surfaces which were remarkably
smooth (Plates 4 & 5) except for a film which seemed to coat some of the spores or remained in
patches attached to the spore surface (Plate 4, fig. 1; Plate 5, figs 1, 2, 4, 6).

(d) Pycnidia: The occurrence of pycnidia amongst the Australian Ramalina species is not
common. When present, they are always pale in colour, and produce pycnoconidia which are
rod-shaped, usually 3-5 x 0-5 jam in size. It was observed that they mainly occurred on sterile
thalli rather than on thalli bearing either apothecia or soredia. This suggests that they may
provide an alternative means of propagation for the species. Their irregular occurrence and
apparent uniformity prevented their taxonomic use.

(e) Pseudocyphellae: Many lichenologists have discussed these morphological characters (Du
Reitz, 1924; Culberson & Culberson, 1968; Duncan, 1970; Beltman, 1978; Bowler, 1981; and
Hale, 1981). It is generally accepted that these features are cortical pores in the surface of the
thallus which allow the exchange of gases. They are areas which have direct contact with the
medullary hyphae. In the Ramalinae they vary in shape and size and several names have been
applied to such surface features: striae, striations, tubercules, and papillae.

Pseudocyphellae appear as shallow depressions (striae) in the cortex (Plate 3, figs 5, 6) or as
white linear markings (striations) on the surface (Stevens, 19830: figs 1B-D, 2A-B). In the
Ramalina subfraxinea complex in Australia some of the taxa produce tubercles resembling
raised warts, the tips of these structures containing small pseudocyphellae (Plate 3, fig. 8). The
term 'tuberculate pseudocyphellae' was used by Krog & Swinscow (1975) to describe this form
of pseudocyphellae. Similar surface features occur in several North American taxa, but are
referred to as papillae, as in R. sinaloensis (Bowler & Rundel, 19726) and R. complanata
(Landron, 1972).

Most sorediate Ramalina species produce pseudocyphellae of one kind or another; in the case
of the R. farinacea taxa from the tropics, it was found that the elongate pseudocyphellae were
precursors to soralia formation (Stevens, 19830).

Bowler (1981) reported that pseudocyphellae are mainly absent in the Fistulariella taxa and
this was found to be so in the Australian fistulose material. As a diagnostic character for the
separation of taxa, pseudocyphellae have been found to be of little value.

Anatomy

The anatomical structure of all Ramalina species found in Australia comprises (1) a cortex
(which I regard as both the outer cortex plus the inner supportive tissue) and (2) a medulla
(which includes the phycobiont).

Plate 4 SEM photographs showing spore shapes in the Australian Ramalinae. Fig. 1 R. subfraxinea var.
leiodea- single spore. Scale 10mm = l-2jum(x 8000). Fig. 2R. australiensis spores. Scale 10mm = 4/xm
(x 2400). Fig. 3 R. pacifica spores. Scale 10 mm = 4 /am (x 2400). Fig. 4 R. subfraxinea var. confirmata
spores. Scale 10 mm = 4 /im (x 2400). Fig. 5 R. subfraxinea var. norstictica spores. Scale 10 mm = 4 /im
(x 2400). Fig. 6 R. subfraxinea var. subfraxinea spores. Scale 10 mm = 4 /mi (x 2400).

RAMALINA IN AUSTRALIA

127

128

G. NELL STEVENS

RAMALINA IN AUSTRALIA 129

(1) Cortex

In the Australian taxa the outer cortical layer is usually 20 /am in thickness and is composed of
thick walled, transverse or randomly oriented hyphae; it may be indistinct or even absent. The
inner layer serves as supportive tissue and varies in thickness. It is composed of bundles of more
or less periclinally arranged thick walled hyphae which make up a continuous or interrupted
sheath around the internal medullary hyphae.

The detail in the outer cortex is often difficult to detect under light microscopy when yellowish
granules are present; a problem pointed out by Bowler (1981). To obtain as much detail as
possible, sections were examined under TEM. Both cortical layers of hyphae are highly
gelatinized and a uniform distribution of lumina is apparent; each lumen is surrounded by a thick
matrix of fibrillar material arranged concentrically (Plate 6, fig. 1). Peveling (1974) commented
that it is difficult to determine whether the different layers around hyphae are part of the proper
cell wall or if they are extracellular additions to the wall.

The change in orientation of the hyphae is apparent under TEM. In transverse section the
randomly orientated outer layer hyphae appear elongate whereas the longitudinally oriented
hyphae appear circular (Plate 6, fig. 2). Scattered between the hyphae are outlines of crystals,
which are more dense in the outer region and may be usnic acid crystals (Plate 6, fig. 3).

(a) Anatomical structure as a means of separating taxa. Krog & 0sthagen (1980) divided the
species of Ramalina found in the Canary Islands into four anatomical types based on the position
of the supportive tissue: the farinacea type, the decipiens type, the bourgaeana type, and the
duriaei type. The latter two types occur in species now placed in the genera Niebla and Dievernia
and are therefore not relevant to Australian material.

The majority of the Australian Ramalinae could be classed as belonging to the farinacea type
(Figs 3A, B), as they possess a thin outer layer of tissue and a continuous inner layer of
mechanical tissue of uneven thickness. Variations can occur in the arrangement of this inner
supportive tissue if it becomes partly discontinuous, leaving sections of outer tissue unsupported
(Fig. 3C). However, this is not comparable with the 'decipiens type' where some of the strands
anastomose across the medulla. None of the Australian taxa could be separated on this trait.

Bowler (1981) pointed out that the Ramalina taxa could be divided arbitrarily into two groups
based on the degree of specialization in the cortical tissue. One group with a narrow, indistinct
outer cortex intergrading with the underlying supportive tissue and the other group with a
distinctly double-structured cortex, but he commented that intergradation could be seen
between the two forms.

The Australian taxa can be divided in this way generally, but thalli within one taxon can often
show differences in thickness of the outer cortical layer. Those taxa with a sparse or indistinct
outer cortex are R. caespitella, R. celastri complex, R. pacifica, R. nervulosa, R. exiguella (Plate
6, fig. 4), R. peruviana, R. reducta, and R. unilateralis. Those taxa with a comparatively distinct
double-structured cortex are R. australiensis, R. canariensis, R. filicaulis, R. fimbriata, R.
glaucescens, R. inflata complex, R. litorea, R. subfraxinea complex (Plate 6, fig. 5), and R.
tenella.

(b) Variation in the anatomical structure within the thallus. It was noticed when serial sectioning
thalli that differences in anatomical structure sometimes occurred depending on the portion of
the branch cut. Two forms of variation were found. The first form of variation is (i) variation in
the alignment of the hyphae in the supportive tissue in basal sections compared to the hyphal
arrangement in the mid and apical sections. Basal sections of R. celastri subsp. ovalis show a
typical thin outer cortex merging with the longitudinally oriented hyphae in the supportive
tissue, as found in the anatomy of R. celastri subsp. celastri (Plate 6, fig. 7). Sections cut through
the middle of the branch and towards the apex showed only a wide band of randomly oriented

Plate 5 SEM photographs showing spore shapes in the Australian Ramalinae. Fig. 1 R. exiguella spores.
Scale 10 mm = 4/u.m(x 2400). Fig.2/?. inflata subsp. inflata spores. Scale 10 mm = 4ju,m(x 2400). Fig. 3
R. inflata subsp. perpusilla (inland taxon) spores. Scale 10 mm = 4 /xm (x 2400). Fig. 4 R. inflata subsp.
perpusilla (coastal taxon) spores. Scale 10 mm = 4 /x,m (x 2400). Fig. 5 R. celastri subsp. celastri spores
within the ascus. Scale 10mm = 8-3//,m(x 1163). Fig. 6R. litorea spores. Scale 10 mm = 4/Ltm(x 2400).

130

G. NELL STEVENS

Fig. 3 Anatomical structures of Australian Ramalina thalli : A . Cross section showing uneven thickness of
supportive tissue in typical '/armacea-type' anatomy; B. Cross section of R. filicaulis terete branch
showing typical 'farinacea-type' anatomy; C. Cross section of/?, inflata subsp.perpusilla thallus showing
variation from Fig. A - the supportive tissue being partly discontinuous; D. Algal clusters on the upper
side only, as found in some thalli of the R. inflata group; E. Cross section of R. whinrayi showing thick
outer cortex and irregular arrangement of supportive tissue which extends into the medullary region,
with isolated rafts of supportive tissue also present.

hyphae forming a network (Plate 6, fig. 6). This anomaly may be explained by the number of
laminal apothecia that this taxon produces on both sides of the branch. The lack of supportive
tissue and the random arrangement of hyphae could be due to the repeated formation of
apothecial tissue from the middle of the branch to the apex. Brandt (1906) found that variations
occurred within a single thallus, and illustrated this with serial sections along the branch of
Ramalina curnowii.

The second form of variation is (ii) variation in cortical structure between the upper side of the
branch and the lower side. This was noticed by Brandt (1906) in the species R. populina. In
Australian material such variation was found to occur in some specimens of R. inflata subsp.
inflata and R. inflata subsp. perpusilla, where the inner cortex on the lower side is narrow,
continuous and of even thickness, without an algal layer, but on the upper side it is partly
discontinuous , of uneven thickness with algal clusters interspersed between the ridges (Fig. 3D) .
This formation appears to be a modification of the normal 'farinacea type' structure, to enable
optimal functioning of the phycobiont under particular environmental conditions. It is not a
constant feature of these taxa, so cannot be used taxonomically.

Another taxon which has an atypical and complex anatomy (Fig. 3E) is R. whinrayi. The
cortex consists of a thick outer cortex and a thick, discontinuous sheath of supportive tissue
made up of bundles with ridges extending across the rn,edulla or occurring as isolated rafts in the
medulla; such intrusions produce islands of medullary tissue with algal clumps. This form of
anatomy has been sighted in the northern hemisphere taxon R. capitata (= R. strepsilis).

(2) Medulla

The central portion of the thallus is occupied by medullary hyphae which are randomly oriented,

either tightly compacted or loosely woven and arachnoid. In the fistulose taxa the hyphae may be

RAMALINA IN AUSTRALIA

131

$?''

'llflQl

Plate 6 TEM photographs showing anatomical features. External edge of outer cortex indicated by black
arrow. Fig. 1 Lumen with surrounding fibrillar matrix. Scale 10 mm = 1-5 /xm (x 6770). Fig. 2 Section
from outer to inner cortex - R. exiguella. Scale 10 mm = 1 /im (x 10450). Fig. 3 Usnic acid? crystals in
outer cortex - R. peruviana. Scale 10 mm = 1-8 jum (x 5682). Fig. 4 Section from outer to inner cortex
showing narrow outer cortex. Scale 10 mm = 5 //.m (x 2125). Fig. 5 Section from ouer to inner cortex
showing broad outer cortex. Scale 10mm = l-2ju,m (x 5000). Fig. 6L.S. of mid-section of branch of R.
celastri subsp. ovalis. Scale 1 mm = 10 /x,m (x 105). Fig. 7 L.S. of mid-section of branch of R. celastri
subsp. celastri. Scale 1 mm = 19 /xm (x 53). (Photographs for Figs 6 & 7 by courtesy of A. Henssen.)

almost totally lacking, or occur in patches appressed to the inner cortex. Ramalinafissa produces
very little medullary hyphae, whereas in some R. inflata subsp. inflata specimens the central
cavity is criss-crossed by arachnoid hyphae to the same degree as R. glaucescens, a species
regarded as having a solid, compressed thallus.

132 G. NELL STEVENS

The sorediate species belonging to the R. farinacea group in Australia have a thick felty
medulla compared to the R. farinacea taxa in the northern hemisphere which have a loosely
compacted but continuous medullary layer. Other sorediate species, R. canariensis, R.
fimbriata, R. reducta, and R. unilateralis, reveal large areas of exposed medulla which produce
soralia. The most compacted medullary hyphae are found in R. whinrayi (Fig. 3E).

Chemistry

Usnic acid is found in the cortex of the Ramalinae in varying amounts but it is not used
taxonomically as a diagnostic property. Atranorin occurs only rarely and in trace amounts so it is
not used for taxonomic purposes either. It is the medullary substances produced by the
Ramalinae - the depsides and the depsidones - which are important.

(a) The orcinol depsides: These comprise both para- and meta-depsides. The /?ara-depside
divaricatic acid is particularly common and is found in ten Australian Ramalina species. It may
occur as the sole medullary constituent or be accompanied by biogenetically closely related
accessory acids such as stenosporic acid, nordivaricatic acid (bothpara-depsides) or by sekikaic
acid (a raeta-depside).

Evernic acid, another para-depside occurs in one saxicolous species endemic to Australia, and
lecanoric acid has been found as an accessory acid in this species. Evernic acid is rare in southern
hemisphere Ramalinae, whereas in the northern hemisphere it is a relatively common con-
stituent in this genus.

The greatest number of lichen acids found in the Australian Ramalinae belong to the orcinol
raeta-depside group. Fourteen substances have been identified in this study. The most common
metabolite is sekikaic acid which is found in nine species. In marked contrast it is found in only a
few northern hemisphere Ramalina species. Sekikaic acid may occur as the sole medullary
constituent or be accompanied by a number of minor acids viz. homosekikaic, ramalinolic,
4'-0-demethylsekikaic, 4'-0-methylnorhomosekikaic, and 4'-0-methylnorsekikaic acids.

Cryptochlorophaeic acid is another orcinol raeta-depside which has been observed alone or
may occur accompanied by paludosic and 4'-0-methylcryptochlorophaeic acids. Boninic acid
too, occurs as a major acid with the minor metabolites 2-0-methylsekikaic, 2,4'-di-0-
methylnorsekikaic, 4'-0-methylpaludosic, and 4,4'-di-0-methylcryoptochlorophaeic acids
(Chester & Elix, 1978).

(b) The fi-orcinol depsidones: Only four (3-orcinol depsidones have been observed in Austra-
lian Ramalinae. Salazinic acid is present in four species and may be accompanied by trace
amounts of protocetraric acid, but protocetraric acid does not occur as a major component in
any of the species. Norstictic acid occurs in one coastal species and psoromic acid is present in
one inland species.

The number of depsidones found in the Australian Ramalinae is less than the number found in
Ramalaina species in Europe or North America, where fumarprotocetraric acid, hypopro-
tocetraric acid, stictic acid, protocetraric acid, norstictic acid, psoromic acid, and salazinic acid
all occur.

Salazinic acid occurs in both sorediate and fertile Ramalina species which inhabit the
Australian coastline (viz. R. pacifica, R. tenella, and R. fissa). In R. pacifica and R. tenella trace
amounts of protocetraric acid are sometimes present accompanying the salazinic acid. The
occurrence of these two acids in the same thallus is probably due to incomplete enzymic
oxidation of the side chain methyl group in the conversion of protocetraric acid to salazinic acid
(Elix, inlitt.).

Triterpenoids, derived from the mevalonic pathway occur in specimens of R. tenella, and
terpenes have been found in several species but neither of these groups was utilized for
taxonomic purposes.

Chemical variation and species concept

Since the chemistry of a particular species is usually constant it has proved a useful property in

RAMALINA IN AUSTRALIA 133

combination with morphological characters to define a lichen species. However, in some
instances, morphologically identical taxa are found to contain different acids and the problem
arises of how to treat them taxonomically. Elix (1982) pointed out that there are three common
patterns of chemical variation (a) replacement type compounds (b) accessory type compounds
and (c) chemosyndromic variation.

(a) Replacement compounds: Replacement compounds occur in a number of Australian
Ramalina taxa including R. whinrayi, R. glaucescens, the R. inflata complex, the R. farinacea
complex, and the R. subfraxinea complex. The most common acids involved are divaricatic acid,
sekikaic acid, and salazinic acid.

The thalli of R. whinrayi produce either divaricatic acid nordivaricatic acid or the
replacement compound sekikaic acid with minor metabolites (4'-0-demethylsekikaic acid and
trace amounts of 4'-0-methyl norhomosekikaic acid). On the Tasmanian coastline both of these
races occur in the one population, their morphology is indistinguishable, and one thallus was
found to contain both divaricatic and sekikaic acids.

On the Bass Strait islands only the divaricatic acid race occurs on the islands in the Hogans
Group (the most northerly islands) , whereas on the Kents Group of islands and on Craggy Island
only the sekikaic acid race was found, although most thalli contain trace amounts of divaricatic
acid as well. Because of this dual occurrence of the acids in some thalli, both acid races are
regarded as belonging to the one taxon, viz. R. whinrayi.

An intensive study of the habitats in which each acid race occurs would be necessary to
understand and interpret the segregation of the two acid races on the different islands. A survey
similar to that carried out by Culberson (1969) on taxa in the R. siliquosa complex may solve
this problem if it could be found that the Tasmanian population occupied separate niches
according to the acids present. In this case the chemical races could be regarded as taxonomically
significant (Culberson, 1967, 1969ft) and the explanation of the mixed acids would infer
hybridization (Brodo, 1978) . But the islands are small and inaccessible and such a survey was not
possible during this study.

The occurrence of replacement compounds was also observed in the R. inflata group, where
lichen populations which are morphologically indistinguishable may contain either divaricatic
acid or sekikaic acid as medullary substances. Here again such taxa have been regarded as
chemical strains within the species; a decision strengthened by the presence of trace amounts of
sekikaic acid in some thalli which contain divaricatic acid. A third replacement compound,
salazinic acid, also occurs in the R. inflata complex. Biogenetically this depsidone is quite remote
from the two depsides, yet thalli possessing this acid are remarkably similar to those with a
depside chemistry. However, the depsidone taxon has been retained as a separate species (R.
fissa), as it has a geographically different distribution pattern from the depside containing
taxa.

(b) Accessory compounds: Accessory type compounds are also present in the R. inflata
complex, occurring sporadically in addition to the constant constituents, but having no
correlation with any morphological or distributional variations (Elix, 1982). The presence of
nordivaricatic acid in some divaricatic acid taxa and the presence of consalazinic or scabrosin
derivatives in the salazinic taxon have been regarded as instances of accessory acids accompany-
ing the major constituent and thus requiring no taxonomic recognition. Similarly the accessory
acid connorstictic has been found in the norstictic acid taxon in the R. subfraxinea complex.

Another Ramalina species complex comprising taxa with chemical variants of the replace-
ment type and with accessory compounds is the tropical/subtropical R. farinacea complex. The
acids involved are divaricatic acid (+ accessory compound stenosporic acid) and the sekikaic
aggregate of acids and salazinic acid, but this group is further complicated by chemosyndromic
variation occurring within taxa which produce the sekikaic aggregate of acids.

(c) Chemosyndromic variation: A chemosyndrome is a group of biogenetically related metab-
olites where one or two compounds are regularly the major components, and the minor

134 G. NELL STEVENS

biosequentially related constituent of one taxon becomes the major constituent of the other
(Culberson & Culberson, 1977; Elix, 1982).

In the R. farinacea complex, the two tropical/subtropical taxa R. nervulosa var. luciae and R.
nervulosa var. dumeticola contain the same acid components but these occur in different
quantities in each taxon (Stevens, 1983a).

Correlation between secondary-product chemistry and ecogeography in the Ramalina
subfraxinea complex

The importance of chemistry in identifying taxa which have different amplitudes of ecological
tolerance has been stressed by Culberson (1967, 1969b), Culberson & Culberson (1967), and
Sheard (1978). The notable example of correlation between secondary-product chemistry and
ecogeography is found in the R. siliquosa species complex of the northern hemisphere. This
complex contains six sibling species each with a different depsidone chemistry (norstictic acid,
salazinic acid, stictic acid, protocetraric acid, hypoprotocetraric acid, and acid deficient).
Culberson (1969b) maintained that each chemical race was a different species which reflected
the physiological differences between the taxa. However, the R. siliquosa chemistry comprises a
replacement series of medullary depsidones which are able to be ranked by increasing numbers
of oxidation steps in their biosynthesis (Culberson, Culberson & Johnson, 1977).

The maritime R. subfraxinea complex in the southern hemisphere is similar to the maritime R.
siliquosa complex in the northern hemisphere as it is made up of taxa which produce six different
acids; four of the acids are depsides and two are depsidones (viz. sekikaic acid, divaricatic acid,
cryptochlorophaeic acid, boninic acid, norstictic, and salazinic acids), but these acids are not
closely related biosequentially (Elix, in litt.).

Morphologically only the salazinic acid taxon is distinguishable; the other taxa show only
slight morphological variations from one another, none of which is sufficiently unique and
consistent that the chemical constituents of every individual can be infallibly predicted from
appearance alone.

Culberson, Culberson & Johnson (1977) found a correlation between the chemistry in the
R. siliquosa taxa and different amplitudes of ecological tolerance of each taxon. Accordingly,
a study of the thalli belonging to the R. subfraxinea complex was undertaken in order to find
if a particular chemotype could be related to a particular type of habitat based on the
amount of exposure to the sea (e.g. exposed = on trees or rocks facing the open sea; sheltered
= on trees growing on the landward fringe of a mangrove community or on trees in clay pan
habitats).

In Table 3 the type of habitat (exposed or sheltered) at each location site is compared with the
percentage of thalli containing each acid found at the site. The table shows that there is a
difference in the percentage of certain acids at each site. The cryptochlorophaeic acid taxon
occurs in areas experiencing the greatest amount of exposure to the open sea, the boninic acid
taxon is generally found in exposed habitats (usually adjacent to quiet water), whereas the thalli
containing sekikaic acid, divaricatic acid, or norstictic acid are mainly found in sheltered well
protected areas away from salt spray. Salazinic acid containing thalli appear to tolerate a broad
spectrum of habitats from exposed to sheltered.

From Table 3 it can be seen that there is a definite tendency for different chemotypes to
occupy some microhabitats more commonly than others.

This environmental selection of different chemotypes into separate habitats is also apparent
to a larger extent along the more tropical coastline of Australia (e.g. north of latitude 1630'S),
where collections made from any one site always contain only one chemotype. When the type of
habitat was considered it was found that collections from Princess Charlotte Bay (a vast area of
clay pan) contained sekikaic acid, whereas thalli collected from the exposed shorelines of the
small tropical islands of Ingram I., Turtle I., and Sue I. all contained cryptochlorophaeic acid;
thalli collected from Lizard I. all contained salazinic acid.

Although the collections from these northern regions may not have been as numerous as those
to the south, enough material was collected at each site to make these findings plausible.

KAMAL/NA IN AUSTRALIA

135

Table 3 Medullary metabolites expressed as a percentage of total number of specimens collected at
various sites in relation to type of habitat and latitude.

o

I a

.So

Latitude S

!

3

j_j

CTJ

J=
<+-

<u
a.
>>

Boninic

Cryptochlo:

Sekikaic

Norstictic

Salazinic

Divaricatic

Number of
collected

16

Daintree River

sheltered

23

37

40

87

16

Mossman

sheltered

29

63

8

38

17

Mission Beach

exposed

75

10

15

20

18

Murray River

exposed

40

33

27

15

18

Hinchbrook I.

sheltered

1

42

24

15

16

2

82

19

Townsville

exposed

59

29

12

17

20

Bowen

exposed

3

94

3

33

21

Hallidays Bay A.

sheltered

32

68

34

21

HallidaysBayB.

exposed

94

6

35

21

Eimeo

sheltered

25

35

4

36

28

22

St Lawrence

sheltered

9

8

14

69

70

23

Keppel Sands

exposed

96

4

46

23

Gladstone

sheltered

3

42

30

25

60

24

Turkey

sheltered

4

48

48

33

25

Burnett Hds

exposed

10

90

23

26

Hervey Bay

exposed

36

64

22

27
28

Moreton Bay

exposed

1

99

100

29

Clarence R.

exposed

66

34

23

Distribution patterns in relation to climatic indices

The distinct distribution patterns of the Ramalinae indicate that there are differences in the
ecological requirements of each taxon. The composite map of the distribution of all of the taxa
(Fig. 1) shows that geographically the distributions lie in either the tropical or the temperate
zones with some taxa overlapping in the intermediate zones (e.g. subtropical and warm
temperate). In the higher plants it has been recognized that plants growing under tropical
conditions have different growth responses compared to plants growing under temperate
conditions, and that such responses are closely related to thermal optima for photosynthesis
(Specht, 1981c; Nix, 1982).

As a knowledge of the major climatic indices which regulate plant response (e.g. temperature ,
precipitation, and solar radiation) has proved valuable in understanding the distribution of the
phanerogram flora of Australia, it seemed reasonable to examine their value in understanding
the distribution of the Ramalina taxa throughout Australia.

Nix (1982) proposed that Australian plants can be classified in terms of their photosynthetic
response to temperature. His three major groups are relevant to the pattern of Ramalinae
distribution. (1) Megatherm Group - plants with photosynthetic optimum 26-28C; lower
threshold 10C, upper threshold 38C. (2) Mesotherm Group - plants with photosynthetic
optimum 19-22C; lower threshold 5C, upper threshold 33C. (3) Microtherm Groups - plants
with photosynthetic optimum 10-14C; lower threshold 0C, upper threshold 25C. These three
major groupings have geographical equivalents as shown in Table 4.

Nix (1982) integrated the climatic factors, temperature, precipitation, and solar radiation into
a single multif actor Growth Index (G.I.). The Growth Index is different for plants with different
thermal growth responses. Figs 4A-B and 5 A indicate the distribution of megatherm,
mesotherm, and microtherm plants respectively with G.I. values >0-45 and G.I. values >0-30

136 G. NELL STEVENS

Table 4 Thermal divisions of Australian plant biota and their geographical equivalents, as suggested by
Nix (1982).

Thermal divisions

Mean annual
temperature

Geographical
equivalent

Megatherm
Megatherm/Mesotherm
inter/one
Mesotherm

24C

20-24C
14-20C

tropical
subtropical

Mesotherm/Microtherm
interzone

Microtherm

12-14C
<12C

warm temperate
cool temperate

<0-45 in the first two groups and with G.I. values <0-6 to >0-2 in the latter. The multifactor
Growth Index (G.I.) has values ranging from zero to unity and can never exceed the value of the
single most limiting factor. It is apparent from Fig. 4C, that Ramalina taxa occur only in those
areas with a G.I. larger than 0-30 in at least one thermal response category.

The distribution pattern of each Ramalina taxon conforms in a general sense to the patterns of
G.I. for megathermic, mesothermic, and microthermic phanerogams, indicating that the
climatic indices which determine distribution limits for the higher plant species also create
similar limits for the species of Ramalina.

Biogeographical division of the Australian Ramalinae

By superimposing the phanerogam thermal growth response patterns onto the distribution
pattern of each Ramalina taxon, the latter are able to be divided into megatherm, mesotherm,
and microtherm groups quite satisfactorily (Figs 4C, 5B).

(1) The megatherm element

Nine of the Ramalinae can be regarded as confined to a region with a megatherm Growth Index
value >0-30 <0-45. Their southern limits rarely extend below the tropics (latitudes 23S-24S).
These taxa are R. tropica, R. tenella, R. subfraxinea var. subfraxinea, R. subfraxinea var.
norstictica, R. nervulosa var. nervulosa, R. nervulosa var. luciae, and R. litorea. Ramalina
subfraxinea var. confirmata and R. subfraxinea var. leiodea are exceptions as their southern limit
reaches latitudes 33S and 29S respectively (Table 5). The occurrence of R. subfraxinea var.
confirmata on Sue Island (latitude 920'S) confirms its placement in the megatherm group; R.
subfraxinea var. leiodea extends to 18S (Figs 4A, 28).

(2) The mesotherm element

Other Ramalina taxa which have an extended range along the eastern coastline are regarded as
mesotherm elements as they have a southern distribution reaching the warm temperate region of
Australia. These taxa are R. exiguella, R. inflata subsp. perpusilla, R. peruviana, and R. pacifica
(Table 5). Although R. pacifica extends into the tropics to latitude 17S it has been regarded as a
mesotherm element because of its world distribution pattern which indicates its overall range is
in the subtropics (Stevens, 19836).

The presence of R. exiguella, R. peruviana, and R. inflata subsp. perpusilla in the tropical
regions at higher elevations than sea-level is similar to the occurrence of subtropical or even
temperate phanerogams which are reported to find an optimal thermal environment with
increase in elevation at low latitudes (Nix, 1982). Two taxa which have restricted distribution in
the mountains at altitudes exceeding 900 m are R. filicaulis and R. nervulosa var. dumeticola.
Being rare in occurrence it is difficult to establish in what thermal group they belong. They have
been tentatively placed as mesothermic plants as the former occurs in areas where R. celastri
subsp. celastri, R. peruviana, and R. inflata subsp. perpusilla (all mesothermic plants) grow; and
the latter must be a mesothermic offshoot of the otherwise megathermic R. nervulosa group.

The distribution pattern of R. inflata subsp. perpusilla covers a region corresponding to

RAMALINA IN AUSTRALIA

137

DISTRIBUTION PATTERN OF MEGATHERM PHANEROGAMS

Growth index veluet > 0.45
I I Growth index ralu > 0.30 < 0.45
(After Nix, 1981)

B

DISTRIBUTION PATTERN OF MESOTHERM PHANEROGAMS
Hill Growth index valun > 0.45
HUM Growth index veluei > 0.30 < 0.46
(After Nix, 1081)

Fig. 4 A. Distribution patterns of megatherm
phanerogams with G.I. values >O30; B. Dis-
tribution patterns of mesotherm phanerogams
with G.I. values >0-30; C. Ramalina distribu-
tion pattern with megatherm and mesotherm
plant boundaries for plants with G.I. values
>0-30, superimposed.

138

G. NELL STEVENS

5

XX

XX

XXX

$

XX

XX

XXX

$

X XX

XX

XXX X

o

X X

X X

XX XX

OS
co

XX

XX

XX XX

00
CO

X XX

XX

XXXX

r-

CO

XX XX

XX

XXX

CO

X X XX

XX

XXXX

&

X XX XX XX

XX

XXXX

CO

x x xxxx

XX

XX

CO

X X XX XX XX

XX

XX

<N

CO

XXX X X

X

co

X X XX XX X

X

XXXX XXXXX

XX

Os
CN

X XX XX XXX

X

00

XX XXXXXXX X

CN

X XXXXXX X

CS

X XXXXXXX X

8

X XXXX X

3

xxxxxxxxxxxx x

co

xxxxxxxxxxxxx

CN

XXXXX X XXXX

ts

XXXXXXXXX XX

03
X

O
<N

XX XX X XX

03

Q

rH

XX X

"3

OO

1 (

XXXX X XXXXX

1

XXX XX X XXX

c

S

XXXXXX

2

i*

1 1

XXX

<A

T 1

X XXXXX X

CO

X

<

X XX

c

oo

o

a

3
.0

T3

w -. ~

.5 | Q .3

is

.t^

v 'CJ O *- -5J -SJ -^

2

</5

_1

"^o^^ a 3 -2 ^ c5

"a

5

_^

5-* W r "?"* ^ ^5 "^ Q

4*!

Q

03

c

* e ^ -3 ^> Ti'B'^c'

3

o

"3

O3O3-K3 ^" O3 . , ' '"*

a

>>->> -i > CX,Jrt'

Pi

ex

3

Illl i 3 li g |. a |lil|

to
X3

3

'% "3 Q ^ -^ ^

0)

'll ^^> | ^ o | > H 1 1"**! 5 1 1 J[ 1

1^1

Illll 1

3

ii^i^^c^^cS-Scx.!^,^ , u a c -S "5c

Sj -S <.

o o <e, 3 o

H

c^c^oiD^c^ccccc^c<c^(^Qi^(^c^D^a;a;ft;

(^ Q^ Q^

ftj ft; QZ QC oj DC

RAMALINA IN AUSTRALIA 139

mesotherm response patterns with G. I. values above 0-30. The inland areas with G.I. values
>0-30 <0-45, correspond to areas where the sekikaic acid race as well as the divaricatic acid race
occur. Along the coast, the areas with G.I. >0-45 mainly produce thalli with divaricatic acid a
trace of sekikaic acid.

To the south, the mesotherm region with G.I. values >0-45 coincides with the R. inflata subsp.
inflata distribution (divaricatic acid alone or rarely + trace sekikaic acid). No specimens of the
R. inflata complex occur in the region delimited by the mesotherm elements with G.I. >0-30
<0.45 in inland New South Wales. In the temperate southern and south-western regions
of Australia R. inflata subsp. australis occupies this mesotherm region (G.I. >0-30 <0-45)
and again the sekikaic acid race is present at inland sites, as well as the divaricatic acid race. a
Along the coastline only the divaricatic acid race is present which is a similar situation to that
found with the taxon R. inflata subsp. perpusilla. Thus in the R. inflata complex, only taxa
containing divaricatic acid trace sekikaic acid are found along the eastern and southern
coastlines.

In the south-west corner of Western Australia both divaricatic acid and sekikaic acid taxa
occur, reflecting the general aridity of this part of Australia even close to the coast.

There is a continuation of both mesotherm elements into Tasmania (Fig. 4B) and both R.
inflata subsp. inflata and R. inflata subsp. australis occur, but in different regions. The R. inflata
subsp. inflata taxa occur in the inland montane region whereas R. inflata subsp. australis taxa are
found in the drier eastern areas (Figs 21, 22). Sekikaic acid taxa do not occur in Tasmania
(extreme dryness is not a feature of the island). A microtherm influence could be responsible for
the distribution pattern of the R. inflata subspecies in Tasmania and in many instances the
presence of intermediate morphs causes difficulty in delimiting Tasmanian taxa.

Another taxon placed in the mesothermic group is R. celastri subsp. celastri. The majority of
collection sites for this taxon lie within the belt with G.I. values >0-45, with a few collection
localities to the west in the area covered by the mesotherm belt with G.I. >0-30 <0-45.
Specimens from the latter area are always broader than coastal specimens. This taxon extends
south to latitude 38S with lessening frequency. At approximately 38S, 145E the mesotherm
region of plants with G.I. >0-45 is replaced by the region of plants with G.I. >0-30 <0-45 (Fig.
4B). This latter region is occupied by R. celastri subsp. ovalis, a taxon which extends to Western
Australia (Fig. 20). It is also found in northern Tasmania.

Ramalina australiensis occurs coastally in the mesotherm pattern with G.I. value >0-45 (e.g.
Toogoom in Queensland and Yamba to Twofold Bay in New South Wales, latitudes 25S and
29-37S respectively).

The main distribution of R. canariensis is along the southern coastline of Australia in the
region with mesotherm G.I. values >0-30 <0-45 which corresponds to a Mediterranean climate
region. Such environments are found in South Africa, Chile, and the Mediterranean (Raven,
1973) - all areas where this taxon occurs. The presence of/?, canariensis at several sites along the
south-east coastline of Australia may be due to suitable microhabitat conditions.

The two species found on the Bass Strait islands (viz. R. whinrayi and R. caespitella} cannot be
ascribed accurately to any of the three response groups.

(3) The microtherm element

There are four taxa which occur in the mountainous areas of eastern and/or southeastern
Australia at altitudes exceeding 600 m which can be regarded as belonging to the microtherm
response group: (1) R. reducta, (2) R. glaucescens, (3) R. unilateralis , and (4) R. fimbriata.

(1) Ramalina reducta occurs at elevations over 1000 m in open-forests on the New England
tableland. The area experiences very low winter temperatures with snow. Figs 5 A-B show that it
covers a region with G.I. values >0-2 <0-6.

(2) Ramalina glaucescens is classed as a microtherm element with its two chemical races
occupying two different response groups (i) G.I. >0-4 <0-6 and (ii) G.I. >0-2 <0-4 the former
region being occupied by the sekikaic acid race and the latter by the acid-deficient race (Figs 5B,
13).

(3) Ramalina unilateralis occurs inland and along the coast at several sites, but its distribution

140

G. NELL STEVENS

Microtherra

GI > 0.6

> 0.4<0.6

> 0.2<0.4
IGI 4 0.2

Fig. 5 A. Distribution patterns of microtherm phanerogams with G.I. values >0-2 (after Nix, 1981).

pattern corresponds with the microtherm element rather than the mesotherm element. Its range
covers a region where microthermic G.I. values are >0-2 <0-6 (Fig. 5B).

(4) Ramalinafimbriata grows on rock outcrops at high altitudes in the open-forest country of
temperate southeast Australia (Fig. 5B); its range covers a region where microthermic G.I.
values are >0-2 <0-6.

Biogeography

Taxa in the genus Ramalina found in Europe and North America are generally distinct from the
taxa which occur in Africa, South America, Australasia, and the Pacific Islands.

The records of Ramalina taxa which occur in the various regions of the world are scattered
through the literature. Only the few present day revisions of the genus can be relied on to contain
correct identification of the species; many publications record names of doubtful application.
Those regions covered in recent reliable publications are: Canary Islands (Krog & 0sthagen,
1980), East Africa (Krog & Swinscow, 1974, 1975, 1976), Fennoscandia and the British Isles
(Krog & James, 1977), Europe (Poelt, 1969), and the West Indies (Landron, 1972).

Although the Ramalinae of 'North America have not been completely revised, many of the
taxa have been recently investigated by Hale (1978), Rundel (1978a), Bowler (1977), Bowler &
Rundel (19720, 19726, 1973, 1974, 1978), Rundel & Bowler (1974, 1976), Hale & Culberson
(1970), and Moore (1968). Less recent studies cover the Hawaiian lichens (Magnusson &
Zahlbruckner, 1945; Magnusson, 1956); the Chinese Ramalinae (Zahlbruckner, 1930) and the
Japanese Ramalinae (Asahina, 1938, 1939).

The South American Ramalinae are in need of revision but records of taxa collected in various
regions of that continent appear in publications of Vainio (1890), Malme (1934), Follmann

RAMALINA IN AUSTRALIA

141

Fig. 5 B. Four Ramalinataxa with distribution patterns within the microtherm boundaries: 1 = R. reducta,
2 = R. glaucescens (sekikaic acid race), 3 = R. unilateralis , 4 = R. fimbriata.

(1967), Rundel (19786), Osorio (19700, 19706, 1972, 1978), and Osorio, Aquila & Zanette
(1980), as well as others. Papers with Ramalina taxa found on the Pacific Islands are listed in
Appendix A of Lichen Ecology (Hawksworth, 1977). The 'Catalogue of lichens from India,
Nepal, Pakistan and Ceylon' (Awasthi, 1965) was consulted to obtain species numbers in India,
as no information concerning the present revision of Indian Ramalinae was obtainable from
India. Nine Ramalina taxa are described for New Zealand (Galloway, 1985).

From the above mentioned records and personal knowledge, some idea of the global
distribution of the taxa which occur in Australia was obtained (Table 6, Fig. 6). Table 6 shows
that Africa has the greatest number of species in common with Australia (50%). This close
relationship with African lichens was mentioned by Rogers & Stevens (1981). New Zealand and
South America have the next highest percentages of species in common with Australia. It is

142

G. NELL STEVENS

Fig. 6 Global distribution of Ramalina species found in Australia.

interesting that (as far as could be ascertained) there is little similarity of species between India
and Australia (see below). The lack of correlation between Australian taxa and those of North
America, Europe, and Asia, confirms that the greatest similarities are found in countries which
have a former Gondwanaland origin.

Palaeobiogeography

Before presenting an interpretation of the likely palaeobiogeography of the Ramalinae, an
understanding of some aspects of the Earth's geological history has to be appreciated.

(1) Geological history of Gondwanaland

The geological events that took place during the Cretaceous and Tertiary Periods which are so
vital to the understanding of the distribution patterns of plants are summarised in Table 7.

The early ideas of land bridges between continents and the later theory of Continental Drift
have been superseded by the theory of Plate Tectonics. This theory is now generally accepted by
most geologists to account for the separation of continents which show evidence of having been
once grouped together as a Pangaea or super-continent, or as two large continents, Laurasia
(north) and Gondwanaland (south). In this theory, the continents break up by rifting and
separate by sea-floor spreading. New oceanic crust is formed at spreading ridges and consumed
in subduction zones. The theory of an expanding earth (Carey, 1976) obviates the necessity for
subduction zones and produces a more neatly-fitting Pangaea than other models, especially in
the closer fit of northern and north-west Australia with Asia, making the distribution of now
distant but similar biota more easily understood; unfortunately this theory has not had general
acceptance.

(2) Plant distribution related to geological events

The map of the Gondwanaland continent during the Cretaceous Period (130-100 m.y. B.P.)
[Fig. 7] shows the relevant palaeolatitudes at that time (Powell, Johnson & Veevers, 1981).
Raven & Axelrod (1974) stated that direct migration from South America and Africa via

RAMALINA IN AUSTRALIA 143

Table 6 The global distribution of the Ramalina taxa occurring in Australia.

Ramalina species

alia

Au

South America

-c

-3 <= -3

2

U

1 8 8 -8

w

s

J

u

R. australiensis
R. caespitella
R. canariensis
R. exiguella
R. filicaulis
R. fimbriata
R. glaucescens
R. litorea
R. peruviana
R. reducta
R. tenella
R. unilateralis
R. whinrayi
R. celastri

subsp. celastri

subsp. ovalis
R. inflata

subsp. inflata

subsp. perp usilla

subsp. australis
R.fissa
R. nervulosa

var. nervulosa

var. luciae

var. dumeticola
R. pacifica
R. subfraxinea

var. subfraxinea

var. leiodea

var. confirmata

var. norstictica
R. tropica

TOTALS

* *

28 14 8

949331

Antarctica to Australia was possible at this time (Table 7). Even in the late Cretaceous,
Australia still had connections with South America through Antarctica, but the migration of
plants via the southern route probably became progressively reduced until finally stopped by the
break of Australia from Antarctica in the Eocene (53-38 m.y. B.P.). The early break-away of
India from the rest of Gondwanaland (125-100 m.y. B.P.), and its rafting north across the
equator with the loss of some Gondwanaland species, could account for the lack of taxa similar
to the Australian Ramalinae.

Two major pathways of dispersal of the higher plants from west Gondwanaland have been
postulated by plant biogeographers: (1) a temperate migration route via Antarctica from west
Gondwanaland (i.e. South America and Africa), to Australia and New Zealand, and (2) a
tropical route from East Africa to Madagascar, the islands of the Indian Ocean and Indo-
Malaysia to Australia (Croizat, 1952; Raven & Axelrod, 1974). As the Ramalina species found
in Australia can be placed in a range of temperature response groups from cool temperate to

144

G. NELL STEVENS

Table 7 The history of the break-up of Gondwanaland, indicating available migration routes (from
Raven & Axelrod, 1974 and Powell, Johnson & Veevers, 1981).

Million

years BP Geological periods

Event/geographic situation

125

110+10
100

90
80

63

55
49

45
30

20
10
Present

early Cretaceous

mid Cretaceous

late Cretaceous

early Palaeocene
Palaeocene-Eocene
mid Eocene

mid Oligocene
early Miocene
late Miocene
Holocene

Africa and South America begin to split; India begins to split

from Australia/ Antarctica but remains attached to

Madagascar.

Direct migration between west Gondwanaland and Australia

ceases.

Direct migration between Africa and Madagascar still

possible.

India rafts northwards and is completely separated from

Australia and Antarctica; no exchange of species possible.

Migration of species between Africa and Australia via east

Antarctica still possible.

Africa and NE. Brazil separated by a narrow strait; migration

possible at this point.

Africa and South America separated by 800 km of ocean but

numerous islands linked them along the mid- Atlantic Ridge.

Australia ceases to have any migration from Africa.

Australia/ Antarctica extend from 40S to the polar region.

South America closely connected with Antarctica 80-30

million years BP.

Separation of New Zealand and New Caledonia from

Australia.

Africa and Europe connected at Spain and possibly with Asia

at Arabia, allowing both northward and southward

migration.

Oceanic crusts form between Antarctica and Australia;

northern edge of Australia now at 30S and commencement

of separation of Australia from Antarctica.

Separation of the continental margins of Australia and

Antarctica but another 10 m.y. of migration of species from

South America through Antarctica to Tasmania along the

South Tasman Rise before final separation.

Direct migration between India and Asia.

The leading edge of Australia as it rafts northwards still south

of the latitude of the Sunda Arcs.

Initial contact between Sunda Arc and New Guinea

(abundant precipitation).

Collision of Australia with the Sunda Arcs.

Antarctic ice-sheet rapid expansion (lowered precipitation).

Australia moves towards the equator at 66 mm year" 1 .

tropical, it could be hypothesised that their routes of entry to Australia were from both the south
(the microtherm and mesotherm taxa) and from the north (the megatherm taxa).

(3) Dispersal and migration

Any postulates concerning the migration pattern of the Ramalinae from a centre of origin are
purely hypothetical as no fossil records are available and any assumptions made are based on the
migration pathways of the higher plants deduced from fossil records. The approach used in this
study is similar to that of J0rgensen (1983) and Sipman (1983) although the writer does not agree
with all of their conclusions.

Biogeographic evidence suggests that many genera and families of gymnosperms and
angiosperms had evolved before the breakup of Gondwanaland began in the Cretaceous, and
had arrived in Australia from South America and Africa via Antarctica during the middle

RAMALINA IN AUSTRALIA

145

3CfN

Fig. 7 Position of east and west Gondwanaland in the early to mid-Cretaceous (130-100 m.y. B.P.). Ticks
show longitudes which have no fixed origin (Adapted from Powell, Johnson & Veevers, 1981).

Cretaceous (108-100 m.y. B.P.). Fossil evidence indicates that many of the Gondwanan
families and genera had occupied Australia by the early Tertiary (63-55 m.y. B.P.) (Specht,
1981c).

(a) Step by step dispersal: Schuster (1979) stated that migration of land plants normally tends to
be by short range dissemination as members of structured communities, and he proposed that
the dispersal of Hepaticae occurred as part of the dispersal of whole plant communities . It is also
reasonable to hypothesise that the Ramalina species also made this step by step spread across the
continents of Laurasia and Gondwanaland with the migrating structural communities during the
Cretaceous-early Tertiary.

This suggested step by step migration of plant communities appears to be a valid means of
dispersal as this enables plants which have similar habitat requirements to expand as a whole, in
favourable climates, and so migrate when conditions are favourable and disappear from areas
where environmental conditions have deteriorated. It seems highly likely that under the
circumstances lichens which grow within these communities would migrate at the same rate and
in the same direction.

If step by step migration halted with sea-floor spreading of the continents, then another means
of dispersal has to be proposed. Seeds of the higher plants have been found to be carried by birds
and animals over long distances; the lighter seeds being wind dispersed. The Ramalinae produce
both spores and vegetative diaspores and it is feasible that these could be wind dispersed or
carried by birds.

(b) Wind dispersal: If long distance dispersal of spores by the wind is accepted then this means
of dispersion would have been in operation in early times. It is reported by Raven & Axelrod
(1974) that with the separation of Australia and Antarctica in the Eocene, there was a

146 G. NELL STEVENS

strengthening of the circum-Antarctic wind which could have carried spores from South
America to Australia, Tasmania, and New Zealand.

The success of a spore in a new environment would need the availability of a suitable
phycobiont and the lack of competition from plants already in the area. With the vegetative
diaspore both partners arrive together so that provided the environment is hospitable to their
genetic requirements they will readily become established in the new habitat. Spores can
become established after long distance wind dispersal since fertile lichen species have been
found on the islands of Hawaii. J0rgensen (1979, 1983) remarked on the efficiency of wind
dispersal of lichen spores in the population of the remote island of Tristan da Cunha.

If the original dispersal of the Ramalinae is comparable with the original dispersal of
angiosperms and gymnosperms, then their migration from west Gondwanaland may have taken
place by both step by step migration and wind dispersal, with the latter means continuing after
the break up of the continents.

(4) Centres of origin

The centre of origin of a group is often regarded as that area having the greatest concentration of
species. Hale used this method in assessing the centre of origin of the genera Relicina (Hale,
1975), Pseudoparmelia (Hale, 19760), Parmelina (Hale, 19766), and Bulbothrix (Hale, 1976c).

If the numbers of Ramalina taxa occurring in each landmass that once formed Gondwanaland
are estimated (and these numbers must be only approximate as without recent revisions many
names could be synonyms or misidentifications) then it is found that 41 Ramalinae occur in
South America, 42 in Africa, 36 in India, and 30 in Australasia.

South America and Africa (once West Gondwanaland) appear to have the largest number of
Ramalinae and could be regarded as the centre of origin of the southern hemisphere taxa.

The occurrence of high numbers of taxa within the one genus, in a particular region, can,
however be due to a re-radiation of species after the original dispersal of the taxa. When an
environment is free of competitive species, newly evolved taxa belonging to the genus will
become established. This situation has been found in Western Australia amongst the eucalypts,
where secondary radiation of species has occurred in a region where there is little competition
from other genera (Pry or, 1981).

Such a situation may have arisen in Australia with the evolution of the mangrove flora in this
region (Specht, 19816). Mangroves are the major phorophytes for the present day R. subfrax-
inea complex. It is proposed that the ancient stock of the R. subfraxinea group could have
entered Australia via the northern route from East Africa (as this group is regarded as a
megatherm response group) and become established along the Australian tropical coastline on
the newly evolved mangroves. The occurrence of all six chemotypes in the region around
latitude 18S indicates an area of speciation from where secondary radiation could have spread
to the east and west.

(5) Vicariant taxa

The Croizat theory of vicariance biogeography (Croizat, 1952; Croizat, Nelson & Rosen, 1974)
postulates that species which are ecologically similar but occur on different land masses have a
common ancestry. They are regarded as disjunct populations of once continuously distributed
taxa which have undergone speciation as physical disruption occurred and Croizat cited many
Gondwanaland taxa to support his theory. Galloway (1979) employed the same ideas in his
comments on the distribution of species in the genera Usnea, Pseudocyphellaria, Psoroma, and
Stereocaulon.

A number of taxa belonging to the Fistularia group in Australia appear to be part of the
vicarant taxa of Gondwanaland origin. For exampfe the Australian taxon R. inflata subsp.
perpusilla has an ecotype which grows on mangroves and closely resembles the East African
species R. consanguinea which also occurs on mangroves with the main difference in the spore
size; in some habitats in Australia R. inflata subsp. inflata morphology closely resembles some
forms of the East African taxon R. calcarata, but their chemistry differs in the minor
constituents.

RAMALINA IN AUSTRALIA 147

Chemical difference is the only character which separates the Australian taxon R. filicaulis
from the American taxon R. anceps. Material similar to the Australian material, and also lacking
medullary acids, has been cited from South Africa. It is proposed that all three continents
possess species which could be regarded as vicariant taxa belonging to the R. usnea complex, as
they are ecologically and morphologically similar.

On the basis of the evidence discussed above it is postulated that Australia has been populated
by species of Ramalina since the late Cretaceous. Since west Gondwanaland is accepted as the
centre of origin for the angiosperms, based on fossil evidence, there is the possibility of these
acting as phorophytes for ancestral taxa of the present day Ramalina, as that same region is a
likely centre of origin for Ramalina. The step by step spread of the plant communities would
have enabled the Ramalinae to spread by one of two migration routes either (1) the southern
route from South America (after Africa had separated) via the Scotia arc, Antarctica to
Tasmania and Australia or (2) the northern migration route from East Africa via Madagascar,
the Mascarenes, Indo-Malaya to northern Australia, by which means the megathermic Ramali-
nae could have reached tropical Australia. Once established in Australia the megatherm
Ramalina taxa remained a maritime group as inland habitats would have been inhospitable.
Secondary radiation from Australia by some of the megathermic taxa appears to have occurred
with the spread of species to the Pacific and Indian ocean islands.

The microtherm Ramalinae became established in the mountainous region of southern
Australia since these habits would have suited their thermal response patterns as Australia
drifted northwards and the temperatures increased.

The climatic changes that occurred in the late Miocene Period caused the desiccation of the
continent. Subsequent speciation of mesotherm Ramalinae could have occurred producing taxa
which occupied the new and dryer habitats of the south-west and west of Australia.

In this way the present distribution pattern of the Australian Ramalinae reflects their
palaeobiogeographic and palaeoecological history.

Taxonomy

The delimitation of Ramalina species proved difficult owing to the amount of morphological
variation shown by each taxon. A large amount of material had to be examined in order to
ascertain what concepts should be applied in the delimitation of each taxon. Three taxonomic
ranks have been used.

Taxonomic ranks

(1) Species: The rank of species has been given only to those taxa which can be recognized by
several distinct morphological traits which collectively produce a particular thallus type (allow-
ing for an appropriate amount of variation), together with a set of specific anatomical and
chemical properties, and which show a distinct habitat and distribution range.

Difficulties arose when two or more chemotypes produced a single morphotype; this problem
had to be resolved by considering the habitat and distribution range of the taxa, and if the only
difference found was in the chemistry then these chemical races were treated as conspecific.

The concept of species pairs (Artenpaare), as proposed by Poelt (1970, 1972) is that species
which reproduce by vegetative methods (secondary species) have been derived from existing or
extinct fertile species (primary species), and that primary and secondary species have identical
(or closely related) chemical components. This view is held by most lichenologists and species
pairs have been found in many genera e.g. in the Physcia and allied genera (Moberg, 1977),
Parmelia sect. Hypotrachyna (Culberson & Hale, 1973), Dirina (Tehler, 1983), and to a lesser
extent inAlectoria (Brodo & Hawksworth, 1977).

Only a few records of species pairs have been found in the Ramalinae. Rundel & Bowler
(1976) discussed R. leptocarpha and R. sublectocarpha as a fertile-sorediate species pair and
Krog & 0sthagen (1980) have commented that the fertile species R. implectens may be regarded
as the parent morph of the sorediate species R. farinacea.

148 G. NELL STEVENS

None of the Australian sorediate Ramalinae is regarded as derived from any of the fertile taxa
found in Australia today.

(ii) Subspecies: The delimitation of a taxon based on well marked discontinuities in characters
was not always possible. Some Ramalina taxa were found to intergrade into each other, differing
only subtly morphologically yet extending over large areas. When such complexes were
examined carefully the most obvious morphological changes were found to coincide with
different geographical regions.

The term subspecies has therefore been applied to a taxon which could be shown to occupy a
particular region within the total distribution range of a large complex (as defined by Hawks-
worth, 1976). This taxon can be extremely variable within its geographical confines and can
produce several ecotypes which occupy separate habitats.

As subspecies intergrade into each other, their delimitation is much more arbitrary than that
of a species, and depends more on geographical divisions than on chemical or morphological
differences. Imshaug & Brodo (1966) pointed out that the use of the rank of subspecies
permitted a classification which is not only practical but one which indicates a possible
phylogenetic system. The evolutionary relationships that may exist in some of the Australian
taxa have been indicated by the use of the rank of subspecies.

(iii) Variety: The rank of variety has been used when more or less morphologically identical
taxa have been found to produce different chemistries which correlate with distinctive distri-
bution patterns.

The production of either divaricatic acid or sekikaic acid in populations of the one taxon posed
a nomenclatural problem. Several lichenologists have mentioned the same difficulty e.g. in the
genus Dirinaria (J0rgensen, 1974) and in the genus Ramalina (Imshaug, 1972; Rundel, 19780).
The solution has been to regard both acids as representing chemical races within the one taxon.

Hybridization

It is not easy to establish the genetic homogeneity or heterogenity of lichenized fungi. Jahns
(1974) pointed out that thallus or hyphal fusions do occur in lichens, which could enable a form
of hybridization to take place. A specimen of R. leiodea recorded by Elix (1982) as being a
'schizophrenic thallus', had 'left-hand branches containing one acid, the right-hand branches
another and the central branches containing a mixture of both'. In this case the two acids
involved were two depsides, not closely related (cryptochlorophaeic acid and boninic acid).
Another such combination of two acids in one specimen belonging to the R. subfraxinea
complex has been found during the present study; the specimen was collected at Hinchinbrook I.
where all acid races of the R. subfraxinea complex are present. The thallus concerned contained
both boninic acid and salazinic acid, one a depside and the other a depsidone. As these two acids
are not biosequential metabolites it appears likely that this is a similar form of hybridization to
that found by Elix (1982), and that both could be regarded as 'chimeras' produced by the fusion
of hyphae from two different spore types. It is not known what chemistry the thalli produced
from spores of this plant would have. No other thalli with this combination of acids has been
found.

The occurrence of sekikaic acid in the divaricatic acid race ofR. subfraxinea var. subfraxinea is
regarded as different from the above. Sekikaic acid and divaricatic acid are reasonably closely
related biosequentially and apparently can interconvert (Elix in litt.) which would result in
chemical combinants if the conversion was incomplete. However the hypothesis of interspecific
hybridization (Brodo, 1978) remains feasible and has to be considered beside the hypothesis that
incomplete conversion of a precursor acid occurs during the biosynthetic steps from one acid to
another (Bowler & Rundel, 1978). If hybridization is accepted as the cause of this mixture of
acids then it would be natural to expect the proportions of each acid to be equal in the thallus.
However, this was not the case with the taxa examined; in all instances divaricatic acid occurred
in far larger quantities than did the sekikaic acid. It is therefore suggested that incomplete
conversion of one acid to another has taken place.

RAMALINA IN AUSTRALIA

Table 8 Taxonomically important characters in the Australian Ramalina taxa.

149

substrate

thallus

major acid constituents

orcinol depsides (3-orcinol
depsidones
para- meta-

1

&
_o

M

o

T3

u

.22

_o

1 .a

8 1 1 i 1 4

8

ft

3

e s

2 -*3 H 'S "^ O y

Ramalina species

'

8

u

T3 e5

1 1

u

k.

o

V5

S g

O <U

3 & 's J ^ s *

flS CT O "3 O tin r3
35 u O ca C fii @

R. australiensis

X

X

X

X

R. caespitella

X

X

X

X

R. canariensis

X

X

X

X

R. exiguella

X

X

X

R. filicaulis

X

X

X

R. fimbriata

X

X

X

X

R. glaucescens

X

X

X

X

R. litorea

X

X

X

R. peruviana

X

X

X

X

R. reducta

X

X

X

X

R. tenella

X

X

X

X

R. unilateralis

X

X

X

X

X

R. whinrayi

X

X

X

X

Species complexes

R. celastri

subsp. celastri

X

X

X

X

subsp. ovalis

X

X

X

R. inflata

subsp. inflata

X

X

X

subsp. perpusilla

X

X

X

X

subsp. australis

X

X

X

X

X

R. fissa

X

X

X

X

R. nervulosa

var. nervulosa

X

X

X

X

var. luciae

X

X

X

X

var. dumeticola

X

X

X

X

R. pacifica

X

X

X

X

X

R. subfraxinea

var. subfraxinea

X

X

X

X

var. leiodea

X

X

X

var. confirmata

X

X

X

X

var. norstictica

X

X

X

R. tropica

X

X

X

X

Description of the genus Ramalina

Ramalina Ach., Lichenogr. Univ.: 122,598 (1810), nom. cons.

Thallus fruticose, greenish-grey, pale green or greenish-yellow, caespitose, erect, subpendulous
to pendulous; branching dichotomous, subdichotomous to irregular or dense intricate; branches
few to many, twig-like, strap-like to palmate, flat, subterete or terete, solid or inflated; apices
forked, attenuate, rounded; surface shiny or matt, smooth or rugose, pseudocyphellate;
holdfast delimited or dispersed; soralia small and punctiform to large and round or elongate;

150 G. NELL STEVENS

cortex composed of two layers, the outer cortex hyphae more or less anticlinal or randomly
oriented, the inner cortex hyphae periclinally arranged; hyphae surrounded by a large amount
of matrix; medullary hyphae loosely interwoven between the upper and lower cortex or confined
to the algal zone.

Apothecia terminal, subterminal, marginal, lateral, laminal; convex, plane or concave,
sometimes incised; disc pale to flesh coloured; margin concolourous with thallus; asci clavate,
containing 8 spores; spores hyaline, oblong, ellipsoid or fusiform, straight or curved, 1-septate.
Pycnidia inconspicuous, pale, usually rare in Australian species.

The algae are Trebouxioid, globose, forming an irregular ring along the edge of the inner
cortex; when confined to the upper side only the thallus becomes dichroic with a white lower
surface.
Type species (conserved): Ramalina fraxinea (L.) Ach.

Enumeration of the Australian taxa

The Australian Ramalinae comprise 28 taxa which can be grouped into 17 fertile and 11
sorediate taxa. Sixteen species have no subspecific taxa recognized, and four species contain a
total of 12 subspecific taxa.

Table 8 lists the taxonomically important attributes of each of the 28 taxa; those belonging to
species complexes are placed after the alphabetically arranged single species, as is also done in
the description of the taxa. The disposition of the other Ramalina taxa recorded by Wetmore
(1963) and Weber & Wetmore (1972) is shown in Appendix 1.

The specimens examined are cited with location, latitude and longitude in degrees, type of
substrate, date, the collector's name and the herbarium abbreviation (Holmgren, Keuken &
Schofield, 1981). Collectors initials have been used for those collectors whose specimens are
cited most frequently in this work. These names have been abbreviated as follows:

AA A. Archer JC J. Cashin

ED E. Dahl JW J. S. Whinray

GB G. C. Bratt NCS N. C. Stevens

GK G. Kantvilas NS N. Sammy

GNS G. N. Stevens RF R. B. Filson

HS H. Streimann RS R. D. Seppelt

JAE J. A. Elix RWR R. W. Rogers

Only one specimen is cited from each location for most of the taxa; where the taxon grows
profusely over a wide area, only specimens from selected sites are listed: viz, R. celastri subsp.
celastri, R. celastri subsp. ovalis, R. inflata subsp. inflata, R. inflata subsp. perpusilla, R. inflata
subsp. aus trails, and R. fissa.

Key to Ramalina in Australia

la. Thallus inflated (partially or totally) 35

Ib. Thallus not inflated 2

2a (Ib). Plant saxicolous 3

2b. Plant corticolous 12

3a (2a). Thallus sorediate 4

3b. Thallus not sorediate 5

4a (3a) . Soredia eruption through eroded lower surface and at apices 6. 1?. fimbriata (p. 161)

4b. Soredia contained in well defined soralia, which are marginal and sometimes laminal

2. R. caespitella (p. 155)

5a (3b). Thallus made up of a few loose branches 6

5b . Thallus composed of numerous branches forming a dense cushion 8

6a (5a). Branches compressed, flat (apothecia marginal) 14. R. celastri subsp. celastri (p. 180)

6b. Branches subterete to terete 7

7a (6b). Branching dense, distal, K- (no medullary acids) I. R. austraiiensis (p. 152)

7b. Branching sparse, K+ red (salazinic acid) 20. R. tropica (p. 210)

RAMALINA IN AUSTRALIA

151

8a (5b). Branches solid 9

8b. Branches inflated , often split 10

9a (8a). Apothecia terminal, large 3- 10 mm diam 13. R. whinrayi (p. 179)

9b. Apothecia subterminal (subtending narrow pointed branchlets, apothecia 1-2 mm

diam.) 8. R.litorea(p. 169)

lOa (8b). Thallus showing some inflation; splitting and tattering of branches common 11

lOb. Thallus not inflated 12

lla(lOa). Medulla K+ red (salazinic acid) 19. R. fissa(p. 193)

lib. Medulla K or K+ pink (divaricatic acid or sekikaic acid)

15. R. inffata subsp. australis (p. 191)

12a (2b). Thallus sorediate 13

12b. Thallus not sorediate 21

13a(12a). Lower surface of anastomosing strands resting on the medulla 10. R. reducta (p. 175)

13b. Lower surface not of anastomosing strands only 14

14a (13b). Branches few, usually broad at base (soft textured, soralia apical and marginal)

3. R. canariensis (p. 156)
14b. Branches numerous, narrow 15

15a (14b). Branches usually cartilaginous (splitting open at intervals along their length)

12. R. unilateralis (p. 177)
15b. Branches not cartilaginous, complete 16

16a(15b). Medulla K+ red (salazinic acid) 17

16b. Medulla K- or K+ pink 18

17a (16a). Soralia punctiform, mainly apical on branchlets (thallus up to 3 cm high)

11. If. tenella (p. 176)
17b. Soralia ellipsoidal or round, marginal on elongate branches (thallus usually 4-8 cm

long) 18. R.pacifica(p. 201)

18a (16b). Soralia ellipsoidal or round, marginal on elongate branches 19

18b. Soralia punctiform, marginal and apical (branching dense, intricate, thallus resilient in

texture) 9. R. peruviana (p. 170)

19a (18b). Medulla K+ pink (sekikaic aggregate acids) 20

19b. Medulla K- (divaricatic acid and stenosporic acid) 17. R. nervulosa var. nervulosa (p. 195)

20a (19a). Sekikaic aggregate acids faint ramalinolic acid 17. R. nervulosa var. dumeticola (p. 197)
20b. Sekikaic aggregate acids faint homosekikaic acid .... 17. R. nervulosa var. luciae (p. 199)

21a(12b). MedullaK+red 22

21b. Medulla K- or K+ pink 23

22a (21a). Branches flat, sometimes canaliculate, containing norstictic acid (spores 12-16 x 4-5

/urn) 19. R. subfraxinea var. norstictica (p. 208)

22b. Branches usually subterete, never canaliculate, containing salazinic acid (spores 14-22

x4-6/u,m) 20. R. tropica(p. 210)

23a (21b). Branches subterete to terete, containing no medullary acids 24

23b. Branches flat, with or without medullary acids 28

24a(23a). MedullaK- 25

24b. MedullaK+pink 30

25a (24a). Apothecia lateral and subterminal with branch apex appearing as a long spur (some-
times black tipped) 4. R. exiguella (p. 158)

25b. Apothecia never subterminal nor bearing a spur 26

26a (25b). Distal branchlets occur on otherwise unbranched primary branches

1. R. australiensis (p. 152)
26b. Dense distal branchlets not present

27a (26b). Thallus thread-like (with dichotomous branching) 5. R. filicaulis (p. 160)

27b. Thallus fruticose 28

28a (27b) . Apothecia marginal on both edges (branches may be canaliculate , no medullary acids)

14. R. celastri subsp. celastri (p. 180)

152 G. NELL STEVENS

28b. Apothecia not marginal on both edges 29

29a (28b). Apothecia laminal, numerous, and equal in size 14. I?, celastri subsp. ova/is (p. 184)

29b. Apothecia not laminal and numerous and equal in size 30

30a (29b) . Branches broad with apices forked or lacerated , never simple and attenuate [apothecia
in axils of branches (up to 5 mm diam.) or terminal on the broad margins of the
apices, or small and sparsely spread laminally] 7. R. glaucescens (p. 163)

30b. Branches usually narrow with apices simple and attenuate or bent to form a spur below

apothecium 31

31a (30b). Branches narrow with dichotomous branching (apothecia marginal or subterminal or

in the axils of branches, small 0-5-1-5 mm diam.) 7. R. glaucescens (p. 163)

31b. Branches narrow to broad, branching usually from the base only 32

32a(31b). MedullaK+pink 33

32b. MedullaK- 34

33a (32b). Branches elongate, narrow, often tuberculate, containing sekikaic acid

19. R. subfraxinea var. subfraxinea (p. 203)

33b. Branches not elongate, rarely tuberculate, containing cryptochlorophaeic acid

19. R. subfraxinea var. conffrmata (p. 205)

34a (32b) . Thallus containing boninic acid 19. R. subfraxinea var. leiodea (p. 207)

34b. Thallus containing divaricatic acid 19. R. subfraxinea var. subfraxinea (p. 203)

35a (la). Thallus sorediate 6. R. fimbriata (p. 161)

35b. Thallus not sorediate 36

36a(35b). Thallus button-like, bearing apothecia (3 mm diam.) between sterile branches or
totally sterile with eroded apices, medullary reaction K-, saxicolous

15. R. inffata subsp. australis (p. 191)
36b . Thallus not button-like nor bearing apothecia between sterile branches 37

37a (36b). Branches split open and tattered 38

37b. Branches perforate but complete 39

38a (37a). Thallus grossly inflated, few or no perforations, splitting open of branches, medullary

acids divaricatic or sekikaic 15. R. inffata subsp. australis (p. 191)

38b. Thallus grossly inflated or split apart, no perforations, medullary acid salazinic

19. R. fissa (p. 193)
39a (37b). Thallus semi-inflated to almost compressed, apothecia never concave

15. R. inffata subsp. perpusilla (p. 188)
39b. Thallus totally inflated, apothecia concave and innate 15. R. inffata subsp. inffata (p. 185)

1 . Ramalina australiensis Ny 1 .

Plate 7, fig. 1; Plate 13, fig. 1.

in Bull. Soc. Linn. Normandie II, 4: 120 (1870). Type: Australia, West Australia, Swan River, 1846,

Verreaux s.n. (PC! - holotype; H-NYL 37491! - isotype; usnic acid).
Ramalina myrioclada Mull. Arg. in Flora, Jena 66: 20 (1883). Type: Australia, New South Wales, Twofold

Bay, 1882, White s.n. (G! - holotype; usnic acid). Plate 13, fig. 2.

Thallus corticolous, rarely saxicolous, pale green to grey-green, erect to pendulous, 3-10 cm
long, exceptionally to 20 cm; branching sparse in the basal region of the primary branches,
becoming dense and irregular towards the apices where small branchlets form tufts; branch
width 0-2-0-5(-l-0) mm, primary branches usually subterete and fine, or thick, compressed and
coarse, branches never tortulose, often subsidiary short branchlets occur at right angles to the
primary branch, apices of branchlets often hooked; surface matt, smooth or furrowed, longi-
tudinal splitting occurs on the surface in coarser forms; pseudocyphellae often present; holdfast
delimited although several branches arise from it, attachment of branches to the substrate at
intervals along their length may occur; soralia absent.

Apothecia uncommon, lateral on the primary branches, usually towards the base; disc

Plate 7 Thallus growth forms in the Australian Ramalinae. Scale in mm. Fig. 1 R. australiensis. Fig. 2 R.
caespitella. Fig. 3 R. canariensis. Fig. 4 R. canariensis (large form). Fig. 5 R. exiguella. Fig. 6 R. filicaulis.
Fig. 7 R. fimbriata. Fig. 8 R. litorea.

RAMALINA IN AUSTRALIA

153

iiillllllllllilfillffP" 1 *!!!!

miiiimmiimillillllillliilllillllliilllllllitllit

linn

liiillii II 1 1 II II I III I Illl 1 1 III II III

154 G. NELL STEVENS

0-5-l-0(-l-2) mm diam., plane to convex, margin entire; spores ellipsoid to fusiform, straight,
12-16(-20) x 4-6 /urn (Plate 4, fig. 2).

Chemistry. Usnic acid only.

Remarks. The name R. australiensis as the oldest name has been upheld for this taxon,
overriding the more commonly used name R. myrioclada. However the location given for the
type material - Swan River, Western Australia - is dubious. All recent collections have been
made in eastern Australia with no other material being found in the west. Whether the original
material from Western Australia was wrongly recorded was not able to be ascertained.

The type material of R. australiensis has branchlets with black tips but this feature (as with R.
exiguella) is not present in most specimens. The type is a coarse thallus form of this taxon, rather
resembling that of the subtropical Queensland collection. There is not sufficient type material to
obtain an overall picture of the total thallus.

The degree of secondary branching varies from specimen to specimen, those with dense
branching resemble R. peruviana whereas others have sparse branching producing an open
thallus form resembling the morphology of R. filicaulis N. Stevens.

The names R. usneoides and R. usnea have been applied to some specimens of R. australiensis
collected in New South Wales but the Australian taxon can be distinguished by its branching
pattern and subterete branches instead of the characteristic flattened branches of R. usnea
sens. str.

Distribution and habitat. This subtropical/warm temperate maritime species does not have a

Fig. 8 Distribution of R. australiensis and R. filicaulis * .

RAMALINA IN AUSTRALIA 155

wide distribution (Fig. 8). It occurs along the coast from Yamba 29S to Twofold Bay 37S and
has a disjunct occurrence on the Queensland coast at Toogoom (25S). This Queensland
occurrence cannot be adequately explained, although the rainfall in this area is similar to that
which occurs along the New South Wales coastline. The Queensland material is extremely
coarse, it was collected from Callitris columellaris which was growing on exposed sand dunes.
The morphology of several Ramalina species collected from this area was also extremely coarse,
indicating environmental adaptation.

The major phorophyte of R. australienti is Avicennia marina, which grows in sheltered
positions along the bays and rivers of the New South Wales coastline. However this Ramalina
has also been collected from rocks at Port Hacking and Port Macquarie, which indicates it is not
substrate specific.

This taxon occurs on the offshore islands of the North Island of New Zealand which lie at
equivalent latitudes to those where R. australiensis occurs in Australia. The species has not been
recorded anywhere else.

World distribution: Australia and New Zealand.

Queensland: Toogoom, 2515'S, 15240'E, on Callitris columellaris, 1983, GNS (BRIU4217NS). New
South Wales: Yamba, 2927'S, 15320'E, on Avicennia marina, 1977, GNS (BRIU 1978NS); Nambucca
Hds. 3041'S, 15300'E, on Avicennia marina, 1977, GNS (BRIU1998NS); Port Macquarie, 3127'S,
15255'E, on metamorphic rock, 1975, JAE(JAE 1081); Patonga3330'S, 15120'E, on Avicennia marina,
1978, GNS (BRIU2314NS); Gosford, 3330'S, 15120'E, 1978, JAE (JAE 4715); Newport, 3340'S,
15120'E, on mangroves, 1888, F. R. M. Wilson (MEL 9409); Lilli Pilli Beach, 3404'S, 15110'E, on
sandstone, 1975, JAE (JAE 1181); Sussex Inlet, 3510'S, 15035'E, on Avicenna marina, 1977, RWR
(BRIU1902NS); Buckenbowra R. 3542'S, 15006'E, on Avicennia marina 1983, HS (CBG8306040) ;
Shoalhaven R. 3452'S, 15042'E, onAegiceras corniculatum, 1975, P. Saenger (BRIU2137NS).

2. Ramalina caespitella N. Stevens Plate 7, fig. 2.

in Lichenologist 18: 183 (1986). Type: Australia, Long Island (Bass Strait), 1969, Whinray s.n. (MEL! -
holotype; sekikaic acid, 4'-0-demethylsekikaic acid, several terpenes, and usnic acid).

Thallus saxicolous, pale greenish-grey to stramineous, minute, caespitose, rigid, up to 1-0 cm
high; branching subdichotomous, irregular, narrow thalli densely branched, broad thalli
sparsely branched; branch width 0-1-2-0 mm, exceptions to 6 mm, branches solid, flat to
subterete, narrow branches nodular, apices broad and blunt; surface shiny; holdfast delimited or
diffuse, soralia marginal and laminal, round to ellipsoidal, becoming fissural on the upper
surface occurring on eroded areas of the lower surface and at the apices. Apothecia rare, laminal
near the branch apices, rarely terminal, disc 1-0 mm diam, concave, margin thick, entire; spores
not produced.

Chemistry: sekikaic acid (M), 4'-0-demethylsekikaic acid (t), several terpenes and usnic acid.

Remarks. Ramalina caespitella produces two morphotypes which intergrade into one another.
One morph has narrow subterete to terete nodular branches with apical and marginal soralia,
whilst the other morph produces very thick, flat to subterete branches bearing numerous soralia
on both upper and lower surfaces. The extreme button-like morphology of both forms is
indicative of their exposed, saxicolous habitat. No other taxon has been found to have a close
affinity with R. caespitella.

Distribution and habitat. This temperate maritime taxon is found on several of the islands in
Bass Strait and along parts of the Tasm'anian coastline (Fig. 9). It occurs on siliceous rocks (e.g.
granite on the islands and on quartzite or dolerite at Tasmanian locations) and often occupies
cliff -face habitats.

World distribution. Australia.

Bass Strait: Hogan Group: Hogans I, 3913'S, 14659'E, on granite, 1973, JW (MEL1013070); Kents
Group, Dover I, 3929'S, 14717'E, on granite, 1970, JW (MEL 10 12702); Deal I, 3930'S, 14722'E, on
granite, 1972,/W(MEL1012940); Furneaux Group: West Sister 1, 3945'S, 14756'E, on granite, 1966, JW

156

G. NELL STEVENS

120

140

10 -

20

30

40

Fig. 9 Distribution of R. caespitella.

(MEL1026132); Cape Barren I, 4025'S, 14810'E, on granite, 1969, JW (MEL1018083); Big Chalky I,
4006'S, 14754'E, on granite, 1972,7W(MEL1018053); Isabella Reef, 4025'S, 14830'E, on granite 1973,
JW (MEL522220); Flinders I, 3945'S, 14756'E, on rock, 1965, RF (MEL1026188); Swan I, 4041'S,
14805'E,onrock, 1974, 7W(MEL10191 19); Tasmania: West Point, 4055'S, 14515'E,onquartzite, 1981,
GK (362/81); Rocky Cape, 4051'S, 14531'E, on quartzite, 1970, GB (H032637); Eddystone Pt., 4100'S,
14823'E, on granite, 1973, GB (H032640); Arthurs River, 4107'S, 14442'E, on quartzite, 1970, GB
(H032639); Bruny I., 4329'S, 14709'E, on granite, 1973, GB (H032636).

3. Ramalina canariensis Steiner Plate 7, figs 3-4.

in Ost. hot. Z. 9: 8 (1904). Type: Gran Canaria, Tafira, Bornmuller, PI. Canar. 3500 (W! - isotype;
divaricatic acid and usnic acid).

Thallus eorticolous, pale green to grey green, erect to subpendulous, l-3(-5) cm long;
branching palmate or irregular; branch width l-3(-10) mm, apices blunt, usually split with
margins separated; surface matt, smooth to rugose, sometimes coarsely reticulately ridged by
chondroid strands, pitted distally; holdfast delimited; soralia marginal and apical occurring
through the separation of the upper and lower surfaces.

Apothecia not seen. Reported as rare.

Chemistry. Divaricatic acid, usnic acid.

Remarks. Fine thalli of this taxon come close to resembling immature thalli of R. unilateralis

RAMALINA IN AUSTRALIA 157

and both species contain divaricatic acid, but the cartilaginous surface and the finely lacerated
apices of R. unilateralis enable the two to be separated.

Distribution and habitat. The distribution pattern of R. canariensis in Australia is mainly
maritime in areas experiencing a Mediterranean type of climate (Fig. 10). The only inland
collections are two South Australian locations (Springton and Kuitpo Forest) at elevations of
300 m where frequent mists occur.

Krog & 0sthagen (1980) mentioned that in the Canary Islands (28N) this species 'seems to
prefer well lit sites with a certain influence of mist . . . between 400 and 1000 m altitude' . In some
of the drier Australian coastal sites morning fogs are prevalent.

Along the southern coastline of Australia this species grows in well-lit situations protected
from winds, usually on twigs of Leucopogon parviflora, Casuarina, and Melaleuca spp. It has
also been found on mangroves (Avicennia marina) along the eastern coastline south from
latitude 35S. Ramalina canariensis occurs in homogeneous colonies or immixed with R. fissa.

Overseas specimens were examined from South Africa (Cape Province, latitude 33S at 800 m
elevation) and Chile (Valparaiso, latitude 33S at sea level, on dune shrubs). Both of these
regions have climates similar to southern Australia and other Mediterranean regions. Ramalina
canariensis was previously thought to grow only in the northern hemisphere in the Mediterra-
nean region and on the Canary Islands.

World distribution. Mediterranean region, Canary Islands, South Africa, Australia, and South
America.

120

160

*

40>

Fig. 10 Distribution of R. canariensis.

158 G. NELL STEVENS

New South Wales: Sussex Inlet, 3510'S, 15035'E, on Avicennia marina, 1977, RWR (BRIU1903NS);
Buckenbowra R. 3542'S, 15006'E, on Avicennia marina, 1983, HS (CBG8306039); Batemans Bay,
3545'S, 15010'E on Avicennia marina, 1978, G7VS(BRIU2171NS);Batehaven Beach, 3545'S, 15013'E,
on Casuarina sp., 1975, JAE (JAE 1061); Burrenwarra Pt., 3549'S, 15014'E, on Casuaria sp., 1975, JAE
(JAE 1246). Victoria: Cape Conran, 3748'S, 14844'E, on Leucopogon sp., 1983, CMS (BRIU3701NS);
Westernport Bay, 3813'S, 14518'E, on Avicennia marina, 1978, GNS (BRIU2336NS); Gunnamatta
Beach, 3821'S, 14445'E, on Monotoca elliptica, 1983, CMS (BRIU3692NS); Warrnambool, 3823'S,
14229'E, 1886, F. R. M. Wilson (NSW.L4311); Portland, 3820'S, 14136'E, 1896, F. R. M. Wilson (MEL
9419); Portsea, 3825'S, 14442'E, on tree on sea cliff, 1964, /. Williams (MEL1026126); Somers, 3824'S,
14510'E, 1964, Marie (MEL1026173). South Australia: Kuitpo forest, 3507'S, 13916'E, on Pinus sp.,
1968, RWR (BRIU1426NS); Kangaroo I. Lockwood Corner, 3539'S, 13738'E, 1967, G. Jackson
(S. A. 97647604); Mt Lofty Ra., Carey Gully, 3445', 13900'E, on Exocarpus cupressiformis , 1976, JAE
(JAE 2853); Springton, 3445'S, 13905'E, on Callitris sp., 1976. JAE (JAE 2807); Eyre Peninsula:
Memory Cove, 3440'S, l355Q'E,onMelaleucasp., 1970,flF(MEL1018638);Elliston,3259'S, 13430'E,
1967, N. Donner (SA97648163). Western Australia: Gairdner River, 3414'S, 11917'E, on Hakea sp.,
1980, D. Richardson (WA000943); Recherche Arch. Long I, 3403'S, 12157'E, on Melaleuca sp., 1950,
JW (MEL10020). Bass Strait: Furneaux Group: Flinders I, 4000'S, 14800'E, on shrub, 1977, JW
(MEL10225619); East Sister I, on dune shrub, 1972,/W(MEL1025619); West Sister 1, 3942'S, 14756'E,
on Pomaderris apetala, 1969, JW (MEL521577); Hogan Group: Long I, 3912'S, 14701'E, on Banksia
integrifolia, 1973, JW (MEL1012966).

4. Ramalina exiguella Stirton Plate?, fig. 5; Plate 1.3, fig. 3.

in Trans. Proc. R. Soc. Viet. 17: 68 (1881). Type: Australia, Queensland, Brisbane, Bailey 91 (BRI! -

holotype; BM-isotype; usnic acid).
Ramalina pertenuis Magnusson in Ark. Bot. 32A (2): 7 (1945). Type: Oahu, Keeawa awa Valley, 1922,

Skottsberg s.n. (GB! - part of holotype; usnic acid).

Thallus corticolous, grey-green, caespitose, erect, rigid, up to 3 cm high; branching mainly
from the base, squarrose branchlets sometimes present on main branches; branch width 0-3-1-0
mm, branches subterete, narrow, apices attenuate, sometimes black tipped; cortex shiny,
longitudinally grooved giving a string-like appearance; pseudocyphellae linear; holdfast delimi-
ted; soralia absent.

Apothecia few to many, usually marginal along the branches and/or subterminal causing the
branch apex to become geniculate with a long, attenuate spur; disc. 0-2-2-0 mm diam. , concave,
plane to convex; margin entire, thick on immature apothecia; spores ovoid to ellipsoid or
gibbous and slightly curved, (12-) 14-16 x (5-)6-8 pm (Plate 5, fig. 1).

Chemistry. Usnic acid only.

Remarks. The affinities of this taxon lie with R. gracilis (Pers.) Nyl. ; Nylander (1870) recorded
material from Brazil, Madagascar, and Australia as belonging to R. gracilis. The type of R.
gracilis was not available from either L or PC, so any relationship between these two taxa was
not able to be checked. Some material of R. gracilis held at BM is morphologically similar to R.
exiguella but contains salazinic acid. Krog & Swinscow (1976) reported that R. attenuata (Pers.)
Tuck, from Domingo, resembled R. exiguella but contained psoromic acid. Landron (1972)
stated that R. gracilis contained psoromic acid whereas R. attenuata contained salazinic acid and
that the latter species is synonymous with R. rigida Ach. Howe (1913-14) placed R. rigida and
R. gracilis as synonyms ofR. attenuata.

It could be that this group of taxa is a complex with several chemical races, but until the type
material of all of these species can be located and examined, the relationship between these taxa
cannot be resolved. Stirton stated in his type description that R. exiguella had affinities with R.
melanothrix Laurer, but the latter species is quite distinct and the type material does not
resemble Australian material of R. exiguella.

Distribution and habitat. The distribution of this coastal species is continuous along the eastern
shoreline of Australia from latitudes 23S-35S, with additional occurrences at 18S and 21S. Its
occurrence on the coastal hills is rare, one tropical collection was made from Abbey Peak, 650 m

RAMALINA IN AUSTRALIA

159

elevation at latitude 14S and one subtropical collection from Mt Mothar 320 m elevation at
latitude 26S (Fig. 11).

Ramalina exiguella grows on several species of mangrove (viz. Rhizophora stylosa, Ceriops
tagal, Avicennia marina, and Aegiceras corniculatum) and on other coastal trees (Casuarina
equisitifolia and Callitris columellaris) which grow close to the water. It also occupies very
exposed situations in the dune communities facing the ocean, where it is often the only lichen
present.

This taxon shows a marked difference in size of thallus in different habitats. The thallus may
be minute (1-5 cm high) and narrow (as the name implies) or extremely coarse and ropy and up
to 3 cm high. South of 28S the thalli tend to produce lateral branchlets along the branches.

In sheltered habitats, R. exiguella grows in association with R. inflata subsp. perpusilla, R.
pacifica, R. subfraxinea var. leiodea, and R. subfraxinea var. confirmata.

World distribution. East Africa, Australia, and New Zealand.

Queensland: Abbey Pk 1418'S, 14430'E, on shrubs, 1983, C. McCracken (BRIU4232NS); South Mission
Beach, 1755'S, 14605'E, on Ficus sp., 1983, GNS (BRIU4159NS); Eimeo, 2106'S, 14910'E, on Cerios
tagal, 1979, GNS (BRIU1860NS); Yeppoon, 2308'S, 15045'E, on Casuarina sp., 1980, RWR
(BRIU2221RR); Keppel Sands, 2321'S, 15047'E, on Rhizophora stylosa, 1975, RWR (BRIU890RR);
Rhodds Peninsula, 2402'S, 15140'E, on Rhizophora stylosa, 1975, GNS (BRIU1277NS); Turkey,
2406'S, 15137'E, on Ceriops tagal, 1975, GNS (BRIU1156NS); Round Hill Head, 2410'S, 15154'E, on
Aegiceras corniculatum, 1975, RWR (BRIU1935RR); Burnett Heads, 2445'S, 15225'E, on Casuarina,
1976, GNS (BRIU4203NS); Fraser I, Wathumba Ck, 2450'S, 15313'E, on Ceriops tagal, 1975, GNS

Fig. 11 Distribution of/?, exiguella.

160 G. NELL STEVENS

(BRIU642NS); Urangan, 2519'S, 15255'E, on Rhizophora stylosa, 1975, GNS (BRIU631NS); Boreen
Point, 2618'S, 15258'E, on Callitris columdlaris , 1982, GNS (BRIU4181NS); Cooran, 2617'S,
15250'E, on Araucaria cunninghamii, 1985, NCS (BRIU4335NS); Noosa River mouth, 2624'S,
15304'E, on Casuarina sp., 1980, GNS (BRIU3184NS); Maroochydore, Maroochy R, 2642'S, 15305'E,
onAvicennia marina, 1976, GNS (BRIU1313NS); Currimundi Lake, 2645'S, 15308'E, on Casuarina sp.,
1974, RWR (BRIU3078RR); Bribie I, 2702'S, 15310'E, on Avicennia marina, 1983, GNS
(BRIU3711NS); Moreton I, Koorigal, 2720'S, 15325'E, on Callitris columellaris , 1980, RWR
(BRIU4179NS); Stradbroke I, Myora Springs, 2728'S, 15324'E, on Avicennia marina 1977, GNS
(BRIU1585NS); Thorneside, Tingalpa Ck 2728'S, 15311'E, on Rhizophora stylosa, 1977, GNS
(BRIU1742NS); Wellington Point, 2728'S, 15315'E, on Avicennia marina, 1975, RWR (BRIU5648RR);
Ormiston, 2730'S, 15316'E, on Avicennia marina, 1975, RWR (BRIU5459RR); Coochie Mudlo I,
2735'S, 15320'E, on Callitris columellaris, 1975, GNS (BRIU4579RR); Point Talburpin, 2735'S,
15317'E, on Rhizophora stylosa, 1982, GNS (BRIU4189NS); Little Rocky Point, 2738'S, 15318'E, on
Ceriops tagal, 1977, GNS (BRIU1437NS). New South Wales: Hastings Point, 2826'S, 15332'E, on
Avicennia marina, 1977, GNS (BRIU1929NS); Brunswick Heads, 2833'S, 15333'E, on Callitris columel-
laris, 1982, GNS (BRIU3316NS); Byron Bay, 2840'S, 15336'E, on Cupaniopsis sp., 1979, RWR
(BRIU3179NS); Ballina, 2854'S, 15333'E, on Avicennia marina, 1977, GNS (BRIU1961NS); Yamba,
2927'S, 15320'E, on Avicennia marina, 1977, GNS (BRIU2125NS); Woolgoolga, 3006'S, 15312'E, on
dune shrubs, 1982, GNS (BRIU4178NS); Sawtell, Boambee Ck, 3022'S, 15306'E, onAvicennia marina,
1977, GNS (BRIU2034NS); Macksville, 3041'S, 15300'E, on Avicennia marina, 1982, GNS
(BRIU4177NS); Macleay River, 3054'S, 15303'E, on Casuarina sp., 1982, GNS (BRIU4175NS); Port
Macquarie, 3127'S, 15254'E, on shrub, 1975, JAE (JAE 1260); Port Macquarie, 3127'S, 15254'E, on
Avicennia marina, 1977, GNS (BRIU1983NS); Laurieton, 3120'S, 15255'E, onAvicennia marina, 1977,
GNS (BRIU4339NS); Taree, Saltwater, 3154'S, 15234'E, on tree, 1977, JAE (JAE 4014); Booti Booti,
3215'S, 15231'E, on Casuarina sp., 1982, GNS (BRIU4332NS); Patonga, 3330'S, 15120'E, on
Avicennia marina, 1978, GNS (BRIU2318NS); Towra Point, 3403'S, 15110'E, on Avicennia marina,
1977, C. Scarlett (BRIU641CS); Batehaven, 3545'S, 15010'E, on dead Casuarina sp., 1974, JAE (JAE
417).

5. Ramalina filicauIisN. Stevens, sp. nov. Plate 7, fig. 6.

Thallus corticolus, virido-griseus, pendulus vel decumbens usque ad 20 cm longus, ramificatione pro parte
majore dichotoma; rami lati ad 0-2-0-5 mm, tenues, filicaules, subterete vel teretes, apice attenuati et
filamentosi, saepe uncinatus; pagina laevigata, fere striata; soralia nulla. Apothecia rara, lateralia, disco
piano vel convexo, ad 0-5-1-0 mm in diametro; sporae ellipsoideae vel fusiformes, rectae vel raro
curvatae, 12-16(-20) x 4-6 /urn. Acidum usnicum tantum continens.

Holotype: Australia, Queensland, Lamington Plateau, 9 km N of O'Reillys in rain-forest on Araucaria
cunninghamii, alt. 920 m. 27 November 1983, G. N. Stevens 3730 (MEL 1048085!).

Thallus corticolous, green-grey, pendulous or decumbent, up to 20 cm long; branching
predominantly dichotomous, curving at the axils; branches width 0-2-0-5 mm, narrow, thread-
like, subterete to terete, tapering, apices attenuate and filamentous, often hooked; surface
smooth, with linear pseudocyphellae; holdfast small, delimited but branches often attached to
the substrate along their length; soralia absent.

Apothecia uncommon, lateral sessile, disc 0-5-1-0 mm diam. , plane to convex; margin entire;
spores ellipsoid to fusiform, straight or rarely curved, 12-16(-20) x 4-6 //,m.

Chemistry. Usnic acid only.

Remarks. Ramalina filicaulis may prove to be a member of the R. usnea complex because, apart
from a chemical difference, fine specimens of R. filicaulis (no medullary acids) come close to the
morphology of the type material of R. anceps (norstictic, salazinic acids). Landron (1972)
recorded R. anceps as a species in the Caribbean Islands (10N-21N) that occurs in lower
montane rain-forest, most abundant on trees along the edge of the forest and in areas of high
rainfall.

Rundel (19780) recorded that the R. usnea race with no medullary acids occurs most
abundantly in eastern South America (23S-28S), a distribution pattern closely resembling the
Australian taxon, but the flattened branches of R. usnea do not occur in R. filicaulis. There are
morphological similarities between the Canary Island species R. chondrina and the Australian

RAMALINA IN AUSTRALIA 161

taxon; both are subtropical species which occur in the mountains. Krog & 0sthagen (1980)
recorded R. chondrina as sparse and scattered on trees in laurel forests between 550 and 900 m
elevation at 2730'N to 2830'N. However, the anatomy of these two species differs; R.
chondrina has an even cylinder of supportive tissue, whereas R. filicaulis has an uneven, ribbed
cylinder of supportive tissue.

Distribution and habitat. This species is rare in occurrence, being known from only a few
locations (Fig. 8). It occurs profusely at Jimna State Forest (2640'S) and at Lamington Plateau
(2815'S) at elevations between 600-900 m. Two early collections may have come from lower
elevations in the Rockhampton area (23S), but little information was available on the
herbarium specimens. At both Jimna and Lamington this taxon grows almost exclusively on the
trunks and branches of Araucaria cunninghamii (hoop pine) in the rain-forest. The only other
phorophyte is Citriobatis pauciflora, a shrub which is a remnant of the rain-forest.

The climatic data recorded for the occurrence of the hoop pine is also relevant for R. filicaulis.
Webb & Tracey (1967) record that regions supporting Araucaria cunninghamii have a mean
annual temperature in the subtropics between 12-23C, with occasional frosts; the annual
average rainfall varies from 800-1800 mm with fogs and dew augmenting precipitation in upland
areas. Hoop pine occurs as part of the climax forest only on soils of relatively low fertility (e.g.
Jimna, on metamorphics); on soils of high fertility it does not occur in the climax forest except as
rare veteran trees (e.g. Lamington Plateau, on basalt).

Because of the rarity of this Ramalina it is difficult to assess what environmental factors are
limiting its occurrence elsewhere.

World distribution. Australia.

Queensland: Jimna, 2640'S, 15228'E, on Araucaria cunninghamii, 1980, RWR (BRIU2202RR); Jimna,
Marumba View, 2640'S, 15228'E, on Araucaria cunninghamii, 1980, GNS (BRIU3349NS); Lamington
Plateau, O'Reillys, 2815'S, 15308'E, windfall, 1970, ED (O); O'Reillys, 2815'S, 15308'E, on Citrio-
batus pauciflora, 1984, P. Merottsy (BRIU4344NS).

6. Ramalina fimbriata Krog & Swinscow Plate 7, fig. 7.

in Norw. J. Bot. 21: 117 (1974). Type: Kenya, Central Province, Nyeri District, Krog & Swinscow
2K36/137 (O! - holotype; BM! - isotype; divaricatic acid and usnic acid).

Thallus saxicolous, pale green to stramineous, small, caespitose, usually very compact and
button-like, often minute; up to 1-0 cm high; branching dense and intricate, distally producing
branchlets; branch width 0-3-1-0 mm, branches flat to subterete, variably inflated, some
branches with slit-like perforations; surface shiny to matt, smooth; holdfast diffuse; soralia form
from disintegration of the lower cortex occurring laminally and at the apices (Plate 1, fig. 5).

Apothecia uncommon, but numerous on some thalli, lateral or terminal, disc 0-5-3-0 mm
diam., concave, plane or convex; margin entire or incised at maturity; spores ellipsoid, straight
or rarely curved, 8-13 x 4-5 /xm.

Chemistry. Divaricatic acid (often in trace amounts).

Remarks. Reduced forms of R. unilateralis come close to the morphology of R. fimbriata, but
the former species is corticolous in Australia and shows no inflation of the branches. The
northern hemisphere species R. pollinaria, when growing on rock, produces thalli closely
resembling R. fimbriata, but the former produces evernic acid. It is suggested that convergence
of morphology due to similarity of habitats is the reason for the resemblance.

Distribution and habitat. Ramalina fimbriata has a distribution range from latitude 35S-43S
over most of the Victorian highlands extending into eastern Tasmania (Fig. 12). In Australia this
species is saxicolous only, although in East Africa it is both saxicolous and corticolous (Krog &
Swinscow, 1974).

Its occurrence on several types of rock (viz. granite, sandstone, conglomerate, trachyte, and
dolerite) shows it is not substrate specific. In some localities R. fimbriata occupies sheltered
overhangs and ledges, habitats which would have little moisture available for the thalli, apart

162

G. NELL STEVENS

Fig. 12 Distribution of R. fimbriata.

from vapour or water droplets from the overhang. Yet large monotypic populations are found in
such habitats. Fertile specimens were found at Mt Arapiles under a rock overhang. The
production of apothecia in this sorediate species is rare and the ascospores were produced only
sparingly, so that mature spores were hard to find.

Other habitats where this species occurs are exposed rock surfaces where the small thalli are
further reduced in size, indicating the harsh environmental conditions prevailing (e.g. cold
winter temperatures with frosts and/or snow and hot summer temperatures). Krog & Swinscow
(1974) record R. fimbriata growing on rock in the alpine zone of East Africa at altitudes of
3200-4100 m.

The compact button-like thalli commonly collected in Australia are smaller than the cortico-
lous Kenyan material recorded by Krog & Swinscow (1974), but in all other characteristics they
are identical.

World distribution. Africa, Australia, and New Zealand.

New South Wales: Budawang Ra. 3522'S, 15003'E, on'conglomerate, 1981, W. A. Weber (COLO 616);
Australian Capital Territory: Tharwa, 11 km SW of, 3531'S, 14904'E, on granite, 1919, JAE (JAE 6195);
Booroomba Rocks, 3532'S, 14900'E, on granite, 1979, HS (CBG7910804); Mt Clear, 3543'S, 14907'E,
on granite, 1980, JAE (JAE 9013). Victoria: Mt Arapiles, 3645'S, 14150'E, on sandstone, 1981, RWR
(BRIU2420RR); 1969, A. C. Beauglehole (MEL1013299); Melville Caves Park, 3639'S, 14342'E, on
rock, 1964, RF (MEL1516515); Bogong High Plains, 3644'S, 14718'E, on rock, 1954, C. Skewes
(MEL1025567); Mt Ararat, 3720'S, 14252'E, c. 1875, D. Sullivan (MEL 9471); Hanging Rock, 3723'S,

RAMALINA IN AUSTRALIA 163

14437'E, on trachyte, 1976, GNS (BRIU3127NS); Yea, 3713'S, 14526'E, on granite, 1964, RF
(MEL1026128); You Yangs, 3800'S, 14429'E, on granite, 1966, GB (H032638). Tasmania: Avoca,
4145'S, 14742'E, on granite, 1972, GB (H032552); Freycinet National Park, Mt Amos, 4213'S,
14818'E on granite, 1979, JAE (JAE 5522); Woods Quoin, 4216'S, 14705'E, on dolerite, 1972, GB &
JC (H032336); Lake Tooms Rd, 4220'S, 14728'E, on rock, 1974, GB & JC (H032551).

7. Ramalina glaucescens Krempelh. Plate 8, figs 1-12; Plate 13, fig. 4.

in Verh. zool-bot. Ges. Wien 30: 333 (1880). Type: Australia, Victoria, Mount Ararat, 1875, Sullivan s.n.

(M! - holotype; sekikaic acid and usnic acid).
Ramalina leiodea var.fastigiatula Mull. Arg. in Flora, Jena 66: 21 (1883). Type: Australia, Victoria, The

Grampians, Sullivan 14 (G! - holotype; sekikaic acid (trace) and usnic acid). Plate 13, fig. 6.
Ramalina lacerata Mull. Arg. in Flora, Jena 66: 20 (1883). Type: Australia, Western Australia, Eucla,

Oliver s.n. (G! - holotype; M!, MEL!, UPS! - isotypes; no medullary acids usnic acid only). Plate 13,

fig. 5.
Ramalina brevis F. Wilson in Victorian Nat. 6: 69 (1889). Type: Australia, Victoria, Warrnambool, 1887,

F.R.M. Wilson s.n. (NSW! - holotype (L4091); no medullary acids, usnic acid only). Plate 13, fig. 8.
Ramalina calicaris var. australica Rasanen in Suomal. eldin-ja kasvit. Seur. van. Tiedon. Poytak. 4: 178

(1949). Type: Australia, Victoria, Barwon Heads, 1894, F. R. M. Wilson s.n. (H! (herb. Rasanen) -

holotype; no medullary acids, usnic acid only). Plate 13, fig. 7.

Thallus corticolous, pale green, caespitose, erect to subpendulous, up to 4 cm long, excep-
tions to 6 cm; branching sparse, subdichotomous to irregular, additional side branchlets occur
on narrow branches; branch width (0-5-)l-5(-10) mm, branches compressed, narrow and
canaliculate or broad and flat, apices usually forked, narrow branches with fine pointed apices
broad branches with dissected apices; surface shiny and smooth or matt and rugose, becoming
coarse with chondroid strands showing at the surface, often horny in texture; holdfast delimited
or diffuse (if growing in colonies); soralia absent.

Apothecia common, laminal at the axil of bifurcating branches (to 5 mm diam.) or laminal
towards the branch apices (0-5-1-0 mm diam.), marginal and subterminal on narrow canalicu-
late branches (0-5-1-5 mm diam.), laminal on broad branches (2-10 mm diam.) at or near the
apex margin, with smaller apothecia on the same branch near the centre; disc concave becoming
plane, rarely convex; margin thick to thin, persistent, often inrolled; spores ellipsoid, straight or
curved, 10-12(-16) x 4-5(-6) /tin.

Chemistry. Sekikaic acid ( trace homosekikaic acid or trace divaricatic acid), or usnic acid
only, or divaricatic acid and usnic acid.

Remarks. The type material of R. glaucescens comprises a branch covered with tiny tufted
specimens and a card with four specimens attached to it (which are here designated a, b, c, d).
Two growth forms are represented on the card; a and d have bifurcate branches with small
apothecia lying at the axil of the fork, and b and c have broader branches and bear terminal or
subterminal apothecia, the latter with small subtending spurs. All of the thalli tested contain
sekikaic acid. The specimens are exceptionally small and are immature.

The considerable polymorphism encountered in R. glaucescens presented much difficulty in
interpretation, as there was no correlation with the chemical variation; a situation rather similar
to that encountered by Kristinsson (1969) in dealing with Cetraria islandica in Iceland. Five
different species names already existed for entities now included in the complex: R. glaucescens
Krempelh., R. leiodea var. fastigiatula Mull. Arg., R. brevis F. Wilson, R. lacerata Mull. Arg.,
and R. calicaris var. australica Rasanen. After the examination of large quantities of material
belonging to this complex, both in the field and from herbarium collections, it became obvious
that the four names additional to that of R. glaucescens (the earliest name) could not be
maintained as species, because of the subtle intergradation between each of them (Plate 8, figs
1-15).

An examination of spores from all of the morphotypes failed to show any marked differences
in spore size or shape. Each apothecium produced spores with considerable variation in both

164

G. NELL STEVENS

RAMALINA IN AUSTRALIA 165

characters, so that the ratio of breadth to length was never constant for any particular
morphotype. The ratio of straight to bent spores was also variable with a tendency towards a
greater proportion of bent spores in the coastal thalli.

The taxonomic interpretation of this complexity of morphotypes offered two alternatives: (1)
to regard each existing ecotype as a distinct species which would allow for the five existing
species and would produce at least two other new taxa, or (2) to regard the whole group as one
extremely variable species showing remarkable phenotypic plasticity in response to environ-
mental pressures. The latter would incorporate numerous morphotypes with similar spores and
anatomy, which occupied different habitats, yet showed intergradation between their extreme
morphological forms (Plate 8, figs 1-15). The second alternative was chosen because variation in
chemistry also occurs throughout the group, which made it impossible to satisfactorily find any
point for delimitation based on the acids present.

Consideration was given to applying varietal rank to the different ecotypes, but each ecotype
would then have had a sekikaic acid race and an acid-deficient race. The division of R.
glaucescens into two taxa based on presence or absence of medullary acids could be feasible,
especially as thalli containing sekikaic acid occur mainly east of longitude 147, whereas west of
this latitude the majority of the specimens contain no acids. If a division of taxa was made this
way then a mixed group of morphotypes would be placed in each class. The amount of sekikaic
acid present in R. glaucescens specimens is very variable in quantity, and division of a species on
so fine a difference as presence or absence of traces of sekikaic acid appeared unwarranted.
Usually acid-deficient races are not distinguished taxonomically from their acid-producing
partners.

The extreme variability of this taxon (as described above) indicates it could be a rapidly
expanding species in Australia. As it also occurs in New Zealand, investigation of material here
may provide information to clarify how it should be divided, whether on its chemistry or its
morphology. Until the matter is finally resolved this complex of morphotypes is regarded as a
single, markedly polymorphic species.

Two specimens collected from Walcha Road (31S) contain divaricatic acid; and both sekikaic
and divaricatic acids occur in several thalli collected at Nimmitabel (36S); this anomaly cannot
be explained. Both areas are at altitudes of over 1000 m and such habitats may produce
physiological changes in the metabolism of R. glaucescens. No taxonomic rank was considered
for the atypical divaricatic acid thalli.

Distribution and habitat. Ramalina glaucescens has a wide distribution in the warm to cool
temperate regions of eastern Australia (Fig. 13), occurring at altitudes from sea level to over
1300 m. A large number of morphotypes are found as illustrated in Plate 8, figs 1-15. A general
intergradation of forms can be seen which do not form a continuum but merge from several
directions.

Thalli with a morphology which is larger, but otherwise similar to that of the type of R.
glaucescens, are found along the inland mountains west of the eastern escarpment in New South
Wales and in the mountainous area of central Victoria, as well as in the central and eastern
mountains of Tasmania (Plate 8, figs 2, 3, 4, 5). All of the specimens examined contain sekikaic
acid in varying amounts, except for a few from western Victoria and South Australia which are
acid-deficient.

Intergrading with this morphotype is a narrower, sometimes canaliculate morph which
often has subsidiary branchlets extending from the main branches; it has both terminal and sub-
terminal apothecia. The distribution of this ecotype is in the Eastern Highlands (with a disjunct
appearance in the Grampians - type location oiR. leiodea var.fastigiatula) and in the highlands
of Tasmania (Plate 8, figs 5, 6, 7, 9, 13, 14). In its narrowest form the branches become subterete
and very fine, bearing small apothecia; the apices of the branches are sharply pointed (Plate 8,
figs 11, 12); this spiky form occurs in mountain habitats above 1400 m elevation in the temperate
region, e.g. it occurs at Mt Aggie and the Brindabella Ranges in New South Wales and at Mt
Wombargo in Victoria, and at lower altitudes at Buchan Caves, Victoria, and in Tasmania. All
collections were from twigs of Epacridaceae bushes, so it is not clear whether low temperatures

166

G. NELL STEVENS

Fig. 13 Distribution of R. glaucescens: sekikaic acid race; O acid-deficient race; * divaricatic acid race.

or a particular bark substrate has an effect on this phenotype's distribution. Sekikaic acid was
always present in such thalli.

One ecotype of R. glaucescens grows along the southern Victorian coastline; it has a short
broad thallus with large terminal or laminal apothecia and is acid-deficient. A few specimens
resembling this morphology have been collected from inland in eastern Australia, but that
material contains sekikaic acid and is not so coarsely textured as the coastal thalli. This ecotype
resembles the type of R. brevis (Plate 8, fig. 15).

Another acid-deficient ecotype occurs inland at low altitudes in Victoria and South Australia,
but the thallus is generally narrower than that of the coastal ecotype and the apothecia are not as
large; however, intergradation between the two is evident at some locations (Plate 8, fig. 8). The
presence of small branchlets on the main branches resembles the eastern highlands ecotype. The
type of R. calicaris var. australica belongs to this ecotype. In some habitats the thallus becomes
quite coarse in texture with chondroid strands showing at the surface; such characteristics are
apparent in the material from the arid zone in Western Australia, including the type collection of
R. lacerata.

The largest morphotype in the R. glaucescens complex is found in western Victoria and
eastern South Australia, with a few specimens collected in western New South Wales. The latter
specimens contain sekikaic acid but all other material proved to be acid-deficient. Most of the
ecotypes of R. glaucescens have been found to intergrade with this large morphotype (Plate 8,
fig. 1). It is normally sterile although pycnidia have been detected in some of these specimens.

Four of the type specimens here included in R. glaucescens were initially collected within 100

RAMALINA IN AUSTRALIA 167

km radius of one another in Victoria, which illustrates the ecophenotypic response to different
habitats. Table 9 summarises the morphological variation that is found in the different ecotypes.
This taxon is entirely corticolous, the most common phorophytes being Acacia spp. and
Bursaria spinosa. It occurs as dense colonies along the branches of isolated trees in pastureland
or in open-forest communities. In some localities the entire population may be small and
immature (as found in the type material of both R. glaucescens and R. leiodea var. fastigiatula) ,
whereas in other areas only large mature specimens occur.

World distribution. Australia and New Zealand.

New South Wales: Ben Lomond, 3000'S, 15142'E, on Acacia sp., 1980, GNS (BRIU3249NS); Sherwood
Ck, 3003'S, 15303'E, on lemon tree, 1978, HS (CBG7905095); Walcha, 10 km W of, 3058'S, 15130'E,
on Acacia sp., 1982, CMS (BRIU4100NS); Abercrombie Caves, 3332'S, 14915'S, on Leptospermum sp.,
1978, JAE (JAE 4640); Rockley, 3342'S, 14934'E, E. Boorman (G); Black Springs 10 km N of, 3349'S,
14948'E, on Acacia sp. , 1984, NCS (BRIU4292NS); Black Springs 5 km S of, 3352'S, 14948'E, on Acacia
sp., 1984, NCS (BRIU4283NS); Abercrombie R. 3412'S, 14945'E, on Pinus sp., 1984, NCS
(BRIU4284NS); Wombeyan Caves, 3418'S, 15002'E, E. Cheel (NSW L4294); Taralga, 37 km N of,
3424'S, 14949'E, on fence posts, 1977, JAE (JAE 3165); Crookwell, 3428'S, 14930'E, on Acacia sp.,
1980, GNS (BRIU3489NS); Yass, 26 km S of, 3500'S, 14850'E, on Casuarina sp., 1975, HS
(CBG058278); Little Peppercorn Ck, 3535'S, 14837'E, on Bursaria spinosa, 1979, HS (CBG7906928);
Rules Point, 3543'S, 14831'E, on shrub, 1979, HS (CBG7907045); Lake Eucumbene, 3610'S, 14850'E,
on Hymenanthera sp., 1971, L. Craven (MEL1020182); Adaminaby, 3612'S, 14850'E, on Briar, 1984,
NCS (BRIU4287NS); Countegany, 3612'S, 14929'E, on Acacia melanoxylon, 1976, JAE (JAE 1925);
Cooma 46 km S of, 3614'S, 14905'E, on Acacia sp., 1978, JAE (JAE 5440); Nimmitabel, 3631'S,
14915'E, on Prunus sp., 1980, GNS (BRIU3215NS); Holts Flat, 3640'S, 14915'E, on Acacia sp., 1980,
G7VS(BRIU3310NS); Delegate, 3703'S, 14857'E, on Acacia sp., 1980, G7VS (BRIU3490NS). Australian
Capital Territory: Ginninderra Ck, 3505'S, 14903'E, 1977, HS (CBG066912); Uriarra Crossing, 3508'S,
14857'E, on Casuarina sp., 1974, JAE (JAE 628); Blundells Ck Rd., 3521'S, 14850'E, on Acacia sp.,
1977, D. Verdon (CBG8008538); Mt Aggie, 3527'S, 14846'E, on Hymenanthera dentata, 1979, JAE
(JAE 5870). Victoria: Walwa, 9 km S of, 3555'S, 14745'E, on Acacia sp. , 1984, NCS (BRIU4338NS); SW
of Corryong, 3616'S, 14650'E, on Briar, 1984, NCS (BRIU4288NS); Myrtleford, 19 km S of, 3639'S,
14642'E, on Acacia sp., 1984, NCS (BRIU4290NS); Tawonga, 3641'S, 14708'E, on Prunus sp., 1984,
NCS (BRIU4293NS); Bright, 3644'S, 14658'E, on copper beech, 1984, NCS (BRIU4286NS); Mt
Wombergo, 3636'S, 14811'S, on Hymenanthera sp., 1966, RF. (MEL1026138); Bonang, 3712'S,
14843'E, on Malus, 1980, GNS (BRIU3491NS); Kyneton, 3715'S, 14428'E, 1897, F. R. M. Wilson,
(MEL 9440); Newlyn, 3725'S, 14400'E, on Crataegus sp., 1981, RWR (BRIU2334RR); Buchan Caves,
3730'S, 14810'E, 1965, RF (MEL1026151); Omeo, 7 km N of, 3704'S, 14738'E, on Acacia sp., 1984,
NCS (BRIU4285NS); Skipton, 11 km S of, 3747'S, 14322'E, on Hymenanthera sp., 1984, NCS
(BRIU4291NS); Buninyong, 3742'S, 14340'E, on trees, F. R. M. Wilson (MEL 9479); Steiglitz,
Moorabool R dam, 3753'S, 14411'E, on twigs, 1968, GB (H032786); Yallourn, Storage dam,3810'S,
14620'E, 1975, KF(MEL1013745); Koweerup, 12 km NE of, 3810'S, 14536'E, on Pinus sp., 1984, NCS
(BRIU4336NS); Loy Yang, 3811'S, 14637'E, on Acacia sp., 1984, NCS (BRIU4337NS);